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Boston UniversityCollege <strong>of</strong> LiberalArtsLibraryFrom <strong>the</strong> Pierce Collection


THE GEOGRAPHICALDISTRIBUTIONOF ANIMALS.BOSTON UNIVERSITYCOLLEGE OF»!3ERALARTSLIBRARV


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THE GEOGRAPDISTRIBUTION OF ANIMALS.WITH A STUDY OF THE RELATIONS OF LIVING AND EXTINCTKAUNAS AS ELUCIDATING THE PAST CHANGESOF THE EARTH'S SURFACE.By ALFRED RUSSEL WALLACE,AUTHOR OF THE " MALAY ARCHIPELAGO," ETC.IN TWO VOLUMES.Vol. I.WITH MAPS AND ILLUSTRATIONS.BOSTON UNIVERSITY'COLLEGE OF 'BERAL ARTSHARPER &NEW YORK:BROTHERS, PUBLISHERS,FRANKLIN SQUARE.1876.


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—qLjOl£76PREFACE.<strong>The</strong> present work is an attempt to collect and summarize <strong>the</strong>existing information on <strong>the</strong> Distribution <strong>of</strong> Land Animals;and to explain <strong>the</strong> more remarkable and interesting <strong>of</strong> <strong>the</strong>facts, by means <strong>of</strong> established laws <strong>of</strong> physical and organicchange.<strong>The</strong> main idea, which is here worked out in some detail for<strong>the</strong> whole earth, was stated sixteen years ago in <strong>the</strong> concludingpages <strong>of</strong> a paper on <strong>the</strong> " Zoological Geography <strong>of</strong> <strong>the</strong> MalayArchipelago," which appeared in <strong>the</strong> Journal <strong>of</strong> Proceedings oj<strong>the</strong> Linnean Society for 1860 ; and again, in a paper read before<strong>the</strong> Eoyal Geographical Society in 1863, it was briefly summarizedin <strong>the</strong> following passage :"My object has been to show <strong>the</strong> important bearing <strong>of</strong>researches into <strong>the</strong> natural history <strong>of</strong> every part <strong>of</strong> <strong>the</strong> world,upon <strong>the</strong> <strong>study</strong> <strong>of</strong> its past history. An accurate knowledge <strong>of</strong>any groups <strong>of</strong> birds or <strong>of</strong> insects and <strong>of</strong> <strong>the</strong>ir <strong>geographical</strong> <strong>distribution</strong>,may enable us to map out <strong>the</strong> islands and continents<strong>of</strong> a former epoch,—<strong>the</strong> amount <strong>of</strong> difference that exists between<strong>the</strong> <strong>animals</strong> <strong>of</strong> adjacent districts being closely relatedto preceding geological changes. By <strong>the</strong> collection <strong>of</strong> suchminute facts, alone, can we hope to fill up a great gap in <strong>the</strong>


viPREFACE.past history <strong>of</strong> <strong>the</strong> earth as revealed by geology, and obtainsome indications <strong>of</strong> <strong>the</strong> existence <strong>of</strong> those ancient lands whichnow lie buried beneath <strong>the</strong> ocean, and have left us nothing but<strong>the</strong>se living records <strong>of</strong> <strong>the</strong>irformer existence."<strong>The</strong> detailed <strong>study</strong> <strong>of</strong> several groups <strong>of</strong> <strong>the</strong> birds and insectscollected by myself in <strong>the</strong> East, brought prominently before mesome <strong>of</strong> <strong>the</strong> curious problems <strong>of</strong> Geographical Distribution;but I should hardly have ventured to treat <strong>the</strong> whole subject,had it not been for <strong>the</strong> kind encouragement <strong>of</strong> Mr. Darwin andPr<strong>of</strong>essor Newton, who, about six years ago, both suggested thatI should undertake <strong>the</strong> task. I accordingly set to work ; butsoon became discouraged by <strong>the</strong> great dearth <strong>of</strong> materials inmany groups, <strong>the</strong> absence <strong>of</strong> general systematic works,and <strong>the</strong>excessive confusion that pervaded <strong>the</strong> classification. Nei<strong>the</strong>rwas it easy to decide on any satisfactory method <strong>of</strong> treating<strong>the</strong> subject. ;During<strong>the</strong> next two years, however, several importantcatalogues and systematic treatises appeared, whichinduced me to resume my work ;and during <strong>the</strong> last three yearsit has occupied a large portion <strong>of</strong> my time.After much consideration, and some abortive trials, an outlineplan <strong>of</strong> <strong>the</strong> book was matured ; and as this is, so far as I amaware, quite novel, it will be well to give a few <strong>of</strong> <strong>the</strong> reasonsforadopting it.Most <strong>of</strong> <strong>the</strong> previous writingson Geographical Distributionappeared to me to be unsatisfactory, because <strong>the</strong>y drew <strong>the</strong>irconclusions from a more or less extensive selection <strong>of</strong> facts ;anddid not clearly separate groups <strong>of</strong> facts <strong>of</strong> unequal value, orthose relating to groups <strong>of</strong> <strong>animals</strong> <strong>of</strong> unequal rank. As anexample <strong>of</strong> what is meant, I may refer to Mr. Andrew Murray'slarge and valuable work on <strong>the</strong> Geographical Distribution <strong>of</strong>Mammalia, in which an immense number <strong>of</strong> coloured maps are


;FKEFACE.viiused to illustrate <strong>the</strong><strong>distribution</strong> <strong>of</strong> various groups <strong>of</strong> <strong>animals</strong>.<strong>The</strong>se maps are not confined to groups <strong>of</strong> any fixed rank, butare devoted to a selection <strong>of</strong> groups <strong>of</strong> various grades. Someshow <strong>the</strong> range <strong>of</strong> single species <strong>of</strong> a genus—as <strong>the</strong> lion, <strong>the</strong>tiger, <strong>the</strong> puma, and a species <strong>of</strong> fox ; o<strong>the</strong>rs are devoted tosections <strong>of</strong> genera,—as <strong>the</strong> true wolves ;o<strong>the</strong>rs to genera,—as <strong>the</strong>hyaenas, and <strong>the</strong> bears ; o<strong>the</strong>rs to portions <strong>of</strong> families,—as <strong>the</strong>flying squirrels, and <strong>the</strong> oxen <strong>with</strong> <strong>the</strong> bisons ; o<strong>the</strong>rs to families,— as <strong>the</strong> Mustelidse, and <strong>the</strong> Hystricidse ; and o<strong>the</strong>rs to groups<strong>of</strong> families or to orders,—as <strong>the</strong> Insectivora, and <strong>the</strong> opossums<strong>with</strong> <strong>the</strong> kangaroos. But in no one grade are all <strong>the</strong> groupstreated alike.Many genera are wholly unnoticed, while severalfamilies are only treated in combination <strong>with</strong> o<strong>the</strong>rs, or arerepresented by some <strong>of</strong> <strong>the</strong> more important genera.In making <strong>the</strong>se observations I by no means intend tocriticise Mr. Murray's book, but merely to illustrate by anexample, <strong>the</strong> method which has been hi<strong>the</strong>rto employed, andwhich seems to me not well adapted to enable us to establish<strong>the</strong> foundations <strong>of</strong> <strong>the</strong> science <strong>of</strong> <strong>distribution</strong> on a secure basis.To do this, uniformity <strong>of</strong> treatment appeared to me essential,both as a matter <strong>of</strong> principle, and to avoid all imputation <strong>of</strong> apartial selection <strong>of</strong> facts, which may be made to prove anything.I determined, <strong>the</strong>refore,to take in succession every well-establishedfamily <strong>of</strong> terrestrial vertebrates, and to give an account <strong>of</strong><strong>the</strong> <strong>distribution</strong> <strong>of</strong> all its component genera, as far asmaterialswere available. Species, as such, are systematically disregarded,—firstly, because <strong>the</strong>y are so numerous as to be unmanageableand, secondly, because <strong>the</strong>y represent <strong>the</strong> most recent modifications<strong>of</strong> form, due to a variety <strong>of</strong> <strong>of</strong>ten unknown causes, andare <strong>the</strong>refore not so clearly connected <strong>with</strong> <strong>geographical</strong> changesas are <strong>the</strong> natural groups <strong>of</strong> species termed genera ; which maybe considered to represent <strong>the</strong> average and more permanent


—viiiPREFACE.<strong>distribution</strong> <strong>of</strong> an organic type, and to be more clearly influencedby <strong>the</strong> various known or inferred changes in <strong>the</strong> organic andphysical environment.This systematic review <strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> families andgenera, now forms <strong>the</strong> last part <strong>of</strong> my book—GeographicalZoology; but it was nearly <strong>the</strong> first written, and <strong>the</strong> copiousmaterials collected for it enabled me to determine <strong>the</strong> zoo<strong>geographical</strong>divisions <strong>of</strong> <strong>the</strong> earth (regions and sub-regions) tobe adopted. I next drew up tables <strong>of</strong> <strong>the</strong> families and generafound in each region and sub-region ;and this afforded a basisfor <strong>the</strong> <strong>geographical</strong> treatment <strong>of</strong> <strong>the</strong> subject—Zoological Geography—<strong>the</strong>most novel, and perhaps <strong>the</strong> most useful andgenerally interesting part <strong>of</strong> my work. While this was in progressI found it necessary to make a careful summary <strong>of</strong> <strong>the</strong> <strong>distribution</strong><strong>of</strong> extinct Mammalia. This was a difficult task, owingto <strong>the</strong> great uncertainty that prevails as to <strong>the</strong> affinities <strong>of</strong> many<strong>of</strong> <strong>the</strong> fossils, and my want <strong>of</strong> practical acquaintance <strong>with</strong>Palaeontology; but having carefully examined and combined<strong>the</strong> works <strong>of</strong> <strong>the</strong> best authors, I have given what I believe is<strong>the</strong> first connected sketch <strong>of</strong> <strong>the</strong> relation <strong>of</strong> extinct Mammaliato <strong>the</strong> <strong>distribution</strong> <strong>of</strong> living groups, and have arrived at somevery interesting and suggestive results.It will be observed that man is altoge<strong>the</strong>r omitted from<strong>the</strong> series <strong>of</strong> <strong>the</strong> animal kingdom as here given, and some explanation<strong>of</strong> this omission may perhaps be required. If <strong>the</strong>genus Homo had been here treated like all o<strong>the</strong>r genera, nothingmore than <strong>the</strong> bare statement— " universally distributed"could have been given ;—and this would inevitably have provoked<strong>the</strong> criticism that it conveyed no information. If, on <strong>the</strong>o<strong>the</strong>r hand, I had given an outline <strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong>varieties or races <strong>of</strong> man, I should have departed from <strong>the</strong> plan<strong>of</strong> my work for no sufficient reason. Anthropology is a science


PREFACE.ixby itself; and it seems better to omit it altoge<strong>the</strong>r from azoological work, than to treat it in a necessarily superficialmanner.<strong>The</strong> best method <strong>of</strong> illustrating a work <strong>of</strong> this kind was amatter requiring much consideration. To have had a separatecoloured or shaded map for each family would have made<strong>the</strong> work too costly, as <strong>the</strong> terrestrial vertebrates alonewould have required more than three hundred maps. I hadalso doubts about <strong>the</strong> value <strong>of</strong> this mode <strong>of</strong> illustration, as itseemed ra<strong>the</strong>r to attract attention to details than to favour <strong>the</strong>development <strong>of</strong> general views.T determined <strong>the</strong>refore to adopta plan, suggested in conversation by Pr<strong>of</strong>essor Newton ;and tohave one general map, showing <strong>the</strong> regions and sub-regions,which could be referred to by means <strong>of</strong> a series <strong>of</strong> numbers.<strong>The</strong>se references I give in <strong>the</strong> form <strong>of</strong> diagrammatic headingsto each family ; and, when <strong>the</strong> map has become familiar,<strong>the</strong>se will, I believe, convey at a glance a body <strong>of</strong> importantinformation.Taking advantage <strong>of</strong> <strong>the</strong> recent extension <strong>of</strong> our knowledge<strong>of</strong> <strong>the</strong> depths <strong>of</strong> <strong>the</strong> great oceans, I determined to give upon thismap a summary <strong>of</strong> our knowledge <strong>of</strong> <strong>the</strong> contours <strong>of</strong> <strong>the</strong> oceanbed, by means <strong>of</strong> tints <strong>of</strong> colour increasing in intensity <strong>with</strong><strong>the</strong> depth.Such a map, when it can be made generally accurate,will be <strong>of</strong> <strong>the</strong> greatest service in forming an estimate <strong>of</strong> <strong>the</strong>more probable changes <strong>of</strong> sea and land during <strong>the</strong> Tertiaryperiod ; and it is on <strong>the</strong> effects <strong>of</strong> such changes that any satisfactoryexplanation <strong>of</strong> <strong>the</strong> facts <strong>of</strong> <strong>distribution</strong> must to a greatextent depend.O<strong>the</strong>r important factors in determining <strong>the</strong> actual <strong>distribution</strong><strong>of</strong> <strong>animals</strong> are, <strong>the</strong> zones <strong>of</strong> altitude above <strong>the</strong> sea level, and <strong>the</strong>strongly contrasted character <strong>of</strong> <strong>the</strong> surface as regards vegetation—aprimary condition for <strong>the</strong> support <strong>of</strong> animal life. I


PREFACE.<strong>the</strong>refore designed a series <strong>of</strong> six maps <strong>of</strong> <strong>the</strong> regions, drawn ona uniform scale, on which <strong>the</strong> belts <strong>of</strong> altitude are shown bycontour-shading, while <strong>the</strong> forests, pastures, deserts, and perennialsnows, are exhibited by means <strong>of</strong> appropriate tints <strong>of</strong> colour.<strong>The</strong>se maps will, I trust, facilitate <strong>the</strong> <strong>study</strong> <strong>of</strong> <strong>geographical</strong><strong>distribution</strong> as a science, by showing, in some cases, an adequatecause in <strong>the</strong> nature <strong>of</strong> <strong>the</strong> terrestrial surface for <strong>the</strong> actual <strong>distribution</strong><strong>of</strong> certain groups <strong>of</strong> <strong>animals</strong>. As it is hoped <strong>the</strong>y willbe constantly referred to, double folding has been avoided, and<strong>the</strong>y are consequently ra<strong>the</strong>r small ; but Mr. Stanford, and hisable assistant in <strong>the</strong> map department, Mr. Bolton, have takengreat care in working out <strong>the</strong> details from <strong>the</strong> latest observations;and this, combined <strong>with</strong> <strong>the</strong>clearness and <strong>the</strong> beauty <strong>of</strong> <strong>the</strong>irexecution, will I trust render <strong>the</strong>m both interesting and instructive.In order to make <strong>the</strong> book more intelligible to those readerswho have no special knowledge <strong>of</strong> systematic zoology, and towhom most <strong>of</strong> <strong>the</strong> names <strong>with</strong> which its pages are <strong>of</strong>ten crowdedmust necessarily be unmeaning, I give a series <strong>of</strong> twenty plates,each one illustrating at once <strong>the</strong> physical aspect and <strong>the</strong>specialzoological character <strong>of</strong> some well-marked division <strong>of</strong> a region.Great care has been taken to associate in <strong>the</strong> pictures, such speciesonly as do actually occur toge<strong>the</strong>r in nature ; so that each platerepresents a scene which is,at all events, not an impossible one.<strong>The</strong> species figured all belong to groups which are ei<strong>the</strong>r peculiarto, or very characteristic <strong>of</strong>,<strong>the</strong> region whose zoology <strong>the</strong>yillustrate ;and it is hoped that <strong>the</strong>se pictures will <strong>of</strong> <strong>the</strong>mselvesserve to convey a notion <strong>of</strong> <strong>the</strong> varied types <strong>of</strong> <strong>the</strong> higher<strong>animals</strong> in <strong>the</strong>ir true <strong>geographical</strong> <strong>relations</strong>. <strong>The</strong> artist, Mr. J.B. Zwecker, to whose talent as a zoological draughtsman andoreat knowledge both <strong>of</strong> animal and vegetable forms we areindebted for this set <strong>of</strong> drawings, died a few weeks after he


PREFACE.X ihad put <strong>the</strong> final touches to <strong>the</strong> pro<strong>of</strong>s. He is known to manyreaders by his vigorous illustrations <strong>of</strong> <strong>the</strong> works <strong>of</strong> SirSamuelBaker, Livingstone, and many o<strong>the</strong>r travellers,—but <strong>the</strong>se, hislast series <strong>of</strong> plates, were, at my special request, executed <strong>with</strong>a care, delicacy, and artistic finish, which his o<strong>the</strong>r designsseldom exhibit. It must, however, be remembered, that <strong>the</strong>figures <strong>of</strong> <strong>animals</strong> here given are not intended toshow specificor generic characters for <strong>the</strong> information <strong>of</strong> <strong>the</strong> scientific zoologist,but merely to give as accurate an idea as possible, <strong>of</strong> some<strong>of</strong> <strong>the</strong> more remarkable and more restrictedtypes <strong>of</strong> beast andbird, amid <strong>the</strong> characteristic scenery <strong>of</strong> <strong>the</strong>ir native country ;-'-and in carrying out this object <strong>the</strong>re are probably few artistwho would have succeeded better than Mr. Zwecker hasdone.<strong>The</strong> general arrangement <strong>of</strong> <strong>the</strong> separate parts <strong>of</strong> which <strong>the</strong>work is composed, has been, to some extent, determined by<strong>the</strong> illustrations and maps, which allmore immediately belongto Part III. It was at first intended to place this part last, butas this arrangement would have brought all <strong>the</strong> illustrationsinto <strong>the</strong> second volume, its place was changed,—perhaps ino<strong>the</strong>r respects for <strong>the</strong> better, as it naturally follows Part II.Yet for persons not well acquainted <strong>with</strong> zoology, it will perhapsbe advisable to read <strong>the</strong> more important articles <strong>of</strong> PartIV. (and especially <strong>the</strong> observations at <strong>the</strong> end <strong>of</strong> each order)after Part II., thus making Part III. <strong>the</strong> conclusion <strong>of</strong> <strong>the</strong>work.Part IV. is, in fact, a book <strong>of</strong> reference, in which <strong>the</strong> <strong>distribution</strong><strong>of</strong> all <strong>the</strong> families and most <strong>of</strong> <strong>the</strong> *genera <strong>of</strong> <strong>the</strong> higher<strong>animals</strong>, is given in systematic order. Part III. is treatedsomewhat more popularly; and, although it is necessarilycrowded <strong>with</strong> scientific names (<strong>with</strong>out which <strong>the</strong> inferences


xiiPREFACE.and conclusions would have nothing solid to rest on), <strong>the</strong>se maybe omitted by <strong>the</strong> non-scientific reader, or merely noted as acertain number or proportion <strong>of</strong> peculiar generic types. ManyEnglish equivalents to family and generic names are, however,given; and, assisted by <strong>the</strong>se, it is believed that any readercapable <strong>of</strong> understanding Lyell's "Principles," or Darwin's" Origin," will have no difficulty in following <strong>the</strong> main argumentsand appreciating <strong>the</strong> chief conclusions arrived at in <strong>the</strong>present work.To those who are more interested in facts than in <strong>the</strong>ories,pie book will serve as a kind <strong>of</strong> dictionary <strong>of</strong> <strong>the</strong> geographyand affinities <strong>of</strong> <strong>animals</strong>. By means <strong>of</strong> <strong>the</strong> copious Index, <strong>the</strong>native country, <strong>the</strong> systematic position, and <strong>the</strong> numericalextent <strong>of</strong> every important and well established genus <strong>of</strong> landanimalmay be at once discovered ;—information now scatteredthrough hundreds <strong>of</strong> volumes.In <strong>the</strong> difficult matters <strong>of</strong> synonymy, and <strong>the</strong> orthography <strong>of</strong>generic names, I have been guided ra<strong>the</strong>r by general utilitythan by any fixed rules. When I have taken a whole familycroup from a modern author <strong>of</strong> repute, I have generally followedhis nomenclature throughout. In o<strong>the</strong>r cases, I use <strong>the</strong> nameswhich are to be found in a majority <strong>of</strong> modern authors, ra<strong>the</strong>rthan follow <strong>the</strong> strict rule <strong>of</strong> priority in adopting some newlydiscovered appellation <strong>of</strong> early date. In orthography I haveadopted all such modern emendations as seem coming intogeneral use, and which do not lead to inconvenience ;but where<strong>the</strong> alteration is such as to completely change <strong>the</strong> pronunciationand appearance <strong>of</strong> a well-known word, I have not adopted it.I have also thought it best to preserve <strong>the</strong> initial letter <strong>of</strong> wellknownand old-establishedto <strong>the</strong> Indices <strong>of</strong> established works.names, for convenience <strong>of</strong> referenceAs an example I may referto Enicurus,—& name which has been in use nearly half a


;PREFACE.xiiicentury, and which is to be found under <strong>the</strong> letter E, in Jerdon'sBirds <strong>of</strong> India, Blyth's Catalogue, Bonaparte's Conspectus, and<strong>the</strong> Proceedings <strong>of</strong> <strong>the</strong> Zoological Society <strong>of</strong> London down to1865. Classicists now write Henicurus as <strong>the</strong> correct formbut this seems to me one <strong>of</strong> those cases in which orthographicalaccuracy should give way to priority, and still more to convenience.In combining and arranging so much detailfrom such variedsources, many errors and omissions must doubtless have occurred.Owing to my residence at a distance from <strong>the</strong> scientificlibraries<strong>of</strong> <strong>the</strong> metropolis, I was placed at a great disadvantage ; and Icould hardly have completed <strong>the</strong> work at all, had I not beenpermitted to have a large number <strong>of</strong> volumes at once, from <strong>the</strong>library <strong>of</strong> <strong>the</strong> Zoological Society<strong>of</strong> London, and to keep <strong>the</strong>mfor months toge<strong>the</strong>r;—a privilege for which I return my bestthanks to Mr. Sclater <strong>the</strong> Secretary, and to <strong>the</strong> Council.Should my book meet <strong>with</strong> <strong>the</strong> approval <strong>of</strong> working naturalists,I venture to appeal to <strong>the</strong>m, to assist me in renderingany future editions more complete, by sending me (to <strong>the</strong> care <strong>of</strong>my publishers) notes <strong>of</strong> any important omissions, or corrections<strong>of</strong> any misstatements <strong>of</strong> fact ;as well as copies <strong>of</strong> any <strong>of</strong> <strong>the</strong>irpapers or essays, and especially <strong>of</strong> any lists, catalogues, andmonographs, containing information on <strong>the</strong> classification or<strong>distribution</strong> <strong>of</strong> living or extinct <strong>animals</strong>.To <strong>the</strong> many friends who have given me information orassistance 1 beg to tender my sincere thanks.Especially am Iindebted to Pr<strong>of</strong>essor Newton, who not only read through much<strong>of</strong> my rough MSS., but was so good as to make numerous correctionsand critical notes.<strong>The</strong>se were <strong>of</strong> great value to me, as<strong>the</strong>y <strong>of</strong>ten contained or suggested important additional matter,or pointed out systematic and orthographical inaccuracies.


::::—::xivPREFACE.Pr<strong>of</strong>essor Flower was so good as toread over my chapters onextinct <strong>animals</strong>, and to point out several errors into which I hadfallen.Dr. Gun<strong>the</strong>r gave me much valuable information on <strong>the</strong>classification <strong>of</strong> reptiles, marking on my lists <strong>the</strong> best establishedand most natural genera, and referring me to reliablesources <strong>of</strong>information.I am also greatly indebted to <strong>the</strong> following gentlemen fordetailed information on special subjects :To Sir Victor Brooke, for a MS. arrangement <strong>of</strong> <strong>the</strong> genera<strong>of</strong> Bovidse, <strong>with</strong> <strong>the</strong> details <strong>of</strong> <strong>the</strong>ir <strong>distribution</strong>To Mr. Dresser, for lists <strong>of</strong> <strong>the</strong> characteristic birds <strong>of</strong> Nor<strong>the</strong>rnand Arctic EuropeTo Dr. Hooker, forinformation on <strong>the</strong> colours and odours <strong>of</strong>New Zealand plantsTo Mr. Kirby, for a list <strong>of</strong> <strong>the</strong> butterflies <strong>of</strong> ChiliTo Pr<strong>of</strong>essor Mivart, for a classification <strong>of</strong> <strong>the</strong> Batrachia, andan early pro<strong>of</strong> <strong>of</strong> his article on " Apes " in <strong>the</strong> EncyclopediaBritannicaTo Mr. Salvin, for correcting my list <strong>of</strong> <strong>the</strong> birds <strong>of</strong> <strong>the</strong>Galapagos, and for o<strong>the</strong>r assistanceTo Mr. Sharpe, for MS. lists <strong>of</strong> <strong>the</strong> birds <strong>of</strong> Madagascarand <strong>the</strong> Cape Verd Islands:To Canon Tristram, for a detailed arrangement <strong>of</strong> <strong>the</strong> difficultfamily <strong>of</strong> <strong>the</strong> warblers,—Sylviidse :To Viscount Walden, for notes on <strong>the</strong> systematic arrangement<strong>of</strong> <strong>the</strong> Pycnonotidse and Timaliidse, and for an early pro<strong>of</strong> <strong>of</strong>his list <strong>of</strong> <strong>the</strong> birds <strong>of</strong> <strong>the</strong> Philippine Islands.I also have to thank many naturalists, both in this countryand abroad, who have sent me copies <strong>of</strong> <strong>the</strong>ir papers;and I trust <strong>the</strong>y will continue to favour me in <strong>the</strong> samemanner.


PREFACE.xvAn author may easily be mistaken in estimating his ownwork. I am well aware that this first outline <strong>of</strong> a greatsubject is, in parts, very meagre and sketchy; and, thoughperhaps overbur<strong>the</strong>ned <strong>with</strong> some kinds <strong>of</strong> detail, yet leavesmany points most inadequately treated. It is <strong>the</strong>refore <strong>with</strong>some hesitation that I venture to express <strong>the</strong> hope that I havemade some approach to <strong>the</strong> standard <strong>of</strong> excellence I have aimedat ;—which was, that my book should bear a similar relation to<strong>the</strong> eleventh and twelfth chapters <strong>of</strong> <strong>the</strong> " Origin <strong>of</strong> Species," asMr. Darwin's " Animals and Plants under Domestication " doesto <strong>the</strong> first chapter <strong>of</strong> that work. Should it be judged worthy<strong>of</strong> such a rank, my long, and <strong>of</strong>ten wearisome labours, will bewell repaid.Makch, 1876.


2—CONTENTS OF THE EIEST VOLUME.PAET I.THE PEINCIPLES AND GENEKAL PHENOMENA OF DISTRIBUTION.CHAPTEE I.INTRODUCTORY.CHAPTER II.THE MEANS OF DISPERSAL AND THE MIGRATIONS OF ANIMALS.Means <strong>of</strong> Dispersal <strong>of</strong> Mammalia (p. 10)—Climate as a Limit to <strong>the</strong> Range <strong>of</strong>Mammals (p. 11)—Valleys and Rivers as Barriers to Mammals (p. 12)—Arms<strong>of</strong> <strong>the</strong> Sea as Barriers to Mammals ([j. 13)—Ice-floes and drift-wood as aiding<strong>the</strong> Dispersal <strong>of</strong> Mammals (p. 14)—Means <strong>of</strong> Dispersal <strong>of</strong> Birds (p. 15)—Dispersal<strong>of</strong> Birds by Winds (p. 16)—Barriers to <strong>the</strong> Dispersal <strong>of</strong> Birds (p. 17)<strong>The</strong> Phenomena <strong>of</strong> Migration (p. 18)—Migrations <strong>of</strong> Birds (p. 19)—Generalremarks on Migration (p. 25)—Means <strong>of</strong> Dispersal <strong>of</strong> Reptiles and Amphibia(p. 28)—Means <strong>of</strong> Dispersal <strong>of</strong> Fishes (p. 29)—Means <strong>of</strong> Dispersal <strong>of</strong> Mollusca(p. 30)— Means <strong>of</strong> Dispersal <strong>of</strong> Insects and <strong>the</strong> Barriers which limit <strong>the</strong>irRange (p. 32) 10—34CHAPTER III.DISTRIBUTION AS AFFECTED BY THE CONDITIONS AND CHANGES OF THE EARTH'SLand and Water (p.SURFACE.35)—Continental Areas (p. 36)—Recent Changes in <strong>the</strong> ContinentalAreas (p. 38)—<strong>The</strong> Glacial Epoch as affecting <strong>the</strong> Distribution <strong>of</strong>Animals (p. 40)—Changes <strong>of</strong> Vegetation as affecting <strong>the</strong> Distribution <strong>of</strong> Animals(p. 43)— Organic Changes as affecting Distribution (p. 44) . 35—49Vol. I.—


—xviiiCONTENTS OF THE FIRST VOLUME.CHAPTER IV.ON ZOOLOGICAL REGIONS.Principles upon which Zoological Regions should be formed (p.53)—Which class<strong>of</strong> Animals is <strong>of</strong> most importance in determining Zoological Regions (p. 56)Various Zoological Regions proposed since 1857 (p. 58)— Discussion <strong>of</strong> proposedRegions (p. 61)—Reasons for adopting <strong>the</strong> Six Regions first proposed by Mr.Sclater (p.63)—Objections to <strong>the</strong> system <strong>of</strong> Circumpolar Zones (p. 67)—Does<strong>the</strong> Arctic Fauna characterise an independent Region (p.68)—Palsearctic Region(p. 71)—Ethiopian Region (p. 73)—Oriental Region (p. 75)—Australian Region(p. 77)—Neotropical Region (p. 78)—Nearctic Region (p. 79)—Observationson <strong>the</strong> series <strong>of</strong> Sub-regions (p. 80) 50—82CHAPTER V.CLASSIFICATION AS AFFECTING THE STUDY OF GEOGRAPHICAL DISTRIBUTION.Classification <strong>of</strong> <strong>the</strong> Mammalia (p. 85)—Classification <strong>of</strong> Birds (p. 92)—Classification<strong>of</strong> Eeptiles (p. 98)—Classification <strong>of</strong> Amphibia (p. 100)—Classification <strong>of</strong>Fishes (p. 101)—Classification <strong>of</strong> Insects (p. 102)—Classification <strong>of</strong> Mollusca104) 83—104(P-PART IION THE DISTRIBUTIONOF EXTINCT ANIMALS.CHAPTER VI.THE EXTINCT MAMMALIA OF THE OLD WORLD.Historic and Post-pliocene Period (p. 110)—Pliocene Period (p. 112)—GeneralConclusions as to <strong>the</strong> Pliocene and Post-pliocene Faunas <strong>of</strong> Europe (p. 113)—Miocene Period (p. 114)—Extinct Animals <strong>of</strong> Greece (p. 115)—Miocene Fauna<strong>of</strong> Central and Western Europe (p. 117)—Upper Miocene Deposits <strong>of</strong> India(p. 121)— General Observations on <strong>the</strong> Miocene Faunas <strong>of</strong> Europe and Asia(p. 123)—Eocene Period (p. 124)—General Considerations on <strong>the</strong> ExtinctMammalian Fauna <strong>of</strong> Europe (p. 126) 107—128


—CONTENTS OF THE FIRST VOLUME.xixCHAPTER VII.EXTINCT MAMMALIA OF THE NEW WORLD.North America—Post-pliocene Period (p. 129)—Remarks on <strong>the</strong> Post-plioceneFauna <strong>of</strong> North America (p. 130)— Tertiary Period (p. 132)—Primates (p. 32)Insectivora (p. 133)—Carnivora (p. 134)—Ungulata (p. 135)—Proboscidea(p.138)—Tillodontia (p. 139)—Eodentia (p. 140)—General Relations <strong>of</strong> <strong>the</strong> ExtinctTertiary Mammalia <strong>of</strong> North America and Europe (p. 140)- South America(p. 143)—Fauna <strong>of</strong> <strong>the</strong> Brazilian Caves (p. 143) — Pliocene Period <strong>of</strong> TemperateSouth America (p. 146)—Pliocene Mammalia <strong>of</strong> <strong>the</strong> Antilles (p. 148)—Eocene Fauna <strong>of</strong> South America (p. 148)—General Remarks on <strong>the</strong> ExtinctMammalian Fauna <strong>of</strong> <strong>the</strong> Old and New Worlds (p. ] 48)—<strong>The</strong> Birth-place anc 1Migrations <strong>of</strong> some Mammalian Families and Genera (p. 153) . 129—156CHAPTER VIII.VARIOUS EXTINCT ANIMALS ;—AND ON THE ANTIQUITY OF THE GENERA OFINSECTS AND LAND-MOLLUSCA.Extinct Mammalia <strong>of</strong> Australia (p. 157)—Mammalian Remains <strong>of</strong> <strong>the</strong> SecondaryFormations (p. 159)—Extinct Birds (p. 160)—Pahearctic Region and NorthIndia (p. 161)—North America (p. 163)— South America, Madagascar, NewZealand (p. 164)—Extinct Tertiary Reptiles (p. 165)—Antiquity <strong>of</strong> <strong>the</strong> Genera<strong>of</strong> Insects (p.(p. 168)166)—Antiquity <strong>of</strong> <strong>the</strong> Genera <strong>of</strong> Land and Fresh-water Shells157-170PART III.ZOOLOGICAL GEOGRAPHY :A REVIEW OF THE CHIEF FORMS OF LIFEIN THE SEVERAL REGIONS AND SUB-REGIONS, WITH THE INDICA-TIONS THEY AFFORD OF GEOGRAPHICAL MUTATIONS.CHAPTER IX.THE ORDER OF SUCCESSION OF THE REGIONS.—COSMOPOLITAN GROUPS OFANIMALS.—TABLES OF DISTRIBUTION.Order <strong>of</strong> succession <strong>of</strong> <strong>the</strong> Regions (p. 173)—Cosmopolitan Groups (p. 175)Tables <strong>of</strong> Distributions <strong>of</strong> Families and Genera (p. 177) . .173—179


xxCONTENTS OF THE FIRST VOLUME.CHAPTER X.THE PAL.EARCTIC REGION.Zoological Characteristics <strong>of</strong> <strong>the</strong> Palsearctic Region (p.181)—Summary <strong>of</strong> PalsearcticVertebrata (p. 186)—Insects (p. 1 87)—Land-shells (p. 190)—<strong>The</strong> PalsearcticSub-regions (p. 190)—Central and Nor<strong>the</strong>rn Europe (p. 191)—NorthEuropean Islands (p. 197)—Mediterranean Sub-region (p. 199)—<strong>The</strong> Mediterraneanand Atlantic Islands (p. 206) —<strong>The</strong> Siberian Sub-region, or Nor<strong>the</strong>rnAsia (p. 216)—Japan and North China, or <strong>the</strong> Manchurian Sub-region (p. 220)— Birds (p. 223)—Insects (p. 227)—Eemarks on <strong>the</strong> General Character <strong>of</strong> <strong>the</strong>Fauna <strong>of</strong> Japan (p. 230)— General Conclusions as to <strong>the</strong> Fauna <strong>of</strong> <strong>the</strong> PalsearcticRegion (p. 231)—Table I. Families <strong>of</strong> Animals inhabiting <strong>the</strong> PalsearcticRegion (p. 234)—Table II. List <strong>of</strong> <strong>the</strong> Genera <strong>of</strong> Terrestrial Mammaliaand Birds <strong>of</strong> <strong>the</strong> Palsearctic Region (p. 239) .... 181—250CHAPTER XLTHE ETHIOPIAN REGION.Zoological Characteristics <strong>of</strong> <strong>the</strong> Ethiopian Region (p. 252)—Summary <strong>of</strong> EthiopianVertebrates (p. 255)—<strong>The</strong> Ethiopian Sub-regions (p. 258)—<strong>The</strong> EastAfrican Sub-region, or Central and East Africa (p. 258)—<strong>The</strong> West AfricanSub-region (p. 262)—Islands <strong>of</strong> <strong>the</strong> West African Sub-region (p. 265)—SouthAfrican Sub-region (p. 266)—Atlantic Islands <strong>of</strong> <strong>the</strong> Ethiopian Region ;—St.Helena (p. 269)—Tristan dAcunha (p. 271)—Madagascar and <strong>the</strong> MascareneIslands, or <strong>the</strong> Malagasy Sub-region (p. 272)—<strong>The</strong> Mascarene Islands (p. 280)—Extinct Fauna <strong>of</strong> <strong>the</strong> Mascarene Islands and Madagascar (p. 282)—GeneralRemarks on <strong>the</strong> Insect Fauna <strong>of</strong> Madagascar (p. 284)— On <strong>the</strong> probable PastHistory <strong>of</strong> <strong>the</strong> Ethiopian Region (p. 285)—Table I. Families <strong>of</strong> Animalsinhabiting <strong>the</strong> Ethiopian Region (p. 294)—Table II. List <strong>of</strong> Genera <strong>of</strong> TerrestrialMammalia and Birds <strong>of</strong> <strong>the</strong> Ethiopian Region (p. 300) . 251—313CHAPTER XII.THE ORIENTAL REGION.Zoological Characteristics <strong>of</strong> <strong>the</strong> Oriental Region (p; 315)—Summary <strong>of</strong> OrientalVertebrata (p. 318)—<strong>The</strong> Oriental Sub-regions (p. 321)—Hindostan, or IndianSub-region (p. 321)—Range <strong>of</strong> <strong>the</strong> Genera <strong>of</strong> Mammalia which inhabit <strong>the</strong> Subregion<strong>of</strong> Hindostan (p. 322)— Oriental, Palsearctic, and Ethiopian Genera <strong>of</strong>Birds in Central India (p. 224)—Sub-region <strong>of</strong> Ceylon and South India(p-. 326)—<strong>The</strong> Past History <strong>of</strong> Ceylon and South India, as indicated by its Fauna(p. 328)—Himalayan or Indo-Chinese Sub-region (p. 329)—Islands <strong>of</strong> <strong>the</strong>


—CONTENTS OF THE FIRST VOLUME.xxiIndo-Chinese Sub-region (p. 333)—Indo-Malaya, or <strong>the</strong> Malayan Sub-region(p. 334)—Malayan Insects (p. 341)—<strong>The</strong> Zoological Relations <strong>of</strong> <strong>the</strong> severalIslands <strong>of</strong> <strong>the</strong> Indo-Malay Sub-region (p. 345)— Philippine Islands (p. 345)—Java (p. 349)—Malacca, Sumatra, and Borneo (p. 353)— Probable recent GeographicalChanges in <strong>the</strong> Indo-Malay Islands (p. 357)—Probable Origin <strong>of</strong> <strong>the</strong>Malayan Fauna (p. 359)—Concluding Remarks on <strong>the</strong> Oriental Region (p. 362)—Table I. Families <strong>of</strong> Animals inhabiting <strong>the</strong> Oriental Region (p. 365)—Table II. Genera <strong>of</strong> Terrestrial Mammalia and Birds in <strong>the</strong> Oriental Region(p. 371) 314-386CHAPTER XIII.THE AUSTRALIAN REGION.General Zoological Characteristics <strong>of</strong> <strong>the</strong> Australian Region (p. 390)—Summary<strong>of</strong> <strong>the</strong> Australian Vertebrata (p. 397)—Supposed Land-connection betweenAustralia and South America (p. 398)—Insects (p. 403)—Land shells (p. 407)—Australian Sub-regions (p. 408)—Austro-Malayan Sub-region (p. 409)—Papua, or <strong>the</strong> New Guinea Group (p. 409)—<strong>The</strong> Moluccas (p. 417)— InsectsPeculiarities <strong>of</strong> <strong>the</strong> Moluccan Fauna (p. 420)—Timor Group (p. 422)—Celebes Group (p. 426)—Origin <strong>of</strong> <strong>the</strong> Fauna <strong>of</strong> Celebes (p. 436)— Australiaand Tasmania, or <strong>the</strong> Australian Sub-region (p. 438)—<strong>The</strong> Pacific Islands, orPolynesian Sub-region (p. 442)—Fiji, Tonga, and Samoa Islands (p. 443)Society and Marquesas Islands (p. 443)—Ladrone and Caroline Islands (p. 444)—New Caledonia and <strong>the</strong> New Hebrides (p. 444)— Sandwich Islands (p. 445)—Reptiles <strong>of</strong> <strong>the</strong> Polynesian Sub-region (p. 448)—New Zealand Sub-region(p. 449)— Islets <strong>of</strong> <strong>the</strong> New Zealand Sub-region (p. 453)—Reptiles, Amphibia,and Fresh-water Fishes (p. 456)—Insects (p. 457)—<strong>The</strong> Ancient Fauna <strong>of</strong> NewZealand (p. 459)—<strong>The</strong> Origin <strong>of</strong> <strong>the</strong> New Zealand Fauna (p. 459)—Causes <strong>of</strong><strong>the</strong> Poverty <strong>of</strong> Insect-life in New Zealand : its Influence on <strong>the</strong> Character <strong>of</strong><strong>the</strong> Flora (p. 462)—Concluding Remarks on <strong>the</strong> Early History <strong>of</strong> <strong>the</strong> AustralianRegion (p. 464)—Table I. Families <strong>of</strong> Animals inhabiting <strong>the</strong> AustralianRegion (p. 468)—Table II. Genera <strong>of</strong> Terrestrial Mammalia and Birds <strong>of</strong> <strong>the</strong>Australian Region (p. 473) 387—485Index to Vol. I 489-503


MAPS AND ILLUSTRATIONS IN VOL. I.1. Map <strong>of</strong> <strong>the</strong> World, showing <strong>the</strong> Zoo-Geographical Regions and <strong>the</strong>contour <strong>of</strong> <strong>the</strong> Ocean-bed FrontispieceTo face page2. Map <strong>of</strong> <strong>the</strong> Pahearctic Region . . . . . . .1813. Plate I. <strong>The</strong> Alps <strong>of</strong> Central Europe <strong>with</strong> Characteristic Animals 1954. Plate II. Characteristic Mammalia <strong>of</strong> Western Tartary . . . 2185. Plate III. Characteristic Animals <strong>of</strong> North China .... 2266. Map <strong>of</strong> <strong>the</strong> Ethiopian Region 2517. Plate IV. Characteristic Animals <strong>of</strong> East Africa . ...2618. Plate V. Scene in West Africa <strong>with</strong> Characteristic Animals . . 2649. Plate VI. Scene in Madagascar <strong>with</strong> Characteristic Animals . . 27810. Map <strong>of</strong> <strong>the</strong> Oriental Region 31511. Plate VII. Scene in Nepaul <strong>with</strong> Characteristic Animals . . . 33112. Plate VIII. A Forest in Borneo <strong>with</strong> Characteristic Mammalia .. 33713. Plate IX. A Malacca Forest <strong>with</strong> some <strong>of</strong> its Peculiar Birds . . 34014. Map <strong>of</strong> <strong>the</strong> Australian Region 38715. Plate X. Scene in New Guinea <strong>with</strong> Characteristic Animals . . 41516. Plate XI. <strong>The</strong> Characteristic Mammalia <strong>of</strong> Tasmania . . 43917. Plate XII. <strong>The</strong> Plains <strong>of</strong> New South Wales <strong>with</strong> Characteristic Animals44218. Plate XIII. Scene in New Zealand <strong>with</strong> some <strong>of</strong> its Remarkable Birds 455


THEGEOGRAPHICAL DISTRIBUTIONOF ANIMALS.PART I.THE PRINCIPLES AND GENERAL PHENOMENAOF DISTRIBUTION.


CHAPTEE I.INTRODUCTORY.It is a fact <strong>with</strong>in <strong>the</strong> experience <strong>of</strong> most persons, that <strong>the</strong>various species <strong>of</strong> <strong>animals</strong> arenot uniformly dispersed over <strong>the</strong>surface <strong>of</strong> <strong>the</strong> country. If we have a tolerable acquaintance<strong>with</strong> any district, be it a parish, acounty, or a larger extent <strong>of</strong>territory, we soon become aware that each well-marked portion<strong>of</strong> it has some peculiarities in its animal productions. If wewant to find certain birds or certain insects, we have not only tochoose <strong>the</strong> right season but to go to <strong>the</strong> right place. If wetravel beyond our district in various directions we shall almostcertainly meet <strong>with</strong> something new to us ; some species whichwe were accustomed to see almost daily will disappear, o<strong>the</strong>rs'which we have never seen before will make <strong>the</strong>ir appearance.If we go very far, so as to be able to measure our journey bydegrees <strong>of</strong> latitude and longitude and to perceive importantchanges <strong>of</strong> climate and vegetation, <strong>the</strong> differences in <strong>the</strong> forms <strong>of</strong>animal life will become greater ;till at length we shall come to acountry where almost everything will be new, all <strong>the</strong> familiarcreatures <strong>of</strong> our own districtless differing from <strong>the</strong>m.If we have been observantbeing replaced by o<strong>the</strong>rs more orduring our several journeys, andhave combined and compared <strong>the</strong> facts we have collected, it willbecome apparent that <strong>the</strong> change we have witnessed has been<strong>of</strong> two distinct kinds.In our own and immediately surroundingdistricts, particular species appeared and disappeared because


4 DISTRIBUTION OF ANIMALS. [part i.<strong>the</strong> soil, <strong>the</strong> aspect, or <strong>the</strong> vegetation, was adapted to <strong>the</strong>m or<strong>the</strong> reverse. <strong>The</strong> marshes, <strong>the</strong> heaths, <strong>the</strong> woods and forests,<strong>the</strong> chalky downs, <strong>the</strong> rocky mountains, had each <strong>the</strong>ir peculiarinhabitants, which reappeared again and again as we came totracts <strong>of</strong> country suitable for <strong>the</strong>m. But as we got fur<strong>the</strong>r awaywe began to find that localities very similar to those we hadleft behind were inhabited by a somewhat different set <strong>of</strong> species;and this difference increased <strong>with</strong> distance, not<strong>with</strong>standingthat almost identical external conditions might be <strong>of</strong>ten met<strong>with</strong>. <strong>The</strong> first class <strong>of</strong> changes is that <strong>of</strong> stations ; <strong>the</strong> secondthat <strong>of</strong> habitats. <strong>The</strong> one is a local, <strong>the</strong> o<strong>the</strong>r a geograiihicalphenomenon. <strong>The</strong> whole area over which a particular animalis found may consist <strong>of</strong> any number <strong>of</strong> stations, but rarely <strong>of</strong>more than one habitat.Stations, however, are <strong>of</strong>ten so extensiveas to include <strong>the</strong> entire range <strong>of</strong> many species. Such are <strong>the</strong>great seas and oceans, <strong>the</strong> Siberian or <strong>the</strong> Amazonian forests,<strong>the</strong> North African deserts, <strong>the</strong> Andean or <strong>the</strong> Himalayanhighlands.<strong>The</strong>re is yet ano<strong>the</strong>r difference in <strong>the</strong> nature <strong>of</strong> <strong>the</strong> changewe have been considering. <strong>The</strong> new <strong>animals</strong> which we meet<strong>with</strong> as we travel in any direction from our starting point, aresome <strong>of</strong> <strong>the</strong>m very much like those we have left behind us,and can be at once referred to familiar types ; while o<strong>the</strong>rsare altoge<strong>the</strong>r unlike anything we have seen at home. Whenwe reach <strong>the</strong> Alps we find ano<strong>the</strong>r kind <strong>of</strong> squirrel, in Sou<strong>the</strong>rnItaly a distinct mole, in Sou<strong>the</strong>rn Europe fresh warblersand unfamiliar buntings. We meet also <strong>with</strong> totally newforms ;as <strong>the</strong> glutton and <strong>the</strong> snowy owl in Nor<strong>the</strong>rn, <strong>the</strong> genetand <strong>the</strong> hoopoe in Sou<strong>the</strong>rn, and <strong>the</strong> saiga antelope andcollared pratincole in Eastern Europe. <strong>The</strong> first series areexamples <strong>of</strong> what are termed representative species, <strong>the</strong> second<strong>of</strong> distinct groups or types <strong>of</strong> <strong>animals</strong>. <strong>The</strong> one represents acomparatively recent modification, and an origin in or near <strong>the</strong>locality where it occurs ; <strong>the</strong> o<strong>the</strong>r is a result <strong>of</strong> very ancientchanges both organic and inorganic, and is connected <strong>with</strong> some<strong>of</strong> <strong>the</strong>to discuss.most curious and difficult <strong>of</strong> <strong>the</strong> problems we shall have


chap, i.] INTRODUCTORY.Having thus defined our subject, let us glance at <strong>the</strong> opinionsthat have generally prevailed as to <strong>the</strong> nature and causes <strong>of</strong><strong>the</strong> phenomena presented by <strong>the</strong> <strong>geographical</strong> <strong>distribution</strong> <strong>of</strong><strong>animals</strong>.It was long thought, and is still a popular notion, that <strong>the</strong>manner in which <strong>the</strong> various kinds <strong>of</strong> <strong>animals</strong> are dispersedover <strong>the</strong> globe is almost wholly due to diversities <strong>of</strong> climate and<strong>of</strong> vegetation. <strong>The</strong>re is indeed much to favour this belief. <strong>The</strong>arctic regions are strongly characterised by <strong>the</strong>ir white bearsand foxes, <strong>the</strong>ir reindeer, ermine, and walruses, <strong>the</strong>ir whiteptarmigan, owls, and falcons ;<strong>the</strong> temperate zone has its foxesand wolves, its rabbits, sheep, beavers, and marmots, its sparrowsand its song birds ;while tropical regions alone produce apes andelephants, parrots and peacocks, and a thousand strange quadrupedsand brilliant birds which arefound nowhere in <strong>the</strong> coolerregions. So <strong>the</strong> camel, <strong>the</strong> gazelle and <strong>the</strong> ostrich live in <strong>the</strong>desert ; <strong>the</strong> bison on <strong>the</strong> prairie ; <strong>the</strong> tapir, <strong>the</strong> deer, and <strong>the</strong>jaguar in forests. Mountains and marshes, plains and rockyprecipices, have each <strong>the</strong>ir animal inhabitants ;•be thought, in <strong>the</strong> absenceand it might well<strong>of</strong> accurate inquiry, that <strong>the</strong>se ando<strong>the</strong>r differences would sufficiently explain why most <strong>of</strong> <strong>the</strong>regions and countriesinto which <strong>the</strong> earth is popularly dividedshould have certain <strong>animals</strong> peculiar to <strong>the</strong>m and should wanto<strong>the</strong>rs which are elsewhere abundant.A more detailed and accurate knowledge <strong>of</strong> <strong>the</strong> productions <strong>of</strong>different portions <strong>of</strong> <strong>the</strong> earth soon showed that this explanationwas quite insufficient ; for it was found that countries exceedinglysimilar in climate and all physicalvery distinct animal populations.features may yet have<strong>The</strong> equatorial parts <strong>of</strong> Africaand South America, for example, are very similar in climateand are both covered <strong>with</strong> luxuriant forests, yet <strong>the</strong>ir animal lifeis widely different; elephants, apes, leopards, guinea-fowlsand touracos in <strong>the</strong> one, are replaced by tapirs, prehensiletailedmonkeys, jaguars, curassows and toucans in <strong>the</strong> o<strong>the</strong>r.Again, parts <strong>of</strong> South Africa and Australia are wonderfullysimilar in <strong>the</strong>ir soil and climate;yet one has lions, antelopes,zebras and giraffes ; <strong>the</strong> o<strong>the</strong>r only kangaroos, wombats, phalan-


6 DISTRIBUTION OF ANIMALS. [part I.gers and mice. In like manner parts <strong>of</strong> North America andEurope are very similar in all essentials <strong>of</strong> soil climate andvegetation, yet <strong>the</strong> former has racoons, opossums, and hummingbirds; while <strong>the</strong> latter possesses moles, hedgehogs and true flycatchers.Equally striking are <strong>the</strong> facts presented by <strong>the</strong><strong>distribution</strong> <strong>of</strong> many large and important groups <strong>of</strong> <strong>animals</strong>.Marsupials (opossums, phalangers &c.) are found from temperateVan Diemen's land to <strong>the</strong> tropical islands <strong>of</strong> New Guinea andCelebes, and in America from Chili to Virginia. No crowsexist inSouth America, while <strong>the</strong>y inhabit every o<strong>the</strong>r part <strong>of</strong><strong>the</strong> world, not excepting Australia. Antelopes are found onlyin Africa and Asia ; <strong>the</strong> sloths only in South America ; <strong>the</strong> truelemurs are confined to Madagascar, and <strong>the</strong> birds-<strong>of</strong>-paradise toNew Guinea.If we examine more closely <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong> inany extensive region, we find that different, though closely alliedspecies, are <strong>of</strong>ten found on <strong>the</strong> opposite sides <strong>of</strong> any considerablebarrier to <strong>the</strong>ir migration.Thus, on <strong>the</strong> two sides <strong>of</strong> <strong>the</strong> Andesand Eocky Mountains in America, almost all <strong>the</strong> mammalia, birds,and insects are <strong>of</strong> distinct species. To a less extent, <strong>the</strong> Alpsand Pyrenees forma similar barrier, and even great rivers andriver plains, as those <strong>of</strong> <strong>the</strong> Amazon and Ganges, separate moreor less distinct groups <strong>of</strong> <strong>animals</strong>. Arms <strong>of</strong> <strong>the</strong> sea are stillmore effective, if <strong>the</strong>y are permanent ; a circumstance in somemeasure indicated by <strong>the</strong>ir depth. Thus islands far away fromland almost alwayselse ;have very peculiar <strong>animals</strong> found nowhereas is strikingly <strong>the</strong> case in Madagascar and New Zealand,and to a less degree in <strong>the</strong> West India islands. But shallowstraits, like <strong>the</strong> English Channel or <strong>the</strong> Straits <strong>of</strong> Malacca, arenot found to have <strong>the</strong> same effect, <strong>the</strong> <strong>animals</strong> being nearly orquite identical on <strong>the</strong>ir opposite shores. A change <strong>of</strong> climate ora change <strong>of</strong> vegetation may form an equally effectivebarrier tomigration. Many tropical and polar <strong>animals</strong> are pretty accuratelylimited by certain iso<strong>the</strong>rmal lines ;and <strong>the</strong> limits <strong>of</strong> <strong>the</strong>great forests in most parts <strong>of</strong> <strong>the</strong> world strictly determine <strong>the</strong>ranges <strong>of</strong> many species.Naturalists have now arrived at <strong>the</strong>conclusion, that by some


chap, i.] INTRODUCTORY.slow process <strong>of</strong> development or transmutation, all <strong>animals</strong> havebeen produced from those which preceded <strong>the</strong>m ; and <strong>the</strong> oldnotion that every species was specially created as <strong>the</strong>y nowexist, at a particular time and in a particular spot, isabandonedas opposed to many striking facts, and unsupported by anyevidence. This modification <strong>of</strong> animal forms took place veryslowly, so that <strong>the</strong> historical period <strong>of</strong> three or four thousandyears has hardly produced any perceptible change in a singlespecies. Even <strong>the</strong> time since <strong>the</strong> last glacial epoch, which on<strong>the</strong> very lowest estimate must be from 50,000 to 100,000 years,has only served to modify a few <strong>of</strong> <strong>the</strong> higher <strong>animals</strong> into veryslightly different species. <strong>The</strong> changes <strong>of</strong> <strong>the</strong> forms <strong>of</strong> <strong>animals</strong>appear to have accompanied, and perhaps to have dependedon,changes <strong>of</strong> physical geography, <strong>of</strong> climate, or <strong>of</strong> vegetation;since it is evident that an animal which is well adapted to onecondition <strong>of</strong> things will require to be slightly changed in constitutionor habits, and <strong>the</strong>refore generally in form, structure, orcolour, in order to be equally well adapted to a changedcondition <strong>of</strong> surrounding circumstances. Animals multiply sorapidly, that we may consider <strong>the</strong>m as continually trying toextend <strong>the</strong>ir range ; and thus any new land raised above <strong>the</strong>sea by geological causes becomes immediately peopled by acrowd <strong>of</strong> competing inhabitants, <strong>the</strong> strongest and best adapted<strong>of</strong> which alone succeed in maintaining <strong>the</strong>ir position.If we keep in view <strong>the</strong>se facts— that <strong>the</strong> minor features <strong>of</strong> <strong>the</strong>earth's surface are everywhere slowly changing ;that <strong>the</strong> forms,and structure, and habits <strong>of</strong> all living things are also slowlychanging ; while <strong>the</strong> great features <strong>of</strong> <strong>the</strong> earth, <strong>the</strong> continents,and oceans, and l<strong>of</strong>tiest mountain ranges, only change after verylong intervals and <strong>with</strong> extreme slowness ; we must see that<strong>the</strong> present <strong>distribution</strong> <strong>of</strong> <strong>animals</strong> upon <strong>the</strong> several parts <strong>of</strong><strong>the</strong> earth's surface is <strong>the</strong> final product <strong>of</strong> all <strong>the</strong>se wonderfulrevolutions in organic and inorganic nature. <strong>The</strong> greatest andmost radical differences in <strong>the</strong> productions <strong>of</strong> any part <strong>of</strong> <strong>the</strong>globe must be dependent on isolation by <strong>the</strong> most effectualand most permanent barriers. That ocean which has remainedbroadest and deepest from <strong>the</strong> most remote geological epochs


8 DISTRIBUTION OF ANIMALS. [part i.will separate countries <strong>the</strong> productions <strong>of</strong> which most widelyand radically differ; while <strong>the</strong> most recently-depressed seas,or <strong>the</strong> last-formed mountain ranges, will separate countries<strong>the</strong> productions <strong>of</strong> which are almost or quite identical. Itwill be evident, <strong>the</strong>refore, that <strong>the</strong> <strong>study</strong> <strong>of</strong> <strong>the</strong> <strong>distribution</strong><strong>of</strong> <strong>animals</strong> and plants may add greatly to our knowledge<strong>of</strong> <strong>the</strong> past history <strong>of</strong> our globe. It may reveal to us, in amanner which no o<strong>the</strong>r evidence can, which are <strong>the</strong> oldestand most permanent features <strong>of</strong> <strong>the</strong> earth's surface, and which<strong>the</strong> newest. It may indicate <strong>the</strong> existence <strong>of</strong> islands or continentsnow sunk beneath <strong>the</strong> ocean, and which have left norecord <strong>of</strong> <strong>the</strong>ir existence save <strong>the</strong> animal and vegetable productionswhich have migrated to adjacent lands. It thusbecomes an important adjunct to geology, which can rarely domore than determine what lands have been raised above <strong>the</strong>waters, under what conditions and at what period; but canseldom ascertain anything <strong>of</strong> <strong>the</strong> position or extent <strong>of</strong> thosewhich have sunk beneath it. Our present <strong>study</strong> may <strong>of</strong>tenenable us, not only to say where lands must have recentlydisappeared, but also to form some judgment as to <strong>the</strong>ir extent,and <strong>the</strong> time that has elapsed since <strong>the</strong>ir submersion.Having thus briefly sketched <strong>the</strong> nature and objects <strong>of</strong> <strong>the</strong>subject we have to <strong>study</strong>, it will be necessary—before enteringon a detailed examination <strong>of</strong> <strong>the</strong> zoological features <strong>of</strong> <strong>the</strong>different parts <strong>of</strong> <strong>the</strong> earth, and <strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> orders,families, and genera <strong>of</strong> <strong>animals</strong>— to examine certain preliminaryfacts and principles essential for our guidance. We must firstinquire what are <strong>the</strong> powers <strong>of</strong> multiplication and dispersal <strong>of</strong><strong>the</strong> various groups <strong>of</strong> <strong>animals</strong>, and <strong>the</strong> nature <strong>of</strong> <strong>the</strong> barriersthat most effectually limit <strong>the</strong>ir range. We have <strong>the</strong>n toconsider <strong>the</strong> effects <strong>of</strong> changes in physical geography and inclimate ;to examine <strong>the</strong> nature and extent <strong>of</strong> such changes ashave been known to occur ; to determine what o<strong>the</strong>rs are possibleor probable ; and to ascertain <strong>the</strong> various modes in which suchchanges affect <strong>the</strong> structure, <strong>the</strong> <strong>distribution</strong>, or <strong>the</strong> very existence<strong>of</strong> <strong>animals</strong>.


3chap, i.] INTRODUCTORY. 9Two subjects <strong>of</strong> a different nature must next engage ourattention. We have to deal <strong>with</strong> two vast masses <strong>of</strong> facts,each involving countlessdetails, and requiring subdivision andgrouping to be capable <strong>of</strong> intelligible treatment. All <strong>the</strong> continentsand <strong>the</strong>ir chief subdivisions, and all <strong>the</strong> more importantislands <strong>of</strong> <strong>the</strong> globe, have to be compared as regards <strong>the</strong>ir variousanimal forms. To do this effectively we require a naturaldivision <strong>of</strong> <strong>the</strong> earth especially adapted to our purpose ; and weshall have to discuss at some length <strong>the</strong> reasons for <strong>the</strong> particularsystem adopted,—a discussion which must to some extentanticipate and summarize <strong>the</strong> conclusions <strong>of</strong> <strong>the</strong> whole work.We have also to deal <strong>with</strong> many hundreds <strong>of</strong> families and manythousands <strong>of</strong> genera <strong>of</strong><strong>animals</strong>, and here too a true and naturalclassification is <strong>of</strong> great importance. We must <strong>the</strong>refore give aconnected view <strong>of</strong> <strong>the</strong> classification adopted in <strong>the</strong> variousclasses <strong>of</strong> <strong>animals</strong> dealt <strong>with</strong>.And lastly, as <strong>the</strong> existing <strong>distribution</strong> <strong>of</strong> <strong>animals</strong> is <strong>the</strong>result and outcome <strong>of</strong>all preceding changes <strong>of</strong> <strong>the</strong> earth and <strong>of</strong>itsinhabitants, we require as much knowledge as we can get <strong>of</strong><strong>the</strong> <strong>animals</strong> <strong>of</strong> each country during past geological epochs, inorder to interpret <strong>the</strong> facts we shall accumulate. We shall,<strong>the</strong>refore, enter upon a somewhat detailed sketch <strong>of</strong> <strong>the</strong> variousforms <strong>of</strong> extinct <strong>animals</strong> that have lived upon <strong>the</strong> earthduring<strong>the</strong> Tertiary period ; discuss <strong>the</strong>ir migrations at various epochs,<strong>the</strong> changes <strong>of</strong> physical geography that <strong>the</strong>y imply, and <strong>the</strong>extent to which <strong>the</strong>y enable us to determine <strong>the</strong> birthplace<strong>of</strong> certain families and genera.<strong>The</strong> preliminary studies above enumerated will, it is believed,enable us to see <strong>the</strong> bearing <strong>of</strong> many facts in <strong>the</strong> <strong>distribution</strong> <strong>of</strong><strong>animals</strong> that would o<strong>the</strong>rwise be insoluble problems ;and, whatis hardly less valuable, will teach us to estimate <strong>the</strong> comparativeimportance <strong>of</strong> <strong>the</strong> various groups <strong>of</strong> <strong>animals</strong>, and to avoid<strong>the</strong> common error <strong>of</strong> cutting <strong>the</strong> gordian knot <strong>of</strong> each difficultyby vast hypo<strong>the</strong>tical changes in existing continents and oceans—probably <strong>the</strong> most permanent features <strong>of</strong> our globe.Vol. I—


CHAPTEE II.THE MEANS OF DISPERSAL AND THE MIGRATIONS OF ANIMALS.All <strong>animals</strong> are capable <strong>of</strong> multiplying so rapidly, that if asingle pair were placed ina continent <strong>with</strong> abundance <strong>of</strong> foodand no enemies, <strong>the</strong>y might fully stock it in a very short time.Thus, a bird which produces ten pairs <strong>of</strong> young during its lifetime(and this is far below <strong>the</strong> fertility <strong>of</strong> many birds) will, ifwe take its life at five years, increase to a hundred millions inabout forty years, a number sufficient to stock a large country.Many fishes and insects are capable <strong>of</strong> multiplying severalthousandfold each year, so that in a few years <strong>the</strong>y would reachbillions and trillions. Even large and slow breeding mammals,which have only one at a birth but continue to breed from eightto ten successive years, may increase from a single pair to tenmillions in less than forty years.But as <strong>animals</strong> rarely have an unoccupied country to breedin, and as <strong>the</strong> food in any one district is strictly limited, <strong>the</strong>irnatural tendency is to roam in every direction in search <strong>of</strong> freshpastures, or new hunting grounds.In doing so, however, <strong>the</strong>ymeet <strong>with</strong> many obstacles. Kocks and mountains have to beclimbed, rivers or marshes to be crossed, deserts or forests to betraversed ;while narrow straits or wider arms <strong>of</strong> <strong>the</strong> sea separateislands from <strong>the</strong> main land or continents from each o<strong>the</strong>r.Wehave now to inquire what facilities <strong>the</strong> different classes <strong>of</strong><strong>animals</strong> have for overcoming <strong>the</strong>se obstacles, and what kind o£barriers are most effectual in checking <strong>the</strong>ir progress.Means <strong>of</strong> Dispersal <strong>of</strong> Mammalia.—Many <strong>of</strong> <strong>the</strong> largest mammaliaare able to roam over whole continents and are hardly


chap, ii.] DISPERSAL AND MIGRATION. 11stopped by any physical obstacles. <strong>The</strong> elephant is almostequally at home on plainsand mountains, and it even climbs to<strong>the</strong> highest summit <strong>of</strong> Adam's Peak in Ceylon, which isand rocky as to be very difficult <strong>of</strong> ascent for man.so steepIt traversesrivers <strong>with</strong> great ease and forces its way through <strong>the</strong> densestjungle. <strong>The</strong>re seems <strong>the</strong>refore to be no limit to its powers <strong>of</strong>wandering, but <strong>the</strong><strong>of</strong> enduring changes <strong>of</strong> climate.great powers <strong>of</strong> dispersal.necessity <strong>of</strong> procuring food and its capacity<strong>The</strong> tiger is ano<strong>the</strong>r animal <strong>with</strong>It crosses rivers and sometimes evenswims over narrow straits <strong>of</strong> <strong>the</strong> sea, and it can endure <strong>the</strong>severe cold <strong>of</strong> North 'China and Tartary as well as <strong>the</strong> heats <strong>of</strong><strong>the</strong> plains <strong>of</strong> Bengal. <strong>The</strong> rhinoceros, <strong>the</strong> lion, and many <strong>of</strong> <strong>the</strong>ruminants have equal powers <strong>of</strong> dispersal ;so that wherever <strong>the</strong>reis land and sufficient food, <strong>the</strong>re are no limits to <strong>the</strong>ir possiblerange.O<strong>the</strong>r groups <strong>of</strong> <strong>animals</strong> are more limited in <strong>the</strong>ir migrations.<strong>The</strong> apes, lemurs, and many monkeys are so strictlyadapted to an arboreal lifethat <strong>the</strong>y can never roam far beyond<strong>the</strong> limits <strong>of</strong> <strong>the</strong> forest vegetation. <strong>The</strong> same may be said <strong>of</strong><strong>the</strong> squirrels, <strong>the</strong> opossums, <strong>the</strong> arboreal cats, and <strong>the</strong> sloths, <strong>with</strong>many o<strong>the</strong>r groups <strong>of</strong> less importance. Deserts or open countryare equally essential to <strong>the</strong> existence <strong>of</strong> o<strong>the</strong>rs. <strong>The</strong> camel, <strong>the</strong>hare, <strong>the</strong> zebra, <strong>the</strong> giraffeexist in a forestprairie marmots.and many <strong>of</strong> <strong>the</strong> antelopes could notcountry any more than could <strong>the</strong> jerboas or <strong>the</strong><strong>The</strong>re are o<strong>the</strong>r <strong>animals</strong> which are confined to mountains, andcould not extend <strong>the</strong>ir range into lowlands or forests. <strong>The</strong> goatsand <strong>the</strong> sheep are <strong>the</strong> most striking group <strong>of</strong> this kind, inhabitingmany <strong>of</strong> <strong>the</strong> highest mountains <strong>of</strong> <strong>the</strong> globe ;<strong>of</strong> which <strong>the</strong>European ibex and mouflon are striking examples. Eivers areequally necessary to <strong>the</strong> existence <strong>of</strong> o<strong>the</strong>rs, as <strong>the</strong> beaver, otter,water-vole and capybara ; and to such <strong>animals</strong> high mountainrangesor deserts must form an absolutely impassable barrier.Climate as a Limit to <strong>the</strong> Range <strong>of</strong> Mammals.—Climate appearsto limit <strong>the</strong> range <strong>of</strong> many <strong>animals</strong>, though <strong>the</strong>re is some reasonto believe that in many cases it is not <strong>the</strong> climate itself so muchas <strong>the</strong> change <strong>of</strong> vegetation consequent on climate which produces<strong>the</strong> effect. <strong>The</strong> quadrumana appear to be limited by climate,


;12 DISTRIBUTION OF ANIMALS. [part i.since <strong>the</strong>y inhabit almost all <strong>the</strong> tropical regions but do notrange more than about 10° beyond <strong>the</strong> sou<strong>the</strong>rn and 12° beyond<strong>the</strong> nor<strong>the</strong>rn tropic, while <strong>the</strong> great bulk <strong>of</strong> <strong>the</strong> species arefound only <strong>with</strong>in an equatorial belt about 30° wide. But as<strong>the</strong>se <strong>animals</strong> are almost exclusively fruit-eaters, <strong>the</strong>ir <strong>distribution</strong>depends as much on vegetation as on temperature ;and thisis. strikingly shown by <strong>the</strong> fact that <strong>the</strong> Semnopi<strong>the</strong>eus schistaceusinhabits <strong>the</strong> Himalayan mountains to a height <strong>of</strong> 11,000feet, where it has been seen leaping among fir-trees loaded <strong>with</strong>snow-wreaths ! Some nor<strong>the</strong>rn <strong>animals</strong> are bounded by <strong>the</strong>iso<strong>the</strong>rmal <strong>of</strong> 32°. Such are <strong>the</strong> polar bear and <strong>the</strong> walrus,which cannot live in a state <strong>of</strong> nature far beyond <strong>the</strong> limits <strong>of</strong><strong>the</strong> frozen ocean ;but as <strong>the</strong>y live in confinement in temperatecountries, <strong>the</strong>ir range is probably limited by o<strong>the</strong>r conditionsthan temperature.We must not <strong>the</strong>refore be too hasty in concluding, that <strong>animals</strong>which we now see confined to a very hot or a very cold climateare incapable <strong>of</strong> living in any o<strong>the</strong>r. <strong>The</strong> tiger was once considereda purely tropical animal, but it inhabits permanently <strong>the</strong>cold plains <strong>of</strong> Manchuria and <strong>the</strong> Amoor, a country <strong>of</strong> an almostarctic winter climate. Few <strong>animals</strong> seem to us more truly inhabitants<strong>of</strong> hot countries than <strong>the</strong> elephants and rhinocerosesyet in Post-tertiary times <strong>the</strong>y roamed over <strong>the</strong> whole <strong>of</strong> <strong>the</strong>nor<strong>the</strong>rn continents to <strong>with</strong>in <strong>the</strong> arctic circle ;and we know that<strong>the</strong> climate was <strong>the</strong>n as cold as it is now, from <strong>the</strong>ir entire bodiesbeing preserved in ice. Some change must recently haveoccurred ei<strong>the</strong>r in <strong>the</strong> climate, soil, or vegetation <strong>of</strong> Nor<strong>the</strong>rnAsia which led to <strong>the</strong> extinction <strong>of</strong> <strong>the</strong>se forerunners <strong>of</strong> existingtropical species ;and we must always bear in mind that similarchanges may have acted upon o<strong>the</strong>r species which we now findrestricted <strong>with</strong>in narrow limits, but which may once have roamedover a wide and varied territory.Valleys and Rivers as Barriers to Mammals.—To <strong>animals</strong> whichthrive best in dry and hilly regions, a broad level and marshyvalley must <strong>of</strong>ten prove an effectual barrier. <strong>The</strong> difference <strong>of</strong>vegetation and <strong>of</strong> insect life,toge<strong>the</strong>r <strong>with</strong> an unhealthy atmosphere,no doubt <strong>of</strong>ten checks migration if it is attempted. Thus


;chap, ii.] DISPERSAL AND MIGRATION. 13many <strong>animals</strong> are restricted to <strong>the</strong> slopes <strong>of</strong> <strong>the</strong> Himalayasor to <strong>the</strong> mountains <strong>of</strong> Central India, <strong>the</strong> fiat valley <strong>of</strong> <strong>the</strong>Ganges forming a limit to <strong>the</strong>ir range. In o<strong>the</strong>r cases, however,it is <strong>the</strong> river ra<strong>the</strong>r than <strong>the</strong> valley which is <strong>the</strong> barrier. In<strong>the</strong> great Amazonian plains many species <strong>of</strong> monkeys, birds, andeven insects are fonnd up tonot cross to <strong>the</strong> o<strong>the</strong>r.<strong>the</strong> river banks on one side but doThus in <strong>the</strong> lower part <strong>of</strong> <strong>the</strong> Rio Negrotwo monkeys, <strong>the</strong> Jacchus bicolor and <strong>the</strong> Brachiurus couxiou, arefound on <strong>the</strong> north bank <strong>of</strong> <strong>the</strong> river but never on <strong>the</strong> south,where a red-whiskered Pi<strong>the</strong>cia is alone found.Higher up Atelcspaniscus extends to <strong>the</strong> north bank <strong>of</strong> <strong>the</strong> river while Lagothrixhumboldtii comes down to <strong>the</strong> south bank :native <strong>of</strong> Guiana, <strong>the</strong> latter <strong>of</strong> Ecuador.<strong>of</strong> <strong>the</strong> genus Psophia or trumpeters, isAmazon, Madeira, Eio Negro and some o<strong>the</strong>rs ;<strong>the</strong> former being a<strong>The</strong> range <strong>of</strong> <strong>the</strong> birdsalso limited by <strong>the</strong> riversso that in <strong>the</strong>secases we are able to define <strong>the</strong> limits <strong>of</strong> <strong>distribution</strong> <strong>with</strong> anunusual degree <strong>of</strong> accuracy, and <strong>the</strong>re is little doubt <strong>the</strong> samebarriers also limit a large number <strong>of</strong> o<strong>the</strong>r species.Arms <strong>of</strong> <strong>the</strong> Sea as Barriers to Mammals.—Very few mammalscan swim over any considerable extent <strong>of</strong> sea, although many canswim well for short distances. <strong>The</strong> jaguar traverses <strong>the</strong> wideststreams in South America, and <strong>the</strong> bear and bison cross <strong>the</strong>Mississippi ;and <strong>the</strong>re can be no doubt that <strong>the</strong>y could swim overequal widths <strong>of</strong> salt water, and ifaccidentally carried out to seamight sometimes succeed in reaching islands many miles distant.Contrary to <strong>the</strong> common notion pigs can swim remarkably well.Sir Charles Lyell tells us in his ."Principles <strong>of</strong> Geology" thatduring <strong>the</strong> floods in Scotland in 1829, some pigs only six monthsold that were carried out to sea, swam five miles and got onshore again. He also states, on <strong>the</strong> authority <strong>of</strong> <strong>the</strong> late EdwardForbes, that a pig jumped overboard to<strong>the</strong> Grecian Archipelago, and swam safely toescape from a terrier inshore many milesdistant. <strong>The</strong>se facts render it probable that wild pigs, from<strong>the</strong>ir greater strength and activity, might under favourable circumstancescross arms <strong>of</strong> <strong>the</strong> sea twenty or thirty miles wideand <strong>the</strong>re are facts in <strong>the</strong> <strong>distribution</strong> <strong>of</strong> this tribe <strong>of</strong> <strong>animals</strong>which seem to indicate that <strong>the</strong>y have sometimes done so.Deer


14 DISTRIBUTION OF ANIMALS. [part i.take boldly to <strong>the</strong> water and can swim considerable distances,but we have no evidence to show how long <strong>the</strong>y could live atsea or how many miles <strong>the</strong>y could traverse.Squirrels, rats, andlemmings <strong>of</strong>ten migrate from nor<strong>the</strong>rn countries in bands <strong>of</strong>thousands and hundreds <strong>of</strong> thousands, and pass over rivers, lakesand even arms <strong>of</strong> <strong>the</strong> sea, but <strong>the</strong>y generally perish in <strong>the</strong> saltwater.Admitting, however, <strong>the</strong> powers <strong>of</strong> most mammals toswim considerable distances, we have no reason to believe thatany <strong>of</strong> <strong>the</strong>m could traverse <strong>with</strong>out help straits <strong>of</strong> upwards <strong>of</strong>twenty miles in width, while in most cases a channel <strong>of</strong> halfthat distance would prove an effectual barrier.Ice-floes and Driftwood as Aiding <strong>the</strong> Dispersal <strong>of</strong> Mammals.—In <strong>the</strong> arctic regions icebergs originate in glaciers which descendinto <strong>the</strong> sea, and <strong>of</strong>teneven some vegetation on <strong>the</strong>ir surfaces ;bear masses <strong>of</strong> gravel, earth, andand extensive level icefieldsbreak away and float southwards. <strong>The</strong>se might <strong>of</strong>tencarry <strong>with</strong> <strong>the</strong>m such arctic quadrupeds as frequent <strong>the</strong> ice, oreven on rare occasions true land-<strong>animals</strong>, which might sometimesbe stranded on distant continents or islands.But a moreeffectual because a more wide-spread agent, is to be found in<strong>the</strong> uprooted trees and rafts <strong>of</strong> driftwood <strong>of</strong>ten floated downgreat rivers and carried out to sea. Such rafts or islands aresometimes seen drifting a hundred miles from <strong>the</strong> mouth <strong>of</strong> <strong>the</strong>Ganges <strong>with</strong> living trees erect upon <strong>the</strong>m ;and <strong>the</strong> Amazon, <strong>the</strong>Orinoco, Mississippi, Congo, and most great rivers producesimilar rafts.Spix and Martius declare that <strong>the</strong>y saw at differenttimes on <strong>the</strong> Amazon, monkeys, tiger-cats, and squirrels,being thus carried down <strong>the</strong> stream. On <strong>the</strong> Parana, pumas,squirrels, and many o<strong>the</strong>r quadrupeds have been seen on <strong>the</strong>serafts ; and Admiral W. H. Smyth informed Sir C. Lyell thatamong <strong>the</strong> Philippine islands after a hurricane, he met <strong>with</strong>floating masses<strong>of</strong> wood <strong>with</strong> trees growing upon <strong>the</strong>m, so that<strong>the</strong>y were at first mistaken for islands till it was found that <strong>the</strong>ywere rapidly drifting along. Here <strong>the</strong>refore, we have amplemeans for carrying all <strong>the</strong> smaller and especially <strong>the</strong> arborealmammals out to sea ; and although in most cases <strong>the</strong>y wouldperish <strong>the</strong>re, yet in some favourable instances strong winds or


;chap, ii.] DISPERSAL AND MIGRATION. 15unusual tidal currents might carry <strong>the</strong>m safely to shores perhapsseveral hundred miles from <strong>the</strong>ir native country.<strong>The</strong> fact<strong>of</strong> green trees so <strong>of</strong>ten having been seen erect on <strong>the</strong>se rafts ismost important ;for <strong>the</strong>y would act as a sail by which <strong>the</strong> raftmight be propelled in one direction for several days in succession,and thus at last reach a shore to which a current alone wouldnever have carried it.<strong>The</strong>re are two groups <strong>of</strong> mammals whichhave quite exceptionalmeans <strong>of</strong> dispersal— <strong>the</strong> bats which.fly, and <strong>the</strong> cetacea,seals, &c, which swim. <strong>The</strong> former are capable <strong>of</strong> traversingconsiderable spaces <strong>of</strong> sea, since two North American speciesei<strong>the</strong>r regularly or occasionally visit <strong>the</strong> Bermudas, a distance<strong>of</strong> 600 miles from <strong>the</strong> mainland. <strong>The</strong> oceanic mammals (whalesand porpoises) seem to have no barrier but temperature; <strong>the</strong>polar species being unable to cross <strong>the</strong> equator, while <strong>the</strong> tropicalforms are equally unfitted for <strong>the</strong> cold polar waters.<strong>The</strong> shorefeedingmanatees, however, can only live where <strong>the</strong>y findfoodand a long expanse <strong>of</strong> rocky coast would probably be as completea barrier to <strong>the</strong>m as a few hundred miles <strong>of</strong> open ocean.<strong>The</strong> amphibious seals and walruses seem many <strong>of</strong> <strong>the</strong>m to becapable <strong>of</strong> making long sea journeys, some <strong>of</strong> <strong>the</strong> species beingfound on islands a thousand miles apart, but none <strong>of</strong> <strong>the</strong> arcticare identical <strong>with</strong> <strong>the</strong> antartic species.<strong>The</strong> otters <strong>with</strong> one exception are freshwater <strong>animals</strong>, and wehave no reason to believe <strong>the</strong>y could or would traverse any greatdistances <strong>of</strong> salt water. In fact, <strong>the</strong>y would be less liable todispersal across arms <strong>of</strong> <strong>the</strong> sea than purely terrestrial species,since <strong>the</strong>ir powers <strong>of</strong> swimming would enable <strong>the</strong>m to regain<strong>the</strong> shore ifaccidentally carried out to sea by a sudden flood.Means <strong>of</strong> Dispersal <strong>of</strong> Birds.—It would seem at first sight thatno barrierscould limit <strong>the</strong> range <strong>of</strong> birds, and that <strong>the</strong>y oughtto be <strong>the</strong> most ubiquitous <strong>of</strong> living things, and little fitted <strong>the</strong>reforeto throw any light on <strong>the</strong>, laws or causes <strong>of</strong> <strong>the</strong> <strong>geographical</strong><strong>distribution</strong> <strong>of</strong> <strong>animals</strong>. This, however, is far from being <strong>the</strong>case ; many groups <strong>of</strong> birds are almost as strictly limited bybarriers as <strong>the</strong> mammalia ; and from <strong>the</strong>ir larger numbers and<strong>the</strong> avidity <strong>with</strong> which <strong>the</strong>y have been collected, <strong>the</strong>y furnish


;16 DISTRIBUTION OF ANIMALS. [part i.materials <strong>of</strong> <strong>the</strong> greatest value for our present <strong>study</strong>. <strong>The</strong>different groups <strong>of</strong> birds <strong>of</strong>fer remarkable contrasts in <strong>the</strong> extent<strong>of</strong> <strong>the</strong>ir range, some being <strong>the</strong> most cosmopolite <strong>of</strong> <strong>the</strong> higher<strong>animals</strong>, while o<strong>the</strong>rs are absolutely confined to single spots on<strong>the</strong> earth's surface. <strong>The</strong> petrels (Procellariidce) and <strong>the</strong> gulls(Laridce) are among <strong>the</strong> greatest wanderers ; but most <strong>of</strong> <strong>the</strong>species are confined to one or o<strong>the</strong>r <strong>of</strong> <strong>the</strong> great oceans, or to <strong>the</strong>arctic or antarctic seas, a few only being found <strong>with</strong> scarcelyany variation over almost <strong>the</strong> whole globe.plovers wander along <strong>the</strong> shores as far as<strong>The</strong> sandpipers anddo <strong>the</strong> petrels over <strong>the</strong>ocean. Great numbers <strong>of</strong> <strong>the</strong>m breed in <strong>the</strong> arctic regions andmigrate as far as India and Australia, or down to Chili andBrazil ; <strong>the</strong> species <strong>of</strong> <strong>the</strong> old and new worlds, however, beinggenerally distinct. In striking contrast to <strong>the</strong>se wide rangeswe find many <strong>of</strong> <strong>the</strong> smaller perching birds, <strong>with</strong> some <strong>of</strong> <strong>the</strong>parrots and pigeons, confined to small islands <strong>of</strong> a few squaremiles in extent, or to single valleys or mountains on <strong>the</strong> mainland.Dispersal <strong>of</strong> Birds by Winds.—Those groups <strong>of</strong> birds whichpossess no powers <strong>of</strong> flight, such as <strong>the</strong> ostrich, cassowary, andapteryx, are in exactly <strong>the</strong> same position as mammalia as regards<strong>the</strong>ir means <strong>of</strong> dispersal, or are perhaps even inferior to <strong>the</strong>msince, although <strong>the</strong>y are able to cross rivers by swimming, it isdoubtful if<strong>the</strong>y eould remain so long in <strong>the</strong> water as most landquadrupeds. A very large number <strong>of</strong> short-winged birds, suchas toucans, pittas, and wrens, are perhaps worse <strong>of</strong>f; for <strong>the</strong>y canfly very few miles at a time, and on falling into <strong>the</strong> water wouldsoon be drowned. It is only <strong>the</strong> strong-flying species that canventure to cross any great width <strong>of</strong> sea ;and even <strong>the</strong>se rarely doso unless compelled by necessity to migrate in search <strong>of</strong> food, orto a more genial climate.Small and weak birds are, however,<strong>of</strong>ten carried accidentally across great widths <strong>of</strong> ocean by violentgales. This is well exemplified by <strong>the</strong> large numbers <strong>of</strong>stragglers from North America, which annually reach <strong>the</strong>Bermudas. No less than sixty-nine species <strong>of</strong> American birdshave occurred in Europe, most <strong>of</strong> <strong>the</strong>m in Britain and Heligoland.<strong>The</strong>y consist chiefly <strong>of</strong> migratory birds which in autumn


chap, ii.] BIRDS. 17return along <strong>the</strong> eastern coasts <strong>of</strong> <strong>the</strong> United States, and <strong>of</strong>tenfly from point to point across bays and inlets. <strong>The</strong>y are <strong>the</strong>nliable to be blown out tosea by storms, which are prevalent atthis season ; and it is almost always at this time <strong>of</strong> year that<strong>the</strong>ir occurrence has been noted on <strong>the</strong> shores <strong>of</strong> Europe. Itmay, however, be doubted whe<strong>the</strong>r this is not an altoge<strong>the</strong>rmodern phenomenon, dependent on <strong>the</strong> number <strong>of</strong> vessels constantlyon <strong>the</strong> Atlantic which afford resting-places to <strong>the</strong> wanderers; as it is hardly conceivable that such birds as titlarks,cuckoos, wrens, warblers, and rails, could remain on <strong>the</strong> wing<strong>with</strong>out food or rest, <strong>the</strong> time requisite to pass over 2,000 miles<strong>of</strong> ocean. It is somewhat remarkable that no European birdsreach <strong>the</strong> American coast but a few which pass by way <strong>of</strong>Iceland and Greenland ; whereas a considerable number doreach <strong>the</strong> Azores, fully half way across ; so that <strong>the</strong>ir absencecan hardly be due to <strong>the</strong> prevailing winds being westerly.<strong>The</strong>case <strong>of</strong> <strong>the</strong> Azores is, however, an argument for <strong>the</strong> unassistedpassage <strong>of</strong> birds for that distance ; since two <strong>of</strong> <strong>the</strong> finches arepeculiar ' species,' but closely allied to European forms, so that<strong>the</strong>ir progenitors must, probably, have reached <strong>the</strong> islands before<strong>the</strong> Atlantic was a commercial highway.Barriers to <strong>the</strong> Dispersal <strong>of</strong> Birds.—We have seen that, as arule, wide oceans are an almost absolute barrier to <strong>the</strong> passage <strong>of</strong>most birds from one continent to ano<strong>the</strong>r ; but much narrowerseas and straits are also very effectualbarriers where <strong>the</strong> habits<strong>of</strong> <strong>the</strong> birds are such as to preserve <strong>the</strong>m from being carriedaway by storms. All birds which frequent thickets and forests,and which feed near or on <strong>the</strong> ground, are secure from suchaccidents ; and <strong>the</strong>y are also restricted in <strong>the</strong>ir range by <strong>the</strong>extent <strong>of</strong> <strong>the</strong> forests <strong>the</strong>y inhabit. In South America a largenumber <strong>of</strong> <strong>the</strong> birds have <strong>the</strong>ir ranges determined by <strong>the</strong> extent<strong>of</strong> <strong>the</strong> forest country, while o<strong>the</strong>rs are equally limited to <strong>the</strong>open plains.Such species are also bounded by mountain rangeswhenever <strong>the</strong>se rise above <strong>the</strong> woody region.Great rivers, suchas <strong>the</strong> Amazon, also limit <strong>the</strong> range <strong>of</strong> many birds, even when<strong>The</strong><strong>the</strong>re would seem to be no difficulty in <strong>the</strong>ir crossing <strong>the</strong>m.supply <strong>of</strong> food, and <strong>the</strong> kind <strong>of</strong> vegetation, soil, and climate


18 DISPERSAL AND MIGRATION. [part i.best suited to a bird's habits, are probably <strong>the</strong> causes which markout <strong>the</strong> exact limits <strong>of</strong> <strong>the</strong> range <strong>of</strong> each species ; to which mustbe added <strong>the</strong> prevalence <strong>of</strong> enemies <strong>of</strong> ei<strong>the</strong>r <strong>the</strong> parent birds,<strong>the</strong> eggs, or <strong>the</strong> young.most where monkeys do not occur ;In <strong>the</strong> Malay Archipelago pigeons aboundand in South America <strong>the</strong>same birds are- comparatively scarce in <strong>the</strong> forest plains wheremonkeys are very abundant, while <strong>the</strong>y are plentiful on <strong>the</strong> openplainsand campos, and on <strong>the</strong> mountain plateaux, where <strong>the</strong>senest-hunting quadrupeds are rarely found. Some birds areconfined to swamps, o<strong>the</strong>rs to mountains ; some can only live onrocky streams, o<strong>the</strong>rs on deserts or grassy plains.<strong>The</strong> Phenomena <strong>of</strong> Migration.—<strong>The</strong> term " migration " is <strong>of</strong>tenapplied to <strong>the</strong> periodical or irregular movements <strong>of</strong> all <strong>animals</strong> ;but it may be questioned whe<strong>the</strong>r <strong>the</strong>re are any regular migrantsbut birds and fishes. <strong>The</strong> annual or periodical movements<strong>of</strong> mammalia are <strong>of</strong> a different class. Monkeys ascend <strong>the</strong>Himalayas in summer to a height <strong>of</strong> 10,000 to 12,000 feet, anddescend again in winter.Wolves everywhere descend from <strong>the</strong>mountains to <strong>the</strong> lowlands in severe wea<strong>the</strong>r. In dry seasonsgreat herds <strong>of</strong> antelopes move southwards towards <strong>the</strong> Cape <strong>of</strong>Good Hope. <strong>The</strong> well-known lemmings, in severe winters, atlong intervals, move down from <strong>the</strong> mountains <strong>of</strong> Scandinavia inimmense numbers, crossing lakes and rivers, eating <strong>the</strong>ir waythrough haystacks, and surmounting every obstacle till <strong>the</strong>yreach <strong>the</strong> sea, whence very few return. <strong>The</strong> alpine hare, <strong>the</strong>arctic fox, and many o<strong>the</strong>r <strong>animals</strong>, exhibit similar phenomenaon a smaller scale ; and generally it may be said, that whenevera favourable succession <strong>of</strong> seasons has led to a great multiplication<strong>of</strong> any species, it must on <strong>the</strong> pressure <strong>of</strong> hunger seekfood in fresh localities.For such movements as <strong>the</strong>se we haveno special term. <strong>The</strong> summer and winter movements bestcorrespond to true migration, but <strong>the</strong>y arealways on a smallscale, and <strong>of</strong> limited extent; <strong>the</strong> o<strong>the</strong>r movements are ra<strong>the</strong>rtemporary incursions than true migrations.<strong>The</strong> annual movements <strong>of</strong> many fishes are more strictlyanalogous to <strong>the</strong> migration <strong>of</strong> birds, since <strong>the</strong>y take placein large bodies and <strong>of</strong>ten to considerable distances, and are


;chap. 11.] BIRDS. 19immediately connected <strong>with</strong> <strong>the</strong> process <strong>of</strong> reproduction.as <strong>the</strong> salmon, enter rivers ;approach <strong>the</strong> coast in <strong>the</strong> breeding season ;Some,o<strong>the</strong>rs, as <strong>the</strong> herring and mackerel,but <strong>the</strong> exact course<strong>of</strong> <strong>the</strong>ir migrations is unknown, and owing to our completeignorance <strong>of</strong> <strong>the</strong> area each species occupies in <strong>the</strong> ocean, and <strong>the</strong>absence <strong>of</strong> such barriers and <strong>of</strong> such physical diversities as occuron <strong>the</strong> land, <strong>the</strong>y are <strong>of</strong> far less interest and less connected <strong>with</strong>our present <strong>study</strong> than <strong>the</strong> movements <strong>of</strong> birds, to which weshall now confine ourselves.Migrations <strong>of</strong> Birds.—In all <strong>the</strong> temperate parts <strong>of</strong> <strong>the</strong> globe<strong>the</strong>re are a considerable number <strong>of</strong> birds which reside only apart <strong>of</strong> <strong>the</strong> year, regularly arriving and leaving at tolerably fixedepochs. In our own country many nor<strong>the</strong>rn birds visit us inwinter, such as <strong>the</strong> fieldfare, redwing, snow-bunting, turnstone,and numerous ducks and waders ;<strong>with</strong> a few, like <strong>the</strong> black redstart,and (according to Rev. C. A. Johns) some <strong>of</strong> <strong>the</strong> woodcocksfrom <strong>the</strong> south. In <strong>the</strong> summer a host <strong>of</strong> birds appear—<strong>the</strong>cuckoo, <strong>the</strong> swifts and swallows, and numerous warblers, being<strong>the</strong> most familiar,—which stay to build <strong>the</strong>ir nests and rear <strong>the</strong>iryoung, and <strong>the</strong>n leave us again. <strong>The</strong>se are true migrants ; buta number <strong>of</strong> o<strong>the</strong>r birds visit us occasionally, like <strong>the</strong> waxwing,<strong>the</strong> oriole, and <strong>the</strong> bee-eater, and can only be classed asstragglers, which, perhaps from too rapid multiplication one yearand want <strong>of</strong> food <strong>the</strong> next, are driven to extend <strong>the</strong>ir ordinaryrange <strong>of</strong> migration to an unusual degree.We will now endeavourto sketch <strong>the</strong> chief phenomena <strong>of</strong> migration in differentcountries.Europe.—It is well ascertained that most <strong>of</strong> <strong>the</strong> birds thatspend <strong>the</strong>ir spring and summer in <strong>the</strong> temperate parts <strong>of</strong> Europepass <strong>the</strong> winter in North Africa and Western Asia. <strong>The</strong> wintervisitants, on <strong>the</strong> o<strong>the</strong>r hand, pass <strong>the</strong> summer in <strong>the</strong> extremenorth<strong>of</strong> Europe and Asia, many <strong>of</strong> <strong>the</strong>m having been found tobreed in Lapland. <strong>The</strong> arrival <strong>of</strong> migratory birds from <strong>the</strong>south is very constant as to date, seldom varying more than aweek or two, <strong>with</strong>out any regard to <strong>the</strong> wea<strong>the</strong>r at <strong>the</strong> timebut <strong>the</strong> departure is less constant, and more dependent on <strong>the</strong>wea<strong>the</strong>r.Thus <strong>the</strong> swallow always comes to us about <strong>the</strong> middle


20 DISPERSAL AND MIGRATION. [part i.<strong>of</strong> April, however cold it may be, while its departure may takeplace from <strong>the</strong> end <strong>of</strong> September to late in October, and is said byForster to occur on <strong>the</strong> first N. or KE. wind after <strong>the</strong> 20th <strong>of</strong>September.Almost all <strong>the</strong> migratory birds <strong>of</strong> Europe go southward to<strong>the</strong> Mediterranean, move alongover in three places only ;its coasts east or west, and crossei<strong>the</strong>r from <strong>the</strong> south <strong>of</strong> Spain, in <strong>the</strong>neighbourhood <strong>of</strong> Gibraltar, from Sicily over Malta, or to <strong>the</strong>east by Greece and Cyprus. <strong>The</strong>y are thus always in sight <strong>of</strong>land.<strong>The</strong> passage <strong>of</strong> most small birds (and many <strong>of</strong> <strong>the</strong> largerones too) takes place at night ;and <strong>the</strong>y only cross <strong>the</strong> Mediterraneanwhen <strong>the</strong> wind is steady from near <strong>the</strong> east or west,and when <strong>the</strong>re is moonlight.It is a curious fact, but one that seems to be well au<strong>the</strong>nticated,that <strong>the</strong> males <strong>of</strong>ten leave before <strong>the</strong> females, and bothbefore <strong>the</strong> young birds,later and alone.old ones ;which in considerable numbers migrate<strong>The</strong>se latter, however, seldom go so far as <strong>the</strong>and numbers <strong>of</strong> young birds do not cross <strong>the</strong> Mediterranean,but stay in <strong>the</strong> south <strong>of</strong> Europe.<strong>The</strong> same rule appliesto <strong>the</strong> northward migration; <strong>the</strong> young birds stopping short<strong>of</strong> <strong>the</strong> extreme arctic regions, to which <strong>the</strong> old birds migrate. 1When old and young go toge<strong>the</strong>r, however, <strong>the</strong> old birds take<strong>the</strong> lead. In <strong>the</strong> south <strong>of</strong> Europe few <strong>of</strong> <strong>the</strong> migratory birdsstay to breed, but pass on to more temperate zones ;thus, in <strong>the</strong>south <strong>of</strong> France, out <strong>of</strong> 350 species only 60 breed <strong>the</strong>re. <strong>The</strong>same species is<strong>of</strong>ten sedentary in one part <strong>of</strong> Europe and migratoryin ano<strong>the</strong>r ; thus, <strong>the</strong> chaffinch is a constant resident inEngland, Germany, and <strong>the</strong> middle <strong>of</strong> France ;<strong>the</strong> south <strong>of</strong> France and in Holland :<strong>the</strong>but a migrant inrook visits <strong>the</strong> south<strong>of</strong> France in winter only : <strong>the</strong> Falco tinnunculus is both aresident and a migrant in <strong>the</strong> south ot France, according toM. Marcel de Serres, <strong>the</strong>re being two regular passages everyyear, while a certain number always remain.1Marcel de Serres states this as a general fact for wading and swimmingbirds. He says that <strong>the</strong> old birds arrive in <strong>the</strong> extreme north almost alone,<strong>the</strong> young remaining on <strong>the</strong> shores <strong>of</strong> <strong>the</strong> Baltic, or on <strong>the</strong> lakes <strong>of</strong> Austria,Hungary, and Russia. See his prize 'essay, Des Causes des Migrations, &c.2nd ed., Paris, 1845, p. 121.


chap, ii.] BIRDS. 21We see, <strong>the</strong>n, that migration is governed by certain intelligiblelaws ; and that it varies in many <strong>of</strong> its details, even in <strong>the</strong> samespecies, according to changed conditions. It may be lookedupon as an exaggeration <strong>of</strong> a habit common to all locomotive<strong>animals</strong>, <strong>of</strong> moving about in search <strong>of</strong> food. This habit is greatlyrestricted in quadrupeds by <strong>the</strong>ir inability to cross <strong>the</strong> sea oreven to pass through <strong>the</strong> highly-cultivated valleys <strong>of</strong> suchcountries as Europe ; but <strong>the</strong> power <strong>of</strong> flight in birds enables<strong>the</strong>m to crossevery kind <strong>of</strong> country, and even moderate widths<strong>of</strong> sea ; and as <strong>the</strong>y mostly travel at night and high in <strong>the</strong> air,<strong>the</strong>ir movements are difficult to observe, and are supposed to bemore mysterious than <strong>the</strong>y perhaps are. In <strong>the</strong> tropics birdsmove about to different districts according as certain fruitsbecome ripe, certain insects abundant, or as flooded tracts dryup. On <strong>the</strong> borders <strong>of</strong> <strong>the</strong> tropics and <strong>the</strong> temperate zoneextends a belt <strong>of</strong> country <strong>of</strong> a more or less arid character, andliable to be parched at <strong>the</strong> summer solstice. In winter andearly spring its nor<strong>the</strong>rn margin is verdant, but it soon becomesburnt up, and most <strong>of</strong> its birds necessarily migrate to <strong>the</strong> morefertile regions to <strong>the</strong> north <strong>of</strong> <strong>the</strong>m.<strong>The</strong>y thus follow <strong>the</strong> springor summer as it advances from <strong>the</strong> south towards <strong>the</strong> pole, feedingon <strong>the</strong> young flower buds,<strong>the</strong> abundance <strong>of</strong> juicy larvae, and on<strong>the</strong> ripening fruits ; and as soon as <strong>the</strong>se become scarce <strong>the</strong>yretrace <strong>the</strong>ir steps homewards to pass <strong>the</strong> winter. O<strong>the</strong>rs whosehome is nearer <strong>the</strong> pole are driven south by cold, hunger, anddarkness, to more hospitable climes, returning northward in <strong>the</strong>early summer. As a typical example <strong>of</strong> a migratory bird, let ustake <strong>the</strong> nightingale. During <strong>the</strong> winter this bird inhabitsalmost all North Africa, Asia Minor, and <strong>the</strong> Jordan Valley.Early in April it passes into Europe by <strong>the</strong> three routes alreadymentioned,and spreads over France, Britain, Denmark, and <strong>the</strong>south <strong>of</strong> Sweden, which it reaches by <strong>the</strong> beginning <strong>of</strong> May.does not enter Brittany, <strong>the</strong> Channel Islands, or <strong>the</strong> western part<strong>of</strong> England, never visitingWales, except <strong>the</strong> extreme south <strong>of</strong>Glamorganshire, and rarely extending far<strong>the</strong>r north than Yorkshire.It spreads over Central Europe, through Austria andHungary to Sou<strong>the</strong>rn Russia and <strong>the</strong> warmer parts <strong>of</strong> Siberia,It


;22 DISPERSAL AND MIGRATION. [part i.but it never<strong>the</strong>less breeds in <strong>the</strong> Jordan Valley, so that insome places it is only <strong>the</strong> surplus population that migrates.In August and September, all who can- return to <strong>the</strong>ir winterquarters.Migrations <strong>of</strong> this type probably dateback from at least <strong>the</strong>period when <strong>the</strong>re was continuous land along <strong>the</strong> routepassedover; and it is a suggestive fact that this land connection isknown to have existed in recent geological times. Britain wasconnected <strong>with</strong> <strong>the</strong> Continent during,and probably before, <strong>the</strong>glacial epoch ; and Gibraltar, as well as Sicily and Malta, werealso recently united <strong>with</strong> Africa, as is proved by <strong>the</strong> fossilelephants and o<strong>the</strong>r large mammalia found in <strong>the</strong>ir caverns,<strong>the</strong> comparatively shallow water stillbyexisting in this part <strong>of</strong> <strong>the</strong>Mediterranean while <strong>the</strong> remainder is <strong>of</strong> oceanic pr<strong>of</strong>undity,and by <strong>the</strong> large amount <strong>of</strong> identity in <strong>the</strong> species <strong>of</strong> land <strong>animals</strong>still inhabiting <strong>the</strong> opposite shores <strong>of</strong> <strong>the</strong> Mediterranean. <strong>The</strong>submersion <strong>of</strong> <strong>the</strong>se two tracts <strong>of</strong> land (which were perhaps <strong>of</strong>considerable extent) would be a slow process, and from year toyear <strong>the</strong> change might be hardly perceptible. It is easy to seehow <strong>the</strong> migration that had once taken place over continuousland would be kept up, first over lagoons and marshes, <strong>the</strong>n overa narrow channel, and subsequently over a considerable sea,no one generation <strong>of</strong> birds ever perceiving any difference in <strong>the</strong>route.<strong>The</strong>re is, however, no doubt that <strong>the</strong> sea-passage is now verydangerous, to many birds. Quails cross in immense flocks, andgreat numbers are drowned at sea whenever <strong>the</strong> wea<strong>the</strong>r is unfavourable.Some individuals always stay through <strong>the</strong> winterin <strong>the</strong> south <strong>of</strong> Europe, and a few even in England and Irelandand were <strong>the</strong>sea to become a little wider <strong>the</strong> migration wouldcease, and <strong>the</strong> quail, like some o<strong>the</strong>r birds, would remaindivided between south Europe and north Africa. Aquaticbirds are observed to follow <strong>the</strong> routes <strong>of</strong> great rivers andlakes, and <strong>the</strong> shores <strong>of</strong> <strong>the</strong> sea. One great body reachescentral Europe by way <strong>of</strong> <strong>the</strong> Danube from <strong>the</strong> shores <strong>of</strong> <strong>the</strong>Black Sea; ano<strong>the</strong>r ascends <strong>the</strong> Rhone Valley from <strong>the</strong> Gulf<strong>of</strong> Lyons.


;;chap, ii.] BIRDS. 23India and China.—In <strong>the</strong> peninsula <strong>of</strong> India and in Chinagreat numbers <strong>of</strong> nor<strong>the</strong>rn birds arrive during September andOctober, and leave from March to May. Among <strong>the</strong> smallerbirds are wagtails, pipits, larks, stonechats, warblers, thrushes,buntings, shrikes, starlings, hoopoes, and quails. Some species<strong>of</strong> cranes and storks, many ducks, and greatnumbers <strong>of</strong> Scolopacidcealso visit India in winter ;and to prey upon <strong>the</strong>secome a band <strong>of</strong> rapacious birds— <strong>the</strong> peregrine falcon, <strong>the</strong> hobby,kestrel, common sparrowhawk, harrier,and <strong>the</strong> short-eared owl.<strong>The</strong>se birds are almost all natives <strong>of</strong> Europe and "Western Asia<strong>the</strong>y spread over all nor<strong>the</strong>rn and centralIndia, mingling <strong>with</strong><strong>the</strong> sedentary birds <strong>of</strong> <strong>the</strong> oriental fauna, and give to<strong>the</strong> ornithology<strong>of</strong> Hindostan at this season quite a European aspect.<strong>The</strong> peculiar species <strong>of</strong> <strong>the</strong> higher Himalayas do not as a ruledescend to <strong>the</strong> plains in winter, but merely come lower down <strong>the</strong>mountains; and in sou<strong>the</strong>rn India and Ceylon comparativelyfew <strong>of</strong> <strong>the</strong>se migratory birds appear.In China <strong>the</strong> migratory birds follow generally <strong>the</strong> coast line,coming southwards in winter from eastern Siberia and nor<strong>the</strong>rnJapan ; while a few purely tropical forms travel northwards insummer to Japan, and on <strong>the</strong> mainland as far as <strong>the</strong> valley <strong>of</strong><strong>the</strong> Amoor.North America.—<strong>The</strong> migrations <strong>of</strong> birds in North Americahave been carefully studied by resident naturalists, and presentsome interesting features. <strong>The</strong> birds <strong>of</strong> <strong>the</strong> eastern parts <strong>of</strong>North America are pre-eminently migratory, a much smaller proportionbeing permanent residents than in corresponding latitudesin Europe.Thus, in Massachusetts <strong>the</strong>re are only about 30 species<strong>of</strong> birds which are resident all <strong>the</strong> year, while <strong>the</strong> regularsummer visitors are 106. Comparing <strong>with</strong> this our own country,though considerably fur<strong>the</strong>r north,<strong>the</strong> proportions are reversed<strong>the</strong>re being 140 residents and 63 summer visitors. This differenceis clearly due to <strong>the</strong> much greater length and severity <strong>of</strong><strong>the</strong> winter, and <strong>the</strong> greater heat <strong>of</strong> summer, in America than<strong>with</strong> us. <strong>The</strong> number <strong>of</strong> permanent residents increases prettyregularly as we go southward ; but <strong>the</strong> number <strong>of</strong> birds at anylocality during <strong>the</strong> breeding season seems to increase as we go


24 DISPERSAL AND MIGRATION. [part I.northward as far as Canada, where, according to Mr. Allen, morespecies breed than in <strong>the</strong> warm Sou<strong>the</strong>rn States. Even in <strong>the</strong>extreme north, beyond <strong>the</strong> limit <strong>of</strong> forests, <strong>the</strong>re are no less than60 species which breed ; in Canada about 1 60 ; while in<strong>The</strong> extentCarolina <strong>the</strong>re are only 135, and in Louisiana, 130.<strong>of</strong> <strong>the</strong> migration varies greatly, some species only going a fewdegrees north and south, while o<strong>the</strong>rs migrate annually from<strong>the</strong> tropics to <strong>the</strong> extreme north <strong>of</strong> <strong>the</strong> continent ; and everygradation occurs between <strong>the</strong>se extremes. Among those whichmigrate fur<strong>the</strong>st are <strong>the</strong> species <strong>of</strong> Dendrceca, and o<strong>the</strong>rAmericanflycatching warblers (Mniotiltidce), many <strong>of</strong> which breedon <strong>the</strong> shores <strong>of</strong> Hudson's Bay, and spend <strong>the</strong> winter in Mexicoor <strong>the</strong> West Indian islands.<strong>The</strong> great migratory movement <strong>of</strong> American birds is almostwholly confined to <strong>the</strong> east coast ; <strong>the</strong> birds <strong>of</strong> <strong>the</strong> high centralplains and <strong>of</strong> California being for <strong>the</strong> most part sedentary, oronly migrating for short distances.All <strong>the</strong> species which reachSouth America, and most <strong>of</strong> those which winter in Mexicoand Guatemala, are exclusively eas'tern species ; though a fewEocky Mountain birds range southward along <strong>the</strong> plateaux <strong>of</strong>Mexico and Guatemala, but probably not as regular annualmigrants.In America as in Europe birds appear in spring <strong>with</strong> greatregularity, while <strong>the</strong> time <strong>of</strong> <strong>the</strong> autumnal return is less constant.More curious is <strong>the</strong> fact, also observed in both hemispheres,that <strong>the</strong>y do not all return by <strong>the</strong> same route followed ingoing northwards, some species being constant visitors to certainlocalities in spring but not in autumn, o<strong>the</strong>rs in autumn but notin spring.Some interesting cases have been observed in America <strong>of</strong> agradual alteration in <strong>the</strong> extent <strong>of</strong> <strong>the</strong> migration <strong>of</strong> certain birds.A Mexican swallow (Hirundo klnifrons) first appeared in Ohioin 1815. Year by year it increased <strong>the</strong> extent <strong>of</strong> its range tillby 1845 it had reached Maine and Canada; and it is now quotedby American writers as extending its annual migrations toHudson's Bay. An American wren (Troglodytes ludovicianus)is ano<strong>the</strong>r bird which has spread considerably northwards since


4chap, ii.] BIRDS. 25<strong>the</strong> time <strong>of</strong> <strong>the</strong> ornithologist Wilson ;and <strong>the</strong> rice-bird, or " Bobo'-link,"<strong>of</strong> <strong>the</strong> Americans, continually widens its range as riceand wheat are more extensively cultivated. Tins bird wintersin Cuba and o<strong>the</strong>r West Indian Islands, and probably also inMexico. In April it enters <strong>the</strong> Sou<strong>the</strong>rn States and passesnorthward, till in June it reaches Canada and extends west to<strong>the</strong> Saskatchewan Eiver in 54° north latitude.South Temperate America.—<strong>The</strong> migratory birds <strong>of</strong>this part<strong>of</strong> <strong>the</strong> world have been observed by Mr. Hudson at BuenosAy res. As in Europe and North America, <strong>the</strong>re are winter andsummer visitors, from Patagonia and <strong>the</strong> tropics respectively.Species <strong>of</strong> Pyrocephalus, Milvulus, swallows, and a hummingbird,are among <strong>the</strong> most regular <strong>of</strong> <strong>the</strong> summer visitors.' <strong>The</strong>yare all insectivorous birds. From Patagonia species <strong>of</strong> Tcenioptera,Cinclodes, and Centrites, come in winter, <strong>with</strong> two gulls,two geese, and six snipes and plovers.appear atFive species <strong>of</strong> swallowsBuenos Ayres in spring, some staying to breed, o<strong>the</strong>rspassing on to more temperate regions far<strong>the</strong>r south. As a rule<strong>the</strong> birds which come late and leave early are <strong>the</strong> most regular.Some are very irregular in <strong>the</strong>ir movements, <strong>the</strong> Molothrus bonariensis,for example, sometimes leaves early in autumn, sometimesremains all <strong>the</strong> winter. Some resident birds also move inwinter to districts where <strong>the</strong>y are never seen in summer.General Remarks on Migration.—<strong>The</strong> preceding summary <strong>of</strong><strong>the</strong> main facts <strong>of</strong> migration(which might have been almost indefinitelyextended, owing to <strong>the</strong> groat mass <strong>of</strong> detailed informationthat exists on <strong>the</strong> subject) appears to accord <strong>with</strong> <strong>the</strong>view already suggested, that <strong>the</strong> "instinct" <strong>of</strong> migration hasarisen from <strong>the</strong> habit <strong>of</strong> wandering in search <strong>of</strong> food common toall <strong>animals</strong>, but greatly exaggerated in <strong>the</strong> case <strong>of</strong> birds by <strong>the</strong>irpowers <strong>of</strong> flight and by <strong>the</strong> necessity for procuring a largeamount <strong>of</strong> s<strong>of</strong>t insect food for <strong>the</strong>ir unfledged young. Migrationin its simple form may be best studied in North America,where it takes place over a continuous land surface <strong>with</strong> a considerablechange <strong>of</strong> climate from south to north. We have here(as probably in Europe and elsewhere) every grade <strong>of</strong> migration,from species which merely shift <strong>the</strong> nor<strong>the</strong>rn and sou<strong>the</strong>rnVol. I.—


26 DISPERSAL AND MIGRATION. [part j.limits <strong>of</strong> <strong>the</strong>ir range a few hundred miles, so that in <strong>the</strong> centralparts <strong>of</strong> <strong>the</strong> area <strong>the</strong> species is a permanent resident, to o<strong>the</strong>rswhich move completely over 1,000 miles <strong>of</strong> latitude, so that inall <strong>the</strong> intervening districts <strong>the</strong>y are only known as birds <strong>of</strong>passage. Now, just as <strong>the</strong> rice-bird and <strong>the</strong> Mexican swallowhave extended <strong>the</strong>irinduced by human agency ;so we may presume that large numbers<strong>of</strong> speciesmigrations, owing to favourable conditionswould extend <strong>the</strong>ir range where favourable conditionsarose through natural causes. If we go back only asfar as <strong>the</strong> height<strong>of</strong> <strong>the</strong> glacial epoch, <strong>the</strong>re is reason to believethat all North America, as far south as about 40° north latitude,was covered <strong>with</strong> an almost continuous and perennial ice-sheet.At this time <strong>the</strong> migratory birds would extend up to this barrier(which would probably terminate in <strong>the</strong> midst <strong>of</strong> luxuriantvegetation, just as <strong>the</strong> glaciers <strong>of</strong> Switzerland now <strong>of</strong>ten terminateamid forests and corn-fields), and as <strong>the</strong> cold decreased and<strong>the</strong> ice retired almost imperceptibly year by year, would followit up far<strong>the</strong>r and far<strong>the</strong>r according as <strong>the</strong> peculiarities <strong>of</strong> vegetationand insect-food were more or less suited to <strong>the</strong>ir severalconstitutions. It is an ascertained fact that many individualbirds return year after year to build <strong>the</strong>ir nests in <strong>the</strong> samespot. This shows a strong local attachment, and is, in fact,<strong>the</strong> faculty or feeling on which <strong>the</strong>ir very existence probablydepends. For were <strong>the</strong>y to wander at random each year, <strong>the</strong>ywould almost certainly not meet <strong>with</strong> places so well suited to<strong>the</strong>m, and might even get into districts where <strong>the</strong>y or <strong>the</strong>iryoung would inevitably perish. It is also a curious fact that inso many cases <strong>the</strong> old birds migrate first,leaving <strong>the</strong> young onesbehind, who follow some short time later, but do not go so far as<strong>the</strong>ir parents. This is very strongly opposed to <strong>the</strong> notion <strong>of</strong>an imperative instinct. <strong>The</strong> old birds have been before, <strong>the</strong>young have not ; and it is only when <strong>the</strong> old ones have all ornearly all gone that <strong>the</strong>young go too, probably following some<strong>of</strong> <strong>the</strong> latest stragglers. <strong>The</strong>y wander, however, almost at random,and <strong>the</strong> majority are destroyed before <strong>the</strong> next spring.This is proved by <strong>the</strong> fact that <strong>the</strong> birds which return in springare as a rule not more numerous than those which came <strong>the</strong>


chap, ii.] BIRDS. 27preceding spring, whereas those which went away in autumnwere two or three times as numerous. Those young birds thatdo get back, however, have learnt by experience, and <strong>the</strong> nextyear <strong>the</strong>y take care to go <strong>with</strong> <strong>the</strong> old ones. <strong>The</strong> most strikingfact in favour <strong>of</strong> <strong>the</strong> " instinct " <strong>of</strong> migration is <strong>the</strong> " agitation,"or excitement, <strong>of</strong> confined birds at <strong>the</strong> time when <strong>the</strong>ir wildcompanions are migrating. It seems probable, however, thatthis is what may be called a social excitement, due to <strong>the</strong>anxious cries <strong>of</strong> <strong>the</strong> migrating birds ; a view supported by <strong>the</strong>fact stated by Marcel de Serres, that <strong>the</strong> black swan <strong>of</strong> Australia,when domesticated in Europe, sometimes joins wild swans in<strong>the</strong>ir northward migration.We must remember too that migrationat <strong>the</strong> proper time isin many cases absolutely essential to<strong>the</strong> existence <strong>of</strong> <strong>the</strong> species ; and it is <strong>the</strong>refore not improbablethat some strong social emotion should have been graduallydeveloped in <strong>the</strong> race, by <strong>the</strong> circumstance that all who forwant <strong>of</strong> such emotion did not join <strong>the</strong>ir fellows inevitablyperished.<strong>The</strong> mode by which a passage originallyoverland has been<strong>with</strong>out apparently stopping on <strong>the</strong> way isconverted into one over <strong>the</strong> sea <strong>of</strong>fers no insuperable difficulties,as has already been pointed out. <strong>The</strong> long flights <strong>of</strong> some birdsthought to be inexplicable,as well as <strong>the</strong>ir finding <strong>the</strong>ir nesting-place <strong>of</strong> <strong>the</strong>previous year from a distance <strong>of</strong> many hundreds or even athousand miles.But <strong>the</strong> observant powers <strong>of</strong> <strong>animals</strong> are verygreat ; and birds flying high in <strong>the</strong> air may be guided by <strong>the</strong>physical features <strong>of</strong> <strong>the</strong> country spread out beneath <strong>the</strong>m in away that would be impracticable to purely terrestrial <strong>animals</strong>.It is assumed by some writers that <strong>the</strong> breeding-place <strong>of</strong> aspecies is to be considered as its true home ra<strong>the</strong>r than that towhich it retires in winter ; but this can hardly be accepted as arule <strong>of</strong> universal application. A bird can only breed successfullywhere it can find sufficient food for its young ;and <strong>the</strong>reason probably why so many <strong>of</strong> <strong>the</strong> smaller birds leave <strong>the</strong>warm sou<strong>the</strong>rn regions to breed in temperate or even cold latitudes,is because caterpillars and o<strong>the</strong>r s<strong>of</strong>t insect larvae are<strong>the</strong>re abundant at <strong>the</strong> proper time, while in <strong>the</strong>ir winter home <strong>the</strong>


;28 DISPERSAL AND MIGRATION. [part i.larvse have all changed into winged insects. But this favourablebreeding district will change its position <strong>with</strong> change <strong>of</strong> climateand as <strong>the</strong> last great change has been one <strong>of</strong> increased warmthin all <strong>the</strong> temperate zones, it is probable that many <strong>of</strong> <strong>the</strong> migratorybirds are comparatively recent visitors. O<strong>the</strong>r changes mayhowever have taken place, affecting <strong>the</strong> vegetation and consequently<strong>the</strong> insects <strong>of</strong> a district ; and we have seldom <strong>the</strong> means<strong>of</strong> determining in any particular caseextension <strong>of</strong> range occurred.in what direction <strong>the</strong> lastFor <strong>the</strong> purposes <strong>of</strong> <strong>the</strong> <strong>study</strong> <strong>of</strong><strong>geographical</strong> <strong>distribution</strong> <strong>the</strong>refore, we must, except in specialcases, consider <strong>the</strong> true range <strong>of</strong> a species to comprise all <strong>the</strong>area winch it occupies regularly for any part <strong>of</strong> <strong>the</strong> year, whileall those districts which it only visits at more or less distantintervals,apparently driven by storms or by hunger, and whereit never regularly or permanently settles, should not be includedas forming part <strong>of</strong> its area <strong>of</strong> <strong>distribution</strong>.Means <strong>of</strong> Dispersal <strong>of</strong> Reptiles and Amphibia.—If we leaveout <strong>of</strong> consideration <strong>the</strong> true marine groups—<strong>the</strong> turtles and seasnakes—reptilesare scarcely more fitted for traversing seas andoceans than are mammalia. We accordingly find that in thoseoceanic islands which possess no indigenous mammals, land reptilesare also generally wanting.<strong>The</strong> several groups <strong>of</strong> <strong>the</strong>se <strong>animals</strong>,however, differ considerably both in <strong>the</strong>ir means <strong>of</strong> dispersaland in <strong>the</strong>ir power <strong>of</strong> resisting adverse conditions. Snakes aremost dependent on climate,becoming very scarce in temperateand cold climates and entirely ceasing at 62° north latitude, and<strong>the</strong>y do not ascend very l<strong>of</strong>ty mountains, ceasing at 6,000 feetelevation in <strong>the</strong> Alps. Some inhabit deserts, o<strong>the</strong>rs swamps andmarshes, while many are adapted for a life in forests. <strong>The</strong>yswim rivers easily, but apparently have no means <strong>of</strong> passing<strong>the</strong> sea, since <strong>the</strong>y are very rarely found on oceanic islands.Lizards are also essentially tropical, but <strong>the</strong>y go somewhatfar<strong>the</strong>r north than snakes, and ascend higher on <strong>the</strong> mountains,reaching 10,000 feet in <strong>the</strong> Alps. <strong>The</strong>y possess too someunknown means (probably in <strong>the</strong> egg-state) <strong>of</strong> passing over <strong>the</strong>ocean, since <strong>the</strong>y are found to inhabit many islands where <strong>the</strong>reare nei<strong>the</strong>r mammalia nor snakes.


chap. II.] REPTILES AND FISHES. 29<strong>The</strong> amphibia are much less sensitive to cold than are truereptiles, and <strong>the</strong>y accordingly extend much far<strong>the</strong>r north, frogsbeing found <strong>with</strong>in <strong>the</strong> arctic circle. <strong>The</strong>ir semi-aquatic lifealso gives <strong>the</strong>m facilities for dispersal, and <strong>the</strong>ir eggs are no doubtsometimes carried by aquatic birds from one pond or stream toano<strong>the</strong>r.Salt water is fatal to <strong>the</strong>m as well as to <strong>the</strong>ir eggs, andhence it arises that <strong>the</strong>y are seldom found in those oceanicislands from which mammalia are absent. Deserts and oceanswould probably form <strong>the</strong> most effectual barriers to <strong>the</strong>ir dispersal;whereas both snakes and lizards abound in deserts, andhave some means <strong>of</strong> occasionally passing <strong>the</strong> ocean which frogsand salamanders do not seem to possess.Means <strong>of</strong> Dispersal <strong>of</strong> Fishes.—<strong>The</strong> fact that <strong>the</strong> same species<strong>of</strong> freshwater fish <strong>of</strong>ten inhabit distinct river systems, provesthat <strong>the</strong>y have some means <strong>of</strong> dispersal over land. <strong>The</strong> manyau<strong>the</strong>ntic accounts <strong>of</strong> fish falling from <strong>the</strong> atmosphere, indicateone <strong>of</strong> <strong>the</strong> means by which <strong>the</strong>y may be transferred from oneriver basin to ano<strong>the</strong>r, viz., by hurricanes and whirlwinds, which<strong>of</strong>ten carry up considerable quantities <strong>of</strong> water and <strong>with</strong> it fishes<strong>of</strong> small size. In volcanic countries, also, <strong>the</strong> fishes <strong>of</strong> subterraneanstreams may sometimes be thrown up by volcanic explosions,as Humboldt relates happened in South America.Ano<strong>the</strong>rmode by which fishes may be distributed is by <strong>the</strong>ir eggs beingoccasionally carried away by aquatic birds ; and it is stated byGmelin that geese and ducks during <strong>the</strong>ir migrations feed on <strong>the</strong>eggs <strong>of</strong> fish, and that some <strong>of</strong> <strong>the</strong>se pass through <strong>the</strong>ir bodies<strong>with</strong> <strong>the</strong>ir vitality unimpaired. 1Even water-beetles flying fromone pond to ano<strong>the</strong>r might occasionally carry <strong>with</strong> <strong>the</strong>m some <strong>of</strong><strong>the</strong> smaller eggs <strong>of</strong> fishes.are also enabled toBut it is probable that fresh-water fishmigrate by changes <strong>of</strong> level causing streamsto alter <strong>the</strong>ir course and carry <strong>the</strong>ir waters into adjacent basins.On plateaux <strong>the</strong> sources <strong>of</strong> distinct river systems <strong>of</strong>ten approacheach o<strong>the</strong>r, and <strong>the</strong> same thing occurs <strong>with</strong> lateral tributarieson <strong>the</strong> lowlands near <strong>the</strong>ir mouths. Such changes, althoughsmall in extent, and occurring only at long intervals, would1Quoted in LyelPs Principles <strong>of</strong> Geology (11th ed. vol. ii. p. 374), fromAmozn. Acad. Essay 75.


30 DISPERSAL AND MIGRATION. [part i.act very powerfully in modifying <strong>the</strong> <strong>distribution</strong> <strong>of</strong> fresh-waterfish.Sea fish would seem at first sight to have almost unlimitedmeans <strong>of</strong> dispersal, but this is far from being <strong>the</strong> case. Temperatureforms a complete barrier to a large number <strong>of</strong> species, coldwater being essential to many, while o<strong>the</strong>rs can only dwell in<strong>the</strong> warmth <strong>of</strong> <strong>the</strong> tropics. Deep water is ano<strong>the</strong>r barrier tolarge numbers <strong>of</strong> species which are adapted to shores andshallows ; and thus <strong>the</strong> Atlantic is quite as impassable a gulfto most fishes as it is to birds. MaDy sea fishes migrate to alimited extent for <strong>the</strong> purpose <strong>of</strong> depositing <strong>the</strong>ir spawn infavourable situations. <strong>The</strong> herring, an inhabitant <strong>of</strong> <strong>the</strong> deepsea, comes in shoals to our coast in <strong>the</strong> breeding season ;while<strong>the</strong> salmon quits <strong>the</strong> nor<strong>the</strong>rn seas and enters our rivers, mountingupwards to <strong>the</strong> clear cold water near <strong>the</strong>ir sources to depositits eggs. Keeping in mind <strong>the</strong> essential fact that changes <strong>of</strong>temperature and <strong>of</strong> depth are <strong>the</strong> main barriers to <strong>the</strong> dispersal<strong>of</strong> fish, we shall find little difficulty in tracing <strong>the</strong> causes thathave determined <strong>the</strong>ir <strong>distribution</strong>.Means <strong>of</strong> Dispersal <strong>of</strong> Mollusca.—<strong>The</strong> marine, fresh-water, andland mollusca are three groups whose powers <strong>of</strong> dispersal andconsequent <strong>distribution</strong> are very different, and must be separatelyconsidered. <strong>The</strong> Ptcropoda, <strong>the</strong> lanthina, and o<strong>the</strong>r groups <strong>of</strong>floating molluscs, drift about in mid-ocean, and <strong>the</strong>ir dispersalis probably limited chiefly by temperature, but perhaps also by<strong>the</strong> presence <strong>of</strong> enemies or <strong>the</strong> scarcity <strong>of</strong> proper food. <strong>The</strong>univalve and bivalve mollusca, <strong>of</strong> which <strong>the</strong> whelk and <strong>the</strong>cockle may be taken as types, move so slowly in <strong>the</strong>ir adultstate, that we should expect <strong>the</strong>m to have an exceedingly limited<strong>distribution</strong>; but <strong>the</strong> young <strong>of</strong> all <strong>the</strong>se are free swimmingembryos, and <strong>the</strong>y thus have a powerful means <strong>of</strong> dispersal, andare carried by tidesand currents so as ultimately to spread overevery shore and shoal that <strong>of</strong>fers conditions favourable for <strong>the</strong>irdevelopment. <strong>The</strong> fresh water molluscs, which one might atfirstsuppose could not range beyond <strong>the</strong>ir own river-basin, areyet very widely distributed in common <strong>with</strong> almost all o<strong>the</strong>rfresh water productions ; and Mr. Darwin has shown that this is


chap, ii.] LAND-SHELLS AND INSECTS. 31due to <strong>the</strong> fact, that ponds and marshes are constantly frequentedby wading and swimming birds which arepre-eminently wanderers,and which frequently carry away <strong>with</strong> <strong>the</strong>m <strong>the</strong> seeds <strong>of</strong>plants, and <strong>the</strong> eggs <strong>of</strong> molluscs and aquatic insects. Freshwater molluscs just hatched were found toa duck's foot suspended in an aquarium ;attach <strong>the</strong>mselves toand <strong>the</strong>y would thus beeasily carried from one lake or river to ano<strong>the</strong>r, and by <strong>the</strong> help<strong>of</strong> different species <strong>of</strong> aquatic birds, might soon spread all over<strong>the</strong> globe. Even a water-beetle has been caught <strong>with</strong> a smallliving shell (Ancylus) attached to it ;and <strong>the</strong>se fly long distancesand are liable to be blown out to sea, one having been caught onboard <strong>the</strong> Beagle when forty-five miles from land. Althoughfresh water molluscs and <strong>the</strong>ir eggs must frequently be carriedout to sea, yet this cannot lead to <strong>the</strong>ir dispersal, since saltwater is almost immediately fatal to <strong>the</strong>m ;forced toand we are <strong>the</strong>reforeconclude that <strong>the</strong> apparently insignificant and uncertainmeans <strong>of</strong> dispersal above alluded to are really what haveled to <strong>the</strong>ir wide <strong>distribution</strong>.<strong>The</strong> true land-shells <strong>of</strong>fer a stillmore difficult case, for <strong>the</strong>y are exceedingly sensitive to <strong>the</strong>influence <strong>of</strong> salt water; <strong>the</strong>y are not likely to be carried byaquatic birds, and yet <strong>the</strong>y are more or less abundant all over<strong>the</strong> globe, inhabiting <strong>the</strong> most remote oceanic islands. It hasbeen found, however, that land-shells have <strong>the</strong> power <strong>of</strong> lyingdormant a long time. Some have lived two years and a halfshut up in pill boxes ;and one Egyptian desert snail came to lifeafter having been glued down to a tablet in <strong>the</strong> British Museumfor four years !We are indebted to Mr. Darwin for experiments on <strong>the</strong> power<strong>of</strong> land shells to resist seawater, and he found that when <strong>the</strong>yhad formed a membranous diaphragm over <strong>the</strong> mouth <strong>of</strong> <strong>the</strong>shell <strong>the</strong>y survived many days'immersion (in one case fourteendays) ; and ano<strong>the</strong>r experimenter, quoted by Mr. Darwin, found thatout <strong>of</strong> one hundred land shells immersed for a fortnight in <strong>the</strong> sea,twenty-seven recovered.It is <strong>the</strong>refore quite possible for <strong>the</strong>m tobe carried in <strong>the</strong> chinks <strong>of</strong> drift wood for many hundred milesacross <strong>the</strong> sea, and this is probably one <strong>of</strong> <strong>the</strong> most effectualmodes <strong>of</strong> <strong>the</strong>ir dispersal. Very young shells would also some-


32 DISPERSAL AND MIGRATION. [part i.times attach <strong>the</strong>mselves to <strong>the</strong> feet <strong>of</strong> birds walking or restingon <strong>the</strong> ground, and as many <strong>of</strong> <strong>the</strong> waders <strong>of</strong>ten go far inland,this may have been one <strong>of</strong> <strong>the</strong> methods <strong>of</strong> distributing species<strong>of</strong> land shells ;for it must always be remembered that nature canafford to wait, and that if but once in a thousand years a singlebird should convey two or three minute snails to a distant island,thisis all that is required for us to find that island well stocked<strong>with</strong> a great and varied population <strong>of</strong> land shells.Means <strong>of</strong> Dispersal <strong>of</strong> Insects a?id <strong>the</strong> Barriers luhich Limit<strong>the</strong>ir Range.—Winged insects, as a whole, have perhaps morevaried means <strong>of</strong> dispersal over <strong>the</strong> globe than any o<strong>the</strong>r highlyorganised <strong>animals</strong>. Many <strong>of</strong> <strong>the</strong>m can fly immense distances,and <strong>the</strong> more delicate ones are liable to be carried by stormsand hurricanes over a wide expanse <strong>of</strong> ocean. <strong>The</strong>y are <strong>of</strong>tenmet <strong>with</strong> far out at sea. Hawk-moths frequently fly on boardships as <strong>the</strong>y approach <strong>the</strong> shores <strong>of</strong> tropical countries, and <strong>the</strong>yhave sometimes been captured more than 250 miles from <strong>the</strong>nearest land. Dragon-flies came on board <strong>the</strong> Adventure frigatewhen fifty miles <strong>of</strong>f <strong>the</strong> coast <strong>of</strong> South America. A sou<strong>the</strong>rlywind brought flies in myriads to Admiral Smyth's ship in <strong>the</strong>Mediterranean when he was 100 miles distant from <strong>the</strong> coast <strong>of</strong>Africa. A large Indian beetle (Chrysochroa ocellata) was quiterecently caught alive in <strong>the</strong> Bay <strong>of</strong> Bengal by Captain Payne <strong>of</strong><strong>the</strong> barque William Mansoon, 273 miles from <strong>the</strong> nearest land.Darwin caught a locust 370 miles from land; and in 1844swarms <strong>of</strong> locusts several miles in extent, and as thick as <strong>the</strong>flakes in a heavy snowstorm, visited Madeira.come <strong>with</strong> perfect safety more than 300 miles ;<strong>The</strong>se must haveand as <strong>the</strong>y continuedflying over <strong>the</strong> island for a long time, <strong>the</strong>y could evidentlyhave travelled to a much greater distance, Numbers <strong>of</strong> livingbeetles belonging to seven genera, some aquatic and some terrestrial,were caught by Mr. Darwin in <strong>the</strong> open sea, seventeenmiles from <strong>the</strong> coast <strong>of</strong> South America, and <strong>the</strong>y did not seeminjured by <strong>the</strong> salt water.Almost all <strong>the</strong> accidental causes thatlead to <strong>the</strong> dispersal <strong>of</strong> <strong>the</strong> higher <strong>animals</strong> would be still morefavourable for insects. Floating trees could carry hundreds <strong>of</strong>insects for one bird or mammal ;and so many <strong>of</strong> <strong>the</strong> larvre, eggs,


;chap, ii.] DISPERSAL AND MIGRATION. 33and pupse <strong>of</strong> insects have <strong>the</strong>ir abode in solidmight survive being floated immense distances.timber, that <strong>the</strong>yGreat numbers<strong>of</strong> tropical insects have been captured in <strong>the</strong> London docks,where <strong>the</strong>y have been brought in foreign timber ;and some haveemerged from furniture after remaining torpid for many years.Most insects have <strong>the</strong> power <strong>of</strong> existing weeks or months <strong>with</strong>outfood, and some are very tenacious <strong>of</strong> life. Many beetleswill survive immersion for hours in strong spirit ; and water afew degrees below <strong>the</strong> boiling point will not always kill <strong>the</strong>m.We can <strong>the</strong>refore easily understand how, in <strong>the</strong> course <strong>of</strong> agesinsects may become dispersed by means which would be quiteinadequate in <strong>the</strong> case <strong>of</strong> <strong>the</strong> higher <strong>animals</strong>.<strong>The</strong> drift-wood andtropical fruits that reach Ireland and <strong>the</strong> Orkneys ; <strong>the</strong> doublecocoa-nuts that cross <strong>the</strong> Indian ocean from <strong>the</strong> Seychelle Islandsto <strong>the</strong> coast <strong>of</strong> Sumatra; <strong>the</strong> winds that carry volcanic dust andashes for thousands <strong>of</strong> miles ;revolving course over wide oceans ;<strong>the</strong> hurricanes that travel in <strong>the</strong>irall indicate means by whicha few insects may, at rare intervals be carried to remote regions,and become <strong>the</strong> progenitors <strong>of</strong> a group <strong>of</strong> allied forms.But <strong>the</strong> dispersal <strong>of</strong> insects requires to be looked at fromano<strong>the</strong>r point <strong>of</strong> view. <strong>The</strong>y are, <strong>of</strong> all <strong>animals</strong>, perhaps <strong>the</strong>most wonderfully adapted for special conditions ;and are so <strong>of</strong>tenfitted to fill one place in nature and one only, that <strong>the</strong> barriersagainst <strong>the</strong>ir permanent displacement are almost as numerousand as effective as <strong>the</strong>ir means <strong>of</strong> dispersal.Hundreds <strong>of</strong> species<strong>of</strong> lepidoptera, for example, can subsist in <strong>the</strong> larva state only onone species <strong>of</strong> plant ; so that even if <strong>the</strong> perfect insects werecarried to a new country, <strong>the</strong> continuance <strong>of</strong> <strong>the</strong> race would dependupon <strong>the</strong> same or a closely allied plant being abundant<strong>the</strong>re. O<strong>the</strong>r insects require succulent vegetable food all <strong>the</strong>year round, and are <strong>the</strong>refore confined to tropical regionssome can live only in deserts, o<strong>the</strong>rs in forests ; some are dependenton water-plants, some on mountain-vegetation.Manyare so intimately connected <strong>with</strong> o<strong>the</strong>r insects during somepart <strong>of</strong> <strong>the</strong>ir existence that <strong>the</strong>y could not live <strong>with</strong>out <strong>the</strong>msuch are <strong>the</strong> parasitical hymenoptera and diptera, and thosemimicking species whose welfare depends upon <strong>the</strong>ir being


34 DISTRIBUTION OF ANIMALS. [part i.mistaken for something else. <strong>The</strong>n again, insects have enemiesin every stage <strong>of</strong> <strong>the</strong>ir existence—<strong>the</strong> egg, <strong>the</strong> larva, <strong>the</strong> pupa,ind <strong>the</strong> perfect form ; and <strong>the</strong> abundance <strong>of</strong> any one <strong>of</strong> <strong>the</strong>seanemies may render <strong>the</strong>ir survival impossible in a country o<strong>the</strong>rwisewell suited to <strong>the</strong>m. Ever bearing in mind <strong>the</strong>se twoopposing classes <strong>of</strong> facts, we shall not be surprised at <strong>the</strong>enormous range <strong>of</strong> some groups <strong>of</strong> insects, and at <strong>the</strong> extremelocalization <strong>of</strong> o<strong>the</strong>rs ; and shall be able to give a rational account<strong>of</strong> many phenomena <strong>of</strong> <strong>distribution</strong> that would o<strong>the</strong>rwise seemquite unintelligible


-CHAPTEK III.DISTRIBUTION AS AFFECTED BY THE CONDITIONS AND CHANGES OFTHE EARTH'SSURFACE.<strong>The</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong> over <strong>the</strong> earth's surface, is evidentlydependent in great measure upon those grand and importantcharacteristics <strong>of</strong> our globe, <strong>the</strong> <strong>study</strong> <strong>of</strong> which is termed physicalgeography. <strong>The</strong> proportion <strong>of</strong> land and water ; <strong>the</strong> outlines and<strong>distribution</strong> <strong>of</strong> continents; <strong>the</strong> depth <strong>of</strong> seas and oceans; <strong>the</strong>position <strong>of</strong> islands ;<strong>the</strong> height, direction, and continuity <strong>of</strong> mountainchains ;<strong>the</strong> position and extent <strong>of</strong> deserts, lakes, and forests ;<strong>the</strong> direction and velocity <strong>of</strong> ocean currents, as well as <strong>of</strong> prevalentwinds and hurricanes ; and lastly, <strong>the</strong> <strong>distribution</strong> <strong>of</strong> heat andcold, <strong>of</strong> rain, snow, and ice, both in <strong>the</strong>ir means and in <strong>the</strong>irextremes, have all to be considered when we endeavour toaccount for <strong>the</strong> <strong>of</strong>ten unequal and unsymmetrical manner inwhich <strong>animals</strong> are dispersed over <strong>the</strong> globe. But even thisknowledge is insufficient unless we inquire fur<strong>the</strong>r as to <strong>the</strong>evidence <strong>of</strong> permanence possessed by each <strong>of</strong> <strong>the</strong>sefeatures, inorder that we may give due weight to <strong>the</strong> various causes thathave led to <strong>the</strong> existing facts <strong>of</strong> animal <strong>distribution</strong>.Land and Water.—<strong>The</strong> well-known fact that nearly threefourths <strong>of</strong> <strong>the</strong> surface <strong>of</strong> <strong>the</strong> earth isoccupied by water, and buta little more than one-fourth by land, is important as indicating<strong>the</strong> vast extent <strong>of</strong> ocean by which many <strong>of</strong> <strong>the</strong> continents andislands are separated from each o<strong>the</strong>r. But <strong>the</strong>re is ano<strong>the</strong>r fact


36 DISTRIBUTION OF ANIMALS. [part r.which greatly increases its importance, namely, that <strong>the</strong> meanheight <strong>of</strong> <strong>the</strong> land is very small compared <strong>with</strong> <strong>the</strong> mean depth<strong>of</strong> <strong>the</strong> sea. It has been estimated by Humboldt that <strong>the</strong> meanheight <strong>of</strong> all <strong>the</strong> land surface does not exceed a thousand feet,owing to <strong>the</strong> comparative narrowness <strong>of</strong> mountain ranges and <strong>the</strong>great extent <strong>of</strong> alluvial plains and valleys ;<strong>the</strong> ocean bed, on <strong>the</strong>contrary, not only descends deeper than <strong>the</strong> tops <strong>of</strong> <strong>the</strong> highestmountains rise above its surface, but <strong>the</strong>se pr<strong>of</strong>ound depths arebroad sunken plains, while <strong>the</strong> shallows correspond to <strong>the</strong> mountainranges, so that its mean depth is, as nearly as can be estimated,twelve thousand feet. 1Hence, as <strong>the</strong> area <strong>of</strong> water is threetimes that <strong>of</strong> <strong>the</strong> land, <strong>the</strong> total cubical contents <strong>of</strong> <strong>the</strong> land,above <strong>the</strong> sea level, would be only -^ that <strong>of</strong> <strong>the</strong> waters whichare below that level. <strong>The</strong> important result follows, that whereasit is scarcely possible that in past times <strong>the</strong> amount <strong>of</strong> land surfaceshould ever greatly have exceeded that which now exists, it isjust possible that all <strong>the</strong> land may have been at some timesubmerged ; and <strong>the</strong>refore in <strong>the</strong> highest degree probable thatamong <strong>the</strong> continual changes <strong>of</strong> land and sea that have beenalways going on, <strong>the</strong> amount <strong>of</strong> land surface has <strong>of</strong>ten beenmuch less than it is now. For <strong>the</strong> same reason it is probablethat <strong>the</strong>re have been times when large masses <strong>of</strong> land have beenmore isolated from <strong>the</strong> rest than <strong>the</strong>y are at present; just asNorth America were submerged, orSouth America would be ifas Australia would become if <strong>the</strong> Malay Archipelago were tosink beneath <strong>the</strong> ocean. It is also very important to bear inmind <strong>the</strong> fact insisted on by Sir CharlesLyell, that <strong>the</strong> shallowparts <strong>of</strong> <strong>the</strong> ocean are almost always in <strong>the</strong> vicinity <strong>of</strong> land ;that an amount <strong>of</strong> elevation that would make little differenceand<strong>the</strong> bed <strong>of</strong> <strong>the</strong> ocean, would raise up extensive tracts <strong>of</strong> dry landin <strong>the</strong> vicinity <strong>of</strong> existing continents. It is almost certain,<strong>the</strong>refore, that changes in <strong>the</strong> <strong>distribution</strong> <strong>of</strong> land and seamust have taken place more frequently by additions to, ortoiThis estimate has been made for me by Mr. Stanford from <strong>the</strong> materialsused in delineating <strong>the</strong> contours <strong>of</strong> <strong>the</strong> ocean-bed on our general map. Itembodies <strong>the</strong> result <strong>of</strong> all <strong>the</strong> soundings <strong>of</strong> <strong>the</strong> Challenger, Tuscarora, ando<strong>the</strong>r vessels; obtainable up to August, 1875.


chap, in.] CONDITIONS AFFECTING DISTRIBUTION. 37modifications <strong>of</strong> pre-existing land, than by <strong>the</strong> upheaval <strong>of</strong>entirely new Continents in mid-ocean. <strong>The</strong>se two principleswill throw light upon two constantly recurring groups <strong>of</strong> factsin <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong>,—<strong>the</strong> restriction <strong>of</strong> peculiar formsto areas not at present isolated,—and on <strong>the</strong> o<strong>the</strong>r hand, <strong>the</strong>occurrence <strong>of</strong> allied forms in lands situated on opposite<strong>of</strong> <strong>the</strong> great oceans.shoresContinental Areas.—Although <strong>the</strong> dry land <strong>of</strong> <strong>the</strong> earth'ssurface is distributed <strong>with</strong> so much irregularity, that <strong>the</strong>re ismore than twice as much north <strong>of</strong> <strong>the</strong> equator as<strong>of</strong> it,<strong>the</strong>re is southand about twice as much in <strong>the</strong> Asiatic as in <strong>the</strong> Americanhemisphere; and, what is still more extraordinary, that on ahemisphere <strong>of</strong> which a point in St. George's Channel betweenEngland and Ireland is <strong>the</strong> centre, <strong>the</strong> land is nearly equal inextent to <strong>the</strong> water, while in <strong>the</strong> opposite hemisphere it is in<strong>the</strong> proportion <strong>of</strong> only one-eighth,— yet <strong>the</strong> whole <strong>of</strong> <strong>the</strong> land isalmost continuous. It consists essentially <strong>of</strong> only three masses:<strong>the</strong> American, <strong>the</strong> Asia-African, and <strong>the</strong> Australian. <strong>The</strong> tw<strong>of</strong>ormer are only separated by thirty-six miles <strong>of</strong> shallow seaat Behring's Straits, so that it is possible to go from Cape Hornto Singapore or <strong>the</strong> Cape <strong>of</strong> Good Hope <strong>with</strong>out ever beingout <strong>of</strong> sight <strong>of</strong> land ; and owing to <strong>the</strong> intervention <strong>of</strong> <strong>the</strong>numerous islands <strong>of</strong> <strong>the</strong> Malay Archipelago <strong>the</strong> journey mightbe continued under <strong>the</strong> same conditions as far as Melbourne andHobart Town.This curious fact, <strong>of</strong> <strong>the</strong> almost perfect continuity<strong>of</strong> all <strong>the</strong> great masses <strong>of</strong> land not<strong>with</strong>standing <strong>the</strong>irextremelyirregular shape and <strong>distribution</strong>, is no doubt dependent on <strong>the</strong>circumstances just alluded to ;that <strong>the</strong> great depth <strong>of</strong> <strong>the</strong> oceansand <strong>the</strong> slowness <strong>of</strong> <strong>the</strong> process <strong>of</strong> upheaval, has almost alwaysproduced <strong>the</strong> new lands ei<strong>the</strong>r close to, or actually connected<strong>with</strong> pre-existing lands ;and this has necessarily led to a muchgreater uniformity in <strong>the</strong> <strong>distribution</strong> <strong>of</strong> organic forms, thanwould have prevailed had <strong>the</strong> continents been more completelyisolated from each o<strong>the</strong>r.<strong>The</strong> isthmuses which connect Africa <strong>with</strong> Asia,and North<strong>with</strong> South America, are, however, so small and insignificantcompared <strong>with</strong> <strong>the</strong> vast extent <strong>of</strong> <strong>the</strong> countries <strong>the</strong>y unite that


38 DISTRIBUTION OF ANIMALS. [part i.we can hardly consider <strong>the</strong>m to form more than a nominalconnection. <strong>The</strong> Isthmus <strong>of</strong> Suez indeed, being* itself a desert,and connecting districts which for a great distance are more orless desert also, does not effect any real union between <strong>the</strong> luxuriantforest-clad regions <strong>of</strong> intertropical Asia and Africa. <strong>The</strong>Isthmus <strong>of</strong> Panama is a more effectual line<strong>of</strong> union, since it ishilly, well watered, and covered <strong>with</strong> luxuriant vegetation ;andwe accordingly find that <strong>the</strong> main features <strong>of</strong> South Americanzoology are continued into Central America and Mexico. InAsia a great transverse barrier exists, dividing that continentinto a nor<strong>the</strong>rn and sou<strong>the</strong>rn portion ; and as <strong>the</strong> lowlands occuron <strong>the</strong> south and <strong>the</strong> highlands on <strong>the</strong> north <strong>of</strong> <strong>the</strong> great mountainrange, which ischange <strong>of</strong> climate is produced ;situated not far beyond <strong>the</strong> tropic, an abruptso that a belt <strong>of</strong> about a hundredmiles wide, is all that intervenes between a luxuriant tropicalregion and an almost arctic waste.Between <strong>the</strong> nor<strong>the</strong>rn part<strong>of</strong> Asia, and Europe, <strong>the</strong>re is no barrier <strong>of</strong> importance ;and it isimpossible to separate <strong>the</strong>se regions as regards <strong>the</strong> main features<strong>of</strong> animal life. Africa, like Asia, has a great transverse barrier,but it is a desert instead <strong>of</strong> a mountain chain ; and it is foundthat this desert is a more effectual barrier to <strong>the</strong> diffusion <strong>of</strong><strong>animals</strong> than <strong>the</strong> Mediterranean Sea; partly because it coincides<strong>with</strong> <strong>the</strong> natural division <strong>of</strong> a tropical from a temperate climate,but also on account <strong>of</strong> recent geological changes which we shallpresently allude to. It results <strong>the</strong>n from this outline sketch <strong>of</strong><strong>the</strong> earth's surface, that <strong>the</strong> primary divisions <strong>of</strong> <strong>the</strong> geographercorrespond approximately <strong>with</strong> those <strong>of</strong> <strong>the</strong> zoologist. Somelarge portion<strong>of</strong> each <strong>of</strong> <strong>the</strong> popular divisions forms <strong>the</strong> nucleus<strong>of</strong> a zoological region ; but <strong>the</strong> boundaries are so changed that<strong>the</strong> geographer would hardly recognise <strong>the</strong>m : it has, <strong>the</strong>refore,been found necessary to give <strong>the</strong>m those distinct "nameswill be fully explained in our next chapter.Recent Changes in <strong>the</strong>whichContinental Areas.—<strong>The</strong> important facthas been now ascertained, that a considerable portion <strong>of</strong> <strong>the</strong>Sahara south <strong>of</strong> Algeria and Morocco was under water at a veryrecent epoch. Over much <strong>of</strong> this area sea-shells, identical <strong>with</strong>those now living in <strong>the</strong> Mediterranean, are abundantly scattered,


chap, m.] CONDITIONS AFFECTING DISTRIBUTION. 39not only in depressions below <strong>the</strong> level <strong>of</strong> <strong>the</strong> sea but up to aheight <strong>of</strong> 900 feet above it. Borings for water made by <strong>the</strong>French government have shown, that <strong>the</strong>se shells occur twentyfeet deep in <strong>the</strong> sand ;and <strong>the</strong> occurrence <strong>of</strong> abundance <strong>of</strong> salt,sometimes even forming considerable hills, is<strong>of</strong> <strong>the</strong> disappearance <strong>of</strong> a large body <strong>of</strong> salt water.an additional pro<strong>of</strong>cockle is one <strong>of</strong> <strong>the</strong> most abundant <strong>of</strong> <strong>the</strong> shells found ;<strong>The</strong> commonand <strong>the</strong>Eev. H. B. Tristram discovered a new fish, in a salt lake nearly300 miles inland, but which has since been found to inhabit <strong>the</strong>Gulf <strong>of</strong> Guinea.Connected <strong>with</strong> this pro<strong>of</strong> <strong>of</strong> recent elevationin <strong>the</strong> Sahara, we have most interesting indications <strong>of</strong> subsidencein <strong>the</strong> area <strong>of</strong> <strong>the</strong> Mediterranean, which were perhaps contemporaneous.Sicily and Malta are connected <strong>with</strong> Africa by asubmerged bank from 300 to 1,200 feet below <strong>the</strong> surface ;while<strong>the</strong> depth <strong>of</strong> <strong>the</strong> Mediterranean, both to <strong>the</strong> east and west, isenormous, in some parts more than 13,000 feet ; and ano<strong>the</strong>rsubmerged bank <strong>with</strong> a depth <strong>of</strong> 1,000 feet occurs at <strong>the</strong> straits<strong>of</strong> Gibraltar. In caves in Sicily, remains <strong>of</strong> <strong>the</strong> living Africanelephant have been found by Baron Anca ;and in o<strong>the</strong>r caves Dr.Falconer discovered remains <strong>of</strong> <strong>the</strong> Elephas antiquus and <strong>of</strong> twospecies <strong>of</strong> Hippopotamus. In Malta, three species <strong>of</strong> elephanthave been discovered by Captain Spratt ;a large one closely alliedto E. antiquus and two smaller ones not exceeding five feet highwhen adult. <strong>The</strong>se facts clearly indicate, that when NorthAfrica was separated by a broad arm <strong>of</strong> <strong>the</strong> sea from <strong>the</strong> rest oi<strong>the</strong> continent, it was probably connected <strong>with</strong> Europe ;and thisexplains why zoologists find <strong>the</strong>mselves obliged to place it along<strong>with</strong> Europe in <strong>the</strong> same zoological region.Besides this change in <strong>the</strong> level <strong>of</strong> <strong>the</strong> Saharaand <strong>the</strong> Mediterraneanbasin, Europe has undergone many fluctuations in itsphysical geography in very recent times. In Wales, abundance<strong>of</strong> sea-shells <strong>of</strong> living species have been found at an elevation<strong>of</strong> 1,300 feet; and in Sardinia <strong>the</strong>re is pro<strong>of</strong> <strong>of</strong> an elevation<strong>of</strong> 300 feet since <strong>the</strong> human epoch ; and <strong>the</strong>se are only samples<strong>of</strong> many such changes <strong>of</strong> level.But <strong>the</strong>se changes, though veryimportant locally and as connected <strong>with</strong> geological problems,need not be fur<strong>the</strong>r noticed here ; as <strong>the</strong>y were not <strong>of</strong> a


40 DISTRIBUTION OF ANIMALS. [part i.nature to affect <strong>the</strong> larger features <strong>of</strong> <strong>the</strong> earth's surface or todetermine <strong>the</strong> boundaries <strong>of</strong> great zoological regions.<strong>The</strong> only o<strong>the</strong>r o<strong>the</strong>r recent change <strong>of</strong> great importance whichcan be adduced to illustrate our present subject, is that whichhas taken place between North and South America.marine<strong>The</strong> livingshells <strong>of</strong> <strong>the</strong> opposite coasts <strong>of</strong> <strong>the</strong> isthmus <strong>of</strong> Panama,as well as <strong>the</strong> corals and fishes, are generally <strong>of</strong> distinct species,but some are identical and many are closely allied ; <strong>the</strong> WestIndian fossil shells and corals <strong>of</strong> <strong>the</strong> Miocene period, however,are found to be largely identical <strong>with</strong> those <strong>of</strong> <strong>the</strong> Pacificcoast.<strong>The</strong> fishes <strong>of</strong> <strong>the</strong> Atlantic and Pacific shores <strong>of</strong> America areas a rule very distinct ; but Dr. Giin<strong>the</strong>r has recently shownthat a considerable number <strong>of</strong> species inhabiting <strong>the</strong> seas onopposite sides <strong>of</strong> <strong>the</strong> isthmus are absolutely identical. <strong>The</strong>sefacts certainly indicate, that during <strong>the</strong> Miocene epoch a broadchannel separated North and South America ; and it seems probablethat a series <strong>of</strong> elevations and subsidences have takenplace uniting and separating <strong>the</strong>m at different epochs ;<strong>the</strong> mostrecent submersion having lasted but a short time, and thus,while allowing <strong>the</strong> passage <strong>of</strong> abundance <strong>of</strong> locomotive fishes,not admitting <strong>of</strong> much change in <strong>the</strong> comparatively stationarymollusca.<strong>The</strong> Glacial Epoch as affecting <strong>the</strong> Distribution <strong>of</strong> Animals.—<strong>The</strong> remarkable refrigeration <strong>of</strong> climate in <strong>the</strong> nor<strong>the</strong>rn hemisphere<strong>with</strong>in <strong>the</strong> epoch, <strong>of</strong> existingspecies, to which <strong>the</strong> termGlacial epoch is applied, toge<strong>the</strong>r <strong>with</strong> <strong>the</strong> changes <strong>of</strong> level thataccompanied and perhaps assisted to produce it, has been one <strong>of</strong><strong>the</strong> chief agents in determining many <strong>of</strong> <strong>the</strong> details <strong>of</strong> <strong>the</strong> existing<strong>distribution</strong> <strong>of</strong> <strong>animals</strong> in temperate zones. A comparison<strong>of</strong> <strong>the</strong> effects produced by existing glaciers <strong>with</strong> certain superficialphenomena in <strong>the</strong> temperate parts <strong>of</strong> Europe and NorthAmerica, renders it certain that between <strong>the</strong> Newer Pliocene and<strong>the</strong> Eecent epochs, a largeportion <strong>of</strong> <strong>the</strong> nor<strong>the</strong>rn hemispheremust have been covered <strong>with</strong> a sheet <strong>of</strong> ice several thousandfeet thick, like that which now envelopes <strong>the</strong> interior <strong>of</strong> Greenland.Much fur<strong>the</strong>r south <strong>the</strong> mountains were covered <strong>with</strong>perpetual snow, and sent glaciers down every valley ;and all <strong>the</strong>


5;chap, in.] CONDITIONS AFFECTING DISTEIBUTION. 41great valleys on <strong>the</strong> sou<strong>the</strong>rn side <strong>of</strong> <strong>the</strong> Alps poured downstreams <strong>of</strong> ice which stretched farout into <strong>the</strong> plains <strong>of</strong> Nor<strong>the</strong>rnItaly, and have left <strong>the</strong>ir debris in <strong>the</strong> form <strong>of</strong> hugemountainous moraines, in some cases more than a thousand feethigh.In Canada and New Hampshire <strong>the</strong> marks <strong>of</strong> moving iceare found on <strong>the</strong> tops <strong>of</strong> mountains from 3,000 to 5,000 feethigh ; and <strong>the</strong> whole surface <strong>of</strong> <strong>the</strong> country around and to <strong>the</strong>north <strong>of</strong> <strong>the</strong> great lakes is scored by glaciers. Wherever <strong>the</strong>land was submerged during a part <strong>of</strong> this cold period, a depositcalled boulder-clay, or glacial-drift has been formed. This is amass <strong>of</strong> sand, clay, or gravel, full <strong>of</strong> angular or rounded stones<strong>of</strong> all sizes, up to huge blocks as large as a cottage ; and especiallycharacterized by <strong>the</strong>se stones being distributed confusedlythrough it, <strong>the</strong> largest being as <strong>of</strong>ten near <strong>the</strong> top as near <strong>the</strong>bottom, and never sorted into layers <strong>of</strong> different sizes as inmaterials carried by water. Such deposits are known to beformed by glaciers and icebergs ; when deposited on <strong>the</strong> land byglaciers<strong>the</strong>y form moraines, when carried into water and thusspread <strong>with</strong> more regularity over a wider area <strong>the</strong>y form drift.This drift is rarely found except where <strong>the</strong>re iso<strong>the</strong>r evidence <strong>of</strong>ice-action, and never south <strong>of</strong> <strong>the</strong> 40th parallel <strong>of</strong> latitude, towhich in <strong>the</strong> nor<strong>the</strong>rn hemisphere signs <strong>of</strong> ice-action extend.In <strong>the</strong> sou<strong>the</strong>rn hemisphere, inPatagonia and in New Zealand,exactly similar phenomena occur.A very interesting confirmation <strong>of</strong> <strong>the</strong> reality <strong>of</strong> this coldepoch is derived from <strong>the</strong> <strong>study</strong> <strong>of</strong> fossil remains. Both <strong>the</strong>plants and <strong>animals</strong> <strong>of</strong> <strong>the</strong> Miocene period indicate that <strong>the</strong>climate <strong>of</strong> Central Europe was decidedly warmer or more equablethan it is now ; since <strong>the</strong> flora closely resembled that <strong>of</strong> <strong>the</strong>Sou<strong>the</strong>rn United States, <strong>with</strong> a likeness also to that <strong>of</strong> EasternAsia and Australia. Many <strong>of</strong> <strong>the</strong> shells were <strong>of</strong> tropical generaand <strong>the</strong>re were numbers <strong>of</strong> large mammalia allied to <strong>the</strong>elephant, rhinoceros, and tapir. At <strong>the</strong> same time, or perhapssomewhat earlier, atemperate climate extended into <strong>the</strong> arcticregions, and allowed a magnificent vegetation <strong>of</strong> shrubs andforest trees, some <strong>of</strong> <strong>the</strong>m evergreen, ta 'flourish <strong>with</strong>in twelvedegrees <strong>of</strong> <strong>the</strong> Pole. In <strong>the</strong> Pliocene period we find ourselvesVol. I.—


42 DISTRIBUTION OF ANIMALS. [part iamong forms implying a climate very little different from <strong>the</strong>present ; and our own Crag formation furnishes evidence <strong>of</strong> agradual refrigeration <strong>of</strong> climate ; since its three divisions, <strong>the</strong>Coralline, Ked, and Norwich Crags, show a decreasing number<strong>of</strong> sou<strong>the</strong>rn, and an increasing number <strong>of</strong> nor<strong>the</strong>rn species, as weapproach <strong>the</strong> Glacial epoch. Still later than <strong>the</strong>se we have <strong>the</strong>shells <strong>of</strong> <strong>the</strong> drift, almost all <strong>of</strong> which are nor<strong>the</strong>rn and many<strong>of</strong> <strong>the</strong>m arctic species. Among <strong>the</strong> mammalia indicative <strong>of</strong>cold, are <strong>the</strong> mammoth and <strong>the</strong> reindeer.In gravels and cavedeposits<strong>of</strong> Post-Pliocene date we find <strong>the</strong> same two <strong>animals</strong>,which soon disappear as<strong>the</strong> climate approached its present condition;and Pr<strong>of</strong>essor Forbes has given a list <strong>of</strong> fifty shellswhich inhabited <strong>the</strong> British seas before <strong>the</strong> Glacial epoch andinhabit it still, but are all wanting in <strong>the</strong> glacial deposits. <strong>The</strong>whole <strong>of</strong> <strong>the</strong>se are found in <strong>the</strong> Newer Pliocene strata <strong>of</strong> Sicilyand <strong>the</strong> south <strong>of</strong> Europe, where <strong>the</strong>y escaped destruction during<strong>the</strong> glacial winter.<strong>The</strong>re are also certain facts in <strong>the</strong> <strong>distribution</strong> <strong>of</strong> plants, whichare so well explained by <strong>the</strong> Glacial epoch that <strong>the</strong>y may be saidto give an additional confirmation to it. All over <strong>the</strong> nor<strong>the</strong>rnhemisphere <strong>with</strong>in <strong>the</strong> glaciated districts, <strong>the</strong> summits <strong>of</strong> l<strong>of</strong>tymountains produce plants identical <strong>with</strong> those <strong>of</strong> <strong>the</strong> polarregions. In <strong>the</strong> celebrated case <strong>of</strong> <strong>the</strong> "White Mountains an NewHampshire, United States (latitude 45°), all <strong>the</strong> plants on <strong>the</strong>summit are arctic species, none <strong>of</strong> which exist in <strong>the</strong> lowlandsfor near a thousand miles fur<strong>the</strong>r north. It has also been remarked.that <strong>the</strong> plants <strong>of</strong> each mountain are more especiallyrelated to those <strong>of</strong> <strong>the</strong> countries directly north <strong>of</strong> it. Thus,those <strong>of</strong> <strong>the</strong> Pyrenees and <strong>of</strong> Scotland are Scandinavian, andthose <strong>of</strong> <strong>the</strong> White Mountains are all species found in Labrador.Now, remembering that we have evidence <strong>of</strong> an exceedinglymild and uniform climate in <strong>the</strong> arctic regions during <strong>the</strong>Miocene period and aevident that <strong>with</strong> each degreegradual refrigeration from that time, it is<strong>of</strong> change more and more hardyplants would be successively driven southwards ;till at last <strong>the</strong>plains <strong>of</strong> <strong>the</strong> temperate zone would be inhabited by plants, whichwere once confined to alpine heights or to <strong>the</strong> arctic regions.


chap, ill.] CONDITIONS AFFECTING DISTRIBUTION. 43As <strong>the</strong> icy mantle gradually melted <strong>of</strong>f <strong>the</strong> face <strong>of</strong> <strong>the</strong> earth<strong>the</strong>se plants would occupy <strong>the</strong> newly exposed soil, and wouldthus necessarily travel in two directions, back towards <strong>the</strong> arcticcircle and up towards <strong>the</strong> alpine peaks. <strong>The</strong> facts are thusexactly explained by a cause which independent evidence hasproved to be a real one, and every such explanation is an additionalpro<strong>of</strong> <strong>of</strong> <strong>the</strong> reality <strong>of</strong> <strong>the</strong> cause. But this explanation implies,that in cases where <strong>the</strong> Glacial epoch cannot have so actedalpine plants should not be nor<strong>the</strong>rn plants ;and a striking pro<strong>of</strong><strong>of</strong> this is to be found on <strong>the</strong> Peak <strong>of</strong> Teneriffe, a mountain12,000 feet high. In <strong>the</strong> uppermost 4,500 feet <strong>of</strong> this mountainabove <strong>the</strong> limit <strong>of</strong> trees, Von Buch found only eleven species <strong>of</strong>plants, eight <strong>of</strong> which were peculiar ; but <strong>the</strong> whole were alliedto those found at lower elevations. On <strong>the</strong> Alps or Pyrenees atthis elevation, <strong>the</strong>re would be a rich flora comprising hundreds<strong>of</strong> arctic plants ; and <strong>the</strong> absence <strong>of</strong> anything corresponding to<strong>the</strong>m in this case, in which <strong>the</strong>ir ingress was cut <strong>of</strong>f by <strong>the</strong> sea,is exactly what <strong>the</strong> <strong>the</strong>ory leads us to expect.Changes <strong>of</strong> Vegetation as affecting <strong>the</strong> Distribution <strong>of</strong> Animals.—As so many <strong>animals</strong> are dependent on vegetation, its changesimmediately affect <strong>the</strong>ir <strong>distribution</strong>. A remarkable example <strong>of</strong>this is afforded by <strong>the</strong> pre-historic condition <strong>of</strong> Denmark, asinterpreted by means <strong>of</strong> <strong>the</strong> peat-bogs and kitchen-middens.This country is now celebrated for itsbeech-trees; oaks and pinesbeing scarce ; and it is known to have had <strong>the</strong> same vegetation in<strong>the</strong> time <strong>of</strong> <strong>the</strong> Eomans. In <strong>the</strong> peat-bogs, however, are founddeposits <strong>of</strong> oak trees ; and deeper still pines alone occur. Now<strong>the</strong> kitchen-middens tell us much <strong>of</strong> <strong>the</strong> natural history <strong>of</strong>Denmark in <strong>the</strong> early Stone period ; and a curious confirmation<strong>of</strong> <strong>the</strong> fact that Denmark like Norway was <strong>the</strong>n chiefly covered<strong>with</strong> pine forests is obtained by <strong>the</strong> discovery, that <strong>the</strong> Capercailziewas <strong>the</strong>n abundant, a bird which feeds almost exclusivelyon <strong>the</strong> young shoots and seeds <strong>of</strong> pines and allied plants. <strong>The</strong>cause <strong>of</strong> this change in <strong>the</strong> vegetation is unknown ; but from <strong>the</strong>known fact that when forests are destroyed trees, <strong>of</strong> a differentkind usually occupy <strong>the</strong> ground, we may suppose that some suchchange as a temporary submergence might cause an entirely


44 DISTRIBUTION OF ANIMALS. [part i.different vegetation and a considerably modified fauna to occupy<strong>the</strong> country.Organic Changes as affecting Distribution.—We have now brieflytouched on some <strong>of</strong> <strong>the</strong> direct effects <strong>of</strong> changes in physicalgeography, climate, and vegetation, on <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong>; but <strong>the</strong> indirect effects <strong>of</strong> such changes are probably <strong>of</strong>quite equal, if not <strong>of</strong> greater importance. Every changebecomes <strong>the</strong> centre <strong>of</strong> an ever-widening circle <strong>of</strong> effects. <strong>The</strong>different members <strong>of</strong> <strong>the</strong> organic world are so bound toge<strong>the</strong>r bycomplex <strong>relations</strong>, that any one change generally involvesnumerous o<strong>the</strong>r changes, <strong>of</strong>ten <strong>of</strong> <strong>the</strong> most unexpected kind.We know comparatively little <strong>of</strong> <strong>the</strong> way in which one animalor plant is bound up <strong>with</strong> o<strong>the</strong>rs, but we know enough to assureus that groups <strong>the</strong> most apparently disconnected are <strong>of</strong>tendependent on each o<strong>the</strong>r. We know, for example, that <strong>the</strong>introduction <strong>of</strong> goats into St. Helena utterly destroyed a wholeflora <strong>of</strong> forest trees ; and <strong>with</strong> <strong>the</strong>m all <strong>the</strong> insects, mollusca, andperhaps birds directly or indirectly dependent on <strong>the</strong>m. Swine,which ran wild in Mauritius, exterminated <strong>the</strong> Dodo. <strong>The</strong> same<strong>animals</strong> are known to be <strong>the</strong> greatest enemies <strong>of</strong> venomousserpents. Cattle will, in many districts, wholly prevent <strong>the</strong>growth <strong>of</strong> trees ;and <strong>with</strong> <strong>the</strong> trees <strong>the</strong> numerous insects dependenton those trees, and <strong>the</strong> birds which fed upon <strong>the</strong> insects,must disappear, as well as <strong>the</strong> small mammalia which feed on<strong>the</strong> fruits, seeds, leaves, or roots. Insects again have <strong>the</strong> mostwonderful influence on <strong>the</strong> "range <strong>of</strong> mammalia.In Paraguay acertain species <strong>of</strong> fly abounds which destroys new-born cattleand horses ;and thus nei<strong>the</strong>r <strong>of</strong> <strong>the</strong>se <strong>animals</strong> have run wild inthat country, although <strong>the</strong>y abound both north and south<strong>of</strong> it.This inevitably leads to a great difference in <strong>the</strong> vegetation <strong>of</strong>Paraguay, and through that toa difference in its insects, birds,reptiles, and wild mammalia. On what causes <strong>the</strong> existence <strong>of</strong><strong>the</strong> fly depends we do not know, but it is not improbable that somecomparatively slight changes in <strong>the</strong> temperature or humidity <strong>of</strong><strong>the</strong> air at a particular season, or <strong>the</strong> introduction <strong>of</strong> some enemymight lead to its extinction or banishment. <strong>The</strong> whole face <strong>of</strong><strong>the</strong> country would <strong>the</strong>n soon be changed : new species would


chap, in.] CONDITIONS AFFECTING DISTRIBUTION. 46come in, while many o<strong>the</strong>rs would be unable to live <strong>the</strong>re ; and<strong>the</strong> immediate cause <strong>of</strong> this great alteration would probably bequite imperceptible to us, even if we could watch it in progressyear by year. So, in South Africa, <strong>the</strong> celebrated Tsetse flyinhabits certain districts having well defined limits ; and whereit abounds no horses, dogs, or cattle can live. Yet asses,zebras, and antelopes are unaffected by it. So long as this flycontinues tc exist, <strong>the</strong>re is a living barrier to <strong>the</strong> entrance <strong>of</strong>certain <strong>animals</strong>, quite as effectual as a l<strong>of</strong>ty mountain rangeor a wide arm <strong>of</strong> <strong>the</strong> sea. <strong>The</strong> complex <strong>relations</strong> <strong>of</strong> one form<strong>of</strong> life <strong>with</strong> o<strong>the</strong>rs is nowhere better illustrated than in Mr.Darwin's celebrated case <strong>of</strong> <strong>the</strong> cats and clover, as given in hisOrigin <strong>of</strong> Species, 6th ed., p. 57. He has observed that bothwild heartsease and red-clover are fertilized in this country byhumble-bees only, so that <strong>the</strong> production <strong>of</strong> seed depends on<strong>the</strong> visits <strong>of</strong> <strong>the</strong>se insects. A gentleman who has speciallystudied humble-bees finds that <strong>the</strong>y are largely kept down byfield-mice, which destroy <strong>the</strong>ir combs and nests. Field-micein <strong>the</strong>ir turn are kept down by cats ;and probably also by owls;so that <strong>the</strong>se carnivorous <strong>animals</strong> are really <strong>the</strong> agents in renderingpossible <strong>the</strong> continued existence <strong>of</strong> red-clover and wildheartsease. For if <strong>the</strong>y were absent, <strong>the</strong> field-mice having noenemies, would multiply to such an extent as to destroy, all <strong>the</strong>humble-bees ; and <strong>the</strong>se two plants would <strong>the</strong>n produce noseed and soon become extinct.Mr. Darwin has also shown that one species <strong>of</strong>ten exterminatesano<strong>the</strong>r closely allied to it, when <strong>the</strong> two are brought intocontact. One species <strong>of</strong> swallow and thrush are known tohave increased at <strong>the</strong> expense <strong>of</strong> allied species. Rats, carriedall over <strong>the</strong> world by commerce, are continually extirpatingo<strong>the</strong>r species <strong>of</strong> rats. <strong>The</strong> imported hive-bee is, in Australia,rapidly exterminating a native stingless bee. Any slight change,<strong>the</strong>refore, <strong>of</strong> physical geography or <strong>of</strong> climate, which allowsallied species hi<strong>the</strong>rto inhabiting distinct areas to come intocontact, will <strong>of</strong>ten lead to <strong>the</strong> extermination <strong>of</strong> one <strong>of</strong> <strong>the</strong>m; andthis extermination will be effected by no external force, by noactual enemy, but merely because <strong>the</strong> one is slightly better


—46 DISTRIBUTION OF ANIMALS. [part i.adapted to live, to increase, and to maintain itself under adversecircumstances, than <strong>the</strong> o<strong>the</strong>r.Now if we consider carefully <strong>the</strong> few suggestive facts herereferred- to (and many o<strong>the</strong>rs <strong>of</strong> like import are to be found inMr. Darwin's various works), we shall be led to conclude that<strong>the</strong> several species, genera, families, and orders, both <strong>of</strong> <strong>animals</strong>and vegetables which inhabit any extensive region, are boundtoge<strong>the</strong>r by a series <strong>of</strong> complex <strong>relations</strong> ;so that <strong>the</strong> increase,diminution, or extermination <strong>of</strong> any one, may set in motion aseries <strong>of</strong> actions and reactions more or less affecting a largeportion <strong>of</strong> <strong>the</strong> whole, and requiring perhaps centuries <strong>of</strong> fluctuationbefore <strong>the</strong> balance is restored. <strong>The</strong> range <strong>of</strong> any speciesor group in such a region, will in many cases (perhaps in most)be determined, not by physical barriers, but by <strong>the</strong> competition<strong>of</strong> o<strong>the</strong>r organisms. Where barriers have existed from a remoteepoch, <strong>the</strong>y will at first have kept back certain <strong>animals</strong> fromcoming in contact <strong>with</strong> each o<strong>the</strong>r; but when <strong>the</strong> assemblage<strong>of</strong> organisms on <strong>the</strong> two sides <strong>of</strong> <strong>the</strong> barrier have, after manyages, come to form a balanced organic whole, <strong>the</strong> destruction<strong>the</strong> barrier may lead to a very partial intermingling <strong>of</strong> <strong>the</strong>peculiar forms <strong>of</strong> <strong>the</strong> two regions. Each will have becomemodified in special ways adapted to <strong>the</strong> organic and physicalconditions <strong>of</strong> <strong>the</strong> country, and will form a living barrier to <strong>the</strong>entrance <strong>of</strong> <strong>animals</strong> less perfectly adapted to those conditions.Thus while <strong>the</strong> abolition <strong>of</strong> ancient barriers will always leadtomuch intermixture <strong>of</strong> forms, much extermination and widespreadalteration in some families <strong>of</strong> <strong>animals</strong> ; o<strong>the</strong>r importantgroups will be unable materially to alter <strong>the</strong>ir range ;or <strong>the</strong>ymay make temporary incursions into <strong>the</strong> new territory, and beultimately driven back to very near <strong>the</strong>ir ancient limits.In order to make thissomewhat difficult subject more intelligible,it may be well to consider <strong>the</strong> probable effects <strong>of</strong> certainhypo<strong>the</strong>tical conditions <strong>of</strong> <strong>the</strong> earth's surface :1. If <strong>the</strong> dry land <strong>of</strong> <strong>the</strong> globe had been from <strong>the</strong> firstcontinuous, and nowhere divided up by such boundaries as l<strong>of</strong>tymountain ranges, wide deserts, or arms <strong>of</strong> <strong>the</strong> sea, it seemsprobable that none <strong>of</strong> <strong>the</strong> larger groups (as orders, tribes, or<strong>of</strong>;


chap, in.] CONDITIONS AFFECTING DISTRIBUTION. 47families) would have a limited range ; but, as is to some extent<strong>the</strong> case in tropical America* east <strong>of</strong> <strong>the</strong> Andes, every suchgroup would be represented over <strong>the</strong> whole area, by countlessminute modifications <strong>of</strong> form adapted to local conditions.2. One great physical barrier would, however, even <strong>the</strong>nexist ;<strong>the</strong> hot equatorial zone would divide <strong>the</strong> faunas and floras<strong>of</strong> <strong>the</strong> colder regions <strong>of</strong> <strong>the</strong> nor<strong>the</strong>rn and sou<strong>the</strong>rn hemispheresfrom any chance <strong>of</strong> intermixture. This one barrier would bemore effectual than it is now, since <strong>the</strong>re would be no l<strong>of</strong>tymountain ranges to serve as a bridge for <strong>the</strong> partial interchange<strong>of</strong> nor<strong>the</strong>rn and sou<strong>the</strong>rn forms.3. If such a condition <strong>of</strong> <strong>the</strong> earth as here supposed continuedfor very long periods, we may conceive that <strong>the</strong> actionand reaction <strong>of</strong> tjie various organisms on each o<strong>the</strong>r, combined<strong>with</strong> <strong>the</strong> influence <strong>of</strong> very slowly changing physical conditions,would result in an almost perfect organic balance, which wouldbe manifested by a great stability in <strong>the</strong> average numbers, <strong>the</strong>local range, and <strong>the</strong> peculiar characteristics <strong>of</strong> every species.4. Under such a condition <strong>of</strong> things it is not improbable that<strong>the</strong> total number <strong>of</strong> clearly differentiated specific forms might bemuch greater than it is now, though <strong>the</strong> number <strong>of</strong> generic andfamily types might perhaps be less ;for dominant species wouldhave had ample time to spread into every locality where <strong>the</strong>ycould exist, and would <strong>the</strong>n become everywhere modified int<strong>of</strong>orms best suited to <strong>the</strong> permanent local conditions.5. Now let us consider what would be <strong>the</strong> probable effect <strong>of</strong><strong>the</strong> introduction <strong>of</strong> a barrier, cutting <strong>of</strong>f a portion <strong>of</strong> this homogeneousand well-balanced world. Suppose, for instance, that asubsidence took place, cutting <strong>of</strong>f by a wide arm <strong>of</strong> <strong>the</strong> sea alarge and tolerably varied island. <strong>The</strong> first and most obviousresult would be that <strong>the</strong> individuals <strong>of</strong> a number <strong>of</strong> specieswould be divided into two portions, while o<strong>the</strong>rs, <strong>the</strong> limits <strong>of</strong>whose range agreed approximately <strong>with</strong> <strong>the</strong> line <strong>of</strong> subsidence,would exist in unimpaired numbers on <strong>the</strong> new island or on <strong>the</strong>main land. But <strong>the</strong> species whose numbers were diminishedand whose original area was also absolutely diminished by <strong>the</strong>portion now under <strong>the</strong> sea, would not be able to hold <strong>the</strong>ir


;48 DISTRIBUTION OF ANIMALS. [part i.ground against <strong>the</strong> rival forms whose numbers were intact.Some would probably diminish and rapidly die out; o<strong>the</strong>rswhich produced favourable varieties, might be so modified bynatural selection as to maintain <strong>the</strong>ir existence under a differentform ; and such changes would take place in varying modes on<strong>the</strong> two sides <strong>of</strong> <strong>the</strong> new strait.6. But <strong>the</strong> progress <strong>of</strong> <strong>the</strong>se changes would necessarily affect<strong>the</strong> o<strong>the</strong>r species in contact <strong>with</strong> <strong>the</strong>m. New places would beopened in <strong>the</strong> economy <strong>of</strong> nature which many would struggleobtain ; and modification would go on in ever-widening circleand very long periods <strong>of</strong> time might be required to bring <strong>the</strong>whole again into a state <strong>of</strong> equilibrium.7. A new set <strong>of</strong> factors would in <strong>the</strong> meantime have comeinto play.<strong>The</strong> sinking <strong>of</strong> land and <strong>the</strong> influx, <strong>of</strong> a large body<strong>of</strong> water could hardly take place <strong>with</strong>out producing importantclimatal changes.<strong>The</strong> temperature, <strong>the</strong> winds, <strong>the</strong> rains, mightall be affected, and more or less changed in duration and amount.This would lead to a quite distinct movement in <strong>the</strong> organicworld.Vegetation would certainly be considerably affected, andthrough this <strong>the</strong> insect tribes. We have. seen how closely <strong>the</strong>life <strong>of</strong> <strong>the</strong> higher <strong>animals</strong> is <strong>of</strong>ten bound up <strong>with</strong> that <strong>of</strong> insectsand thus a set <strong>of</strong> changes might arise that would modify <strong>the</strong>numerical proportions, and even <strong>the</strong> forms and habits <strong>of</strong> a greatnumber <strong>of</strong> species, would completely exterminate some, and raiseo<strong>the</strong>rs from a subordinate to a dominant position. And all <strong>the</strong>sechanges would occur differently on oppositetosides <strong>of</strong> <strong>the</strong> strait,since <strong>the</strong> insular climate could not fail to differ considerablyfrom that <strong>of</strong> <strong>the</strong> continent.8. But <strong>the</strong> two sets <strong>of</strong> changes, as above indicated, producedby different modes <strong>of</strong> action <strong>of</strong> <strong>the</strong> same primary cause, wouldact and react on each o<strong>the</strong>r ;and thus lead to such a far-spreadingdisturbance <strong>of</strong><strong>the</strong> organic equilibrium as ultimately perhapsto affect in one way or ano<strong>the</strong>r, every form <strong>of</strong> life upon <strong>the</strong>earth.This hypo<strong>the</strong>tical case is useful as enabling us better to realizehow wide-spreading might be <strong>the</strong> effects <strong>of</strong> one <strong>of</strong> <strong>the</strong> simplestchanges <strong>of</strong> physical geography, upon a compact mass <strong>of</strong> mutually


chap, in.] CONDITIONS AFFECTING DISTRIBUTION. 49adapted organisms. In <strong>the</strong> actual state <strong>of</strong> things, <strong>the</strong> physicalchanges that occur and have occurred through all geological epochsare larger and more varied. Almost every mile <strong>of</strong> land surfacehas been again and again depressed beneath <strong>the</strong> ocean ;most <strong>of</strong><strong>the</strong> great mountain chains have ei<strong>the</strong>r originated or greatlyincreased in height during <strong>the</strong> Tertiary period ; marvellousalterations <strong>of</strong> climate and vegetation have taken place over half<strong>the</strong> land-surface <strong>of</strong> <strong>the</strong> earth ; and all <strong>the</strong>se vast changes haveinfluenced a globe so cut up by seas and oceans, by desertsand snow-clad mountains, that in many <strong>of</strong> its more isolatedland-masses ancient forms <strong>of</strong> life have been preserved, which,in <strong>the</strong> more extensive and more varied continents have longgiven way to higher types. How complex <strong>the</strong>n must have been<strong>the</strong> actions and reactions such a state <strong>of</strong> things would bringabout ; and how impossible must it be for us to guess, in mostcases, at <strong>the</strong> exact nature <strong>of</strong> <strong>the</strong> forces that limit <strong>the</strong> range <strong>of</strong>some species and cause o<strong>the</strong>rs to be rare or to become extinct !All that we can in general hope to do is, to trace out, more orless hypo<strong>the</strong>tically, some <strong>of</strong> <strong>the</strong> larger changes in physicalgeography that have occurred during <strong>the</strong> ages immediately preceedingour own, and to estimate <strong>the</strong> effect <strong>the</strong>y will probablyhave produced on animal <strong>distribution</strong>.We may <strong>the</strong>n, by <strong>the</strong>aid <strong>of</strong> such knowledge as to past organic mutations as <strong>the</strong> geologicalrecord supplies us <strong>with</strong>, be able to determine <strong>the</strong> probablebirthplace and subsequent migrations <strong>of</strong> <strong>the</strong> more importantgenera and families; and thus obtain some conception <strong>of</strong> thatgrand series <strong>of</strong> co-ordinated changes in <strong>the</strong> earth and its inhabitants,whose final result is seen in <strong>the</strong> forms and <strong>the</strong> <strong>geographical</strong><strong>distribution</strong> <strong>of</strong> existing <strong>animals</strong>.


CHAPTER IV.ON ZOOLOGICAL REGIONS.To <strong>the</strong> older school <strong>of</strong> Naturalists <strong>the</strong> native country <strong>of</strong> an animalwas <strong>of</strong> little importance, except in as far as climates differed.Animals were supposed to be specially adapted to live in certainzones or under certain physical conditions, and it was hardlyrecognised that apart from <strong>the</strong>se conditions <strong>the</strong>re was anyinfluence in locality which could materially affect <strong>the</strong>m. Itwas believed that, while<strong>the</strong> <strong>animals</strong> <strong>of</strong> tropical, temperate, andarctic climates, essentially differed ; those <strong>of</strong> <strong>the</strong> tropics wereessentially alike all over <strong>the</strong> world. A group <strong>of</strong> <strong>animals</strong> wassaid to inhabit <strong>the</strong> "Indies;" and important differences <strong>of</strong>structure were <strong>of</strong>ten overlooked from <strong>the</strong> idea, that creaturesequally adapted to live in hot countries and <strong>with</strong> certaingeneral resemblances, would naturally be related to each o<strong>the</strong>r.Thus <strong>the</strong> Toucans and Hornbills, <strong>the</strong> Humming- Birds and Sun-Birds, and even <strong>the</strong> Tapirs and <strong>the</strong> Elephants, came to bepopularly associated as slightly modified varieties <strong>of</strong> tropicalforms <strong>of</strong> life; while to naturalists, who were acquainted <strong>with</strong><strong>the</strong> essential differences <strong>of</strong> structure, it was a never-failingsource <strong>of</strong> surprise, that under climates and conditions soapparently identical, such strangely divergent forms shouldbe produced.To <strong>the</strong> modern naturalist, on <strong>the</strong> o<strong>the</strong>r hand, <strong>the</strong> nativecountry (or " habitat " as it is technically termed) <strong>of</strong> an animal


chap, iv.] ZOOLOGICAL KEGIONS. 51or a group <strong>of</strong> <strong>animals</strong>, is a matter <strong>of</strong> <strong>the</strong> first importance ; and,as regards <strong>the</strong> general history <strong>of</strong> life upon <strong>the</strong> globe, may beconsidered to be one <strong>of</strong> its essential characters. <strong>The</strong> structure,affinities, and habits <strong>of</strong> a species, now form only a part <strong>of</strong> itsnatural history. We require also to know its exact range at<strong>the</strong> present day and in prehistoric times, and to have someknowledge <strong>of</strong> its geological age, <strong>the</strong> place <strong>of</strong> its earliest appearanceon <strong>the</strong> globe, and <strong>of</strong> <strong>the</strong> various extinct forms most nearlyallied to it. To those who accept <strong>the</strong> <strong>the</strong>ory <strong>of</strong> development asworked out by Mr. Darwin, and <strong>the</strong> views as to <strong>the</strong> generalpermanence and immense antiquity <strong>of</strong> <strong>the</strong> great continentsandoceans so ably developed by Sir Charles Lyell, it ceases to be amatter <strong>of</strong> surprise that <strong>the</strong> tropics <strong>of</strong> Africa, Asia,and Americashould differ in <strong>the</strong>ir productions, but ra<strong>the</strong>r that <strong>the</strong>y shouldhave anything in common.<strong>The</strong>ir similarity, not <strong>the</strong>ir diversity,is <strong>the</strong> fact that must frequently puzzles us.<strong>The</strong> more accurate knowledge we have <strong>of</strong> late years obtained<strong>of</strong> <strong>the</strong> productions <strong>of</strong> many remote regions, combined <strong>with</strong> <strong>the</strong>greater approaches that have been made to a naturalclassification<strong>of</strong> <strong>the</strong> higher <strong>animals</strong>, has shown, that every continent orwell-marked division <strong>of</strong> a continent, every archipelago andeven every island, presents problems <strong>of</strong> more or less complexityto <strong>the</strong> student <strong>of</strong> <strong>the</strong> <strong>geographical</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong>. Ifwe take up <strong>the</strong> subject from <strong>the</strong> zoological side, and <strong>study</strong> anyfamily, order, or even extensive genus, we are almost sure tomeet <strong>with</strong> some anomalies ei<strong>the</strong>r in <strong>the</strong> present or past<strong>distribution</strong><strong>of</strong> <strong>the</strong> various forms. Let us adduce a few examples <strong>of</strong><strong>the</strong>se problems.Deer have a wonderfully wide range, over <strong>the</strong> whole <strong>of</strong> Europe,Asia, and North and South America ;yet in Africa south <strong>of</strong><strong>the</strong> great desert <strong>the</strong>re are none.Bears range over <strong>the</strong> whole <strong>of</strong>Europe, Asia, and North America, and true pigs <strong>of</strong> <strong>the</strong> genusSus, over all Europe and Asia and as far as New Guinea ;yetboth bears and pigs, like deer, are absent from Tropical andSouth Africa.Again, <strong>the</strong> West Indian islands possess very few Mammalia,all <strong>of</strong> small size and allied to those <strong>of</strong> America, except oneBOSTON UNIVERSITY~'COLLEGE Or r*Al ARTS


;52 DISTRIBUTION OF ANIMALS. [part i.genus ; and that belongs to an Order, " Insectivora," entirelyabsent from South America, and to a family, " Centetidae," all<strong>the</strong> o<strong>the</strong>r species <strong>of</strong> which inhabit Madagascar only. And asif to add force to this singular correspondence we have oneMadagascar species <strong>of</strong> a beautiful day-flying Moth, Urania, all<strong>the</strong> o<strong>the</strong>r species <strong>of</strong> which inhabit tropical America. <strong>The</strong>seinsects are gorgeously arrayed in green and gold, and areunlike any o<strong>the</strong>r Lepidoptera upon <strong>the</strong> globe.<strong>The</strong> island <strong>of</strong> Ceylon generally agrees in itsquiteproductions <strong>with</strong><strong>the</strong> Sou<strong>the</strong>rn part <strong>of</strong> India;yet it has several birds which areallied to Malayan and not to Indian groups, and a fine<strong>of</strong> <strong>the</strong> genus HcsHa, as well as severalare purely Malayan.butterflygenera <strong>of</strong> beetles, whichVarious important groups <strong>of</strong> <strong>animals</strong> are distributed in away not easy to explain. <strong>The</strong> anthropoid apes in West Africaand Borneo ; <strong>the</strong> tapirs in Malaya and South America ; <strong>the</strong>camel tribe in <strong>the</strong> deserts <strong>of</strong> Asia and <strong>the</strong> Andes ; <strong>the</strong> trogonsin South America and Tropical Asia, <strong>with</strong> one species in Africa;<strong>the</strong> marsupials in Australia and America, are examples.<strong>The</strong> cases here adduced (and <strong>the</strong>y might be greatly multiplied)are merely to show <strong>the</strong> kind <strong>of</strong> problems <strong>with</strong> which <strong>the</strong>naturalist now has to deal ; and in order to do so he requiressome system <strong>of</strong> <strong>geographical</strong> arrangement, which shall serve<strong>the</strong> double purpose <strong>of</strong> affording a convenient subdivisionsubject, and at <strong>the</strong> same time <strong>of</strong> giving<strong>of</strong> hisexpression to <strong>the</strong> mainresults at which he has arrived. Hence <strong>the</strong> recent discussionson " Zoological Eegions," or, what are <strong>the</strong> most naturalprimary divisions <strong>of</strong> <strong>the</strong> earth as regards its forms <strong>of</strong> animallife.<strong>The</strong> divisions in use till quite recently were <strong>of</strong> two kindsei<strong>the</strong>r those ready made by geographers, more especially <strong>the</strong>quarters or continents <strong>of</strong> <strong>the</strong> globe ; or those determined byclimate and marked out by certain parallels <strong>of</strong> latitude or byiso<strong>the</strong>rmal lines. Ei<strong>the</strong>r <strong>of</strong> <strong>the</strong>se methods was better thannone at all; but from <strong>the</strong> various considerations explained in<strong>the</strong> preceding chapters, it will be evident, that such divisionsmust have <strong>of</strong>ten been very unnatural, and have disguised many


chap, iv.] ZOOLOGICAL REGIONS. 53<strong>of</strong> <strong>the</strong> most important and interesting phenomena which a<strong>study</strong> <strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong> presents to us.<strong>The</strong> merit <strong>of</strong> initiating a more natural system, that <strong>of</strong> determiningzoological regions, not by any arbitrary or a priori considerationbut by <strong>study</strong>ing <strong>the</strong> actualranges <strong>of</strong> <strong>the</strong> more importantgroups <strong>of</strong> <strong>animals</strong>, is due to Mr. Sclater, who, in 1857, establishedsix primary zoological regions from a detailed examination <strong>of</strong><strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> chief genera and families <strong>of</strong> Birds.Before stating what <strong>the</strong>se regions are, what objections havebeen made to <strong>the</strong>m, what o<strong>the</strong>r divisions have been sinceproposed, and what are those which we shall adopt in thiswork, it will be well to consider <strong>the</strong> general principles whichshould guide us in <strong>the</strong> choice between rival systems.Principles on which Zoological Regions should he formed.—It will be evident in <strong>the</strong> first place that nothing like a perfectzoological division <strong>of</strong> <strong>the</strong> earth is possible. <strong>The</strong> causesthat have led to <strong>the</strong> present <strong>distribution</strong> <strong>of</strong> animal life are sovaried, <strong>the</strong>ir action and reaction have been so complex, thatanomalies and irregularities are sure to exist which will mar <strong>the</strong>symmetry <strong>of</strong> any rigid system. On two main points every systemyet proposed, or that probably can be proposed, is open toobjection ; <strong>the</strong>y are,—lstly, that <strong>the</strong> several regions are not <strong>of</strong>equal rank ;—2ndly, that <strong>the</strong>y are not equally applicable toclasses <strong>of</strong> <strong>animals</strong>. As to <strong>the</strong> first objection, it will be foundimpossible to form any three or more regions, each <strong>of</strong> which differsfrom <strong>the</strong> rest in an equal degree or in <strong>the</strong> same manner. Onewill surpass all o<strong>the</strong>rs in <strong>the</strong> possession <strong>of</strong> peculiar families;ano<strong>the</strong>r will have many characteristic genera ;be mainly distinguished by negative characters.allwhile a third will<strong>The</strong>re will alsobe found many intermediate districts, which possess some <strong>of</strong> <strong>the</strong>characteristics <strong>of</strong> two well-marked regions, <strong>with</strong> a few specialfeatures <strong>of</strong> <strong>the</strong>ir own, or perhaps <strong>with</strong> none; and it will be adifficult question to decide in all cases which region shouldpossess this doubtful territory, or whe<strong>the</strong>r it should be formedinto a primary region itself. Again, two regions which havenow well-marked points <strong>of</strong> difference, may be shown to have beenmuch more alike at a comparatively recent geological epoch;


54 DISTRIBUTION OF ANIMALS. [part i.and this, it may be said, proves <strong>the</strong>ir fundamental unity andthat <strong>the</strong>y ought to form but one primary region. To obviatesome <strong>of</strong> <strong>the</strong>se difficulties a binary or dichotomous division issometimes proposed; that portion <strong>of</strong> <strong>the</strong> earth which differsmost from <strong>the</strong> rest being cut <strong>of</strong>f as a region equal in rankto all that remains, which is subjected again and again to<strong>the</strong> same process.To decide <strong>the</strong>se various points it seems advisable that convenience,intelligibility, and custom, should largely guide us.<strong>The</strong> first essential is, a broadly marked and easily rememberedset <strong>of</strong> regions ;which correspond, as nearly as truth to naturewill allow, <strong>with</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> most important groups<strong>of</strong> <strong>animals</strong>.What <strong>the</strong>se groups are we shall presently explain.In determining <strong>the</strong> number, extent, and boundaries <strong>of</strong> <strong>the</strong>seregions, we must be guided by a variety <strong>of</strong> indications, since<strong>the</strong> application <strong>of</strong> fixed rules is impossible. <strong>The</strong>y should evidentlybe <strong>of</strong> a moderate number, corresponding as far aspracticable <strong>with</strong> <strong>the</strong> great natural divisions <strong>of</strong> <strong>the</strong> globe markedout by nature, and which have always been recognized bygeographers. <strong>The</strong>re should be some approximation to equality<strong>of</strong> size, since <strong>the</strong>re is reason to believe that a tolerably extensivearea has been an essential condition for <strong>the</strong> development <strong>of</strong>most animal forms ; and it is found that, o<strong>the</strong>r things beingequal, <strong>the</strong> numbers, variety and importance <strong>of</strong> <strong>the</strong> forms <strong>of</strong>animal and vegetable life, do bear some approximate relationto extent <strong>of</strong> area. Although <strong>the</strong> possession <strong>of</strong> peculiar familiesor genera is<strong>the</strong> main character <strong>of</strong> a primary zoological region,yet <strong>the</strong> negative character <strong>of</strong> <strong>the</strong> absence <strong>of</strong> certain familiesor genera is <strong>of</strong> equal importance, when this absence does notmanifestly depend on unsuitability to <strong>the</strong> support <strong>of</strong> <strong>the</strong> group,and especially when <strong>the</strong>re is now no physical barrier preventing<strong>the</strong>ir entrance.This will become evident when we consider that<strong>the</strong> importance <strong>of</strong> <strong>the</strong> possession <strong>of</strong> a group by one region dependson its absence from <strong>the</strong> adjoining regions ; and if <strong>the</strong>re isnow no barrier to its entrance, we may be sure that <strong>the</strong>re hasonce been one ;and that <strong>the</strong> possession <strong>of</strong> <strong>the</strong> area by a distinctand well balanced set <strong>of</strong> organisms, which must have been slowly


chap, iv.] ZOOLOGICAL REGIONS. 55developed and adjusted, is<strong>the</strong> living barrier that now keeps outintruders.When it is ascertained that <strong>the</strong> chief differences whichnow obtain between two areas did not exist in Miocene orPliocene times, <strong>the</strong> fact is one <strong>of</strong> great interest, and enables usto speculate <strong>with</strong> some degree <strong>of</strong> probability as to <strong>the</strong> causes thathave brought about <strong>the</strong> present state <strong>of</strong> things ; but it is not areason for uniting <strong>the</strong>se two areas into one region. Our objectis to represent as nearly as possible <strong>the</strong> main features <strong>of</strong> <strong>the</strong><strong>distribution</strong> <strong>of</strong> existing <strong>animals</strong>, not those <strong>of</strong> any or all pastgeological epochs. Should we ever obtain sufficient informationas to <strong>the</strong> geography and biology <strong>of</strong> <strong>the</strong> earth at past epochs, wemight indeed determine approximately what were <strong>the</strong>Plioceneor Miocene or Eocene zoological regions ; but any attempt toexhibit all <strong>the</strong>se in combination <strong>with</strong> those <strong>of</strong> our own period,must lead to confusion.<strong>The</strong> binary or dichotomous system, although it brings out<strong>the</strong> fundamental differences <strong>of</strong> <strong>the</strong> respective regions, is aninconvenient one in its application, and ra<strong>the</strong>r increases thanobviates, <strong>the</strong> difficulty as to equality or inequality <strong>of</strong> regions;for although a, b, c, and d, may be areas <strong>of</strong> unequal zoologicalrank, a being <strong>the</strong> most important, and d <strong>the</strong> least, yet thisinequality will probably be still greater if we first divide<strong>the</strong>m into a, on one side, and b, c, and d, on <strong>the</strong> o<strong>the</strong>r,and <strong>the</strong>n, by ano<strong>the</strong>r division, make b, an area <strong>of</strong> <strong>the</strong> second,and c, and d, <strong>of</strong> <strong>the</strong> third rank only.Coming to <strong>the</strong> second objection, <strong>the</strong> <strong>of</strong>ten incompatible<strong>distribution</strong> <strong>of</strong> different groups <strong>of</strong> <strong>animals</strong>, affords ground foropposition to any proposed scheme <strong>of</strong> zoological regions.<strong>The</strong>reis first <strong>the</strong> radical difference between laud and sea <strong>animals</strong>;<strong>the</strong> most complete barriers to <strong>the</strong> dispersal <strong>of</strong> <strong>the</strong> one, sometimes<strong>of</strong>fering <strong>the</strong> greatest facilities for <strong>the</strong> emigration <strong>of</strong> <strong>the</strong> o<strong>the</strong>r,and vice versa. A large number <strong>of</strong> marine <strong>animals</strong>, however,frequent shallow water only ;and <strong>the</strong>se, keeping near <strong>the</strong> coasts,will agree generally in <strong>the</strong>ir <strong>distribution</strong> <strong>with</strong> those inhabiting<strong>the</strong> land. But among land <strong>animals</strong> <strong>the</strong>mselves <strong>the</strong>re are verygreat differences <strong>of</strong> <strong>distribution</strong>, due to certain specialities


56 DISTRIBUTION OF ANIMALS. [part i.in <strong>the</strong>ir organization or mode <strong>of</strong> life. <strong>The</strong>se act mainly intwo ways,— lstly, by affecting <strong>the</strong> facilities <strong>with</strong> which ;— <strong>the</strong>ycan be dispersed, ei<strong>the</strong>r voluntarily or involuntarily 2ndly,by <strong>the</strong> conditions which enable <strong>the</strong>m to multiply and establish<strong>the</strong>mselves in certain areas and not in o<strong>the</strong>rs. When both<strong>the</strong>se means <strong>of</strong> diffusion are at a maximum, <strong>the</strong> dispersal <strong>of</strong>a group becomes universal, and ceases to have much interestfor us. This is <strong>the</strong> case <strong>with</strong> certain groups <strong>of</strong> fungi andlichens, as well as <strong>with</strong> some <strong>of</strong> <strong>the</strong> lower <strong>animals</strong> ; and ina less degree, as has been shown by Mr. Darwin, <strong>with</strong> manyfresh-water plants and <strong>animals</strong>.At <strong>the</strong> o<strong>the</strong>r extreme we mayplace certain arboreal vertebrata such as sloths and lemurs,which have no means <strong>of</strong> passing such barriers as narrowstraits or moderately high mountains, and whose survival inany new country <strong>the</strong>y might reach, would be dependent on <strong>the</strong>presence <strong>of</strong> suitable forests and <strong>the</strong> absence <strong>of</strong> dangerous enemies.Almost equally, or perhaps even more restricted, are <strong>the</strong> means<strong>of</strong> permanent diffusion <strong>of</strong> terrestrial molluscs; since <strong>the</strong>se are<strong>with</strong>out any but very rare and accidental means <strong>of</strong> being safelytransported across <strong>the</strong> sea ; <strong>the</strong>ir individual powers <strong>of</strong> locomotionare highly restricted ; <strong>the</strong>y are especially subject to <strong>the</strong>attacks <strong>of</strong> enemies; and <strong>the</strong>y <strong>of</strong>ten depend not only on apeculiar vegetation, but on <strong>the</strong> geological character <strong>of</strong> <strong>the</strong> country,<strong>the</strong>ir abundance being almost in direct proportion to <strong>the</strong>presence <strong>of</strong> some form <strong>of</strong> calcaxeous rocks. Between <strong>the</strong>seextremes we find <strong>animals</strong> possessed <strong>of</strong> an infinite gradation <strong>of</strong>powers to disperse and to maintain <strong>the</strong>mselves ;and it will evidentlybe impossible that <strong>the</strong> limits which best define <strong>the</strong><strong>distribution</strong> <strong>of</strong> one group, should be equally true for all o<strong>the</strong>rs.Which class <strong>of</strong> Animals is <strong>of</strong> most importance, in determiningZoological Regions.—To decide this question we have to considerwhich groups <strong>of</strong> <strong>animals</strong> are best adapted to exhibit,by <strong>the</strong>ir existing <strong>distribution</strong>, <strong>the</strong> past changes and presentphysical condition <strong>of</strong> <strong>the</strong> earth's surface; and at <strong>the</strong> sametime, by <strong>the</strong> abundance <strong>of</strong> <strong>the</strong>ir remains in<strong>the</strong> various tertiaryformations will best enable us to trace out <strong>the</strong> more recent<strong>of</strong> <strong>the</strong> series <strong>of</strong> changes, both <strong>of</strong> <strong>the</strong> earth's surface and


6;chap, iv.] ZOOLOGICAL REGIONS. 57<strong>of</strong> its inhabitants, by which <strong>the</strong> present state <strong>of</strong> things hasbeen brought about. For this purpose we require a groupwhich shall be dependent for its means <strong>of</strong> dispersal on <strong>the</strong> <strong>distribution</strong><strong>of</strong> land and water, and on <strong>the</strong> presence or absence<strong>of</strong> l<strong>of</strong>ty mountains, desert plains or plateaux, and great forestssince <strong>the</strong>se are <strong>the</strong> chief physical features <strong>of</strong> <strong>the</strong> earth's surfacewhose modifications at successive periods we wish to discover.It is also essential that <strong>the</strong>y should not be subject to dispersalby many accidental causes ; as this would inevitably in timetend to obliterate <strong>the</strong> effect <strong>of</strong> natural barriers, and produce ascattered <strong>distribution</strong>, <strong>the</strong> causes <strong>of</strong> which we could only guess at.Again, it is necessary that <strong>the</strong>y should be so highly organized asnot to be absolutely dependent on o<strong>the</strong>r groups <strong>of</strong> <strong>animals</strong>, and<strong>with</strong> so much power <strong>of</strong> adaptation as to be able to exist in oneform or ano<strong>the</strong>r over <strong>the</strong> whole globe. And lastly, it is highlyimportant that <strong>the</strong> whole group should be pretty well known,and that a fairly natural classification, especially <strong>of</strong> its minordivisions such asfamilies and genera, should have been arrivedat ; <strong>the</strong> reason for which last proviso is explained in our nextchapter, on classification.Now in every one <strong>of</strong> <strong>the</strong>se points <strong>the</strong> mammalia arepreeminent; and <strong>the</strong>y possess <strong>the</strong> additional advantage <strong>of</strong> being <strong>the</strong>most highly developed class <strong>of</strong> organized beings, and that towhich we ourselves belong. We should <strong>the</strong>refore construct ourtypical or standard Zoological Regions in <strong>the</strong> first place, from aconsideration <strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> mammalia, only bringing toour aid <strong>the</strong> <strong>distribution</strong> <strong>of</strong> o<strong>the</strong>r groups to determine doubtfulpoints. Regions so established will be most closely in accordance<strong>with</strong> those long-enduring features <strong>of</strong> physical geography,on which <strong>the</strong> <strong>distribution</strong> <strong>of</strong> all forms <strong>of</strong> life fundamentallydepend ; and all discrepancies in <strong>the</strong> <strong>distribution</strong> <strong>of</strong> o<strong>the</strong>rclasses <strong>of</strong> <strong>animals</strong> must be capable <strong>of</strong> being explained, ei<strong>the</strong>rby <strong>the</strong>ir exceptional means <strong>of</strong> dispersion or by special conditionsaffecting <strong>the</strong>ir perpetuation and increase in each locality.If <strong>the</strong>se considerations are well founded, <strong>the</strong> objections <strong>of</strong>those who <strong>study</strong> insects or molluscs, for example,— that ourregions are not true for <strong>the</strong>ir departments <strong>of</strong> nature—cannot beVol. I.—


58 DISTRIBUTION OF ANIMALS. [part i.maintained. For <strong>the</strong>y will find, that a careful consideration <strong>of</strong><strong>the</strong> exceptional means <strong>of</strong> dispersal and conditions <strong>of</strong> existence <strong>of</strong>each group, will explain most <strong>of</strong> <strong>the</strong> divergences from <strong>the</strong> normal<strong>distribution</strong> <strong>of</strong> higher <strong>animals</strong>.We shall thus be led to an intelligent comprehension <strong>of</strong> <strong>the</strong>phenomena <strong>of</strong> <strong>distribution</strong> in all groups, which would not be<strong>the</strong> case if every specialist formed regions for his own particular<strong>study</strong>. In many cases we should find that no satisfactorydivision <strong>of</strong> <strong>the</strong> earth could be made tocorrespond <strong>with</strong> <strong>the</strong> <strong>distribution</strong>even <strong>of</strong> an entire class ; but we should have <strong>the</strong> coleopteristand <strong>the</strong> lepidopterist each <strong>with</strong> his own Geography.Andeven this would probably not suffice, for it is very doubtful if<strong>the</strong> detailed <strong>distribution</strong> <strong>of</strong> <strong>the</strong> Longicornes, so closely dependenton woody vegetation, could be made to agree <strong>with</strong> that <strong>of</strong> <strong>the</strong>Staphylinidse or <strong>the</strong> Carabidse which abound in many <strong>of</strong> <strong>the</strong>most barren regions, or <strong>with</strong> that <strong>of</strong> <strong>the</strong> Scarabeidse, largely dependenton <strong>the</strong> presence <strong>of</strong> herbivorous mammalia.And wheneach <strong>of</strong> <strong>the</strong>se enquirers had settled a division <strong>of</strong> <strong>the</strong> earth into" regions "which exhibited <strong>with</strong> tolerable accuracy <strong>the</strong> phenomena<strong>of</strong> <strong>distribution</strong> <strong>of</strong> his own group, we should have gainednothing whatever but a very complex mode <strong>of</strong> exhibiting<strong>the</strong> bare facts <strong>of</strong> <strong>distribution</strong>. We should <strong>the</strong>n have to beginto work out <strong>the</strong> causes <strong>of</strong> <strong>the</strong> divergence <strong>of</strong> one group fromano<strong>the</strong>r in this respect ; but as each worker would refer to hisown set <strong>of</strong> regions as <strong>the</strong> type, <strong>the</strong> whole subject would becomeinvolved in inextricable confusion. <strong>The</strong>se considerations seemto make it imperative that one set <strong>of</strong> "regions" should beestablished as typical for Zoology ; and it is hoped <strong>the</strong> reasonshere advanced will satisfy most naturalists that <strong>the</strong>se regionscan be best determined, in <strong>the</strong> first place, by a <strong>study</strong> <strong>of</strong> <strong>the</strong> <strong>distribution</strong><strong>of</strong> <strong>the</strong> mammalia, supplemented in doubtful cases bythat <strong>of</strong> <strong>the</strong> o<strong>the</strong>r vertebrates.We will now proceed to a discussion<strong>of</strong> what <strong>the</strong>se regions are.Various Zoological Regions proposed since 1857.— It has alreadybeen pointed out that a very large number <strong>of</strong> birds are limitedby <strong>the</strong> same kind <strong>of</strong> barriers as mammalia ;it will <strong>the</strong>reforenot be surprising that a system <strong>of</strong> regions formed to suit <strong>the</strong>


—CHAr. iv.] • ZOOLOGICAL REGIONS. 59one, should very nearly represent <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> o<strong>the</strong>r.Mr. Sclater's regions are as follows :1. <strong>The</strong> Palsearctic Region ; including Europe, Temperate Asia,and N. Africa to <strong>the</strong> Atlas mountains.2. <strong>The</strong> Ethiopian Region ;Africa south <strong>of</strong> <strong>the</strong> Atlas, Madagascar,and <strong>the</strong> Mascarene Islands, <strong>with</strong> Sou<strong>the</strong>rn Arabia.3. <strong>The</strong> Indian Region ; including India south <strong>of</strong> <strong>the</strong> Himalayas,to South China, and to Borneo and Java.4. <strong>The</strong> Australian Region ; including Celebes and Lombock,eastward to Australia and <strong>the</strong> Pacific Islands.5. <strong>The</strong> Nearctic Region; including Greenland, and N.America, to Nor<strong>the</strong>rn Mexico.6. <strong>The</strong> Neotropical Region ; including South America, <strong>the</strong>Antilles, and Sou<strong>the</strong>rn Mexico.This division <strong>of</strong> <strong>the</strong> earth received great support from Dr.Giin<strong>the</strong>r, who, in <strong>the</strong> Proceedings <strong>of</strong> <strong>the</strong> Zoological Society for1858, showed that <strong>the</strong> <strong>geographical</strong> <strong>distribution</strong> <strong>of</strong> Reptilesagreed <strong>with</strong> it very closely, <strong>the</strong> principal difference being that<strong>the</strong> reptiles <strong>of</strong> Japan have a more Indian character than <strong>the</strong>birds, this being especially <strong>the</strong> case <strong>with</strong> <strong>the</strong> snakes. In <strong>the</strong>volume for 1868 <strong>of</strong> <strong>the</strong> same work, Pr<strong>of</strong>essor Huxley discussesat considerable length <strong>the</strong> primary and secondary zoologicaldivisions <strong>of</strong> <strong>the</strong> earth. He gives reasons for thinking that <strong>the</strong>most radical primary division, both as regards birds and mammals,is into a Nor<strong>the</strong>rn and Sou<strong>the</strong>rn hemisphere (Arctogreaand Notogaea), <strong>the</strong> former, however, embracing all Africa, while<strong>the</strong> latter includes only Australasia and <strong>the</strong> Neotropical orAustro-Columbian region. Mr. Sclater had grouped his regionsprimarily into Palasogaea and Neogaea, <strong>the</strong> Old and NewWorlds <strong>of</strong> geographers ; a division which strikingly accords<strong>with</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> passerine birds, but not so well<strong>with</strong> that <strong>of</strong> mammalia or reptiles. Pr<strong>of</strong>essor Huxley pointsout that <strong>the</strong> Nearctic, Palsearctic, Indian, and Ethiopianregions <strong>of</strong> Mr. Sclater have a much greater resemblance toeach o<strong>the</strong>r than any one <strong>of</strong> <strong>the</strong>m has to Australia or to SouthAmerica ;and he fur<strong>the</strong>r suggests that New Zealand alone haspeculiarities which might entitle it to rank as a primary region


;GO DISTRIBUTION OF ANIMALS. [part i.along <strong>with</strong> Australasia and South America ;and that a CircumpolarProvince might be conveniently recognised as <strong>of</strong> equalrank <strong>with</strong> <strong>the</strong> Palsearctic and Nearctic provinces.In 1866, Mr. Andrew Murray published a large and copiouslyillustrated volume on <strong>the</strong> Geograjihical Distribution <strong>of</strong> Mammals,in which he maintains that <strong>the</strong> great and primarymammalian regions are only four : 1st. <strong>The</strong> Palrearctic region<strong>of</strong> Mr. Sclater, extended to include <strong>the</strong> Sahara and Nubia2nd. <strong>the</strong> Indo-African region, including <strong>the</strong> Indian and Ethiopianregions <strong>of</strong> Mr. Sclater; 3rd. <strong>the</strong> Australian region (unaltered);4th. <strong>the</strong> American region, including both North and SouthAmerica. <strong>The</strong>se are <strong>the</strong> regions as described by Mr. Murray,but his coloured map <strong>of</strong> " Great Mammalian Regions " showsall Arctic America to a little south <strong>of</strong> <strong>the</strong> Iso<strong>the</strong>rmal <strong>of</strong> 32°Fahr. as forming <strong>with</strong> Europe and North Asia one great region.At <strong>the</strong> meeting <strong>of</strong> <strong>the</strong> British Association at Exeter in 1869,Mr. W. T.Blanford read a paper on <strong>the</strong> Fauna <strong>of</strong> British India,in which he maintained that a large portion <strong>of</strong> <strong>the</strong> peninsula<strong>of</strong> India had derived its Fauna mainly from Africa ;and that <strong>the</strong>term "Indian region" <strong>of</strong> Mr. Sclater was misleading, becauseIndia proper, if it belongs to it at all, is <strong>the</strong> least typical portion<strong>of</strong> it. He <strong>the</strong>refore proposes to call it <strong>the</strong> " Malayan region,"because in <strong>the</strong> Malay countries it is most highly developed.Ceylon and <strong>the</strong> mountain ranges <strong>of</strong> Sou<strong>the</strong>rn India have markedMalay affinities.In 1871 Mr. E. Blyth published in Nature "A suggested newDivision <strong>of</strong> <strong>the</strong> Earth into Zoological Regions," in which heindicates seven primary divisions or regions, subdivided intotwenty-six sub -regions. <strong>The</strong> seven regions are defined asfollows: 1. <strong>The</strong> Boreal region; including <strong>the</strong> whole <strong>of</strong> <strong>the</strong>Palsearctic and Nearctic regions <strong>of</strong> Mr. Sclater along <strong>with</strong> <strong>the</strong>West Indies, Central America, <strong>the</strong> whole chain <strong>of</strong> <strong>the</strong> Andes,<strong>with</strong> Chili and Patagonia. 2. <strong>The</strong> Columbian region ; consisting<strong>of</strong> <strong>the</strong> remaining part <strong>of</strong> South America. 3. <strong>The</strong> Ethiopianregion ;comprising besides that region <strong>of</strong> Mr. Sclater, <strong>the</strong> valley<strong>of</strong> <strong>the</strong> Jordan, Arabia, and <strong>the</strong> desert country towards India,<strong>with</strong> all <strong>the</strong> plains and table lands <strong>of</strong> India and <strong>the</strong> nor<strong>the</strong>rn


;chap, iv.] ZOOLOGICAL REGIONS. 61half <strong>of</strong> Ceylon. 4. <strong>The</strong> Lemurian region ;consisting <strong>of</strong> Madagascarand its adjacent islands. 5. <strong>The</strong> Austral-Asian regionwhich is <strong>the</strong> Indian region <strong>of</strong> Mr. Sclater <strong>with</strong>out <strong>the</strong> portiontaken to be added to <strong>the</strong> Ethiopian region. 6. <strong>The</strong> Melanesianregion ; which is <strong>the</strong> Australian region <strong>of</strong> Mr. Sclater <strong>with</strong>outNew Zealand and <strong>the</strong> Pacific Islands, which form 7. <strong>the</strong>Polynesian region. Mr. Blyth thinks this is " a true classification<strong>of</strong> zoological regions as regards mammalia and birds."In an elaborate paper on <strong>the</strong> birds <strong>of</strong> Eastern North America,<strong>the</strong>ir <strong>distribution</strong> and migrations (Bulletin <strong>of</strong> Museum <strong>of</strong> ComparativeZoology, Cambridge, Massachusetts, Vol. 2), Mr. J. A.Allen proposes a division <strong>of</strong> <strong>the</strong> earth in accordance <strong>with</strong> wha<strong>the</strong> terms, " <strong>the</strong> law <strong>of</strong> circumpolar <strong>distribution</strong> <strong>of</strong>life in zones,"as follows : 1. Arctic realm. 2. North temperate realm. 3.American tropical realm. 4. Indo-African tropical realm.5. South American tropical realm. 6. African temperate realm.7. Antarctic realm. 8. Australian realm. Some <strong>of</strong> <strong>the</strong>se aresubdivided into regions; (2) consisting <strong>of</strong> <strong>the</strong> American and<strong>the</strong> Europaeo-Asiatic regions ; (4) into <strong>the</strong> African and Indianregions; (8) into <strong>the</strong> tropical Australian region, and one comprising<strong>the</strong> sou<strong>the</strong>rn part <strong>of</strong> Australia and New Zealand. <strong>The</strong>o<strong>the</strong>r realms each form a single region.Discussion <strong>of</strong> proposed Regions.—Before proceeding to define<strong>the</strong> regions adopted in this work, it may be as well to makea few remarks on some <strong>of</strong> <strong>the</strong> preceding classifications, andto give <strong>the</strong> reasons which seem to render it advisable to adoptvery few <strong>of</strong> <strong>the</strong> suggested improvements on Mr. Sclater'soriginal proposal. Mr. Blyth's scheme is one <strong>of</strong> <strong>the</strong> leastnatural, and also <strong>the</strong> most inconvenient. <strong>The</strong>re can be littleuse in <strong>the</strong> knowledge that a group <strong>of</strong> <strong>animals</strong> is found in <strong>the</strong>Boreal Eegion, if <strong>the</strong>ir habitat might still be ei<strong>the</strong>r Patagonia,<strong>the</strong> West Indies, or Japan ; and it is difficult to see on whatprinciple <strong>the</strong> Madagascar group <strong>of</strong> islands is made <strong>of</strong> equalrank <strong>with</strong> this enormous region, seeing that its forms <strong>of</strong> lifehave marked African affinities. Nei<strong>the</strong>r does it seem advisableto adopt <strong>the</strong> Polynesian Eegion, or that comprising New Zealandalone (as hinted at by Pr<strong>of</strong>essor Huxley and since adopted by


;G2 DISTRIBUTION OF ANIMALS. [part i.Mr. Sclater in his Lectures on Geographical Distribution at <strong>the</strong>Zoological Gardens in May 1874), because it is absolutely <strong>with</strong>outindigenous mammalia and very poor in all forms <strong>of</strong> life,and <strong>the</strong>refore by no means prominent or important enough t<strong>of</strong>orm a primary region <strong>of</strong> <strong>the</strong> earth.It may be as well here to notice what appears tobe a seriousobjection to making New Zealand, or any similar isolateddistrict, one <strong>of</strong> <strong>the</strong> great zoological regions, comparable to SouthAmerica, Australia, or Ethiopia ;which is, that its claim to thatdistinction rests on grounds which are liable to fail. It isbecause New Zealand, in addition to its negative merits, possessesthree families <strong>of</strong> birds (Apterygidse living, Dinornithidse andPalapterygidae extinct), and a peculiar lizard-like reptile,Hatteria, which has to be classed in a distinct order, Ehynchocephalina,that <strong>the</strong> rank <strong>of</strong> a Eegion is claimed for it. Butsupposing, what is not at all improbable, that o<strong>the</strong>r Ehynchocephalinashould be discovered in <strong>the</strong> interior <strong>of</strong> Australia orin New Guinea, and that Apterygidre or Palapterygi'dre shouldbe found to have inhabited Australia in Post-Pliocene times,(as Dinornithidae have already been proved to have done) <strong>the</strong>claims <strong>of</strong> New Zealand would entirely fail, and it would beuniversally acknowledged to be a part <strong>of</strong> <strong>the</strong> great Australianregion. No such reversal can take place in <strong>the</strong> case <strong>of</strong> <strong>the</strong>o<strong>the</strong>r regions ;because <strong>the</strong>y rest, not upon one or two, but upon alarge number <strong>of</strong> peculiarities, <strong>of</strong> such a nature that <strong>the</strong>re isroom upon <strong>the</strong> globe for discoveries that can seriously modify<strong>the</strong>m. Even if one or two peculiar types, like Apterygidre ornoHatteria, should permanently remain characteristic <strong>of</strong> New Zealandalone, we can account for <strong>the</strong>se by <strong>the</strong> extreme isolation<strong>of</strong><strong>the</strong> country, and <strong>the</strong> absence <strong>of</strong> enemies, which have enabled<strong>the</strong>se defenceless birds and reptiles to continue <strong>the</strong>ir existencejust as <strong>the</strong> isolation and protection <strong>of</strong> <strong>the</strong> caverns <strong>of</strong> Carniolahave enabled <strong>the</strong> Proteus to survive in Europe.But supposingthat <strong>the</strong> Proteus was <strong>the</strong> sole representative <strong>of</strong> an order <strong>of</strong>Batrachia, and that two or three o<strong>the</strong>r equally curious andisolated forms occurred <strong>with</strong> it,no one would propose that <strong>the</strong>secaverns or <strong>the</strong> district containing <strong>the</strong>m, should form one <strong>of</strong> <strong>the</strong>


chap, iv.] ZOOLOGICAL REGIONS. 63primary divisions <strong>of</strong> <strong>the</strong> earth. Nei<strong>the</strong>r can much stress belaid on <strong>the</strong> negative characteristics <strong>of</strong> New Zealand, since <strong>the</strong>yare found to an almost equal extent in every oceanic island.Again, it is both inconvenient and misleading to pick outcertain tracts from <strong>the</strong> midst <strong>of</strong> one region or sub-region and toplace <strong>the</strong>m in ano<strong>the</strong>r, on account <strong>of</strong> certain isolated affinitieswhich may <strong>of</strong>ten be accounted for by local peculiarities.Evenif <strong>the</strong> resemblance <strong>of</strong> <strong>the</strong> fauna <strong>of</strong> Chili and Patagonia to that<strong>of</strong> <strong>the</strong> Palsearctic and Nearctic regions was much greater than itis, this mode <strong>of</strong> dealing <strong>with</strong> it would be objectionable ; but itis still more so, when we find that <strong>the</strong>se countries have astrongly marked South American character, and that <strong>the</strong> nor<strong>the</strong>rnaffinities are altoge<strong>the</strong>r exceptional. <strong>The</strong> Eodentia, whichcomprise a large portion <strong>of</strong> <strong>the</strong> mammalia <strong>of</strong> <strong>the</strong>se countries,are wholly South American in type, and <strong>the</strong> birds are almost allallied to forms characteristic <strong>of</strong> tropical America.For analogous reasons <strong>the</strong> Ethiopian must not be made toinclude any part <strong>of</strong> India or Ceylon ;for although <strong>the</strong> Fauna <strong>of</strong>Central India has some African affinities, <strong>the</strong>se do not preponderate;and it will not be difficult to show that to follow Mr.Andrew Murray in uniting bodily <strong>the</strong> Ethiopian and Indianregions <strong>of</strong> Mr. Sclater, is both unnatural and inconvenient.<strong>The</strong>resemblances between <strong>the</strong>m are <strong>of</strong> <strong>the</strong> same character as thosewhich would unite <strong>the</strong>m both <strong>with</strong> <strong>the</strong>Palasarctic and Nearcticregions ; and although it may be admitted, that, as Pr<strong>of</strong>essorHuxley maintains, this group forms one <strong>of</strong> <strong>the</strong> great primarydivisions <strong>of</strong> <strong>the</strong> globe, it is far too extensive and too heterogeneousto subserve <strong>the</strong> practical uses for which we require adivision <strong>of</strong> <strong>the</strong> world into zoological regions.Reasons for adopting <strong>the</strong> six Regions first proposed by Mr. Sclater.—So that we do not violate any clear affinities or produce anyglaring irregularities, it is a positive, and by no means an unimportant,advantage to have our named regionsapproximatelyequal in size, and <strong>with</strong> easily defined, and <strong>the</strong>refore easily remembered,boundaries. All elaborate definitions <strong>of</strong> interpenetratingfrontiers, as well as regions extending over three-fourths<strong>of</strong> <strong>the</strong> land surface <strong>of</strong> <strong>the</strong> globe, and including places which are


G4 DISTRIBUTION OF ANIMALS. [part i.<strong>the</strong> antipodes <strong>of</strong> each o<strong>the</strong>r, would be most inconvenient, evenif <strong>the</strong>re were not such difference <strong>of</strong> opinion about <strong>the</strong>m.<strong>The</strong>recan be little doubt, for example, that <strong>the</strong> most radical zoologicaldivision <strong>of</strong> <strong>the</strong> earth is made by separating <strong>the</strong> Australian regionfrom <strong>the</strong> rest; but although it is something useful anddefinite to know that a group <strong>of</strong> <strong>animals</strong> is peculiar to Australia,it is exceedingly vague and unsatisfactory to say <strong>of</strong> any o<strong>the</strong>rgroup merely that it is extra-Australian. Nei<strong>the</strong>r can it be saidthat, from any point <strong>of</strong> view, <strong>the</strong>se two divisions are <strong>of</strong> equalimportance. <strong>The</strong> next great natural division that can be madeis <strong>the</strong> separation <strong>of</strong> <strong>the</strong> Neotropical Region <strong>of</strong> Mr. Sclater from<strong>the</strong> rest <strong>of</strong> <strong>the</strong> world. We thus have three primary divisions,which Pr<strong>of</strong>essor Huxley seems inclined to consider as <strong>of</strong>tolerably equal zoological importance. But a consideration <strong>of</strong>all <strong>the</strong> facts, zoological and palseontological, indicates, that <strong>the</strong>great nor<strong>the</strong>rn division (Arctogsea) is fully as much more importantthan ei<strong>the</strong>r Australia or South America,as its four componentparts are less important ; and if so, convenience requiresus to adopt <strong>the</strong> smaller ra<strong>the</strong>r than <strong>the</strong> larger divisions.This question, <strong>of</strong> comparative importance or equivalence <strong>of</strong>value, is very difficult to determine.It may be considered from<strong>the</strong> point <strong>of</strong> view <strong>of</strong> speciality or isolation, or from that <strong>of</strong>richness and variety <strong>of</strong> animal forms.In isolation and speciality,determined by what <strong>the</strong>y want as well as what <strong>the</strong>y possess, <strong>the</strong>Australian and Neotropical regions are undoubtedly each comparable<strong>with</strong> <strong>the</strong> rest <strong>of</strong> <strong>the</strong> earth (Arctogsea). But in richnessand variety <strong>of</strong> forms, <strong>the</strong>y are both very much inferior, and aremuch more nearly comparable <strong>with</strong> <strong>the</strong> separate regions whichcompose it.Taking <strong>the</strong> families <strong>of</strong> mammalia as established by<strong>the</strong> best authors, and leaving out <strong>the</strong> Cetacea and <strong>the</strong> Bats,which are almost universally distributed, and about whoseclassification <strong>the</strong>re is much uncertainty, <strong>the</strong> number <strong>of</strong> familiesrepresented in each <strong>of</strong> Mr. Sclater's regions is as follows :I. Palsearctic region has 31 families <strong>of</strong> terrestrial mammalia.II. Ethiopian „ „ 40 „ „III. Indian „ „ 31 „ „IV. Australian „ „ 14 „ „V. Neotropical „ „ 26 „ „VI. Nearctic „ ,,23 „ „


;chap. iv.] ZOOLOGICAL REGIONS. 65We see, <strong>the</strong>n, that even <strong>the</strong> exceedingly rich and isolated Neotropicalregion is less rich and diversified in its forms <strong>of</strong> mammalianlife than <strong>the</strong> very much smaller area <strong>of</strong> <strong>the</strong> Indian region,or <strong>the</strong> temperate Palcearctic, and very much less so than <strong>the</strong>Ethiopian region ; while even <strong>the</strong> comparatively poor Nearcticregion, is nearly equal to it in <strong>the</strong> number <strong>of</strong> its family types. If<strong>the</strong>se were united <strong>the</strong>y would possess fifty-five families, a numbervery disproportionate to those <strong>of</strong> <strong>the</strong> remaining two.Ano<strong>the</strong>rconsideration is, that although <strong>the</strong> absence <strong>of</strong> certain forms <strong>of</strong>life makes a region more isolated, it does not make itzoologicallymore important ; for we have only to suppose some five or sixfamilies, now common to both, to become extinct ei<strong>the</strong>r in <strong>the</strong>Ethiopian or <strong>the</strong> Indian regions, and <strong>the</strong>y would become asstrongly differentiatedwhile still remaining as rich in family types.from all o<strong>the</strong>r regions as South America,In birds exactly <strong>the</strong>same phenomenon recurs, <strong>the</strong> family types being less numerous inSouth America than in ei<strong>the</strong>r <strong>of</strong> <strong>the</strong> o<strong>the</strong>r tropical regions <strong>of</strong> <strong>the</strong>earth, but a larger proportion <strong>of</strong> <strong>the</strong>m are restricted to it.It willbe shown fur<strong>the</strong>r on, that <strong>the</strong> Ethiopian and Indian, (or, as I proposeto call it in this work, Oriental) regions, are sufficiently differentiatedby very important groups <strong>of</strong> <strong>animals</strong> peculiar toeachand that, on strict zoological principles <strong>the</strong>y are entitled torank asregions <strong>of</strong> equal value <strong>with</strong> <strong>the</strong> Neotropical and Australian.It is perhaps less clear whe<strong>the</strong>r <strong>the</strong> Palsearctic shouldbe separated from <strong>the</strong> Oriental region, <strong>with</strong> which it has undoubtedlymuch in common ; but <strong>the</strong>re are many and powerfulreasons for keeping it distinct.facies in <strong>the</strong> animal forms <strong>of</strong> <strong>the</strong> two regions ;<strong>The</strong>re is an unmistakably differentand although n<strong>of</strong>amilies <strong>of</strong> mammalia or birds, and not many genera, are whollyconfined to <strong>the</strong> Palsearctic region, a very considerable number<strong>of</strong> both have <strong>the</strong>ir metropolis in it,and are very richly represented.<strong>The</strong> distinction between <strong>the</strong> characteristic forms <strong>of</strong> life in tropicaland cold countries is,on <strong>the</strong> whole, very strongly marked in <strong>the</strong>nor<strong>the</strong>rn hemisphere ; and to refuse to recognise this in a subdivision<strong>of</strong> <strong>the</strong> earth which is established for <strong>the</strong> very purpose <strong>of</strong>expressing such contrasts more clearly and concisely than byordinary <strong>geographical</strong>terminology, would be both illogical and


;GG DISTRIBUTION OF ANIMALS. [part i.inconvenient. <strong>The</strong> one question <strong>the</strong>n remains, whe<strong>the</strong>r <strong>the</strong>Nearctic region should be kept separate, or whe<strong>the</strong>r it shouldform part <strong>of</strong> <strong>the</strong> Paleearctic or <strong>of</strong> <strong>the</strong> Neotropical regions.Pr<strong>of</strong>essorHuxley and Mr. Blyth advocate <strong>the</strong> former course ;Mr.Andrew Murray (for mammalia) and Pr<strong>of</strong>essor Newton (for birds)think <strong>the</strong> latter would be more natural. No doubt much is tobe said for both views, but both cannot be right ; and it will beshown in <strong>the</strong> latter part <strong>of</strong> this chapter that <strong>the</strong> Nearctic regionis, on <strong>the</strong> whole, fully as well defined as <strong>the</strong> PalaBarctic, by positivecharacters which differentiate it from both <strong>the</strong> adjacentregions. More evidence in <strong>the</strong> same direction will be found in<strong>the</strong> Second Part <strong>of</strong> this work, in which <strong>the</strong> extinct faunas <strong>of</strong> <strong>the</strong>several regions are discussed.A confirmation <strong>of</strong> <strong>the</strong> general views here set forth, as to <strong>the</strong>distinctness and approximate equivalence <strong>of</strong> <strong>the</strong> six regions, isto be found in <strong>the</strong> fact, that if any two or more <strong>of</strong> <strong>the</strong>m are combined<strong>the</strong>y <strong>the</strong>mselves become divisions <strong>of</strong><strong>the</strong> next lower rank,or " sub-regions " —and <strong>the</strong>se will be very much more important,;both zoologically and <strong>geographical</strong>ly, than <strong>the</strong> subdivisions <strong>of</strong><strong>the</strong> remaining regions,.It is admitted <strong>the</strong>n that <strong>the</strong>se six regionsare by no means <strong>of</strong> precisely equal rank, and that some <strong>of</strong> <strong>the</strong>mare far more isolated and better characterized than o<strong>the</strong>rs ; butit is maintained that, looked at from every point <strong>of</strong> view, <strong>the</strong>yare more equal in rank than any o<strong>the</strong>rs that can be formedwhile in <strong>geographical</strong> equality, compactness <strong>of</strong> area, and facility<strong>of</strong> definition, <strong>the</strong>y are beyond all comparison better than anyo<strong>the</strong>rs that have yet been proposed for<strong>the</strong> purpose <strong>of</strong> facilitating<strong>the</strong> <strong>study</strong> <strong>of</strong> <strong>geographical</strong> <strong>distribution</strong>. <strong>The</strong>y may be arrangedand grouped as follows, so as to exhibit <strong>the</strong>ir various<strong>relations</strong> and affinities.Regions.jt ( Neotropical ... Austral zone Notogsea.°') Nearctic ) -n ,Boreal zonePai^arctic ...P1I Ethiopian P1 , . .Palaeogaea' 0riental jPalaeotropical zone\IJArctogaaa.\ Australian ... Austral zone Notogsea.<strong>The</strong> above table shows <strong>the</strong> regions placed in <strong>the</strong> order followedin <strong>the</strong> Fourth Part <strong>of</strong> this work, and <strong>the</strong> reasons for which are


chap, iv.] ZOOLOGICAL REGIONS. 07explained in Chapter IX. As a matter <strong>of</strong> convenience, and foro<strong>the</strong>r reasons adduced in <strong>the</strong> same chapter, <strong>the</strong> detailed exposition<strong>of</strong> <strong>the</strong> <strong>geographical</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> <strong>animals</strong> <strong>of</strong> <strong>the</strong> severalresions in Part III. commences <strong>with</strong> <strong>the</strong> Palsearctic and terminates<strong>with</strong> <strong>the</strong> Nearctic region.Objections to <strong>the</strong> system <strong>of</strong> Circumpolar Zones.—Mr. Allen'ssystem <strong>of</strong> " realms " founded on climatic zones (given atp. 61), having recently appeared in an ornithological work<strong>of</strong> considerable detail and research, calls for a few remarks.<strong>The</strong> author continually refers to <strong>the</strong> " law <strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong>life in circumpolar zones," as if it were one generally acceptedand that admits <strong>of</strong> no dispute. But this supposed " law " onlyapplies to <strong>the</strong> smallest details <strong>of</strong><strong>distribution</strong>—to <strong>the</strong> range andincreasing or decreasing numbers <strong>of</strong> species as we pass fromnorth to south, or <strong>the</strong> reverse ;while it has little bearing on <strong>the</strong>great features <strong>of</strong> zoological geography—<strong>the</strong> limitation <strong>of</strong> groups<strong>of</strong> genera and families to certain areas. It is analogous to<strong>the</strong> " law <strong>of</strong> adaptation " in <strong>the</strong> organisation <strong>of</strong> <strong>animals</strong>, bywhich members <strong>of</strong> various groups are suited for an aerial, anaquatic, a desert, or an arboreal life ;are herbivorous, carnivorous,or insectivorous; are fitted to live underground, or in freshwaters, or on polar ice.It was once thought that <strong>the</strong>se adaptivepeculiarities were suitable foundations for a classification,—thatwhales were fishes, and bats birds ; and even to this clay <strong>the</strong>reare naturalists who cannot recognise <strong>the</strong> essential diversity<strong>of</strong> structure in such groups as swifts and swallows, sun-birds andhumming-birds, under <strong>the</strong> superficial disguise caused by adaptationto a similar mode <strong>of</strong> life.<strong>The</strong> application <strong>of</strong> Mr. Allen'sprinciple leads to equally erroneous results, as may be well seenby considering his separation <strong>of</strong> " <strong>the</strong> sou<strong>the</strong>rn third <strong>of</strong> Australia" to unite it <strong>with</strong> New Zealand as one <strong>of</strong> his secondaryzoological divisions.If <strong>the</strong>re is one country in <strong>the</strong> world whosefauna is strictly homogeneous, that country is Australia ;whileNew Guinea on <strong>the</strong> one hand, and New Zealand on <strong>the</strong> o<strong>the</strong>r,are as sharply differentiated from Australia as any adjacent parts<strong>of</strong> <strong>the</strong> same primary zoological division can possibly be. Yet<strong>the</strong> " laiv <strong>of</strong> circumpolar <strong>distribution</strong> " leads to <strong>the</strong> division <strong>of</strong>


;68 DISTRIBUTION OF ANIMALS. [part i.Australia by an arbitrary east and west line, and a union <strong>of</strong> <strong>the</strong>nor<strong>the</strong>rn two-thirds <strong>with</strong> New Guinea, <strong>the</strong> sou<strong>the</strong>rn third <strong>with</strong>New Zealand. Hardly less unnatural is <strong>the</strong> supposed equivalence<strong>of</strong> South Africa (<strong>the</strong> African temperate realm) to alltropical Africa and Asia, including Madagascar (<strong>the</strong> Indo-African tropical realm). South Africa has, it is true, somestriking peculiarities ; but <strong>the</strong>y are absolutely unimportant ascompared <strong>with</strong> <strong>the</strong> great and radical differences between tropicalAfrica and tropical Asia. On <strong>the</strong>se examples we may fairlyrest our rejection <strong>of</strong> Mr. Allen's scheme.We must however say a few words on <strong>the</strong> zoo-<strong>geographical</strong>nomenclature proposed in <strong>the</strong> same paper, which seems alsovery objectionable. <strong>The</strong> following terms are proposed : realm,region, province, district, fauna and flora ; <strong>the</strong> first being <strong>the</strong>highest, <strong>the</strong> last <strong>the</strong> lowest and smallest sub-division. Consideringthat most <strong>of</strong> <strong>the</strong>se terms have been used in very differentsenses already, and that no means <strong>of</strong> settling <strong>the</strong>ir equivalencein different parts <strong>of</strong> <strong>the</strong> globe has been even suggested,such acomplex system must lead to endless confusion. Until <strong>the</strong>whole subject is far better known and its firstprinciples agreedupon, <strong>the</strong> simpler and <strong>the</strong> fewer <strong>the</strong> terms employed <strong>the</strong> better ;and as " region " was employed for <strong>the</strong> primary divisions byMr. Sclater, eighteen years ago, and again by Mr. AndrewMurray, in his Geographical Distribution <strong>of</strong> Mammals ;nothingbut obscurity can result from each writer using some new, anddoubtfully better, term. For <strong>the</strong> sub-divisions <strong>of</strong> <strong>the</strong> regionsno advantage is gained by <strong>the</strong> use <strong>of</strong> a distinct term— " province"—whichhas been used (by Swainson) for <strong>the</strong> primarydivisions, and which does not itself tell you what rank it holdswhereas <strong>the</strong> term " sub-region " speaks for itself as being unmistakablynext in subordination to region, and this clearness <strong>of</strong>meaning gives it <strong>the</strong> preference over any independent term.As to minor named sub-divisions, <strong>the</strong>y seem at present uncalledfor; and till <strong>the</strong> greater divisions are <strong>the</strong>mselves generallyagreed on, itseems better to adopt no technical names for whatmust, for a long time to come, be indeterminate.Does <strong>the</strong> Arctic Fauna characterize an independentRegion.—


chap, iv.] ZOOLOGICAL REGIONS. 69<strong>The</strong> proposal to consider <strong>the</strong> Arctic regions as constituting one <strong>of</strong><strong>the</strong> primary zoological divisions <strong>of</strong> <strong>the</strong> globe, has been advocatedby many naturalists. Pr<strong>of</strong>essor Huxley seems to consider itadvisable, and Mr. Allen unhesitatingly adopts it, as well as an" antarctic " region to balance it in <strong>the</strong> sou<strong>the</strong>rn hemisphere.<strong>The</strong> reason why an " Arctic Eegion " finds no place in this workmay <strong>the</strong>refore be here stated.No species or group <strong>of</strong> <strong>animals</strong> can properly be classed as" arctic," which does not exclusively inhabit or greatly preponderatein arctic lauds. For <strong>the</strong> purpose <strong>of</strong> establishing <strong>the</strong>need <strong>of</strong> an " arctic "zoological region, we should consider chieflysuch groups as are circumpolar as well as arctic ; because, if<strong>the</strong>y are confined to, or greatly preponderate in, ei<strong>the</strong>r <strong>the</strong>eastern or western hemispheres, <strong>the</strong>y can be at once allocated to<strong>the</strong> Nearctic or Palasarctic regions, and can <strong>the</strong>refore afford nojustification for establishing a new primary division <strong>of</strong> <strong>the</strong>globe.Thus restricted, only three genera <strong>of</strong> land mammalia are trulyarctic : Gulo, Myodes, and Rangifcr. Two species <strong>of</strong> widelydispersed genera are also exclusively arctic, Ursits maritimusand Vilifies lagopus.Exclusively arctic birds are not much more numerous. Ofland birds <strong>the</strong>re are only three genera (each consisting<strong>of</strong> but asingle species), Pinicola, Nyctea, and Surnia. Lagopus is circumpolar,but <strong>the</strong> genus has too wide an extension in <strong>the</strong>temperate zone to be considered arctic. Among aquatic birdswe have <strong>the</strong> genus <strong>of</strong> ducks, Somateria ; three genera <strong>of</strong> Uriidse,Uria, Catarractes, and Mergulus ; and <strong>the</strong> small family Alcidse rconsisting <strong>of</strong> <strong>the</strong> genera Alca and Fratercula. Our total <strong>the</strong>nis, three genera <strong>of</strong> mammalia, three <strong>of</strong> land, and six <strong>of</strong> aquaticbirds, including one peculiar family.In <strong>the</strong> sou<strong>the</strong>rn hemisphere <strong>the</strong>re is only <strong>the</strong> single genusAptenodytes that can be classed as antarctic ; and even that ismore properly south temperate.In dealing <strong>with</strong> this arctic fauna we have two courses opento us ; we must ei<strong>the</strong>r group <strong>the</strong>m <strong>with</strong> <strong>the</strong> o<strong>the</strong>r species andgenera which are common to <strong>the</strong> two nor<strong>the</strong>rn regions, or we


70 DISTRIBUTION OF ANIMALS. [part r.must form a separate primary region for <strong>the</strong>m. As a matter <strong>of</strong>convenience <strong>the</strong> former plan seems <strong>the</strong> best ; and it is thatwhich is in accordance <strong>with</strong> our treatment <strong>of</strong> o<strong>the</strong>r intermediatetracts which contain special forms <strong>of</strong> life. <strong>The</strong> great desertzone, extending from <strong>the</strong> Atlantic shores <strong>of</strong> <strong>the</strong> Sahara acrossArabia to Central Asia, is a connecting link between <strong>the</strong> Palsearctic,Ethiopian, and Oriental regions, and contains a number<strong>of</strong> " desert " forms wholly or almost wholly restricted to it ; but<strong>the</strong> attempt to define it as a separate region would introducedifficulty and confusion. Nei<strong>the</strong>r to <strong>the</strong> " desert " nor to <strong>the</strong>" arctic " regions could any defined limits, ei<strong>the</strong>r <strong>geographical</strong>or zoological, be placed ; and <strong>the</strong> attempt to determine whatspecies or genera should be allotted to <strong>the</strong>m would prove aninsoluble problem. <strong>The</strong> reason perhaps is, that both are essentiallyunstable, to a much greater extent than those great masses<strong>of</strong> land <strong>with</strong> more or less defined barriers, which constitute oursix regions.<strong>The</strong> Arctic Zone has been, <strong>with</strong>in a recent geologicalperiod, both vastly more extensive and vastly lessextensivethan it is at present. At a not distant epoch it extended overhalf <strong>of</strong> Europe and <strong>of</strong> North America. At an earlier date itappears to have vanished altoge<strong>the</strong>r ; since a luxuriant vegetation<strong>of</strong> tall deciduous trees and broad-leaved evergreensflourished <strong>with</strong>in ten degrees <strong>of</strong> <strong>the</strong> Pole ! <strong>The</strong> great desertshave not improbably been equally fluctuating; hence nei<strong>the</strong>r<strong>the</strong> one nor <strong>the</strong> o<strong>the</strong>r can present that marked individualityin <strong>the</strong>ir forms <strong>of</strong> life, which seems to have arisen only whenextensive tracts <strong>of</strong> land have retained some considerable stabilityboth <strong>of</strong> surface and climatal conditions, during periodssufficient for <strong>the</strong> development and co-adaptation <strong>of</strong> <strong>the</strong>ir severalassemblages <strong>of</strong> plants and <strong>animals</strong>.We must also consider that <strong>the</strong>re isno <strong>geographical</strong> difficultyin dividing <strong>the</strong> Arctic Zone between <strong>the</strong> two nor<strong>the</strong>rn regions.<strong>The</strong> only debateable lands, Greenland and Iceland, are generallyadmitted to belong respectively to America and Europe.Nei<strong>the</strong>r is <strong>the</strong>re any zoological difficulty ; for <strong>the</strong> land mammaliaand birds are on <strong>the</strong> whole wonderfully restricted to <strong>the</strong>irrespective regions even in high latitudes ; and <strong>the</strong> aquatic forms


;chap, iv.] ZOOLOGICAL EEGIONS. 71are, for our present purpose, <strong>of</strong> much less importance. As aprimary division <strong>the</strong> " Arctic region " would be out <strong>of</strong> all proportionto <strong>the</strong> o<strong>the</strong>r six, whe<strong>the</strong>r as regards its few peculiartypes or <strong>the</strong> limited number <strong>of</strong> forms and species actually inhabitingit ; but it comes in well as a connecting link betweentwo regions, where <strong>the</strong> peculiar forms <strong>of</strong> both are specially modified; and is in this respect quite analogous to <strong>the</strong> great desertzone above referred to.I now proceed to characterize briefly <strong>the</strong> six regions adoptedin <strong>the</strong> present work, toge<strong>the</strong>r <strong>with</strong> <strong>the</strong> sub-regions into which<strong>the</strong>y may be most conveniently and naturally divided, as shownin our general map.Palcearctic Region.—Thisvery extensive region comprises alltemperate Europe and Asia, from Iceland to Behring's Straits andfrom <strong>the</strong> Azores to Japan. Its sou<strong>the</strong>rn boundary is somewhatindefinite, but it seems advisable to comprise in it all<strong>the</strong> extra- tropical part <strong>of</strong> <strong>the</strong> Sahara and Arabia, and allPersia, Cabul, and Beloochistan to <strong>the</strong> Indus. It comes downto a little below <strong>the</strong> upper limit <strong>of</strong> forests in <strong>the</strong> Himalayas,and includes <strong>the</strong> larger nor<strong>the</strong>rn half <strong>of</strong> China, not quite s<strong>of</strong>ar down <strong>the</strong> coast as Amoy. It has been said that thisregion differs from <strong>the</strong> Oriental by negative characters only ; ahost <strong>of</strong>tropical families and genera being absent, while <strong>the</strong>re islittle or nothing but peculiar species to characterize it absolutely.This however is not true. <strong>The</strong> Paleearctic region is wellcharacterized by possessing 3 families <strong>of</strong> vertebrata peculiarto it, as well as 35 peculiar genera <strong>of</strong> mammalia, and 57<strong>of</strong> birds, constituting about one-third <strong>of</strong> <strong>the</strong> total number itpossesses. <strong>The</strong>se are amply sufficient to characterize a regionpositively ; but we must also consider <strong>the</strong> absence <strong>of</strong> many importantgroups <strong>of</strong> <strong>the</strong> Oriental, Ethiopian, and Nearctic regionsand we shall <strong>the</strong>n find, that taking positive and negativecharacters toge<strong>the</strong>r,and making some allowance for <strong>the</strong> necessarypoverty <strong>of</strong> a temperate as compared <strong>with</strong> tropical regions,<strong>the</strong> Palsearctic is almost asany o<strong>the</strong>r.strongly marked and well defined asSub-divisions <strong>of</strong> <strong>the</strong> Palmarctic Region.—<strong>The</strong>se are by no means


72 DISTRIBUTION OF ANIMALS. [part I.so clearly indicated as in some <strong>of</strong> <strong>the</strong> o<strong>the</strong>r regions, and <strong>the</strong>y areadopted more for convenience than because <strong>the</strong>y are very naturalor strongly marked.<strong>The</strong> first, or European sub-region, comprises Central andNor<strong>the</strong>rn Europe as far South as <strong>the</strong> Pyrenees, <strong>the</strong> Maritimeand Dinaric Alps, <strong>the</strong> Balkan mountains, <strong>the</strong> Black Sea, and <strong>the</strong>Caucasus. On <strong>the</strong> east <strong>the</strong> Caspian sea and <strong>the</strong> Ural mountainsseem <strong>the</strong> most obvious limit ; but it is doubtful if <strong>the</strong>y form <strong>the</strong>actual boundary, which is perhaps better marked by <strong>the</strong> valley<strong>of</strong> <strong>the</strong> Irtish, where a pre-glacial sea almost certainly connected<strong>the</strong> Aral and Caspian seas <strong>with</strong> <strong>the</strong> Arctic ocean, and formedan effective barrier which must still, to some extent, influence<strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong>.<strong>The</strong> next, or Mediterranean sub-region, comprises SouthEurope, North Africa <strong>with</strong> <strong>the</strong> extra-tropical portion <strong>of</strong> <strong>the</strong>Sahara, and Egypt to about <strong>the</strong> first or second cataracts ; andeastward through Asia Minor, Persia, and Cabul, to <strong>the</strong> deserts<strong>of</strong> <strong>the</strong> Indus.<strong>The</strong> third, or Siberian sub-region, consists <strong>of</strong> all north andcentral Asia north <strong>of</strong> Herat, as far as <strong>the</strong> eastern limits <strong>of</strong> <strong>the</strong>great desert plateau <strong>of</strong> Mongolia, and southward to about <strong>the</strong>upper limit <strong>of</strong> trees on <strong>the</strong> Himalayas.<strong>The</strong> fourth, or Manchurian sub-region, consists <strong>of</strong> Japan andNorth China <strong>with</strong> <strong>the</strong> lower valley <strong>of</strong> <strong>the</strong> Amoor ;and it shouldprobably be extended westward in a narrow strip along <strong>the</strong>Himalayas, embracing about 1,000 or 2,000 feet <strong>of</strong> verticaldistance below <strong>the</strong> upper limit <strong>of</strong> trees, till it meets an easternextension <strong>of</strong> <strong>the</strong> Mediterranean sub-region a little beyond Simla.<strong>The</strong>se extensions are necessary to avoid passing from <strong>the</strong> Orientalregion, which isessentially tropical, directly to <strong>the</strong> Siberian subregion,which has an extreme nor<strong>the</strong>rn character ; whereas <strong>the</strong>Mediterranean and Manchurian sub-regions are more temperatein climate. It will be found that between <strong>the</strong> upper limit <strong>of</strong>most <strong>of</strong> <strong>the</strong> typical Oriental groups and <strong>the</strong> Thibetan or Siberianfauna, <strong>the</strong>re is a zone in which many forms occur common totemperate China. This is especially <strong>the</strong> case among <strong>the</strong> pheasantsand finches.


7;chap, iv.] ZOOLOGICAL REGIONS. 73Ethiopian Region.—<strong>The</strong> limits <strong>of</strong> this region have been indicatedby <strong>the</strong> definition <strong>of</strong> <strong>the</strong> Palsearctic region.Besides Africasouth <strong>of</strong> <strong>the</strong> tropic <strong>of</strong> Cancer, and its islands, it comprises <strong>the</strong>sou<strong>the</strong>rn half <strong>of</strong> Arabia.This region has been said to be identical in <strong>the</strong> main characters<strong>of</strong> its mammalian fauna <strong>with</strong> <strong>the</strong> Oriental region, and has<strong>the</strong>refore been united <strong>with</strong> it by Mr. A. Murray. Most importantdifferences have however been overlooked, as <strong>the</strong> followingsummary <strong>of</strong> <strong>the</strong> peculiarities <strong>of</strong> <strong>the</strong> Ethiopian region will, Ithink, show.It possesses 22 peculiar families <strong>of</strong> vertebrates ; 90 peculiargenera <strong>of</strong> mammalia, being two-thirds <strong>of</strong> its whole numberand 179 peculiar genera <strong>of</strong> birds, being three-fifths <strong>of</strong> all itpossesses. It is fur<strong>the</strong>r characterized by <strong>the</strong> absence <strong>of</strong> severalfamilies and genera which range over <strong>the</strong> whole nor<strong>the</strong>rnhemisphere, details <strong>of</strong> which will be found in <strong>the</strong> chaptertreating <strong>of</strong> <strong>the</strong> region. <strong>The</strong>re are, it is true, many points<strong>of</strong> resemblance, not to be wondered at between two tropicalregionsin <strong>the</strong> same hemisphere, and which have evidently beenat one time more nearly connected, both by intervening landsand by a different condition <strong>of</strong> <strong>the</strong> lands that even now connect<strong>the</strong>m.remarkable ;But <strong>the</strong>se resemblances only render <strong>the</strong> differences moresince <strong>the</strong>y show that <strong>the</strong>re has been an ancient andlong-continued separation <strong>of</strong> <strong>the</strong> two regions, developing a distinctfauna in each, and establishing marked specialitieswhich<strong>the</strong> temporary intercommunication and immigration has notsufficed to remove. <strong>The</strong> entire absence <strong>of</strong> such wide-spreadgroups as bears and deer, from a country many parts <strong>of</strong> whichare well adapted to <strong>the</strong>m, and in close proximity to regionswhere <strong>the</strong>y abound, would alone mark out <strong>the</strong> Ethiopian regionas one <strong>of</strong> <strong>the</strong> primary divisions <strong>of</strong> <strong>the</strong> earth, even if it possesseda less number than it actually does <strong>of</strong> peculiar family andgenericgroups.Sub-divisions <strong>of</strong> <strong>the</strong> Ethiopian Region.—<strong>The</strong> African continentsouth <strong>of</strong> <strong>the</strong> tropic <strong>of</strong> Cancer is more homogeneous in itsprominent and superficial zoological features than most <strong>of</strong> <strong>the</strong>o<strong>the</strong>r regions, but <strong>the</strong>re are never<strong>the</strong>less important and deep-Vol. I.—


74 DISTRIBUTION OF ANIMALS. [part i.seated local peculiarities. Two portions can be marked <strong>of</strong>f aspossessing many peculiar forms ; <strong>the</strong> luxuriant forest district<strong>of</strong> equatorial West Africa, and <strong>the</strong> sou<strong>the</strong>rn extremity or Capedistrict. <strong>The</strong> remaining portion has no well-marked divisions,and a large proportion <strong>of</strong> its animal forms range over it fromNubia and Abyssinia, to Senegal on <strong>the</strong> one side and to <strong>the</strong>Zambesi on <strong>the</strong> o<strong>the</strong>r ; this forms our first or East-Africansub-region.<strong>The</strong> second, or West African sub-region extends along <strong>the</strong>coast from Senegal to Angola, and inland to <strong>the</strong> sources <strong>of</strong> <strong>the</strong>Shary and <strong>the</strong> Congo.<strong>The</strong> third, or South African sub-region, comprises <strong>the</strong> CapeColony and Natal, and is roughly limited by a line from DelagoaBay to Walvish Bay.<strong>The</strong> fourth, or Malagasy sub-region, consists <strong>of</strong> Madagascar and<strong>the</strong> adjacent islands, from Eodriguez to <strong>the</strong> Seychelles ;and thisdiffers so remarkably from <strong>the</strong> continent that it has been proposedto form a distinct primary region for its reception. Itsproductions are indeed highly interesting; since it possesses3 families, and 2 sub-families <strong>of</strong> mammals peculiar to itself,while almost all its genera are peculiar. Of <strong>the</strong>se a fewshow Oriental or Ethiopian affinities, but <strong>the</strong> remainder arequite isolated. Turning to o<strong>the</strong>r classes <strong>of</strong> <strong>animals</strong>, we findthat <strong>the</strong> birds are almost as remarkable ; but, as might beexpected, a larger number <strong>of</strong> genera are common to surroundingcountries. More than 30 genera are altoge<strong>the</strong>r peculiar,in separate families or sub-families.and some <strong>of</strong> <strong>the</strong>se are so isolated as to require to be classed<strong>The</strong> African affinity is howeverhere more strongly shown by <strong>the</strong> considerable number (13)<strong>of</strong> peculiar Ethiopian genera which in Madagascar have representativespecies. <strong>The</strong>re can be no doubt <strong>the</strong>refore about Madagascarbeing more nearly related to <strong>the</strong> Ethiopian than to anyo<strong>the</strong>r region ; but its peculiarities are so great, that, were it notfor its small size and <strong>the</strong> limited extent <strong>of</strong> its fauna, its claim torank as a separate region might not seem unreasonable. It istrue that it is not poorer in mammals than Australia ;but thatcountry is far more isolated, and cannot be so decidedly and


chap, iv.] ZOOLOGICAL REGIONS. 75naturally associated <strong>with</strong> any o<strong>the</strong>r region as Madagascar canbe <strong>with</strong> <strong>the</strong> Ethiopian. It is <strong>the</strong>refore <strong>the</strong> better and morenatural course to keep it as a sub-region ; <strong>the</strong> peculiarities itexhibits being <strong>of</strong> exactly <strong>the</strong> same kind as those presented by<strong>the</strong> Antilles, by New Zealand, and even by Celebes and Ceylon,but in a much greater degree.Oriental Region.—On account <strong>of</strong> <strong>the</strong> numerous objectionsthat have been made to naming a regionfrom <strong>the</strong> least characteristicportion <strong>of</strong> it, and not thinking " Malayan," proposed byMr. Blanford, a good term, (as it has a very circumscribed anddefinite meaning, and especially because <strong>the</strong> " Malay " archipelagois half <strong>of</strong> it in <strong>the</strong> Australian region,) I propose to use<strong>the</strong> word " Oriental " instead <strong>of</strong> " Indian," as being <strong>geographical</strong>lyapplicable to <strong>the</strong> whole <strong>of</strong> <strong>the</strong> countries included in <strong>the</strong> regionand to very few beyond it ; as being euphonious, and as beingfree from all confusion <strong>with</strong> terms already used in zoologicalgeography. I trust <strong>the</strong>refore that it may meet <strong>with</strong> generalacceptance.This small, compact, but rich and varied region, consists <strong>of</strong>all India and China from <strong>the</strong> limits <strong>of</strong> <strong>the</strong> Palsearctic region ;all <strong>the</strong> Malay peninsula and islands as far east as Java andBaly, Borneo and <strong>the</strong> Philippine Islands ; and Formosa. It ispositively characterized by possessing 12 peculiar families <strong>of</strong>vertebrata ; by 55 genera <strong>of</strong> land mammalia, and 165 genera<strong>of</strong> land birds, altoge<strong>the</strong>r confined to it ; <strong>the</strong>se peculiar generaforming in each case about one half <strong>of</strong> <strong>the</strong> total number itpossesses.Sub-divisions <strong>of</strong> <strong>the</strong> Oriental region.—First we have <strong>the</strong>Indian sub-region, consisting <strong>of</strong> Central India from <strong>the</strong> foot <strong>of</strong><strong>the</strong> Himalayas in <strong>the</strong> west, and south <strong>of</strong> <strong>the</strong> Ganges to <strong>the</strong>east, as far as a linedrawn from Goa curving south and up to<strong>the</strong> Kistna river; this is <strong>the</strong> portion which has most affinity<strong>with</strong> Africa.<strong>The</strong> second, or Ceylonese sub-region, consists <strong>of</strong> <strong>the</strong>sou<strong>the</strong>rnextremity <strong>of</strong> India <strong>with</strong> Ceylon ; this is a mountainous forestregion, and possesses several peculiar forms as well as someMalayan types not found in <strong>the</strong> firstsub-region.


76 DISTRIBUTION OF ANIMALS. [part i.Next we have <strong>the</strong> Indo-Chinese sub-region, comprising SouthChina and Burmah, extending westward along <strong>the</strong> Himalayanrange to an altitude <strong>of</strong> about 9,000 or 10,000 feet, and southwardto Tavoy or Tenasserim.<strong>The</strong> last is <strong>the</strong> Indo-Malayan sub-region, comprising <strong>the</strong>Peninsula <strong>of</strong> Malacca and <strong>the</strong> Malay Islands to Baly, Borneo,and <strong>the</strong> Philippines.On account <strong>of</strong> <strong>the</strong> absence from <strong>the</strong> first sub-region <strong>of</strong> many <strong>of</strong><strong>the</strong> forms most characteristic <strong>of</strong> <strong>the</strong> o<strong>the</strong>r three, and <strong>the</strong> number<strong>of</strong> families and genera <strong>of</strong> mammalia and birds which occur in itand also in Africa, it has been thought by some naturalists thatthis part <strong>of</strong>India has at least an equal claim to be classed as apart <strong>of</strong> <strong>the</strong> Ethiopian region.discussed in Chapter XII. devoted toThis question will be found fully<strong>the</strong> Oriental region, whereit is shown that <strong>the</strong> African affinity is far less than has beenrepresented, and that in all itswholly Oriental in its fauna.essential features Central India isBefore leaving this region a few words may be said aboutLemuria, a name proposed by Mr. Sclater for <strong>the</strong> site <strong>of</strong> a supposedsubmerged continent extending from Madagascar to Ceylonand Sumatra, in which <strong>the</strong> Lemuroid type <strong>of</strong> <strong>animals</strong> was developed.This is undoubtedly a legitimate and highly probable supposition,and itis an example <strong>of</strong> <strong>the</strong> way in which a <strong>study</strong> <strong>of</strong> <strong>the</strong><strong>geographical</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong> may enable us to reconstruct<strong>the</strong> geography <strong>of</strong> a bygone age.But we must not, as Mr. Blythproposed, make this hypo<strong>the</strong>tical land one <strong>of</strong> our actual Zoologicalregions. It represents what was probably a primaryZoological region in some past geological epoch ; but what thatepoch was and what were <strong>the</strong> limits <strong>of</strong> <strong>the</strong> region in question, weare quite unable to say. If we are to suppose that it comprised<strong>the</strong> whole area now inhabited by Lemuroid <strong>animals</strong>, we mustmake it extend from West Africa to Burmah, South China, andCelebes ; an area which it possibly did once occupy, but whichcannot be formed into a modern Zoological region <strong>with</strong>out violatingmuch more important affinities. If, on <strong>the</strong> o<strong>the</strong>r hand,we leave out all those areas which undoubtedly belong to o<strong>the</strong>rregions, we reduce Lemuria to Madagascar and its adjacent


chap. IV.] ZOOLOGICAL REGIONS. 77islands, which, for reasons already stated, it is not advisable totreat as a primary Zoological region. <strong>The</strong> <strong>the</strong>ory <strong>of</strong> this ancientcontinent and <strong>the</strong> light it may throw on existing anomalies <strong>of</strong><strong>distribution</strong>, will be more fully considered in <strong>the</strong> <strong>geographical</strong>part <strong>of</strong> this work.Australian Region.—Mr. Sclater's original name seems preferableto Pr<strong>of</strong>essor Huxley's, "Austral- Asian- ; " <strong>the</strong> inconvenience<strong>of</strong> which alteration is sufficiently shown by <strong>the</strong> factthat Mr. Blyth proposed to use <strong>the</strong> very same term as anappropriate substitute for <strong>the</strong> " Indian region " <strong>of</strong> Mr. Sclater.Australia is <strong>the</strong> great central mass <strong>of</strong> <strong>the</strong> region ; it is by far <strong>the</strong>richest in varied and highly remarkable forms <strong>of</strong> life ; and it<strong>the</strong>refore seems in every way fitted to give a name to <strong>the</strong> region<strong>of</strong> which it is <strong>the</strong> essential element. <strong>The</strong> limits <strong>of</strong> this regionin <strong>the</strong> Pacific are somewhat obscure, but as so many <strong>of</strong> <strong>the</strong>Pacific Islands are extremely poor zoologically, this is not <strong>of</strong>great importance.Sub-divisions <strong>of</strong> <strong>the</strong> Australian Region.—<strong>The</strong> firstsub-regionis <strong>the</strong> Austro-Malayan, including <strong>the</strong> islands from Celebes andLombock on <strong>the</strong> west to <strong>the</strong> Solomon Islands on <strong>the</strong> east. <strong>The</strong>Australian sub-region comes next, consisting <strong>of</strong> Australia andTasmania. <strong>The</strong> third, or Polynesian sub-region, will consist<strong>of</strong> all <strong>the</strong> tropical Pacific Islands, and is characterized byseveral peculiar genera <strong>of</strong> birds which are all allied to Australiantypes. <strong>The</strong> fourth, consists <strong>of</strong> New Zealand <strong>with</strong> Auckland,Chatham, and Norfolk Islands, and must be called <strong>the</strong> NewZealand sub-region.<strong>The</strong> extreme peculiarities<strong>of</strong> New Zealand, due no doubt toits great isolation and to its being <strong>the</strong> remains <strong>of</strong> a moreextensive land, have induced several naturalists to suggest thatit ought justly to form a Zoological region by itself. But <strong>the</strong>inconveniences <strong>of</strong> such a procedure have been already pointedout ; and when we look at its birds as a whole (<strong>the</strong>y being <strong>the</strong>only class sufficiently well represented t<strong>of</strong>ound any conclusionupon) we find that <strong>the</strong> majority <strong>of</strong> <strong>the</strong>m belong to Australiangenera, and where <strong>the</strong> genera are peculiar <strong>the</strong>y aremost nearlyrelated to Australian types. <strong>The</strong> preservation in <strong>the</strong>se islands


78 DISTRIBUTION OF ANIMALS. [part i.<strong>of</strong> a single representative <strong>of</strong> a unique order <strong>of</strong> reptiles, is, asbefore remarked, <strong>of</strong> <strong>the</strong> same character as <strong>the</strong> preservation <strong>of</strong><strong>the</strong> Proteus in <strong>the</strong> caverns <strong>of</strong> Carniola ;and can give <strong>the</strong> localitywhere it happens to have survived no claim to form a primaryZoological region, unless supported by a tolerably varied anddistinctly characterized fauna, such as never exists in a veryrestricted and insular area.Neotropical Region.—Mr. Sclater's original name for thisregion is preserved, because change <strong>of</strong> nomenclature is alwaysan evil; and nei<strong>the</strong>r Pr<strong>of</strong>essor Huxley's suggested alteration" Austro-Columbia," nor Mr. Sclater's new term " Dendrogaea,"appear to be improvements. <strong>The</strong> region is essentially a tropicalone, and <strong>the</strong> extra-tropical portion <strong>of</strong> it is not importantenough to make <strong>the</strong> name inappropriate. That proposed byPr<strong>of</strong>essor Huxley is not free from <strong>the</strong> same kind <strong>of</strong> criticism,since it would imply that <strong>the</strong> region was exclusively SouthAmerican, whereas a considerable tract <strong>of</strong> North Americabelongs to it. This region includes South America, <strong>the</strong>Antilles and tropical North America ; and it possesses morepeculiar families <strong>of</strong> vertebrates and genera <strong>of</strong> birds and mammaliathan any o<strong>the</strong>r region.Subdivisions <strong>of</strong> <strong>the</strong> Neotropical Region.—<strong>The</strong> great centralmass <strong>of</strong> South America, from <strong>the</strong> shores <strong>of</strong> Venezuela to Paraguayand Eastern Peru, constitutes <strong>the</strong> chief division, and may betermed <strong>the</strong> Brazilian sub-region. It is on <strong>the</strong> whole a forestcountry; its most remarkable forms are highly developedarboreal types ; and it exhibits all <strong>the</strong> characteristics <strong>of</strong> this richand varied continent in <strong>the</strong>ir highest development.<strong>The</strong> second, or Chilian sub-region, consists <strong>of</strong> <strong>the</strong> open plains,pampas, and mountains <strong>of</strong> <strong>the</strong> sou<strong>the</strong>rn extremity <strong>of</strong> <strong>the</strong> continent;farand we must include in it <strong>the</strong> west side <strong>of</strong> <strong>the</strong> Andes asas <strong>the</strong> limits <strong>of</strong> <strong>the</strong> forest near Payta, and <strong>the</strong> whole <strong>of</strong> <strong>the</strong>high Andean plateaus as far as 4° <strong>of</strong> south latitude; whichmakes it coincide <strong>with</strong> <strong>the</strong> range <strong>of</strong> <strong>the</strong> Camelidse and Chinchillidae.<strong>The</strong> third, or Mexican sub-region, consists <strong>of</strong> Central Americaand Sou<strong>the</strong>rn Mexico, but it has no distinguishing character-


chap, iv.] ZOOLOGICAL REGIONS. 79istics except <strong>the</strong> absence <strong>of</strong> some <strong>of</strong> <strong>the</strong> more highly specializedNeotropical groups. It is, however, a convenient division ascomprising <strong>the</strong> portion <strong>of</strong><strong>the</strong> North American continent whichbelongs zoologically to South America.<strong>The</strong> fourth, or Antillean sub-region, consists <strong>of</strong> <strong>the</strong> WestIndia islands(except Trinidad and Tobago, which are detachedportions <strong>of</strong> <strong>the</strong> continent and must be grouped in<strong>the</strong> first subregion);and <strong>the</strong>se reproduce, in a much less marked degree,<strong>the</strong> phenomena presented by Madagascar.Terrestrial mammalsare almost entirely wanting, but <strong>the</strong> larger islands possess threegenera which are altoge<strong>the</strong>r peculiar to <strong>the</strong>m. <strong>The</strong> birds are<strong>of</strong> South American forms, but comprise many peculiar genera.Terrestrial molluscs are more abundant and varied than inanypart <strong>of</strong> <strong>the</strong> globe <strong>of</strong> equal extent ; and if <strong>the</strong>se alone wereconsidered, <strong>the</strong> Antilles would constitute an important Zoologicalregion.Nearctic Region.—This region comprises all temperate NorthAmerica and Greenland. <strong>The</strong> arctic lands and islands beyond<strong>the</strong> limit <strong>of</strong> trees form a transitional territory to <strong>the</strong> Palsearcticregion, but even here <strong>the</strong>re are some characteristic species.<strong>The</strong> sou<strong>the</strong>rn limit between this region and <strong>the</strong> Neotropical islittle uncertain ; but it may be drawn at about <strong>the</strong> Rio Grandedel Norte on <strong>the</strong> east coast, and a little north <strong>of</strong> Mazatlan on<strong>the</strong> west ; while on <strong>the</strong> central plateau it descends much far<strong>the</strong>rsouth, and should perhaps include all <strong>the</strong> open highlands <strong>of</strong>Mexico and Guatemala.several characteristic Nearctic genera.This would coincide <strong>with</strong> <strong>the</strong> range <strong>of</strong>Distinction <strong>of</strong> <strong>the</strong> Nearctic from <strong>the</strong> Palcearctic Region.—<strong>The</strong>Nearctic region possesses twelve peculiar families <strong>of</strong> vertebratesor one-tenth <strong>of</strong> its whole number. It has also twenty-fourpeculiar genera <strong>of</strong> mammalia and fifty-two <strong>of</strong> birds, in eachcase nearly one-third <strong>of</strong> all it possesses.This proportion is verynearly <strong>the</strong> same as in <strong>the</strong> Palsearctic region, while <strong>the</strong> number <strong>of</strong>peculiar families <strong>of</strong> vertebrata is very much greater.aIt has beenalready seen that both Mr. Blyth and Pr<strong>of</strong>essor Huxley aredisposed to unite this region <strong>with</strong> <strong>the</strong> Palsearctic, while Pr<strong>of</strong>essorNewton, in his article on birds in <strong>the</strong> new edition <strong>of</strong> <strong>the</strong>


;80 DISTRIBUTION OF ANIMALS. [part i.Encyclopaedia Britannica, thinks that as regards that class itcan hardly claim to be more than a sub-region <strong>of</strong> <strong>the</strong> Neotropical.<strong>The</strong>se views are mutually destructive, but it will be shown in<strong>the</strong> proper place, that on independent grounds <strong>the</strong> Nearcticregion can very properly be maintained.Subdivisions <strong>of</strong> <strong>the</strong> Nearctic Region.—<strong>The</strong> sub-regions heredepend on <strong>the</strong> great physical features <strong>of</strong> <strong>the</strong> country, and havebeen in some cases accurately definedby American naturalists.First we have <strong>the</strong> Californiau sub-region, consisting <strong>of</strong> Californiaand Oregon—a narrow tract between <strong>the</strong> Sierra Nevadaand <strong>the</strong> Pacific, but characterized by a number <strong>of</strong> peculiarspecies and by several genera found nowhere else in <strong>the</strong> region.<strong>The</strong> second, or Rocky Mountain sub-region, consists <strong>of</strong> thisgreat mountain range <strong>with</strong> its plateaus, and <strong>the</strong> central plainsand prairies to about 100° west longitude, but including NewMexico and Texas in <strong>the</strong> South.<strong>The</strong> third and most important sub-region, which may betermed <strong>the</strong> Alleghanian, extends eastward to <strong>the</strong> Atlantic, including<strong>the</strong> Mississippi Valley, <strong>the</strong> Alleghany Mountains, and<strong>the</strong> Eastern United States. This is an old forest district, andcontains most <strong>of</strong> <strong>the</strong> characteristic animal types <strong>of</strong> <strong>the</strong> region.<strong>The</strong> fourth, or Canadian sub-region, comprises all <strong>the</strong> nor<strong>the</strong>rnpart <strong>of</strong> <strong>the</strong> continent from <strong>the</strong> great lakes to <strong>the</strong> Arctic oceana land <strong>of</strong> pine-forests and barren wastes, characterized by Arctictypes and <strong>the</strong> absence <strong>of</strong> many <strong>of</strong> <strong>the</strong> genera which distinguish<strong>the</strong> more sou<strong>the</strong>rn portions <strong>of</strong> <strong>the</strong> region.Observations on <strong>the</strong> series <strong>of</strong> Sub-regions.—<strong>The</strong> twenty-four subregionshere adopted were arrived at by a careful consideration<strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> more important genera, and <strong>of</strong> <strong>the</strong>materials, both zoological and <strong>geographical</strong>, available for tbeirdetermination ; and it was not till <strong>the</strong>y were almost finallydecided on, that <strong>the</strong>y were found to be equal in number throughoutall <strong>the</strong> regions—four in each. As this uniformity is <strong>of</strong> greatadvantage in tabular and diagrammatic presentations <strong>of</strong> <strong>the</strong><strong>distribution</strong> <strong>of</strong> <strong>the</strong> several families, I decided not to disturbit unless very strong reasons should appear for adopting a greateror less number in any particular case. Such however have not


CHAP. IV.] ZOOLOGICAL EEGIONS. 81arisen ; and it is hoped that <strong>the</strong>se divisions will prove as satisfactoryand useful to naturalists in general as <strong>the</strong>y have been to<strong>the</strong> author. Of course, in a detailed <strong>study</strong> <strong>of</strong> any region muchmore minute sub-division may be required; but even in thatcase it is believed that <strong>the</strong> sub-regions here adopted, will befound, <strong>with</strong> slight modifications, permanently available for exhibitinggeneral results.I give here a table showing <strong>the</strong> proportionate richness andspeciality <strong>of</strong> each region as determined by itsfamilies <strong>of</strong> vertebratesand genera <strong>of</strong> mammalia and birds ; and also a generaltable <strong>of</strong> <strong>the</strong> regions and sub-regions, arranged in <strong>the</strong> order thatseems best to show <strong>the</strong>ir mutual <strong>relations</strong>.Comparative Richness <strong>of</strong> <strong>the</strong> Six Regions.


82 DISTRIBUTION OF ANIMALS. [part. I.Table <strong>of</strong> Regions and Sub-regions— continued.Regions.


—CHAPTER V.CLASSIFICATION AS AFFECTING THE STUDY OF GEOGRAPHICALDISTRIBUTION.A little consideration will convince us, that no inquiry into<strong>the</strong> causes and laws which determine <strong>the</strong> <strong>geographical</strong> <strong>distribution</strong><strong>of</strong> <strong>animals</strong> or plants can lead to satisfactory results, unlesswe have a tolerably accurate knowledge <strong>of</strong> <strong>the</strong> affinities <strong>of</strong> <strong>the</strong>several species, genera, and families to each o<strong>the</strong>r; in o<strong>the</strong>rwords, we require a natural classification to work upon.for example, take three <strong>animals</strong>Let us,a, b, and c—which have ageneral external resemblance to each o<strong>the</strong>r, and are usuallyconsidered to be really allied ; and let us suppose that a and binhabit <strong>the</strong> same or adjacent districts, while c is found far awayon <strong>the</strong> o<strong>the</strong>r side <strong>of</strong> <strong>the</strong> globe, <strong>with</strong> no <strong>animals</strong> at all resemblingit in any <strong>of</strong> <strong>the</strong> intervening countries.We should here have adifficult problem to solve ; for we should have to show that <strong>the</strong>general laws by which we account for <strong>the</strong> main features <strong>of</strong><strong>distribution</strong>, will explain this exceptional case. But now, supposesome comparative anatomist takes <strong>the</strong>se <strong>animals</strong> in hand,and finds that <strong>the</strong> resemblance <strong>of</strong> c to a and b isonly superficial,while <strong>the</strong>ir internal structure exhibits marked and importantdifferences ; and that c really belongs to ano<strong>the</strong>r group <strong>of</strong><strong>animals</strong>, d, which inhabits <strong>the</strong> very region in which c wasfound— and we should no longer have anything to explain.This is no imaginary case. Up to a very few years ago acurious Mexican animal, Bassaris astuta, was almost alwaysclassed in <strong>the</strong> civet family (Viverrida3), a group entirely con-


84 DISTRIBUTION OF ANIMALS. [part i.fined to Africa and Asia ;but it has now been conclusivelyshown by Pr<strong>of</strong>essor Flower that its real affinities are <strong>with</strong> <strong>the</strong>racoons (Procyonidae), a group confined to North and SouthAmerica. In ano<strong>the</strong>r case, however, an equally careful examinationshows, that an animal peculiar to <strong>the</strong> Himalayas (JZlurusfulgens) has its nearest ally in <strong>the</strong> Cercolejrtes <strong>of</strong> South America.Here, <strong>the</strong>refore, <strong>the</strong> <strong>geographical</strong> difficulty really exists, and anysatisfactory <strong>the</strong>ory <strong>of</strong> <strong>the</strong> causes that have led to <strong>the</strong> existing<strong>distribution</strong> <strong>of</strong> living things, must be able to account, more orless definitely, for this and o<strong>the</strong>r anomalies. From <strong>the</strong>se casesit will be evident, that if any class or order <strong>of</strong> <strong>animals</strong> is veryimperfectly known and its classification altoge<strong>the</strong>r artificial, it isuseless to attempt to account for <strong>the</strong> anomalies its <strong>distribution</strong>may present ; since those anomalies may be, to a great extent,due to false notions as to <strong>the</strong> affinities <strong>of</strong> its component species.According to <strong>the</strong> laws and causes <strong>of</strong><strong>distribution</strong> discussed in<strong>the</strong> preceding chapters, we should find limited and defined<strong>distribution</strong> to be <strong>the</strong> rule, universal or indefinite <strong>distribution</strong> tobe <strong>the</strong> exception, in everynatural group corresponding to whatare usually regarded as families and genera ; and so much isthis <strong>the</strong> case in nature, that when we find a group <strong>of</strong> thisnominal rank scattered as it were at random over <strong>the</strong> earth, wehave a strong presumption that it is not natural ; but is, to aconsiderable extent, a haphazard collection <strong>of</strong> species.Of coursethis reasoning will only apply, in cases where <strong>the</strong>re are nounusual means <strong>of</strong> dispersal, nor any exceptional causes whichmisht determine a scattered <strong>distribution</strong>.From <strong>the</strong> considerations now adduced it becomes evident, thatit is <strong>of</strong> <strong>the</strong> first importance for <strong>the</strong> success <strong>of</strong> our inquiry tosecure a natural classification <strong>of</strong> <strong>animals</strong>, especially as regards<strong>the</strong> families and genera.<strong>The</strong> higher groups, such as classes andorders, are <strong>of</strong> less importance for our purpose ; because <strong>the</strong>y arealmost always widely and <strong>of</strong>ten universally distributed, exceptthose which are so small as to be evidently <strong>the</strong> nearly extinctrepresentatives <strong>of</strong> a once more extensive series <strong>of</strong> forms. Wenow proceed to explain <strong>the</strong> classification to be adopted, as lowdown as <strong>the</strong> series <strong>of</strong> families. To <strong>the</strong>se, equivalent English


.CHAP. V.] CLASSIFICATION. 85names are given wherever <strong>the</strong>y exist, in order that readers possessingno technical knowledge, may form some conception <strong>of</strong><strong>the</strong> meaning <strong>of</strong> <strong>the</strong> term " family " in zoology.<strong>The</strong> primary divisions <strong>of</strong> <strong>the</strong> animal kingdom according totwo eminent modern authorities are as follows :Huxley.Classification <strong>of</strong> Animals (1S69).1. Protozoa2. Infusoria3. Ccelenterata4. Annuloida5. Annulosa6. Molluscoida7. Mollufeca8. VertebrataCarus and Gerstaeker.Handbuch der Zoologie (1868).1. Protozoa.2. Coelenterata.3. Echinoderrnata.( 4. Vermes.( 5. Arthropoda.6. Molluscoida.7. Mollusca.8. Vertebrata.For reasons already stated it is only <strong>with</strong> <strong>the</strong> fifth, seventh,and eighth <strong>of</strong> <strong>the</strong>se groups that <strong>the</strong> present work proposes todeal ; and even <strong>with</strong> <strong>the</strong> fifth and seventh only partially and in<strong>the</strong> most general way.<strong>The</strong> classes <strong>of</strong> <strong>the</strong> vertebrata, according to both <strong>the</strong> authorsabove quoted, are: 1. Mammalia. 2. Aves. 3. Eeptilia. 4.Amphibia. 5. Pisces, in which order <strong>the</strong>y will be taken here.<strong>The</strong> sub-classes and orders <strong>of</strong> mammalia are as follows :MAMMALIA.Huxley (1869), Flower (1870).1. Primates2. Chiroptera3. Insectivora4. Carnivora5. Cetaeea )Monodelpbia... < 6. Siren ia )7. Ungulata8. Proboscidea .9. Hyracoidea ..10. RodentiaEdentataill.Didelphia 12. Marsupialia .Ornithodelphia 13. MonotremataCarus (1868).{ 1. Primates.\ 5. Prosimii.2. Chiroptera.3. Insectivora.5 6. Carnivora.I7. Pinnipedia.12. Natantia.5 10. Artiodactyla.11 Perissodactyla.I9. Proboscidea.8. Lamnungia.4. Rodentia.13. Brut a.14. Marsupialia.15. Monotremata.


86 DISTRIBUTION OF ANIMALS. [part I.<strong>The</strong> above series <strong>of</strong> orders is arranged according to Pr<strong>of</strong>essorFlower's Osteology <strong>of</strong> Mammalia, and <strong>the</strong>y will follow in thissuccession throughout my work. Pr<strong>of</strong>essor Huxley arranges<strong>the</strong> same orders in a different series.In determining <strong>the</strong> manner in which <strong>the</strong> several ordersbe subdivided into families, I have been guided inshallmy choice <strong>of</strong>classifications mainly by <strong>the</strong> degree <strong>of</strong> attention <strong>the</strong> author appearsto have paid to <strong>the</strong> group, and his known ability as aIn many cases it is a matter <strong>of</strong> great doubt whe<strong>the</strong>ra certain group should form several distinct families or be unitedinto one or two ; but one method may bring out <strong>the</strong> peculiarities<strong>of</strong> <strong>distribution</strong> much better than <strong>the</strong> o<strong>the</strong>r, and this is, in ourcase, a sufficient reason for adopting it.For <strong>the</strong> Primates I follow, <strong>with</strong> some modifications, <strong>the</strong>classification <strong>of</strong> Mr. St. George Mivart given in his article" Apes " in <strong>the</strong> new edition <strong>of</strong> <strong>the</strong> Encyclopaedia Britannica, andin his paper in <strong>the</strong> Proceedings <strong>of</strong> <strong>the</strong> Zoological Society <strong>of</strong> London,1865, p. 547. It is as follows :systematic zoologist ; and in a less degree by considerations <strong>of</strong>convenience as regards <strong>the</strong> special purposes <strong>of</strong> <strong>geographical</strong> <strong>distribution</strong>.Order-


CHAP. V.] CLASSIFICATION. 87better to treat <strong>the</strong>m as families, a rank which is claimed for <strong>the</strong>anthropoid apes by many naturalists.As no good systematic work on <strong>the</strong> genera and species <strong>of</strong> batshas been yet published, I adopt <strong>the</strong> five families as generallyused in this country, <strong>with</strong> <strong>the</strong> genera as given in <strong>the</strong> papers <strong>of</strong>Dr. J. E. Gray and Mr. Tomes. A monograph by Dr. Petershas long been promised, and his outline arrangement waspublished in 1865, but this will perhaps be materially alteredwhen <strong>the</strong> work appears.FrugivoraInsectivorailstiophoraGyrunorhiniOrder—CHIROPTERA.Fam....9. Pteropida?{ 10. Phyllostomidse(11. Rhinolophidse(12. Vespertilionidae( 13. NoctilionidseFruit-eating Bats.Leaf-nosed Bats.Horse-shoe Bats.True Bats.Dog-headed Bats.<strong>The</strong> genera <strong>of</strong> Chiroptera are in a state <strong>of</strong> great confusion, <strong>the</strong>names used by different authors being 1 <strong>of</strong>ten not at all comparable,so that <strong>the</strong> few details given <strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong>bats are not trustworthy. We have <strong>the</strong>refore made little use<strong>of</strong> this order in <strong>the</strong> <strong>the</strong>oretical part <strong>of</strong> <strong>the</strong> work.<strong>The</strong> osteology <strong>of</strong> <strong>the</strong> Insectivora has been very carefullyworked out by Pr<strong>of</strong>essor Mivart in <strong>the</strong> Jounral <strong>of</strong> Anatomyand Physiology (Vol. ii., p. 380), and I follow his classificationas given <strong>the</strong>re, and in <strong>the</strong> Proceedings <strong>of</strong> <strong>the</strong> Zoological Society(1871).Fam.14. Galeopi<strong>the</strong>cidae15. Macroscelididse16. Tupaiidae17. Erinaceidse ...18. Centetidse19. Potamogalidse20. Chrysochloridse21. Talpidae22. SoricidseOrder—INSECTIVORA.Flving Lemurs.Elephant Shrews.Squirrel Shrews.Hedgehogs.Tenrecs.Otter Shrew.Golden Moles.Moles.Shrews.<strong>The</strong> next order, Carnivora, has been studied in detail byPr<strong>of</strong>essor Flower ;and I adopt <strong>the</strong> classification given by him in<strong>the</strong> Proceedings <strong>of</strong> <strong>the</strong> Zoological Society, 1869, p. 4.


131..88 DISTRIBUTION OF ANIMALS. [part I.Order—CARNIVORA.iEluroideaFissipedia 'Cynoidea. ArctoideaPinnipediaFam.23. Felidfe ...24. Cryptoproctida?25. Viverridae26. Protelidre^27. Hysenidse28. Canidse ...'29. Mustelidae30. Procyonida;iEluridae...32. Ursidee ...33. Otariidae . .34. Trichechidge35. Phocidse....Cats, Lion, &c.Cryptoprocta.Civets.Aard-wolf.Hyaenas.Dogs, Foxes, &c.Weasels.Racoons.Pandas.Bears.Eared Seals.Walrus.Seals.<strong>The</strong> Cetacea is one <strong>of</strong> those orders <strong>the</strong> classification <strong>of</strong> whichis very unsettled. <strong>The</strong> <strong>animals</strong> comprising it are so huge, and<strong>the</strong>re is so much difficulty in preserving <strong>the</strong>m, that only a veryfew species are known <strong>with</strong> anything like completeness. A considerablenumber <strong>of</strong> genera and species have been described orindicated ;but as many <strong>of</strong> <strong>the</strong>se are founded on imperfect specimens<strong>of</strong> perhaps a single individual, it isnot to be wondered atthat those few naturalists who occupy <strong>the</strong>mselves <strong>with</strong> <strong>the</strong> <strong>study</strong><strong>of</strong> <strong>the</strong>se large <strong>animals</strong>, cannot agree as to <strong>the</strong> proper mode <strong>of</strong> grouping<strong>the</strong>m into natural families.<strong>The</strong>y are, however, <strong>of</strong> but littleimportance to us, as almost all <strong>the</strong> species inhabit <strong>the</strong> ocean, and<strong>of</strong> only a few <strong>of</strong> <strong>the</strong>m can it be said that anything is accuratelyknown <strong>of</strong> <strong>the</strong>ir <strong>distribution</strong>. I <strong>the</strong>refore consider it best to followPr<strong>of</strong>essor Cams, who makes a smaller number <strong>of</strong> families ; butI give also <strong>the</strong> arrangement <strong>of</strong> Dr. Gray in his British Museumcatalogue <strong>of</strong> whales and seals, as modified subsequently in<strong>the</strong> Proceedings <strong>of</strong> Zoological Society, 1870, p. 772. <strong>The</strong> Zeuglodontidse,a family <strong>of</strong> extinct tertiary whales, are classed byPr<strong>of</strong>essors Owen and Carus between Cetacea and Sirenia, whilePr<strong>of</strong>essor Huxley considers <strong>the</strong>m to have been carnivorous andallied to <strong>the</strong> seals.


,:8:CHAP. V.] CLASSIFICATION. 89Order— CETACEA.Sub-order I.-Mystaceti.Fam. (Carus).BalaenidaeBalsenopteridaeC Catodontidae ...HyperoodontidseSu£ ord er l}-\ MonodontidfflfOdontoceti.Extinct familyDelphinidas ...Zeuglodontidae,Fam. (Gray).36. Balaenidae.37. Balaenopteridae.38. CatodontidaB.!Hyperoodontidae.Epiodontklae.Xiphiada?.40. (Part <strong>of</strong> Delphinidae.)Platanistida?.41Iniadae.Delphinidae.Globiocephalidas.Orcadse.Belugidae.Pontoporiadae.Order— SIKENIA.<strong>The</strong> order Sirenia, comprising <strong>the</strong> sea-cows, consists <strong>of</strong> a singlefamilyFamily 42.Manatidae.<strong>The</strong> extensive order Ungulata comprises <strong>the</strong> three ordersPachydermata, Solidungula, and Paiminantia <strong>of</strong> <strong>the</strong> older naturalists.<strong>The</strong> following classification is that now generallyadopted, <strong>the</strong> only difference <strong>of</strong> opinion being as to whe<strong>the</strong>rsome <strong>of</strong> <strong>the</strong> groups should be classed as families or sub-families,a matter <strong>of</strong> little importance for our purposeOrder—UNGULATA.Perissodactyla orOdd-toed UngulatesArtiodactyla orEven-toed UngulatesVol. I.—,Fam.43. Equidae... 44. Tapiridse45. Rhinocerotidae . .Suina( 46. Hippopotamida?(47. SuidseI Tylopoda 48. Camelidae< Tragulina 49. Tragulidae50. CervidaePecora{50. J 51. Canielopardidae52. BovidaeHorses.Tapirs.Rhinoceros.Hippopotamus.Swine.Camels.Chevrotains.Deer.Giraffes.( Cattle, Sheep,( Antelopes, &c.


,62.:90 DISTRIBUTION OF ANIMALS. [part I.<strong>The</strong> two next orders consist <strong>of</strong> but a single family each, viz.OrderPROBOSCIDEAHYRACOIDEAFam.53. Elephantidse54. HyracidseElephants.Bock-rabbits.We now come to <strong>the</strong> Eodentia, a very extensive and difficultorder, in which <strong>the</strong>re is still much difference <strong>of</strong> opinion as to<strong>the</strong> details <strong>of</strong> classification, although <strong>the</strong> main outlines are prettywell settled. <strong>The</strong> foundations <strong>of</strong> a true classification <strong>of</strong> thisorder were laid by Mr. G. E. Waterhouse more than thirty yearsago, and succeeding authors have done little more than followhis arrangement <strong>with</strong> unimportant modifications. Pr<strong>of</strong>essorLilljeborg, <strong>of</strong> Upsala, has however made a special <strong>study</strong> <strong>of</strong> thisgroup <strong>of</strong> <strong>animals</strong>, and has given an original and detailed classification<strong>of</strong> all <strong>the</strong> genera. {Systematisk Ofversigt af de GnagandeDaggdjuren, Glires. Upsala, 1866.) I follow this arrangement<strong>with</strong> a few slight modifications suggested by o<strong>the</strong>r naturalists,and which make it better adapted for <strong>the</strong> purposes <strong>of</strong> this work.SimplicidentatixMurina(Waterhouse)Hystricina(Waterhouse)Duplicidentatij (wSZe)Order—RODENTIA.Fam.55. Muridag56. Spalacidae57. Dipodidse58. Myoxidse59. Saccomyidai ...60. Castoridse61. SciuridreHaploodontida?63. Chinchillidse ...64. Octodontidse ..65. Echimyidse66. Cercolabidse ..,67. Hystricidse68. Caviidse( 69. Lagomyidse\ 70. LeporidseRats.Mole-rats.Jerboas.Dormice.Pouched Rats.Beavers.Squirrels.Sewellels.Chinchillas.Octodons.Spiny Rats.Tree Porcupines.Porcupines.Cavies.Pikas.Hares.'<strong>The</strong> Edentata have been classified by Mr. Turner, in <strong>the</strong>Proceedings <strong>of</strong> <strong>the</strong> Zoological Society (1851, p. 205), by Dr.Gray in <strong>the</strong> British Museum Catalogue, and by Pr<strong>of</strong>essor Camsin his Handhuch. <strong>The</strong> former takes a middle course between


''HAP. V.] CLASSIFICATION. 91<strong>the</strong> numerous families <strong>of</strong> Dr. Gray, seven in number, and <strong>the</strong>two families to which Pr<strong>of</strong>essor Carus restricts <strong>the</strong> existingspecies. I <strong>the</strong>refore follow Mr. Turner.Order—EDENTATA.BradypodaEntomophagaFain.71. Bradypodidae'72. Manididae73. Dasypodidae74. Orycteropodidaa...75. MyrinecophagidaeSloths.Scaly Ant-eaters.Armadillos.Ant-bears.Ant-eaters.<strong>The</strong> Marsupials have been well classified and described byMr. Waterhouse in <strong>the</strong> first volume <strong>of</strong> his Natural History ojMammalia, and his arrangement is here followed. <strong>The</strong> subordersadopted by Pr<strong>of</strong>essor Carus are also given.Order—MARSUPIALIA.Kapacia (Wagner)Poephaga (Owen)Carpophaga (Owen)Rhizophaga (Owen)Fam.76. Didelphidae ...77. Dasyuridae ...78. Myrmecobiidae79. Peramelidae ...80. Macropodidae81. Phalangistidae82. PhascolomyidajOpossums.Native Cats.Native Ant-eater.Bandicoots.Kangaroos.Phalangers.Wombats.Order—MONOTEEMATA.<strong>The</strong> last order, <strong>the</strong> Monotremata, consist <strong>of</strong> two families, whichPr<strong>of</strong>essor Carus combines into one, but which it seems morenatural to keep separate.Fam.83. Ornithorhynchidae84. EchidnidaeDuckbill.Echidna.


.92 DISTRIBUTION OF ANIMALS. [part i.BIRDS.Birds are perhaps <strong>the</strong> most difficult to classify <strong>of</strong> all <strong>the</strong>divisions <strong>of</strong> <strong>the</strong> vertebrata. <strong>The</strong> species and genera areexceedingly numerous, and <strong>the</strong>re is such a great uniformityin oeneral structure and even in <strong>the</strong> details <strong>of</strong> external form,that it isexceedingly difficult to find characters by which ordersand families can be characterised.For a long time <strong>the</strong> system<strong>of</strong> Vigors and Swainson was followed ; but this wholly ignoredanatomical characters and in many cases plainly violated wellmarkedaffinities. Characters derived from <strong>the</strong> form <strong>of</strong> <strong>the</strong>sternum, <strong>the</strong> scutellation <strong>of</strong> <strong>the</strong> tarsi, and <strong>the</strong> arrangement <strong>of</strong><strong>the</strong> fea<strong>the</strong>rs, have all assisted in determining natural groups.More recently Pr<strong>of</strong>essor Huxley has applied <strong>the</strong> variations <strong>of</strong> <strong>the</strong>bony palate to <strong>the</strong> general arrangement <strong>of</strong> birds ;and still morerecently Pr<strong>of</strong>essor Garrod has studied certain leg-muscles for<strong>the</strong> same purpose. <strong>The</strong> condition <strong>of</strong> <strong>the</strong> young as regardsplumage, and even <strong>the</strong> form, texture, and coloration <strong>of</strong> <strong>the</strong> egg,have also been applied to solve doubtful cases <strong>of</strong> affinity;yet<strong>the</strong> problem is not settled, and it will probably remain forano<strong>the</strong>r generation <strong>of</strong> ornithologists to determine <strong>with</strong> anyapproach to accuracy what are <strong>the</strong> most natural divisions <strong>of</strong> <strong>the</strong>class into orders and families. In a work like <strong>the</strong> present it isevidently not advisable to adopt all <strong>the</strong> recent classifications;since experience has shown thatno arrangement in which oneset <strong>of</strong> characters is mainly relied on, long holds its ground.Such modifications <strong>of</strong> <strong>the</strong> old system as seem to be wellestablished will be adopted ; but <strong>the</strong> older groups will be adheredto in cases where <strong>the</strong> most recent classificationsare opento doubt, or seem inconvenient as separating families, which,owing to <strong>the</strong>ir similarity in general structure, form and habitsare best kept toge<strong>the</strong>r for <strong>the</strong> purposes <strong>of</strong> <strong>geographical</strong> <strong>distribution</strong>.<strong>The</strong> old plan <strong>of</strong> putting <strong>the</strong> birds <strong>of</strong> prey at <strong>the</strong> head <strong>of</strong> <strong>the</strong>class, is now almost wholly given up ;both because <strong>the</strong>y are not


chap, v.] CLASSIFICATION. 93<strong>the</strong> most highly organised, but only one <strong>of</strong> <strong>the</strong> most specialisedforms <strong>of</strong> birds, and because <strong>the</strong>ir affinities are not <strong>with</strong> <strong>the</strong>Passeres, but ra<strong>the</strong>r <strong>with</strong> <strong>the</strong> cormorants and some o<strong>the</strong>r<strong>of</strong> <strong>the</strong>aquatic groups. <strong>The</strong> Passeres <strong>the</strong>refore are placed first ; and <strong>the</strong>series <strong>of</strong> families is begun by <strong>the</strong> thrushes, which are certainly<strong>the</strong> most typical and generally well-organised form <strong>of</strong> birds.Instead <strong>of</strong> <strong>the</strong> Scansores and Fissirostres <strong>of</strong> <strong>the</strong> older authors, <strong>the</strong>order Picarise, which includes <strong>the</strong>m both, is adopted, but <strong>with</strong>some reluctance; as <strong>the</strong> former are, generally speaking, well markedand strongly contrasted groups, although certain families havebeen shown to be intermediate.In <strong>the</strong> Picarise are included <strong>the</strong>goat-suckers, swifts, and humming-birds, sometimes separatedas a distinct order, Macrochires. <strong>The</strong> parrots and <strong>the</strong> pigeonsform each a separate order. <strong>The</strong> old groups <strong>of</strong> Grallse andAnseres are preserved, as more convenient than breaking <strong>the</strong>mup into widely separated parts ; for though <strong>the</strong> latter plan mayin some cases more strictly represent <strong>the</strong>ir affinities, its detailsare not yet established, nor is it much used by ornithologists.In accordance <strong>with</strong> <strong>the</strong>se views <strong>the</strong> following is <strong>the</strong> series <strong>of</strong>orders and families <strong>of</strong> birds adopted in this work :Class—AVES.Orders.1 Passeres \ Including <strong>the</strong> great mass <strong>of</strong> <strong>the</strong> smaller birds—Crows,'\ Finches, Flycatchers, Creepers, Honeysuckers, &c, &c.2 Picari-e ""\( ^ncm^m g Woodpeckers, Cuckoos, Toucans, Kingfishers,Swifts, &c, &c.3. Psittaci . . . Parrots only.4. Columbee ... Pigeons and <strong>the</strong> Dodo.5. Gallinaa ... Grouse, Pheasants, Curassows, Mound-builders, &c.6. Opisthocomi <strong>The</strong> Hoazin only.7. Accipitres ... Eagles, Owls, and Vultures.8. Grallse ... Herons, Plovers, Eails, &c.9. Anseres ... Gulls, Ducks, Divers, &c10. Struthiones . . . Ostrich, Cassowary, Apteryx, &c.<strong>The</strong> Passeres consist<strong>of</strong> fifty families, which may be arrangedand grouped in series as follows. It must however be rememberedthat <strong>the</strong> first family in each series is not always thatwhich is most allied to <strong>the</strong> last family <strong>of</strong> <strong>the</strong> preceding series.All extensive natural groups consist <strong>of</strong> divergent or branchingalliances, which renders it impossible to arrange <strong>the</strong> whole inone continuous series.


—94 DISTRIBUTION OF ANIMALS. [part I.A.Typical or Turdoid


—:CHAP. V.] CLASSIFICATION. 95D.— Formicaroid Passeres — continued.46. Pteroptochidae47. Pittidfe48. PaictidsePittas.49. Menuridae50. AtrichidaeE.Anomalous Passeres.Lyre-birds.Scrub-birds.<strong>The</strong> preceding arrangement is a modification <strong>of</strong>that proposedby myself in <strong>the</strong> Ibis (1874, p. 406). <strong>The</strong> principal alterationsare adding <strong>the</strong> families Panuridae and Sittidae in seriesA, commencing series B <strong>with</strong> Dicaeidae ; bringing Vireonidaenext to <strong>the</strong> allied American family Mniotiltidae ; and placingMotacillidae in series C next to Alaudidae.Pr<strong>of</strong>essor Newton I place Menuridae and AtrichidaeAt <strong>the</strong> suggestion <strong>of</strong>apart from<strong>the</strong> o<strong>the</strong>r Passeres, as <strong>the</strong>y both possess striking peculiarities <strong>of</strong>anatomical structure.<strong>The</strong> heterogeneous families constituting <strong>the</strong> order Picariae maybe conveniently arranged as follows


:,Numidinae..:96 DISTRIBUTION OF ANIMALS. [PAET I.<strong>The</strong> Psittaci or parrot tribe are still in a very unsettled state<strong>of</strong> classification ; that recently proposed by Pr<strong>of</strong>essor Garroddiffering widely from <strong>the</strong> arrangement adopted in Dr. Finsch'smonograph <strong>of</strong> <strong>the</strong> order. Taking advantage <strong>of</strong> <strong>the</strong> researches<strong>of</strong> <strong>the</strong>se and o<strong>the</strong>r authors, <strong>the</strong> following families are adopted as<strong>the</strong> most convenient in <strong>the</strong> present state <strong>of</strong> our knowledge76. Cacatuidae ...77. Platycercidas78. Palaeornithidae79. Trichoglossidae80. Conuridse ...81. Psittacidae ...82. Nestoridae ...83. Stringopidae<strong>The</strong> Cockatoos.<strong>The</strong> Broad-tailed Paroquets <strong>of</strong> Australia.<strong>The</strong> Oriental Parrots and Paroquets.<strong>The</strong> Brush-to ngued Paroquets and Lories.<strong>The</strong> Macaws and <strong>the</strong>ir allies.<strong>The</strong> African and South American Parrots.<strong>The</strong> Nestors <strong>of</strong> New Zealand.<strong>The</strong> Owl-parrots <strong>of</strong> New Zealand.<strong>The</strong> Columbre, or pigeons, are also in a very unsatisfactorystate as regards a natural classification. <strong>The</strong> families, subfamilies,and genera proposed by various authors are verynumerous, and <strong>of</strong>ten quite irreconcilable. I <strong>the</strong>refore adoptonly two families ; and generally follow Mr. G. E. Gray's handlistfor <strong>the</strong> genera, except where trustworthy authorities existfor a different arrangement. <strong>The</strong> families are :84. Columbidee85. DididsePigeons and Doves.<strong>The</strong> extinct Dodo and allies.<strong>The</strong> Gallinae, or game-birds, may be divided into sevenfamiliesFam.86. Pteroclidae87. Tetraonidae88. Phasianidae ..89. Turnicidae90. Megapodiida?91. Cracidae92. TinamidaeSub-fam.Pavoninae . .LophophorinaePhasianinae . .\ EuplocaminaeGallinaeMeleagrinae . ....{CracinaePenelopinae .Oreophasina?Sand-grouse.Partridges and Grouse.Peafowl.Tragopans, &c.Pheasants.Fire-backed Pheasants, &c.Jungle-fowl.Turkeys.Guinea-fowl.Hemipodes.Mound-makers.Curassows.Guans.Mountain-pheasant.Tinamous.


—; 91..CHAP. V.] CLASSIFICATION.<strong>The</strong> Opisthocomi consist <strong>of</strong> one family containing a singlespecies, <strong>the</strong> " Hoazin " <strong>of</strong> Guiana.Family 93.Opisthocomidae.<strong>The</strong> Accipitres, or birds <strong>of</strong> prey, which were long consideredto be <strong>the</strong> highest and most perfect order <strong>of</strong> birds, are nowproperly placed lower down in <strong>the</strong> series, <strong>the</strong>ir affinities beingmore <strong>with</strong> <strong>the</strong> aquatic than <strong>with</strong> <strong>the</strong> perching birds. <strong>The</strong>following is <strong>the</strong> arrangement adopted by Mr. Sharpe in hisrecently published British Museum catalogue <strong>of</strong> diurnal birds<strong>of</strong> prey :Sub-orders.Fam.VulturidaeFalcones 95. Serpentariidce1 96. Falconida?Pandiones...97. Pandionida?Striges ...98. StrigidseSub-families.( Vulturinre .. ..\SarcorhamphinrePolyborn iaeAccipitrinse( ButeoninseAquilinte . .k. FalconinreVultures.Turkey-buzzards.Caracaras.Hawks.Buzzards.Eagles.Falcons.Fishing-eagles.Owls.<strong>The</strong> Grallse or Grail atores are in a very unsettled state. <strong>The</strong>following series <strong>of</strong> families is in accordance <strong>with</strong> <strong>the</strong> views <strong>of</strong>some <strong>of</strong> <strong>the</strong> best modern ornithologists :99. Rallidae ...100.' Scolopacidae101. Chionididse102. Thinocoridse103. Parridse ...104. Glareolidae105. Charadriidse106. Otididse ...»107. Gruidse ...108. Cariamidas109. Aramida?...110. Pspphiidse111. Eurypygida?112. Rhinochoetida3113. Ardeida3 ...114. Plataleidae115. Ciconiidse116. Palamedeidse117. Ph£enicopterida3Rails, &c.Sandpipers and SnipesSheath-bills.Quail-snipes.Jacanas.Pratincoles.Plovers.Bustards.Cranes.Cariarnas.Guaraunas.Trumpeters.Sun-bitterns.Kagus.Herons.Spoonbills and Ibis.Storks.Screamers.Flamingoes.


—98 DISTRIBUTION OF ANIMALS. [PART I.<strong>The</strong> Anseres or Natatores are almost equally unsettled. <strong>The</strong>flamingoes are usually placed in this order, but <strong>the</strong>ir habits bestassort <strong>with</strong> those <strong>of</strong> <strong>the</strong> waders.Fam.118. Anatidse119. Larida?120. Procellariidae121. Pelecanidse ...122. Spheniscidae123. Colymbidae124. Podicipidse ...125. AlcidseDuck and Geese.Gulls.Petrels.Pelicans.Penguins.Divers.Grebes.Auks.<strong>The</strong> last order <strong>of</strong> birds is <strong>the</strong> Struthiones or Katitse, consideredby many naturalists to form a distinct sub-class. It consists <strong>of</strong>comparatively few species, ei<strong>the</strong>r living or recently extinct.Fam.?126. Struthionidae127. Casuariidse ...128. Apterygidae ...( 129. DinornithidaeExtinct < 130. Palapterygidse( 131. ^EpyornithidseOstriches.Cassowaries.Apteryx.Dinomis.Palapteryx.iEpyornis.EEPTILES.In reptiles I follow <strong>the</strong> classification<strong>of</strong> Dr. Giin<strong>the</strong>r as givenin <strong>the</strong> Philosojrfiical Transactions, vol. clvii., p. 625. He divides<strong>the</strong> class into five orders as follows :


——CHAP. V.] CLASSIFICATION. 99<strong>The</strong> Ophidia, or Snakes, form <strong>the</strong> first order and are classifiedas follows :Fain.1. Typhlopidae2. Tortricidse3. Xenopeltidse4. Uropeltidae5. Calamaridae6. Oligodontidae.7. Colubridae8. Homalopsidse ••9. Psammophidse ..Innocuous Snakes { 10. Rachiodontidae.11. Dendrophidae ..12. Dryiophidas13. Dipsasidae14. Scytalidae.15. Lycodontidae16. Amblycephalidae17. Pythonidae18. Erycida?19. Acrochordidae ..20. ElapidaeVenomous Colubrine } 21. Dendraspididae.Snakes j 22. Atractaspididae.23. HydrophidaeBurrowing Snakes.Dwarf ground-snakes.Colubrine Snakes.Fresh-water Snakes.Desert-snakes.Tree-snakes.Whip-snakes.Nocturnal tree-snakes.Fanged ground-snakes.Blunt-heads.Pythons.Sand-snakes.Wart-snakes.Cobras, &c.Sea-snakes.Viperine Snakes24. Crotalidae25 ViperidaePit-vipers.True vipers<strong>The</strong> second order, Lacertilia, are arranged as follows :Fam.26. Trogonophidse ...27. Chirotidae28. Amphisbsenidse29. Lepidosternidae30. Varanidae31. Heloderrnidae.32. Teidse33. Lacertidae34. Zonuridae35. Chalcidae.36. Anadiadse.37. ChirocoUdae.38. Iphisadae.39. Cercosaurid82.40. Chamaesauridae.41. Gymnopthalmidae42. Pygopodidae43. Aprasiadae.> Amphisbaenians.Water Lizards.Teguexins> Land LizardsGape-eyed Scinks.Two-legged Lizards.


———100 DISTRIBUTION OF ANIMALS. [PART 1.Fam.44. Lialidae.45. Scincidae46. Ophiomoridae47. Sepidae ...48. Acontiadae.49. Geckotidae50. Iguanidae51. Agamidae52. Chameleonida?Scinks.Snake-lizards.Sand-lizards.Geckoes.Iguanas.Fringed Lizards.Chameleons.<strong>The</strong> third order, Bhyncocephalina consists <strong>of</strong> a single family53. Rhyncocephalidae .. <strong>The</strong> Hatteria <strong>of</strong> New Zealand.<strong>The</strong> fourth order, Crocodilia or Loricata, consists <strong>of</strong> threefamilies :54. Gavialidse55. Crocodilidae56. AlligatoridaeGavials.Crocodiles.Alligators.<strong>The</strong> fifth order, Chelonia, consists <strong>of</strong> four families :57. Testudinidae58. Chelydidee59. Trionychidae60. CheloniidaeLand and fresh -water Tortoises.Fresh-water Turtles.S<strong>of</strong>t Turtles.Sea Turtles.AMPHIBIA.In <strong>the</strong> Amphibia I follow <strong>the</strong> classification <strong>of</strong> Pr<strong>of</strong>essorMivart, as given for a largepart <strong>of</strong> <strong>the</strong> order in <strong>the</strong> Proceedings<strong>of</strong> <strong>the</strong> Zoological Society for 1869. For <strong>the</strong> remainder I followDr. Strauch, Dr. Giin<strong>the</strong>r, and a MSS. arrangement kindlyfurnished me by Pr<strong>of</strong>essor Mivart.<strong>The</strong> class isfirst divided into three groups or orders, and <strong>the</strong>ninto families as follows :


CHAP. V.] CLASSIFICATION. 101Order I.—PSEUDOPHIDIA.Fam.1. CseciliadaeCsecilia.Order II.—BATEACHIA URODELA.2. Sirenidae ...3. Proteidae ...4. Arnphiuniidae5. Menopornidae6. SalainandridaeSiren.Proteus.Amphiuma.Giant Salamanders.Salamanders and Newts.Fam.7. Rhinophrynidae8. Phryniscidae ..9. Hylaplesidse ..10. Bufonidae11. Xenorhinidae ..12. Engystoniidae ..13. Bombinatoridse14. Plectromantidae15. AlytidaeOrder III.>Toads.'.} Frogs.BATRACHIA ANOURA.Fam.16. Pelodryadae17. Hylidae ...18. Polypedatidae19. Rauidae ...23. Discoglossidse21. Pipidae ...22. DactvlethridaeiTree Frogs.( Frogs.I Tongueless] Toads.FISHES.<strong>The</strong>se are arranged according to <strong>the</strong> classification <strong>of</strong> Dr.Giin<strong>the</strong>r, whose great work " <strong>The</strong> British Museum Catalogue <strong>of</strong>Fishes," has furnished almost all <strong>the</strong> material for our account<strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> class.In that work all existing fishes are arranged in six sub-classesand thirteen orders. A <strong>study</strong> <strong>of</strong> <strong>the</strong> extraordinary Ceratodusfrom Australia has induced Dr. Giin<strong>the</strong>r to unite three <strong>of</strong> hissub-classes ; but as his catalogue will long remain a handbookfor every student <strong>of</strong> fishes, it seems better to follow <strong>the</strong> arrangement<strong>the</strong>re given, indicating his later views by bracketingtoge<strong>the</strong>r <strong>the</strong> groups he now thinks should be united.


102 DISTRIBUTION OF ANIMALS. [part I.


;chap, v.] CLASSIFICATION. 103<strong>The</strong> number <strong>of</strong> species <strong>of</strong> Butterflies isabout <strong>the</strong> same as that<strong>of</strong> Birds, while <strong>the</strong> six families <strong>of</strong> Coleoptera selected, comprisemore thau twenty thousand species, far exceeding <strong>the</strong> number <strong>of</strong>all o<strong>the</strong>r vertebrates. <strong>The</strong>se families have all been recently catalogued,so that we have very complete information as to <strong>the</strong>irarrangement and <strong>distribution</strong>.LEPIDOPTERA DIURNA, OR BUTTERFLIES.Fam.Fain.1. Danaidse. 9. Libythseidse.2. Satyrida;. 10. Nerneobiidae.3. Elyinniidae. 11. Eurygonidse.4. Morphidse. 12. Erycinidse.5. Brassolidse. 13. Lycsenidae.6. Acraeidse. 14. Pieridse.7. Heliconidse. 15. Papilionidae.8. Nymphalidse. 16. Hesperidae.COLEOPTERA, OR BEETLES.Fam.Fam.1. Cicindelidae... Tiger-beetles. 4. Cetoniidse ... Rose-chafers.2. Carabidse ... Ground-beetles. 5. Buprestidse . . .Metallic Beetles.3. Lucanidae ... Stag-beetles. 6. Longicornia . . . Long-horned Beetles.<strong>The</strong> above families comprise <strong>the</strong> extensive series <strong>of</strong> groundbeetles (Carabidse) containing about 9,000 species, and <strong>the</strong> Longicorns,which are nearly as numerous and surpass <strong>the</strong>m in variety<strong>of</strong> form and colour as well as in beauty. <strong>The</strong> Cetoniidse andBuprestidse are among <strong>the</strong> largest and most brilliant<strong>of</strong> beetles<strong>the</strong> Lucanidae are pre-eminent for remarkable form, and <strong>the</strong>Cicindelidae for elegance; and all <strong>the</strong> families are especialfavourites <strong>with</strong> entomologists, so that <strong>the</strong> whole earth has beenransacked to procure fresh species.Results deduced from a <strong>study</strong> <strong>of</strong> <strong>the</strong>se will, <strong>the</strong>refore, fairlyrepresent <strong>the</strong> phenomena <strong>of</strong> <strong>distribution</strong> <strong>of</strong> Coleoptera, and,. as <strong>the</strong>y are very varied in <strong>the</strong>ir habits, perhaps <strong>of</strong> insects ingeneraL


——104 DISTRIBUTION OF ANIMALS. [PART I.MOLLUSCA.<strong>The</strong> Mollusca are usually divided into five classes asfollows :Classes.I. CephalopodaII. GasteropodaIII. PteropodaIV. BrachiopodaV. ConchiferaCuttle-fish.Snails and aquatic Univalves.Oceanic Snails.Symmetrical Bivalves.Unsymmetrical Bivalves.<strong>The</strong> Gasteropoda and Conchifera alone contain land andfreshwater forms, and to <strong>the</strong>se we shall chiefly confine ourillustrations <strong>of</strong> <strong>the</strong> <strong>geographical</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> Mollusca.<strong>The</strong> classification followed is that <strong>of</strong> Dr. Pfeiffer for <strong>the</strong>Operculata and Dr. Von Martens for <strong>the</strong> Helicidse. <strong>The</strong>families chiefly referred to are :ClassII.—GASTEROPODA.In- operculataOperculata ..Order 2.—Pulmonifera.Fam.1. Helicidse.2. Lhnacidse.3. Oncidiadse.4. Lirnneeidae.5. Auriculidae.6. Aciculidae.7. Diplommatinidse.8. Cyclostomidse.9. Helicinidse.


9PART IT.ON THE DISTRIBUTION OF EXTINCT ANIMALS.Vol. L—


CHAPTER VI.THE EXTINCT MAMMALIA OF THE OLD WORLD.Although it may seem somewhat out <strong>of</strong> place to begin <strong>the</strong>systematic treatment <strong>of</strong> our subject <strong>with</strong> extinct ra<strong>the</strong>r than<strong>with</strong> living <strong>animals</strong>, it is necessary to do so in order that wemay see <strong>the</strong> meaning and trace <strong>the</strong> causes <strong>of</strong> <strong>the</strong> existing <strong>distribution</strong><strong>of</strong> animal forms. It is true, that <strong>the</strong> <strong>animals</strong> foundfossil in a country are very generally allied to those which stillinhabit it ; but this is by no means universally <strong>the</strong> case. If itwere, <strong>the</strong> attempt to elucidate our subject by Palaeontologywould be hopeless, since <strong>the</strong> past would show us <strong>the</strong> samepuzzling diversities <strong>of</strong> faunas and floras that now exist. "Werind however very numerous exceptions to this rule, and it is<strong>the</strong>se exceptions which tell us <strong>of</strong> <strong>the</strong> past migrations <strong>of</strong> wholegroups <strong>of</strong> <strong>animals</strong>. We are thus enabled to determine whatportion <strong>of</strong> <strong>the</strong> existing races <strong>of</strong> <strong>animals</strong> in a country are descendants<strong>of</strong> its ancient fauna, and which are comparativelymodern immigrants ; and combining <strong>the</strong>se movements <strong>of</strong> <strong>the</strong>forms <strong>of</strong> life <strong>with</strong> known or probable changes in <strong>the</strong> <strong>distribution</strong><strong>of</strong> land and sea, we shall sometimes be able to trace approximately<strong>the</strong> long series <strong>of</strong> changes which have resulted in <strong>the</strong>actual state <strong>of</strong> things. To gain this knowledge is our object in<strong>study</strong>ing <strong>the</strong> " Geographical Distribution<strong>of</strong> Animals," and ourplan <strong>of</strong> <strong>study</strong> must be determined, mainly, by <strong>the</strong> facilities itaffords us for attaining this object. In discussing <strong>the</strong> countlessdetails <strong>of</strong> <strong>distribution</strong> we shall meet <strong>with</strong> in our survey <strong>of</strong> <strong>the</strong>zoological regions, we shall <strong>of</strong>ten find it useful to refer to <strong>the</strong>evidence we possess <strong>of</strong> <strong>the</strong> range <strong>of</strong> <strong>the</strong> group in question in


108 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.past times ; and when we attempt to generalise <strong>the</strong> phenomenaon a large scale, <strong>with</strong> <strong>the</strong> details fresh in our memory, we shallfind a reference to <strong>the</strong> extinct faunas <strong>of</strong> various epochs to beabsolutely necessary.<strong>The</strong> decree <strong>of</strong> our knowledge <strong>of</strong> <strong>the</strong> Palaeontology <strong>of</strong> variousparts <strong>of</strong> <strong>the</strong> world is so unequal, that it will not be advisable totreat <strong>the</strong> subject under each <strong>of</strong> our six regions.Yet some subdivisionmust be made, and it seems best to consider separately<strong>the</strong> extinct <strong>animals</strong> <strong>of</strong> <strong>the</strong> Old and <strong>of</strong> <strong>the</strong> New Worlds. Those <strong>of</strong>Europe and Asia are intimately connected, and throw light on<strong>the</strong> past changes which have led to <strong>the</strong> establishment <strong>of</strong> <strong>the</strong>three great continental Old World regions, <strong>with</strong> <strong>the</strong>ir varioussubdivisions. <strong>The</strong> wonderful extinct fauna recently discoveredin North America, <strong>with</strong> what was previously known from Southtemperate America, not only elucidates <strong>the</strong> past history <strong>of</strong> <strong>the</strong>whole continent, but also gives indications <strong>of</strong> <strong>the</strong> mutual <strong>relations</strong><strong>of</strong> <strong>the</strong> eastern and western hemispheres.<strong>The</strong> materials to be dealt <strong>with</strong> are enormous ; and it will benecessary to confine ourselves to a general summary, <strong>with</strong> fullerdetails on those points which directly bear upon our specialsubject. <strong>The</strong> objects <strong>of</strong> most interest to <strong>the</strong> pure zoologist andto <strong>the</strong> geologist—those strange forms which are far<strong>the</strong>st removedfrom any now living—are <strong>of</strong> least interest to us, since we aimat tracing <strong>the</strong> local origin or birthplace <strong>of</strong> existing genera andfamilies ; and for this purpose <strong>animals</strong> whose affinities <strong>with</strong>living forms are altoge<strong>the</strong>r doubtful, are <strong>of</strong> no value whatever.<strong>The</strong> great mass <strong>of</strong> <strong>the</strong> vertebrate fossils <strong>of</strong> <strong>the</strong> tertiary periodconsist <strong>of</strong> mammalia, and this is precisely <strong>the</strong> class which is <strong>of</strong>most value in <strong>the</strong> determination <strong>of</strong> zoological regions. <strong>The</strong><strong>animals</strong> <strong>of</strong> <strong>the</strong> secondary period, though <strong>of</strong> <strong>the</strong>highest interestto <strong>the</strong> zoologist are <strong>of</strong> little importance to us ; both because <strong>of</strong><strong>the</strong>ir very uncertain affinities for any existing groups, and alsobecause we can form no adequate notion <strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong>land and sea in those remote epochs. Our great object is totrace back, step by step, <strong>the</strong> varying <strong>distribution</strong> <strong>of</strong> <strong>the</strong> chiefforms <strong>of</strong> life ; and to deduce, wherever possible, <strong>the</strong> physicalchanges which must have accompanied or caused such changes.


;chap, vi.] MAMMALIA OF THE OLD WORLD. 109<strong>The</strong> natural division <strong>of</strong> our subject <strong>the</strong>refore is into geologicalperiods. We first go back to <strong>the</strong> Post-Pliocene period, whichincludes that <strong>of</strong> <strong>the</strong> caves and gravels <strong>of</strong> Europe containing flintimplements, and extends back to <strong>the</strong>deposit <strong>of</strong> <strong>the</strong> glacial driftin <strong>the</strong> concluding phase <strong>of</strong> <strong>the</strong> glacial epoch. Next we have<strong>the</strong> Pliocene period, divided into its later portion (<strong>the</strong> NewerPliocene) which includes <strong>the</strong> Glacial epoch <strong>of</strong> <strong>the</strong> nor<strong>the</strong>rnhemisphere ; and its earlier portion (<strong>the</strong> Older Pliocene), representedby <strong>the</strong> red and coralline crag <strong>of</strong> England, and deposits <strong>of</strong>similar age in <strong>the</strong> continent. During this earlier epoch <strong>the</strong>climate was not very dissimilar from that which now prevailsbut we next get evidence <strong>of</strong> a still earlier period, <strong>the</strong> Miocene,when a warmer climate prevailed in Europe, and <strong>the</strong> wholefauna and flora were very different. This is perhaps <strong>the</strong> mostinteresting portion <strong>of</strong> <strong>the</strong> tertiary deposits, and furnishes us<strong>with</strong> <strong>the</strong> most valuable materials for our present <strong>study</strong>.Fur<strong>the</strong>r back still we have <strong>the</strong> Eocene period, <strong>with</strong> apparentlyan almost tropical climate in Europe ; and here we find a clueto some <strong>of</strong> <strong>the</strong> most puzzling facts in <strong>the</strong> <strong>distribution</strong> <strong>of</strong> living<strong>animals</strong>. Our knowledge <strong>of</strong> this epoch is however very imperfect;and we wait for discoveries that will elucidate some<strong>of</strong> <strong>the</strong> mystery that still hangs over <strong>the</strong> origin and migrations<strong>of</strong> many important families. Beyond this <strong>the</strong>re is a great chasmin <strong>the</strong> geological record as regards land <strong>animals</strong> ;and we have togo so far back into <strong>the</strong> past, that when we again meet <strong>with</strong> mammalia,birds,and land-reptiles, <strong>the</strong>y appear under such archaicforms that <strong>the</strong>y cease to have any local or <strong>geographical</strong> significance,and we can only refer <strong>the</strong>m to wide-spread classesand orders.For <strong>the</strong> purpose <strong>of</strong> elucidating <strong>geographical</strong> <strong>distribution</strong>,<strong>the</strong>refore, it is, in <strong>the</strong> present state <strong>of</strong> our knowledge,unnecessary to go back beyond <strong>the</strong> tertiary period <strong>of</strong> geology.<strong>The</strong> remains <strong>of</strong> Mammalia being so much more numerous andimportant than those <strong>of</strong> o<strong>the</strong>r classes, we. shall at first confineourselves almost exclusively to <strong>the</strong>se. What is known <strong>of</strong> <strong>the</strong>birds, reptiles, and fishes <strong>of</strong> <strong>the</strong> tertiary epoch will be bestindicated by a brief connected sketch <strong>of</strong> <strong>the</strong>ir fossils in all parts<strong>of</strong> <strong>the</strong> globe, which we shall give in a subsequent chapter.


—110 DISTRIBUTION OF EXTINCT ANIMALS. Tpart ti.Historic Period.—In tracing back <strong>the</strong> history <strong>of</strong> <strong>the</strong> organicworld we find, even <strong>with</strong>in <strong>the</strong> limits <strong>of</strong> <strong>the</strong> historical period,that some <strong>animals</strong> have become extinct, while <strong>the</strong> <strong>distribution</strong> <strong>of</strong>o<strong>the</strong>rs has been materially changed. <strong>The</strong> Rytina <strong>of</strong> <strong>the</strong> NorthPacific, <strong>the</strong> dodo <strong>of</strong> Mauritius, and <strong>the</strong> great auk <strong>of</strong> <strong>the</strong> NorthAtlantic coasts, have been exterminated almost in our owntimes. <strong>The</strong> kitchen-middens <strong>of</strong> Denmark contain remains <strong>of</strong><strong>the</strong> capercailzie, <strong>the</strong> Bos primigenins, and <strong>the</strong> beaver.<strong>The</strong> firststill abounds far<strong>the</strong>r north, <strong>the</strong> second is extinct, and <strong>the</strong> thirdis becoming so in Europe. <strong>The</strong> great Irish elk, a huge-antlereddeer, probably existed almost down to historic times.Pleistoceneor Post-Pliocene Period.—We first meet <strong>with</strong> pro<strong>of</strong>s<strong>of</strong> important changes in <strong>the</strong> character <strong>of</strong> <strong>the</strong> European fauna,<strong>study</strong>ing <strong>the</strong> remains found in <strong>the</strong> caverns <strong>of</strong> England and France,which have recently been so well explored.in<strong>The</strong>se cave -remainsare probably all subsequent to <strong>the</strong> Glacial epoch, and <strong>the</strong>y allcome <strong>with</strong>in <strong>the</strong> period <strong>of</strong> man's occupation <strong>of</strong> <strong>the</strong> country. Yetwe find clear pro<strong>of</strong>s <strong>of</strong> two distinct kinds <strong>of</strong> change in <strong>the</strong>forms <strong>of</strong> animal life. First we have a change clearly traceableto a difference <strong>of</strong> climate. We find such arctic forms as<strong>the</strong> rein-deer, <strong>the</strong> musk-sheep, <strong>the</strong> glutton, and <strong>the</strong> lemming,<strong>with</strong> <strong>the</strong> mammoth and <strong>the</strong> woolly rhinoceros <strong>of</strong> <strong>the</strong> Siberianice-cliffs, inhabiting this country and even <strong>the</strong> south <strong>of</strong> France.This is held to be good pro<strong>of</strong> that a sub-arctic climate prevailedover all Central Europe ; and this climate, toge<strong>the</strong>r <strong>with</strong><strong>the</strong> continental condition <strong>of</strong> Britain, will sufficiently explainsuch a southward range <strong>of</strong> what are now arcticforms.But toge<strong>the</strong>r <strong>with</strong> this change we have ano<strong>the</strong>r that seems atfirst sight to be in an exactly opposite direction. We meet<strong>with</strong> numerous <strong>animals</strong> which now only inhabit Africa, or SouthEurope, or <strong>the</strong> warmer parts <strong>of</strong> Asia. Such are, large felinessome closely related to <strong>the</strong> lion (Felis spelcca), o<strong>the</strong>rs <strong>of</strong> altoge<strong>the</strong>rextinct type (Machairodus) and forming <strong>the</strong> extreme development<strong>of</strong> <strong>the</strong> feline race ;—hyaenas ; horses <strong>of</strong> two or morespecies ;and a hippopotamus. If we go a little fur<strong>the</strong>r back, to<strong>the</strong> remains furnished by <strong>the</strong> gravels and brick-earths, we stillfind <strong>the</strong> same association <strong>of</strong> forms. <strong>The</strong> reindeer, <strong>the</strong> glutton,


chap, vi.] MAMMALIA OF THE OLD WORLD. Ill<strong>the</strong> musk-sheep, and <strong>the</strong> woolly rhinoceros, are associated <strong>with</strong>several o<strong>the</strong>r species <strong>of</strong> rhinoceros and elephant; <strong>with</strong> numerouscivets, now abundant only in warm countries ; and <strong>with</strong>antelopes <strong>of</strong> several species. We also meet here <strong>with</strong> a greatextension <strong>of</strong> range <strong>of</strong> forms now limited to small areas. <strong>The</strong>Saiga antelope <strong>of</strong> Eastern Europe occurs in France, where wildsheep and goats and <strong>the</strong> chamois were <strong>the</strong>n found, toge<strong>the</strong>r <strong>with</strong>several species <strong>of</strong> deer, <strong>of</strong> bear, and <strong>of</strong> hysena. A few extinctgenera even come down to this late period, such as <strong>the</strong> greatsabre-too<strong>the</strong>d tiger, Machairodus ;Galeo<strong>the</strong>riurn, a form <strong>of</strong> Viverridse; Palceospalax, allied to <strong>the</strong> mole ; and Trogon<strong>the</strong>rium, agigantic form <strong>of</strong> beaver.We find <strong>the</strong>n, that even at so early a stage <strong>of</strong> our inquiries wemeet <strong>with</strong> a problem in <strong>distribution</strong> by no means easy to solve.How are we to explain <strong>the</strong> banishment from Europe in so shorta space <strong>of</strong> time (geologically speaking) <strong>of</strong> so many forms <strong>of</strong> lifenow characteristic <strong>of</strong> warmer countries, and this too during aperiod when <strong>the</strong> climate <strong>of</strong> Central Europe was itselfbecomingwarmer ? Such a change must almost certainly have been dueto changes <strong>of</strong> physical geography, which we shalltobe better ableunderstand when we have examined <strong>the</strong> preceding Plioceneperiod. We may here notice, however, that so far as we yetknow, this great recent change in <strong>the</strong> character <strong>of</strong> <strong>the</strong> fauna isconfined to <strong>the</strong> western part <strong>of</strong> <strong>the</strong> Palaearctic region.In cavesin <strong>the</strong> Altai Mountains examined by Pr<strong>of</strong>. Brandt, a great collection<strong>of</strong> fossil bones was discovered. <strong>The</strong>se comprised <strong>the</strong>Siberian rhinoceros and mammoth, and <strong>the</strong> cave hysena ;but all<strong>the</strong> o<strong>the</strong>rs, more than thirty distinct species, are now living inor near <strong>the</strong> same regions. We may perhaps impute this differenceto <strong>the</strong> fact that <strong>the</strong> migration <strong>of</strong> Sou<strong>the</strong>rn types intothis part <strong>of</strong> Siberia was prevented by <strong>the</strong> great mountain anddesert barrier <strong>of</strong> <strong>the</strong> Central Asiatic plateau ;whereas in Europe<strong>the</strong>re was at this time a land connection <strong>with</strong> Africa. PostpHocenedeposits and caverns in Algeria have yielded remainsresembling <strong>the</strong> more sou<strong>the</strong>rn European types <strong>of</strong> <strong>the</strong> Postplioceneperiod, but <strong>with</strong>out any admixture <strong>of</strong> Arctic forms;showing, as we might expect, that <strong>the</strong> glacial cold did not


112 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.extend so far south. We have here remains <strong>of</strong> Equus, Bos,Antilope, Hippopotamus, Elephas, Rhinoceros, Ursus, Canis,and Hycena, toge<strong>the</strong>r <strong>with</strong> Phacoclmrus, an African type <strong>of</strong>swine which has not occurred in <strong>the</strong> European deposits.It is perhaps to <strong>the</strong> earlier portion <strong>of</strong> this period that <strong>the</strong>Meryco<strong>the</strong>rium <strong>of</strong> <strong>the</strong> Siberian drift belongs. This was ananimal related to <strong>the</strong> living camel, thus supporting <strong>the</strong> view that<strong>the</strong> Camelidce are essentially denizens <strong>of</strong> <strong>the</strong> extra-tropical zone.Pliocene Period.Primates.—We here first meet <strong>with</strong> evidence <strong>of</strong> <strong>the</strong> existence<strong>of</strong> monkeys in Central Europe. Species <strong>of</strong> Macacus have leftremains not only in <strong>the</strong> Newer Pliocene <strong>of</strong> <strong>the</strong> Val d'Arno inItaly, but in beds <strong>of</strong> <strong>the</strong> same age at Grays in Essex ; whileSemnopi<strong>the</strong>cus and Cercopi<strong>the</strong>cus, genera now confined to <strong>the</strong>Oriental and Ethiopian regions respectively, have been found in<strong>the</strong> Pliocene deposits <strong>of</strong> <strong>the</strong> South <strong>of</strong> France and Italy.Carnivora.—Most <strong>of</strong> <strong>the</strong> genera which occurred in <strong>the</strong> PostPliocene are found here also, and many <strong>of</strong> <strong>the</strong> same species. Fewnew forms appear, except Hyamarctos, a large bear w T ith charactersapproaching <strong>the</strong> hyaenas, and Pristiphoca, a new form <strong>of</strong> seal,both from <strong>the</strong> Older Pliocene <strong>of</strong> France ; and Galecynus, a foxlikeanimal intermediate between Canis and Viverra, from <strong>the</strong>Pliocene <strong>of</strong> OZninghen in Switzerland.Cetacea.—Species <strong>of</strong> Balcena, Physeter, and Dclphinus occur in<strong>the</strong> Older Pliocene <strong>of</strong> England and France, and <strong>with</strong> <strong>the</strong>se <strong>the</strong>remains <strong>of</strong> many extinct forms, Balccnodon and Hoplocctus(Balsenidse) ;Belemnoziphius and Chone&iphius (Hyperoodontidae),and Hali<strong>the</strong>rium, an extinct form <strong>of</strong> <strong>the</strong> next order— Sirenia,now confined to <strong>the</strong> tropics, although <strong>the</strong> recently extinct Bytina<strong>of</strong> <strong>the</strong> N. W. Pacific show r s that it is also adapted for temperateclimates.Ungulata.—<strong>The</strong> Pliocene deposits are not very rich in thisorder. <strong>The</strong> horses (Equidai) are represented by <strong>the</strong> genus Equus ;and here we first meet <strong>with</strong> Hipparion, in which small lateraltoes appear. Both genera occur in British deposits <strong>of</strong> this age.


;chat, vi.] MAMMALIA OF THE OLD WORLD. 113A more interesting fact for us is <strong>the</strong> occurrence <strong>of</strong> <strong>the</strong> genusTapirus in <strong>the</strong> Newer Pliocene <strong>of</strong> France and in <strong>the</strong> older beds<strong>of</strong> both France and England, since this genus is now isolatedin <strong>the</strong> remotest parts <strong>of</strong> <strong>the</strong> eastern and western tropics. <strong>The</strong>genera Rhinoceros, Hippopotamus, and Sus, occur here asin <strong>the</strong>preceding epoch.We next come to <strong>the</strong> deer genus {Cervus), which appears tohave been at its maximum in this period, no less than eightspecies occurring in <strong>the</strong> Norwich Crag, and Forest-beds. Among<strong>the</strong> Bovidae, <strong>the</strong> antelopes, ox, and bison, are <strong>the</strong> only forms representedhere, as in <strong>the</strong> Post- Pliocene period. Passing on to <strong>the</strong>Proboscidea, we find three species <strong>of</strong> elephants and two <strong>of</strong> Mastodonpreserved in European beds <strong>of</strong> this period, all distinct fromthose <strong>of</strong> Post-Pliocene times.Rodentia.—In this order we find representatives <strong>of</strong> manyliving European forms ; as Cricetus (hamster), Arvicola (vole),Castor (beaver), Arctomys (marmot), Hystrix (porcupine),Lepus (hare), and Lagomys (pika) ; and a few that are extinct,<strong>the</strong> most important being Chalicomys, allied to <strong>the</strong> beaverand Issiodromys, said to come nearest to <strong>the</strong> remarkablePedetes <strong>of</strong> South Africa, both found in <strong>the</strong> Pliocene formations<strong>of</strong> France.General Conclusions as to Pliocene and Post-Pliocene Faunas <strong>of</strong>Europe.—This completes <strong>the</strong> series <strong>of</strong> fossil forms <strong>of</strong> <strong>the</strong> Pliocenedeposits <strong>of</strong> Europe. <strong>The</strong>y show us that <strong>the</strong> presence <strong>of</strong> numerouslarge carnivora and ungulates (now almost wholly tropical) in<strong>the</strong> Post- Pliocene period, was due to no exceptional or temporarycause, but was <strong>the</strong> result <strong>of</strong> a natural succession from similarraces which had inhabited <strong>the</strong> same countries for long precedingages. In order to understand <strong>the</strong> vast periods <strong>of</strong> time coveredbv <strong>the</strong> Pliocene and Post-Pliocene formations, <strong>the</strong> works <strong>of</strong> SirCharles Lyell must be studied. We shall <strong>the</strong>n come to see, that<strong>the</strong> present condition <strong>of</strong> <strong>the</strong> fauna <strong>of</strong> Europe is wholly new andexceptional. For a long succession <strong>of</strong> ages, various* forms <strong>of</strong>monkeys, hyaenas, lions, horses, hipparions, tapirs, rhinoceroses,hippopotami, elephants, mastodons, deer, and antelopes, toge<strong>the</strong>r


114 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.<strong>with</strong> almost all <strong>the</strong> forms now living, produced a rich and variedfauna such as we now see only in <strong>the</strong> open country <strong>of</strong> tropicalAfrica.During all this period we have no reason to believe that<strong>the</strong> climate or o<strong>the</strong>r physical conditions <strong>of</strong> Europe were morefavourable to <strong>the</strong> existence <strong>of</strong> <strong>the</strong>se <strong>animals</strong> than now. We mustlook upon <strong>the</strong>m, <strong>the</strong>refore, as true indigenes <strong>of</strong> <strong>the</strong>country, and<strong>the</strong>ir comparatively recent extinction or banishment as a remarkablephenomenon for which <strong>the</strong>re must have been some adequatecause. What this cause was we can only conjecture ; but itseems most probable that it was due to <strong>the</strong> combined action <strong>of</strong><strong>the</strong> Glacial period,and <strong>the</strong> subsidence <strong>of</strong> large areas <strong>of</strong> land ouceconnecting Europe <strong>with</strong> Africa. <strong>The</strong> existence, in <strong>the</strong> smallisland <strong>of</strong> Malta, <strong>of</strong> no less than three extinct species <strong>of</strong> elephant(two <strong>of</strong> very small stature), <strong>of</strong> a gigantic dormouse, an extincthippopotamus, and o<strong>the</strong>r mammalia, toge<strong>the</strong>r <strong>with</strong> <strong>the</strong> occurrence<strong>of</strong> remains <strong>of</strong> hippopotamus in <strong>the</strong> caves <strong>of</strong> Gibraltar, indicatevery clearly that during <strong>the</strong> Pliocene epoch, and perhaps duringa considerable part <strong>of</strong> <strong>the</strong> Post-Pliocene, a connection existedbetween South Europe and North Africa in at least <strong>the</strong>se twolocalities. At <strong>the</strong> same time we have every reason to believethat Britain was united to <strong>the</strong> Continent, what is now <strong>the</strong> GermanOcean constituting a great river-valley. During <strong>the</strong> height <strong>of</strong><strong>the</strong> Glacial epoch, <strong>the</strong>se large <strong>animals</strong> would probably retire intothis Mediterranean land and into North Africa, making annualmigrations' northwards during <strong>the</strong> summer.But as <strong>the</strong> connectingland sank and became narrower and narrower, <strong>the</strong> migratingherds would diminish, and at last cease altoge<strong>the</strong>r ;and when <strong>the</strong>glacial cold had passed away would be altoge<strong>the</strong>r prevented fromreturning to <strong>the</strong>ir former haunts.Miocene Peeiod.We now come to a period which was wonderfully rich in allforms <strong>of</strong> life, and <strong>of</strong> which <strong>the</strong> geological record is exceptionallycomplete. Various lacustrine, estuarine, and o<strong>the</strong>r deposits inEurope, North India, and North America, have furnished such a


chap, vi.] MAMMALIA OF THE OLD WORLD. 115vast number <strong>of</strong> remains <strong>of</strong> extinct mammalia, as to solve manyzoological problems, and to throw great light on <strong>the</strong> early <strong>distribution</strong>and centres <strong>of</strong> dispersal <strong>of</strong> various groups <strong>of</strong> <strong>animals</strong>.order to show <strong>the</strong> bearing <strong>of</strong> <strong>the</strong>se remains on our special subject,we will first give an account <strong>of</strong> <strong>the</strong> extinct fauna <strong>of</strong> Greece,<strong>of</strong> <strong>the</strong> Upper Miocene period ;Insince this, being nearest to Africaand Asia, best exhibits <strong>the</strong> <strong>relations</strong> <strong>of</strong> <strong>the</strong> old European faunato those countries. We shall <strong>the</strong>n pass to <strong>the</strong> Miocene fauna <strong>of</strong>France and Central Europe ;and conclude <strong>with</strong> <strong>the</strong> remarkableSiwalik and o<strong>the</strong>r Indian extinct faunas, which throw an additionallight on <strong>the</strong> early history <strong>of</strong> <strong>the</strong> animal lifeold-world continents.Extinct Animals <strong>of</strong> Greece.<strong>of</strong> <strong>the</strong> great<strong>The</strong>se are from <strong>the</strong> Upper Miocene deposits at Pikermi, nearA<strong>the</strong>ns, and were collected by M. Gaudry a few years ago.<strong>The</strong>y comprise ten living and eighteen extinct genera <strong>of</strong> mammalia,<strong>with</strong> a few birds and reptiles.Primates.—<strong>The</strong>se are represented by Mcsopi<strong>the</strong>cus, a genusbelieved to be intermediate between <strong>the</strong>monkeys, Semnopi<strong>the</strong>cus and Macacus.two Indian genera <strong>of</strong>Camivora.—<strong>The</strong>se were abundant. Of Fclis <strong>the</strong>re were fourspecies, ranging from <strong>the</strong> size <strong>of</strong> a cat to that <strong>of</strong> a jaguar, a largehycena, and a large weasel (Mustcla). Besides <strong>the</strong>se <strong>the</strong>re were<strong>the</strong> huge Machairodus, larger than any existing lion or tiger, and<strong>with</strong> enormously developed canine teeth ;extinct forms <strong>of</strong> Hysenidae ;genus <strong>of</strong> Viverridse but <strong>with</strong> resemblances toHymnictis and Lycarna,Thalassictis=Ictitlicrium, an extinct<strong>the</strong> hyaenas, representedby three species, some <strong>of</strong> which were larger than anyexisting Viverridse ;Promephytis, an extinct form <strong>of</strong> Mustelidae,having resemblances to <strong>the</strong> European marten, to <strong>the</strong> otters, andto <strong>the</strong> S. African Zorilla ;carnivore <strong>of</strong> <strong>the</strong> size <strong>of</strong> a small pan<strong>the</strong>r,<strong>of</strong> a cat, <strong>the</strong> molars <strong>of</strong> a dog, and <strong>the</strong> jawsa bear.and lastly, Simocyon, an extraordinarybut having <strong>the</strong> caninesshaped like those <strong>of</strong>Ungulata.—<strong>The</strong>se are numerous and very interesting. <strong>The</strong>Equidse are represented by <strong>the</strong> three-toed Hipparion, which con-


116 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.tinued to exist till <strong>the</strong> Older Pliocene period. <strong>The</strong>re are threelarge species <strong>of</strong> Rhinoceros, as well as a species <strong>of</strong> <strong>the</strong> extinctgenus Leptodon <strong>of</strong> smaller size. Eemains <strong>of</strong> a very large wildboar (Sus) were found. Very interesting is <strong>the</strong> occurrence <strong>of</strong> aspecies <strong>of</strong> giraffe (Camelopardalis) as tall as <strong>the</strong> African speciesbut more slender ; and also an extinct genus Ilcllado<strong>the</strong>rium, notquite so tall as <strong>the</strong> giraffe but much more robust, and showingsome approach to <strong>the</strong> Antilopidse in its dentition. Antelopeswere abundant, ranging from <strong>the</strong> size <strong>of</strong> <strong>the</strong> gazelle to that o<strong>the</strong> largest living species.Three or four seem referable to livinggenera, but <strong>the</strong> majority are <strong>of</strong> extinct types, and are classedin <strong>the</strong> genera Palceotragus, Palceoryx, Tragocerus, and Palceoreas ;while Dremo<strong>the</strong>rium is an ancient generalized form <strong>of</strong> Cervidceor deer.Proboscidea.—<strong>The</strong>se are represented by two species <strong>of</strong> Mastodon,and two <strong>of</strong> Dino<strong>the</strong>rium, an extraordinary extinct form•supposed to be, to some extent, intermediate between <strong>the</strong>elephants and <strong>the</strong> aquatic manatees {Sirenia)Bodentia.—This order is represented by a species <strong>of</strong> Hystrix,larger than living porcupines.Edentata.—This order, now almost confined to South America,was represented in <strong>the</strong> Miocene period by several Europeanspecies. Ancylo<strong>the</strong>rium and Macro<strong>the</strong>rium, belonging to an extinctfamily but remotely allied to <strong>the</strong> African ant-bear (Oryctcropus),occur in Greece.Birds.—Species <strong>of</strong> Phasianus and Gallus were found ; <strong>the</strong>latter especially interesting as being now confined to India.Beptiles.—<strong>The</strong>se are few and unimportant, consisting <strong>of</strong> atortoise {Testudo) and a large lizard allied to Varanus.Summary <strong>of</strong> <strong>the</strong> Miocene Fauna <strong>of</strong> Greece.—Although we cannotconsider that <strong>the</strong> preceding enumeration gives us by anymeans a complete view <strong>of</strong> <strong>the</strong> actual inhabitants <strong>of</strong> this part<strong>of</strong> Europe during <strong>the</strong> later portion <strong>of</strong> <strong>the</strong> Miocene period, weyet obtain some important information. <strong>The</strong> resemblance thatappeared in <strong>the</strong> Pliocene fauna <strong>of</strong> Europe, to that <strong>of</strong> <strong>the</strong> opencountry <strong>of</strong> tropical Africa, is now still more remarkable. We


——chap, vi.] MAMMALIA OF THE OLD WORLD. 117not only find great felines, surpassing in size and destructivepower <strong>the</strong> lions and leopards <strong>of</strong> Africa, <strong>with</strong> hyaenas <strong>of</strong> a sizeand in a variety not to be equalled now, but also huge rhinocerosesand elephants, two forms <strong>of</strong> giraffes, and a host <strong>of</strong>antelopes, which, from <strong>the</strong> sample here obtained, were probablyquite as numerous and varied as <strong>the</strong>y now are in Africa.Joined <strong>with</strong> this abundance <strong>of</strong> antelopes we have <strong>the</strong> absence<strong>of</strong> deer, which probably indicates that <strong>the</strong> country was openand somewhat <strong>of</strong> a desert character, since <strong>the</strong>re were deer ino<strong>the</strong>r parts <strong>of</strong> Europe at this epoch. <strong>The</strong> occurrence <strong>of</strong> but asingle species <strong>of</strong> monkey is also favourable to this view, sincea well-wooded country would most likely have supplied manyforms <strong>of</strong> <strong>the</strong>se <strong>animals</strong>.Miocene Fauna <strong>of</strong> Central and Western Europe."We have now to consider <strong>the</strong> Miocene fauna <strong>of</strong> Europegenerally, <strong>of</strong> which we have very full information from numerousdeposits <strong>of</strong> this age in France, Switzerland, Italy,Germany, and Hungary.Primates.—Three distinct forms <strong>of</strong> monkeys have been foundin Europe—in <strong>the</strong> South <strong>of</strong> France, in Switzerland, and Wurtemberg;one was very like Colobus or Semnopi<strong>the</strong>cus ; <strong>the</strong> o<strong>the</strong>rsPliopi<strong>the</strong>cus and Pryopi<strong>the</strong>cus—were <strong>of</strong> higher type, and belongedto <strong>the</strong> anthropomorphous apes, being nearest to <strong>the</strong> genusHylobates or gibbons.Both have occurred in <strong>the</strong> South <strong>of</strong> France,<strong>The</strong> Dryopi<strong>the</strong>cus was a very large animal (equal to <strong>the</strong> gorilla),and M. Lartet considers that in <strong>the</strong> character <strong>of</strong> itsdentition itapproached nearer to man than any <strong>of</strong> <strong>the</strong> existing anthropoidapes.Insectivora.—<strong>The</strong>se small <strong>animals</strong> are represented by numerousremains belonging to four families and a dozen genera. OfErinaceus (hedgehog) several species are found in <strong>the</strong> UpperMiocene ; and in <strong>the</strong> Lower Miocene <strong>of</strong> Auvergne two extinctgenera <strong>of</strong> <strong>the</strong> same familyAmphechinus andTetracus—havebeen discovered. Several species <strong>of</strong> Talpa (mole) occur in <strong>the</strong>Upper Miocene <strong>of</strong> France, while <strong>the</strong> extinct Dinylus isfrom Germany,and Palonospalax from <strong>the</strong> Lower Miocene <strong>of</strong> <strong>the</strong> Isle <strong>of</strong>


——;118 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.Wight. <strong>The</strong> Malayan family Tupaiidae or squirrel-shrews, isbelieved to be represented by Oxygomphus, a fossil discovered inSouth Germany (Wiesenau) by H. von Meyer. <strong>The</strong> Soricidae orshrews, are represented by several extinct genera Plesiosorex,Mysarachne and Galcospalax ; as well as by Amphisorex and Myogalestill living.to Madagascar, isEcMnogale, a genus <strong>of</strong> Centetidse now confinedsaid to occur in <strong>the</strong> Lower Miocene <strong>of</strong> Auvergne,a most interesting determination, if correct, as it would form atransition to <strong>the</strong> Solcnodon <strong>of</strong> <strong>the</strong> Antilles belonging to <strong>the</strong> samefamily ;but I am informed by Pr<strong>of</strong>. Flower that <strong>the</strong> affinities <strong>of</strong><strong>the</strong> <strong>animals</strong> described under this name are very doubtful.Carnivora.—Besides Felis and Machairodus, which extend backto <strong>the</strong> Upper Miocene, <strong>the</strong>re are two o<strong>the</strong>r genera <strong>of</strong> Felidae,Pseudmlurus in <strong>the</strong> Upper Miocene <strong>of</strong> France, and Hycenodon,which occurs in <strong>the</strong> Upper and Lower Miocene <strong>of</strong> France, namedfrom some resemblance in its teeth to <strong>the</strong> hyaenas, and consideredby some Palaeontologists to form a distinct family, Hyaenodontidae.<strong>The</strong> Viverridae, or civets, were very numerous, consisting <strong>of</strong> <strong>the</strong>living genus Viverra, and three extinct formsIdithcrium, as largeoccurring both in France and Hungary.<strong>the</strong> living genus Hyccna, and <strong>the</strong>Thalassirtis=as a pan<strong>the</strong>r, and Soricidis, a smaller form,Of Hycenidce, <strong>the</strong>re wasextinct Hycenidis, which hasoccurred in Hungary as well as in Greece. <strong>The</strong> Canidae, orwolf and fox family, were represented by Pseudocyon, near toCanis ; Hemicyon, intermediate between dogs and gluttonsand Amphicyon, <strong>of</strong> which several species occur in <strong>the</strong> Upperand Lower Miocene <strong>of</strong> France, some <strong>of</strong> <strong>the</strong>m larger than atiger. <strong>The</strong> Mustelidae, or weasels, were represented by fivegenera, <strong>the</strong> existing genera Zutra (otter) and Musiela (weasel)Potamo<strong>the</strong>rium, an extinct form <strong>of</strong> otter ;Taxodon, allied to <strong>the</strong>badger and otter; Palceomephitis in Germany, and <strong>the</strong> Promephytis(already noticed) in Greece.<strong>The</strong> bears were representedonly by Hyccnarctos, which has been noticed as occurring in<strong>the</strong> Pliocene, and first appears in <strong>the</strong> Upper Miocene <strong>of</strong> France.Seals are represented by a form resembling <strong>the</strong> AntarcticOtaria, remains <strong>of</strong> which occur in <strong>the</strong> Upper Miocene <strong>of</strong>France.


;chap, vi.] MAMMALIA OF THE OLD WORLD. 119Cetacea (whales).—<strong>The</strong>se occur frequently in<strong>the</strong> Miocene deposits,four living, and five extinct genera having been describedbut <strong>the</strong>se marine forms are not <strong>of</strong> much importance for ourpurpose.Sirenia (sea-cows).—<strong>The</strong>se are represented by two extinctgenera, Hali<strong>the</strong>rium and Trachy<strong>the</strong>rium.Several species <strong>of</strong> <strong>the</strong>former have been discovered, but <strong>the</strong> latter has occurred inFrance only, and its affinities are doubtful.Ungulata.—Horses are represented by Hipparion and Anchi<strong>the</strong>rium,<strong>the</strong> latter occurring in both Upper and Lower Mioceneand Eocene ; while Hipparion, which is more nearly allied toliving horses, first appears in <strong>the</strong> Upper Miocene and continuesin <strong>the</strong> Pliocene.Hvppo<strong>the</strong>rium, in <strong>the</strong> Upper Miocene <strong>of</strong> <strong>the</strong> Vienna basin,forms a transition to Paloplo<strong>the</strong>rium, an Eocene genus <strong>of</strong> Tapiridaeor Palseo<strong>the</strong>ridse. Tapirs, allied to living forms, occur inboth Upper and Lower Miocene. Ehinoceroses are still foundin <strong>the</strong> Upper Miocene, and here first appear <strong>the</strong> four-toed hornlessrhinoceros, Acero<strong>the</strong>rium. <strong>The</strong> Suidae (swine) are ra<strong>the</strong>rnumerous. Sus (wild boar) continued as far back as <strong>the</strong> UpperMiocene ; but now <strong>the</strong>re first appear a number <strong>of</strong> extinct formswhich have been named Hyo<strong>the</strong>rium, Palceochcerus,all <strong>of</strong> a small or moderate size ;Chceromorus,Hyopotamus, nearly as large as atapir ; and Anthraco<strong>the</strong>rium, nearly <strong>the</strong> size <strong>of</strong> a hippopotamusand, according to Dr. Leidy, <strong>the</strong> type <strong>of</strong> a distinct family.Listriodon, from <strong>the</strong> Upper Miocene <strong>of</strong> <strong>the</strong> Vienna basin, issometimes classed <strong>with</strong> <strong>the</strong> tapirs.We now come to a well-marked new family <strong>of</strong> Artiodactyleor even-toed Ungulata, <strong>the</strong> Anop>lo<strong>the</strong>riidce, which consisted <strong>of</strong>more slender long-tailed <strong>animals</strong>, allied to <strong>the</strong> swine but <strong>with</strong>indications <strong>of</strong> a transition towards <strong>the</strong> camels. <strong>The</strong> only generathat appear in <strong>the</strong> Miocene formation are, Chalico<strong>the</strong>rium, nearlyas large asa rhinoceros, <strong>of</strong> which three species have been foundin Germany and France ;and Synaphodus, known only from itssomewhat from those <strong>of</strong> <strong>the</strong> Anoplo<strong>the</strong>riumteeth, which differwhich appears earlier in <strong>the</strong> Eocene formation.Ano<strong>the</strong>r extinctfamily, Amphimericidce or Xiphodontidce, is represented by two


120 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.genera, Caino<strong>the</strong>rium and Micro<strong>the</strong>rium, in <strong>the</strong> Miocene <strong>of</strong>France. <strong>The</strong>y were <strong>of</strong> very small size, and are supposed to beintermediate between <strong>the</strong> Suidse and Tragulidse.<strong>The</strong> Camelopardalidse, or giraffes, were represented in Europein Miocene times by <strong>the</strong> gigantic Hellado<strong>the</strong>rium, which hasbeen found in <strong>the</strong> south <strong>of</strong> France, and in Hungary, as well asin Greece. <strong>The</strong> chevrotains (Tragulidse) are represented by<strong>the</strong> extinct genus Hyomoschus.<strong>The</strong> Cervidsedo not seem to have appeared in Europe before<strong>the</strong> Upper Miocene epoch, when <strong>the</strong>y were represented byDorca<strong>the</strong>rium and Amjihimoschus, allied to Moschus, and alsotrue Cervus, as well as by small allied forms, Drano<strong>the</strong>rium,Amphitragalus (in <strong>the</strong> Lower Miocene), Micromeryx, Palceomeryx,and Dicroccrus.<strong>The</strong> Bovidse, or hollow-horned ruminants, were not wellrepresented in Central Europe in Miocene times. <strong>The</strong>re wereno sheep, goats, or oxen, and only a few antelopes <strong>of</strong> <strong>the</strong> genusTragocerus, and one allied to Hippotragus ; and <strong>the</strong>se all livedin <strong>the</strong> Upper Miocene period, as did <strong>the</strong> more numerousforms <strong>of</strong> Greece.Proboscidea.—<strong>The</strong> true elephants do not extend back to <strong>the</strong>Miocene period, but <strong>the</strong>y are represented by <strong>the</strong> Mastodons,which had less complex teeth. <strong>The</strong>se first appear in <strong>the</strong> UpperMiocene <strong>of</strong> Europe, five species being known from France,Germany, Switzerland, and Greece. Dino<strong>the</strong>rium, alreadynoticed as occurring in Greece, extended also to Germany andFrance, where remains <strong>of</strong> three species have been found.Bodentia.—A considerable number <strong>of</strong> generic forms <strong>of</strong> thisorder have been obtained from <strong>the</strong> Miocene strata. <strong>The</strong> principalgenera are Cricetodon, allied to <strong>the</strong> hamsters, numerous inboth <strong>the</strong> Upper and Lower Miocene period <strong>of</strong> France ; Myoxus(<strong>the</strong> dormice) in France, and an allied genus, Brachymys, in Germany.<strong>The</strong> beavers were represented by <strong>the</strong> still living genusCastor, and <strong>the</strong> extinct Steneojiber in France. <strong>The</strong> squirrels by<strong>the</strong> existing Scuirus and Sperrnophilus ; and by extinct forms,Lithomys and Aulacodon, in Germany, <strong>the</strong> latter resembling <strong>the</strong>African genus Autocodes. <strong>The</strong> hares, by Lagomys and anby


chap, vi.] MAMMALIA OF THE OLD WORLD. 121extinct form Titanomys. Besides <strong>the</strong>se, remains referred to <strong>the</strong>South American genera, Cavia (cavy) and Basyprocta (agouti),have been found, <strong>the</strong> former in <strong>the</strong> Upper Miocene <strong>of</strong> Switzerland,<strong>the</strong> latter in <strong>the</strong> Lower Miocene <strong>of</strong> Auvergne.Palceomys,allied to <strong>the</strong> West Indian Cap>romys, has been found in <strong>the</strong> samedeposits ;as well as <strong>The</strong>ridomys, said by Gervais to be allied toAnomalurus and Echim.ys, <strong>the</strong><strong>the</strong> latter in S.America.Edentata.—<strong>The</strong>se areformer now living in W. Africa,only represented by <strong>the</strong> Macro<strong>the</strong>riumand Ancylo<strong>the</strong>rhim <strong>of</strong> <strong>the</strong> Grecian deposits, <strong>the</strong> former occurringalso in France and Germany in Upper Miocene strata.Marsupials.—<strong>The</strong>se consist <strong>of</strong> numerous species related to<strong>the</strong> opossums {Didclphys), but separated by Gervais under <strong>the</strong>name Pera<strong>the</strong>rium. <strong>The</strong>y occur in both Upper and LowerMiocene beds.Upper Miocene Deposits<strong>of</strong> <strong>the</strong> Shvalik Hills and o<strong>the</strong>r Localitiesin N. W. India.<strong>The</strong>se remarkable fresh-water deposits form a range <strong>of</strong> hillsat <strong>the</strong> foot <strong>of</strong> <strong>the</strong> Himalayas, a little south <strong>of</strong> Simla. <strong>The</strong>ywere investigated for many years by Sir P. Cautley and Dr.Falconer, and add greatly to our knowledge <strong>of</strong> <strong>the</strong> early fauna<strong>of</strong> <strong>the</strong> Old World continent.Primates.— Remains <strong>of</strong> <strong>the</strong> genera Semnopi<strong>the</strong>cus and Macacuswere found, <strong>with</strong> o<strong>the</strong>r forms <strong>of</strong> intermediate character; andsome teeth indicated <strong>animals</strong> allied to <strong>the</strong> orang-utan <strong>of</strong> Borneo,and <strong>of</strong>similar size.Camivora.—<strong>The</strong>se consisted <strong>of</strong> species <strong>of</strong> Felis and Machairodus<strong>of</strong> large size ;Hyarna, Canis, Mcllivora, and an allied genusUrsitaxus; Ursus, in <strong>the</strong> deposits <strong>of</strong> <strong>the</strong> Nerbudda valley (<strong>of</strong>Pliocene age) ;Hymnarctos as large as <strong>the</strong> cave bear ;Amphicyon<strong>of</strong> <strong>the</strong> size <strong>of</strong> a polar bear (in <strong>the</strong> deposits <strong>of</strong> <strong>the</strong> Indusvalley, west <strong>of</strong> Cashmere) ; Lutra, and an extinct allied genusEnhydrion.Ungulata.— <strong>The</strong>se are very numerous, and constitute <strong>the</strong> mostimportant feature <strong>of</strong> this ancient fauna. Horses are representedby a species <strong>of</strong> Equus from <strong>the</strong> Siwalik Hills and <strong>the</strong> TrawaddyVol. L—10


122 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.deposits in Burmah, and by two o<strong>the</strong>rs from <strong>the</strong> Pliocene <strong>of</strong> <strong>the</strong>Nerbudda Valley ;while Hippo<strong>the</strong>rium—a slender, antelope-likeanimal, found in <strong>the</strong> Siwalik Hills and in Europe—is supposedto form a transition from <strong>the</strong> Equidse to <strong>the</strong> Tapiridae. <strong>The</strong>selatter are found in <strong>the</strong> Upper Indus deposits, where <strong>the</strong>re is aspecies <strong>of</strong> Tapirus, and one <strong>of</strong> an extinct genus Antelo<strong>the</strong>rium.Of Rhinoceros, five extinct species have been found— in <strong>the</strong>Siwalik Hills, in Perim Island, and one at an elevation <strong>of</strong> 16,000feet in <strong>the</strong> deserts <strong>of</strong> Thibet.Hippopotamus occurs in <strong>the</strong> Pliocene<strong>of</strong> <strong>the</strong>Nerbudda, and is represented in <strong>the</strong> older Miocenedeposits by Hcxapirotodon, <strong>of</strong> which three species have beenfound in various parts <strong>of</strong> India. Ano<strong>the</strong>r remarkable genus,Merycopotamus, connects Hippopotamus <strong>with</strong> Anthraco<strong>the</strong>rium;one <strong>of</strong> <strong>the</strong> extinct European forms allied to <strong>the</strong> swine. <strong>The</strong>selast arerepresented by several large species <strong>of</strong> Sus, and by <strong>the</strong>extinct European genus Chcero<strong>the</strong>rium.<strong>The</strong> extinct Anoplo<strong>the</strong>ridae are represented by a species <strong>of</strong><strong>the</strong> European genus Chalico<strong>the</strong>rium, larger than a horse.An extinct camel, larger than <strong>the</strong> living species, was foundin <strong>the</strong> Siwalik Hills.Three species <strong>of</strong> deer (Cervus) have been found in <strong>the</strong>Siwaliks, and one in <strong>the</strong> Nerbudda deposits.A large and a small species <strong>of</strong> giraffe (Camclopardalis) werefound in <strong>the</strong> Siwalik Hills and at Perim Island.<strong>The</strong> Bovidse are represented by numerous species <strong>of</strong> Bos, andby <strong>the</strong> extinct genera Hemibos and AmpMhos. <strong>The</strong>re are alsothree species <strong>of</strong> antelopes, one <strong>of</strong> which is allied to <strong>the</strong> AfricanAlccphalus.We now come toan extraordinary group <strong>of</strong> extinct <strong>animals</strong>,probably forming a new family intermediate between <strong>the</strong>antelope and <strong>the</strong> giraffe. <strong>The</strong> Siva<strong>the</strong>rium was an enormousfour-horned ruminant, larger than a rhinoceros. It had a shorttrunk like a tapir, <strong>the</strong> lower horns on <strong>the</strong> forehead were simple,<strong>the</strong> upper pair palmated. <strong>The</strong> Brama<strong>the</strong>rium, an allied formfrom Perim Island, showed somewhat more affinity for <strong>the</strong>giraffe.Proboscidca.—No less than seven species <strong>of</strong> elephants and four


chap, vi.] MAMMALIA OF THE OLD WORLD. 123<strong>of</strong> mastodons ranged over India, <strong>the</strong>ir remains being found in all<strong>the</strong> deposits from <strong>the</strong> Siwalik Hills to Burmah. A large Dino<strong>the</strong>riumhas also been found at Perim Island.Beptiles.—Many remains <strong>of</strong> birds were found, but <strong>the</strong>se havenot been determined. Reptiles were numerous and interesting,<strong>the</strong> most remarkable being <strong>the</strong> huge tortoise, Golossochclys, whoseshell was twelve feet long and head and neck eight feet more.O<strong>the</strong>r small tortoises <strong>of</strong> <strong>the</strong> genera Testudo, Emys, Trionyxand Emydida were found, <strong>the</strong> Emys being a living species.<strong>The</strong>re were three extinct and one living species <strong>of</strong> crocodile,and one <strong>of</strong> <strong>the</strong>m was larger than any now living. <strong>The</strong> onlyo<strong>the</strong>r reptile <strong>of</strong> importance was a large lizard <strong>of</strong> <strong>the</strong> genusVaranus.General Observations on <strong>the</strong> Miocene faunas <strong>of</strong> Europe andAsia.— Comparing <strong>the</strong> three faunas <strong>of</strong> approximately <strong>the</strong> sameperiod, and allowing for <strong>the</strong> necessarily imperfect record <strong>of</strong>each, we find a wonderful similarity <strong>of</strong> general type over <strong>the</strong>enormous areabetween France on <strong>the</strong> west and <strong>the</strong> Irawaddyriver in Burmah on <strong>the</strong> east. We may even extend our comparisonto Nor<strong>the</strong>rn China, where remains <strong>of</strong> Hywna, Tapir,Rhinoceros, Chalico<strong>the</strong>rium, and Elephas, have been recentlyfound, closely resembling those from <strong>the</strong> Miocene or Pliocenedeposits <strong>of</strong> Europe or India, and showing that <strong>the</strong> Palasarcticregion had <strong>the</strong>n <strong>the</strong> same great extent from west to eastthat it has now. Of about forty genera comprised in <strong>the</strong>Indian Miocene fauna, no less than twenty-seven inhabitedCentral and Western Europe during <strong>the</strong> same epoch. <strong>The</strong> IndianMiocene fossils are much what we should expect as <strong>the</strong> forerunners<strong>of</strong> <strong>the</strong> existing fauna, <strong>the</strong> giraffes and hippopotamibeing <strong>the</strong> only additions from <strong>the</strong> present Ethiopian fauna.<strong>The</strong> numerous forms <strong>of</strong> <strong>the</strong> restricted bovine type, show that<strong>the</strong>se probably originated in India ; while <strong>the</strong> monkeys appearto be altoge<strong>the</strong>r <strong>of</strong> Oriental types.In Europe, however, we meet <strong>with</strong> a totally different assemblage<strong>of</strong> <strong>animals</strong> from those that form <strong>the</strong> existing fauna. Wefind apes and monkeys, many large Felidse, numerous civets


124 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.and hyaenas, tapirs, rhinoceros, hippopotamus, elephants, giraffes,and antelopes, such as now characterise <strong>the</strong> tropics <strong>of</strong> Africaand Asia. Along <strong>with</strong> <strong>the</strong>se we meet <strong>with</strong> less familiar types,showing <strong>relations</strong> <strong>with</strong> <strong>the</strong> Centetidae <strong>of</strong> Madagascar, <strong>the</strong>Tupaiidaa <strong>of</strong> <strong>the</strong> Malay Islands, <strong>the</strong> Cajpromys, <strong>of</strong> <strong>the</strong> West Indies,and <strong>the</strong> Echimys <strong>of</strong> South America. And besides all <strong>the</strong>seliving types we have a host <strong>of</strong> extinct forms,—ten or twelvegenera allied to swine ; nine genera <strong>of</strong> tapir-like <strong>animals</strong> ; four<strong>of</strong> horses ; nine <strong>of</strong> wolves ; <strong>with</strong> many distinct forms <strong>of</strong> <strong>the</strong>Long-extinct families<strong>of</strong> Anoplo<strong>the</strong>ridae, Xiphodontidae, and <strong>the</strong>edentate Macro<strong>the</strong>ridae. It is almost certain that during <strong>the</strong>Miocene period Europe was not only far richer than it is nowin <strong>the</strong> higher forms <strong>of</strong> life,but not improbably richer than anypart <strong>of</strong> <strong>the</strong> globe now is, not excepting tropical Africa andtropical Asia.Eocene Period.<strong>The</strong> deposits <strong>of</strong> Eocene age are less numerous, and spreadover a far more limited area, than those <strong>of</strong> <strong>the</strong> Miocene period,and only restricted portions <strong>of</strong> <strong>the</strong>m furnish any remains <strong>of</strong>land <strong>animals</strong>.Our knowledge <strong>of</strong> <strong>the</strong> Eocene mammalian faunais <strong>the</strong>refore very imperfect and will not occupy us long, asmost <strong>of</strong> <strong>the</strong> new types it furnishes are <strong>of</strong> more interest to <strong>the</strong>zoologist than to <strong>the</strong> student <strong>of</strong> <strong>distribution</strong>. Some <strong>of</strong> <strong>the</strong>Eocene mammalia <strong>of</strong> Europe are, however, <strong>of</strong> interest in comparison<strong>with</strong> those <strong>of</strong> North America <strong>of</strong> <strong>the</strong> same age ;whileo<strong>the</strong>rs show that ancestral types <strong>of</strong> groups now confined toAustralia or to South America, <strong>the</strong>n inhabited Europe.Primates.— <strong>The</strong> only undoubted Eocene examples <strong>of</strong> thisorder, are <strong>the</strong> Ccenojn<strong>the</strong>cus lemuroides from <strong>the</strong> Jura, which haspoints <strong>of</strong> resemblance to <strong>the</strong> South American marmosets andhowlers, and also to <strong>the</strong> Lemuridae ; and a cranium recently discoveredin <strong>the</strong> Department <strong>of</strong> Lot (S.W. France), undoubtedlybelono-ino- to <strong>the</strong> Lemuridae, and which most resembles that <strong>of</strong><strong>the</strong> West African " Potto " (Pcrodidicus). This discovery hasled to ano<strong>the</strong>r for it is now believed that remains formerly


chap, vi.] MAMMALIA OF THE OLD WORLD. 125referred to <strong>the</strong> Anoplo<strong>the</strong>ridae (Adapts and Aphelo<strong>the</strong>riumfrom <strong>the</strong> Upper Eocene <strong>of</strong> Paris) were also Lemurs. Someremains from <strong>the</strong> Lower Eocene <strong>of</strong> Suffolk were at first supposedto be allied to Macacas, but were subsequently referred to <strong>the</strong>Ungulate, Hyracothcrium. <strong>The</strong>re is still, however, some doubtas to its true affinities.Chiroptera.—In <strong>the</strong> Upper Eocene <strong>of</strong> Paris remains <strong>of</strong> batshave been found, so closely resembling living forms as to bereferred to <strong>the</strong> genus Vesyertilio.Carnivora.—<strong>The</strong> only feline remains, are those <strong>of</strong> Hycenodonin <strong>the</strong> Upper Eocene <strong>of</strong> Hampshire, and Pterodon, an allied formfrom beds <strong>of</strong> <strong>the</strong> same age in France ; <strong>with</strong> yElurogale, foundin <strong>the</strong> South <strong>of</strong> France in deposits <strong>of</strong> phosphate <strong>of</strong> lime <strong>of</strong>uncertain age, but probably belonging to this period.Viverridae(civets) are represented by two genera, Tylodon, <strong>the</strong> size <strong>of</strong> aglutton from <strong>the</strong> Upper Eocene, and Palceonyctis, allied toViverra, from <strong>the</strong> Middle Eocene <strong>of</strong> France. <strong>The</strong> Canidse(wolves and foxes) appear to have been <strong>the</strong> most ancient <strong>of</strong><strong>the</strong> existing types <strong>of</strong> Carnivora, five genera being representedby Eocene remains. Of <strong>the</strong>se, Galethylax and Cyo<strong>the</strong>rium weresmall, and <strong>with</strong> <strong>the</strong> existing genus Canis are found in <strong>the</strong> UpperEocene <strong>of</strong> France. Arctocyon, about <strong>the</strong> size <strong>of</strong> a wolf, isa very ancient and generalised form <strong>of</strong> carnivore which cannot be placed in any existing family. It is found in <strong>the</strong> LoweiEocene <strong>of</strong> France, and is thus <strong>the</strong> oldest known member <strong>of</strong><strong>the</strong> Carnivora.Ungidata.—<strong>The</strong>se are more numerous. Equidae (horses) arerepresented by <strong>the</strong> Miocene Anchithcrium in <strong>the</strong> Lower, and by amore ancient form, Anchilophus, in <strong>the</strong> Middle Eocene <strong>of</strong> France.Tapiridse and Paheo<strong>the</strong>ridae were very numerous.Palceo<strong>the</strong>riumand <strong>the</strong> allied genus Pcdoplo<strong>the</strong>rium, were abundant in France andEngland in Upper Eocene times. <strong>The</strong>y somewhat resembled<strong>the</strong> tapir, <strong>with</strong> affinities for <strong>the</strong> horse and rhinoceros. A newgenus, Cadurco<strong>the</strong>rium, allied to <strong>the</strong> rhinoceros and equallylarge, has been found in <strong>the</strong> same deposits <strong>of</strong> phosphate <strong>of</strong> limeas <strong>the</strong> lemur and JElurogale. In <strong>the</strong> Middle Eocene <strong>of</strong> bothEngland and France are found Lophiodon allied to <strong>the</strong> tapir,


126 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.but in some <strong>of</strong> <strong>the</strong> species reaching a larger size ; Propalaio<strong>the</strong>riumand Pachynolophus <strong>of</strong> smaller size and having affinitiesfor <strong>the</strong> o<strong>the</strong>r genera named ; and Plagiolopfius, a small, slenderanimal which Pr<strong>of</strong>essor Huxley thinks may have been a directancestor <strong>of</strong> <strong>the</strong> horse. In <strong>the</strong> Lower Eocene we meet <strong>with</strong>Coryphodon, much larger than <strong>the</strong> tapir,and armed <strong>with</strong> largecanine teeth ; Pliohphus, a generalised type, allied to <strong>the</strong> tapirand horse ; and Hyraco<strong>the</strong>rium, a small animal from <strong>the</strong> LowerEocene <strong>of</strong> England, remotely allied to <strong>the</strong> tapir.Among <strong>the</strong> Artiodactyla, or even-toed ungulates, <strong>the</strong> swineare represented by several extinct genera, <strong>of</strong> moderate or smalls ize— Aco<strong>the</strong>rium,Chceropotamus, Cebochmrus and Dichobune, allfrom <strong>the</strong> Upper and <strong>the</strong> last also from <strong>the</strong> Middle Eocene <strong>of</strong>France ; but Eutelodon, from <strong>the</strong> phosphate <strong>of</strong> lime deposits islarge. <strong>The</strong> Dichobune was <strong>the</strong> most generalised type, presenting<strong>the</strong> characters <strong>of</strong> many <strong>of</strong> <strong>the</strong> o<strong>the</strong>r genera combined,and was believed by Dr. Falconer to approach <strong>the</strong> musk-deer.<strong>The</strong> Caino<strong>the</strong>rium <strong>of</strong> <strong>the</strong> Miocene also occurs here, and an alliedgenus Plesiom&ryx from <strong>the</strong> same deposits as Euteledon.<strong>The</strong> Eocene Anoplo<strong>the</strong>ridae were numerous. <strong>The</strong> Anoplo<strong>the</strong>riumwas a two-toed, long-tailed Pachyderm, ranging from <strong>the</strong>size <strong>of</strong> a hog to that <strong>of</strong> an ass ; <strong>the</strong> allied Eury<strong>the</strong>rium wasfour-toed ; and <strong>the</strong>re are one or two o<strong>the</strong>rs <strong>of</strong> doubtful affinity.All are from <strong>the</strong> Upper Eocene <strong>of</strong> France and England.Rodentia.—Eemains referred to <strong>the</strong> genera Myoxus (dormouse)and Sciurus (squirrel) have been found in <strong>the</strong> Upper Eocene<strong>of</strong> France ; as well as Plesiarctomys, an extinct genus between<strong>the</strong> marmots and squirrels. <strong>The</strong> Miocene <strong>The</strong>ridomys is als<strong>of</strong>ound here.Marsupials.—<strong>The</strong> Didclphys (opossum) <strong>of</strong> Cuvier, now referredto an extinct genus Pera<strong>the</strong>rium, is<strong>of</strong> France and England.found in <strong>the</strong> Upper EoceneGeneral Considerations on <strong>the</strong> Extinct Mammalian Fauna <strong>of</strong>Europe.— It is a curious fact that no family, and hardly a genus,<strong>of</strong> European mammalia occurs in <strong>the</strong> Pliocene deposits, <strong>with</strong>outextending back also into those <strong>of</strong> Miocene age.<strong>The</strong>re are, how-


;chap, vi.] MAMMALIA OF THE OLD WORLD. 127ever, a few groups which seem to be late developments or recentimportations into <strong>the</strong> Palaearctic region, as <strong>the</strong>y occur only inPost-Pliocene deposits. <strong>The</strong> most important <strong>of</strong> <strong>the</strong>se are <strong>the</strong>badger, glutton, elk, reindeer, chamois, goat, and sheep, whichonly occur in caves and o<strong>the</strong>r deposits <strong>of</strong> Post-Pliocene age.Camels only occur in <strong>the</strong> Post-Pliocene <strong>of</strong> Siberia (Meryco<strong>the</strong>rium),although a true Camelus <strong>of</strong> large size appears to have inhabitedsome part <strong>of</strong> Central Asia in <strong>the</strong> Upper Miocene period, beingfound in <strong>the</strong> Siwalik beds. <strong>The</strong> only exclusively Pliocenegenera in Europe are Ursus, Equits, Hippopotamus, Bos,Elephas,Arvicola, Trogon<strong>the</strong>rium, Arctomys, Hystrix and Lepus ; but <strong>of</strong><strong>the</strong>se Equus, Hippopotamus, Bos, and Elephas are found in <strong>the</strong>Miocene deposits <strong>of</strong> India. Owing, no doubt, in part to <strong>the</strong>superior productiveness <strong>of</strong> <strong>the</strong> various Miocene beds, largenumbers <strong>of</strong> groups appear to have <strong>the</strong>ir origin or earliest appearancehere. Such are Insectivora, Felidae, Hyaenidae, Mustelidae,Ursus, Equidae, Tapirus, Ehinocerotidae, Hippopotamidae, Anthraco<strong>the</strong>ridae(extinct), Sus, Camelopardidae, Tragulidae, Cervidae,Bovidse, Elephantidae, and Edentata.Groups which go back to <strong>the</strong> Eocene period, are, Primatesallied to South American monkeys, as well as some <strong>of</strong> <strong>the</strong>Lernuridae ; bats <strong>of</strong> <strong>the</strong> living genus Vespertilio ; Hyaenodontida?,an ancestral form <strong>of</strong> Carnivore ; Viverridae ; Canidae (to <strong>the</strong>Upper Eocene), and <strong>the</strong> ancestral Arctocyonidae to <strong>the</strong> LowerEocene ; Hijamarctos, an ancestral type <strong>of</strong> bears and hyaenasAnchi<strong>the</strong>ridae, ancestral horses, to <strong>the</strong> Middle Eocene ; Palaeo<strong>the</strong>ridae,comprising numerous generalised forms, ancestors <strong>of</strong> <strong>the</strong>rhinoceros, horse, and tapir ; Suidoe, <strong>with</strong> numerous generalisedforms, to <strong>the</strong> Middle Eocene ;Anoplo<strong>the</strong>ridae and Xiphodontidae,ancestral families <strong>of</strong> even-toed Ungulates, connecting <strong>the</strong> ruminants<strong>with</strong> <strong>the</strong> swine ; and lastly, several groups <strong>of</strong> Eodents,and a Marsupial, in <strong>the</strong> Upper Eocene. We thus find all<strong>the</strong> great types <strong>of</strong> Mammalia well developed in <strong>the</strong> earliestportion <strong>of</strong> <strong>the</strong> tertiary period ; and <strong>the</strong> occurrence <strong>of</strong> Quadrumana, <strong>of</strong> <strong>the</strong> highly specialized bats {Vespertilio), <strong>of</strong> variousforms <strong>of</strong> Carnivora, and <strong>of</strong>Ungulates, clearly differentiated into<strong>the</strong> odd and even-toed series, associated <strong>with</strong> such lower forms a&


128 DISTRIBUTION OF EXTINCT ANIMALS. [part iiLemurs and Marsupials— proves,that we have here hardly madean approach towards <strong>the</strong> epoch when <strong>the</strong> mammalian type itselfbegan to diverge into its various modifications. Some <strong>of</strong> <strong>the</strong>Carnivora and Ungulates do, indeed, exhibit a less specialisedstructure than later forms; yet so far back as <strong>the</strong> UpperMiocene <strong>the</strong> most specialised <strong>of</strong> all carnivora, <strong>the</strong> great sabretoo<strong>the</strong>dMachairodus, makes itsappearance.<strong>The</strong> Miocene is, for our special <strong>study</strong>, <strong>the</strong> most valuable andinstructive <strong>of</strong> <strong>the</strong> Tertiary periods, both on account <strong>of</strong> itssuperior richness, and because we here meet <strong>with</strong> many typesnow confined to separate regions. Such facts as <strong>the</strong> occurrencein Europe during this period <strong>of</strong> hippopotami, tapirs, giraffes,Tragulida?, Edentata, and Marsupials— will assist us in solvingmany <strong>of</strong> <strong>the</strong> problems we shall meet <strong>with</strong> in reviewing <strong>the</strong>actual <strong>distribution</strong> <strong>of</strong> living forms <strong>of</strong> those groups. Still morelight will, however, be thrown on <strong>the</strong> subject by <strong>the</strong> fossil forms<strong>of</strong> <strong>the</strong> American continent, which we will now proceed toexamine.


CHAPTEE VII.EXTINCT MAMMALIA OF THE NEW WORLD.<strong>The</strong> discoveries <strong>of</strong> very richdeposits <strong>of</strong> mammalian remains invarious parts <strong>of</strong> <strong>the</strong> United States have thrown great light on<strong>the</strong> <strong>relations</strong> <strong>of</strong> <strong>the</strong> faunas <strong>of</strong> verv distant regions. NorthAmerica now makes a near approach to Europe in <strong>the</strong> numberand variety <strong>of</strong> itsextinct mammalia, and in no part <strong>of</strong> <strong>the</strong> worldhave such perfect specimens been discovered.In what are called<strong>the</strong> " Mauvaises terres " <strong>of</strong> Nebraska (<strong>the</strong> dried-up mud <strong>of</strong> anancient lake), thousands <strong>of</strong> entire crania and some almost entireskeletons <strong>of</strong> ancient <strong>animals</strong> have been found, <strong>the</strong>ir teeth absolutelyperfect, and altoge<strong>the</strong>r more resembling <strong>the</strong> preparations<strong>of</strong> <strong>the</strong> anatomist, than time-worn fossils such as we are accustomedto see in <strong>the</strong> museums <strong>of</strong> Europe. O<strong>the</strong>r deposits havebeen discovered in Oregon, California, Virginia, South Carolina,Texas, and Utah, ranging over all <strong>the</strong> Tertiary epochs, fromPost-Pliocene to Eocene, and furnishing a remarkable picture<strong>of</strong> <strong>the</strong> numerous strange mammalia which inhabited <strong>the</strong> ancientNorth American continent.North America— Post-Pliocene Period.Insedivora.—<strong>The</strong> only indications <strong>of</strong> this order yet discovered,consists <strong>of</strong> a single tooth <strong>of</strong> some insectivorous animal foundin Illinois, but which cannot be referred to any known group.Carnivora.—<strong>The</strong>se are fairly represented. Two species <strong>of</strong>Felis as large as a lion ; <strong>the</strong> equally large extinct Trucifelis,found only in Texas ; four species <strong>of</strong> Canis, some <strong>of</strong> <strong>the</strong>m larger


;130 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.than wolves ; two species <strong>of</strong> Galera, a genus now confined to <strong>the</strong>Neotropical region; two bears, and an extinct genus, Arctodusan extinct species <strong>of</strong> racoon (Procyon), and an allied extinctgenus, Myxophagus— show, that at a very recent period NorthAmerica was better supplied <strong>with</strong> Carnivora than it is now.Remains <strong>of</strong> <strong>the</strong> walrus (Trichechus) have also been found as farsouth as Virginia.Cetacea.—Three species <strong>of</strong> dolphins belonging to existinggenera, have been found in <strong>the</strong> Eastern States ;<strong>of</strong>Manatus, or sea-cow, in Florida and South Carolina.and two speciesUngulata.—Six extinct horses {Equus), and one Hipparion<strong>the</strong> living South American tapir, and a larger extinct species ; aDicotyles, or peccary, and an allied genus, Platygonus ;<strong>of</strong> <strong>the</strong> South American llamas (Auchenia), and onea species<strong>of</strong> a kind <strong>of</strong>camel, Procamclus ; two extinct bisons ; a sheep, and two musksheep(Ovibos) ;<strong>with</strong> three living and one extinct deer (Cervus),show an important increase in its Herbivora.Proboseidea.—Two elephants and two mastodons, added to thisremarkable assemblage <strong>of</strong> large vegetable-feeding quadrupeds.Bodentia.—<strong>The</strong>se consist mainly <strong>of</strong> genera and species stillliving in North America ;<strong>the</strong> only important exceptions being aspecies <strong>of</strong> <strong>the</strong> South American capybara (Hydrochcerus) in SouthCarolina ;and Prao<strong>the</strong>rium, an extinct form <strong>of</strong> hare, found in abone cave in Pennsylvania.Pde?itata.—Rere we meet <strong>with</strong> a wonderful assemblage, <strong>of</strong> sixspecies belonging to four extinct genera, mostly <strong>of</strong> gigantic size.A species <strong>of</strong> Mega<strong>the</strong>rium, three <strong>of</strong> Megalonyx, and one <strong>of</strong>Mylodon—huge terrestrial sloths as large as <strong>the</strong> rhinocerosor even as <strong>the</strong> largest elephants—ranged over <strong>the</strong> Sou<strong>the</strong>rnStates to Pennsylvania, <strong>the</strong> latter {Mylodon) going as far as <strong>the</strong>great lakes and Oregon. Ano<strong>the</strong>r form, Ereptodon, has beenfound in <strong>the</strong> Mississippi Valley.Marsupialia.—<strong>The</strong> living American genus <strong>of</strong> opossums, Didelphys,has been found in deposits <strong>of</strong> this age in South Carolina.Remarks on <strong>the</strong>Post-Pliocene fauna <strong>of</strong> .North America.—<strong>The</strong>assemblage <strong>of</strong> <strong>animals</strong> proved, by <strong>the</strong>se remains, to have


chap. VII.] MAMMALIA OF THE NEW WORLD. 131inhabited North America at a comparatively recent epoch, ismost remarkable. In Europe, we found a striking changein <strong>the</strong> fauna at <strong>the</strong> same period; but that consisted almostwholly in <strong>the</strong> presence <strong>of</strong> <strong>animals</strong> now inhabiting countriesimmediately to <strong>the</strong> north or south. Here we have <strong>the</strong> appearance<strong>of</strong> two new assemblages <strong>of</strong> <strong>animals</strong>, <strong>the</strong> one now confinedto <strong>the</strong> Old World—horses, camels, and elephants ; <strong>the</strong>o<strong>the</strong>r exclusively <strong>of</strong> South American type—Hamas, tapirs,capybaras, Galera, and gigantic Edentata. <strong>The</strong> age <strong>of</strong> <strong>the</strong>various deposits in which <strong>the</strong>se remains are found is somewhatuncertain, and probably extends over a considerable period <strong>of</strong>time, inclusive <strong>of</strong> <strong>the</strong> Glacial epoch, and perhaps both anteriorand subsequent to it. We have here, as in Europe, <strong>the</strong> presenceand apparent co-existence in <strong>the</strong> same area, <strong>of</strong> Arctic andSou<strong>the</strong>rn forms—<strong>the</strong> walrus and <strong>the</strong> manatee— <strong>the</strong> musksheepand <strong>the</strong> gigantic sloths. Unfortunately, as we shall see,<strong>the</strong> immediately preceding Pliocene deposits<strong>of</strong> North Americaare ra<strong>the</strong>r poor in organic remains ;yet it can hardly be owingto <strong>the</strong> imperfection <strong>of</strong> <strong>the</strong> record <strong>of</strong> this period,<strong>the</strong> South American types above numerated occursthat not one <strong>of</strong><strong>the</strong>re, whilea considerable number <strong>of</strong> Old World forms are represented.Nei<strong>the</strong>r in <strong>the</strong> preceding wonderfully rich Miocene or Eoceneperiods, does any one <strong>of</strong> <strong>the</strong>se forms occur ;or, <strong>with</strong> <strong>the</strong> exception<strong>of</strong> Moro<strong>the</strong>rium, from Pliocene deposits west <strong>of</strong> <strong>the</strong> RockyMountains, any apparent ancestor <strong>of</strong> <strong>the</strong>m ! Wehave hereunmistakable evidence <strong>of</strong> an extensive immigration from Southinto North America, not very long before <strong>the</strong> beginning <strong>of</strong> <strong>the</strong>Glacial epoch. It was an immigration <strong>of</strong> types altoge<strong>the</strong>r newto <strong>the</strong> country, which spread over all <strong>the</strong> sou<strong>the</strong>rn and centralportions <strong>of</strong> it, and established <strong>the</strong>mselves sufficiently to leaveabundance <strong>of</strong> remains in <strong>the</strong> few detached localities where <strong>the</strong>yhave been discovered. How such large yet defenceless <strong>animals</strong>as tapirs and great terrestrial sloths, could have made <strong>the</strong>ir wayinto a country abounding in large felines equal in size anddestructiveness to <strong>the</strong> lion and <strong>the</strong> tiger, <strong>with</strong> numerous wolvesand bears <strong>of</strong> <strong>the</strong> largest size, is a great mystery. But it isnever<strong>the</strong>less certain that <strong>the</strong>y did so ;and <strong>the</strong> fact that no such


132 DISTRIBUTION OF EXTINCT ANIMALS. [part 11.migration had occurred for countless preceding ages, proves thatsome great barrier to <strong>the</strong> entrance <strong>of</strong> terrestrial mammaliawhich had previously existed, must for a time have been removed.We must defer fur<strong>the</strong>r discussion <strong>of</strong> this subject tillwe have examined <strong>the</strong> <strong>relations</strong> <strong>of</strong> <strong>the</strong> existing faunas <strong>of</strong> Northand South America.Tertiary Period.When we get to remains <strong>of</strong> <strong>the</strong>Tertiary age, especially those<strong>of</strong> <strong>the</strong> Miocene and Eocene epochs, we meet <strong>with</strong> so manyinteresting and connected types, and such curious <strong>relations</strong> <strong>with</strong>living forms in Europe, that it will be clearer to trace <strong>the</strong>history <strong>of</strong>each order and family throughout <strong>the</strong> Tertiary period,instead <strong>of</strong> considering each <strong>of</strong> <strong>the</strong> subdivisions <strong>of</strong> that periodseparately.It will be well however first to note <strong>the</strong>few American Post-Pliocene or living genera that are found in <strong>the</strong> Pliocene beds.<strong>The</strong>se consist <strong>of</strong> several species <strong>of</strong> Cants, from <strong>the</strong> size <strong>of</strong> a foxto that <strong>of</strong> a large wolf; a Felis as large as a tiger; an Otter(Lutra) ;several species <strong>of</strong> Hipparion ; a peccary (Dicotylcs) ;a deer (Cervus) ;several species <strong>of</strong> Procamelus ; a mastodon;an elephant; and a beaver (Castor). It thus appears that out<strong>of</strong> nearly forty genera found in <strong>the</strong> Post-Pliocene deposits, onlyten are found in <strong>the</strong> preceding Pliocene period. About twelveadditional genera, however, appear <strong>the</strong>re, as we shall see inaoinff over <strong>the</strong> various orders.Primates.—Among <strong>the</strong> vast number <strong>of</strong> extinct mammaliadiscovered in <strong>the</strong> Tertiary deposits <strong>of</strong> North America, noexample <strong>of</strong> this order had been recognized up to 1872, when<strong>the</strong> discovery <strong>of</strong> more perfect remains showed, that a number<strong>of</strong> small <strong>animals</strong> <strong>of</strong> obscure affinities from <strong>the</strong> Lower Eocene<strong>of</strong> Wyoming, were really allied to <strong>the</strong> lemurs and perhapsalso to <strong>the</strong> marmosets, <strong>the</strong> lowest form <strong>of</strong> American monkeys,but having a larger number <strong>of</strong> teeth than ei<strong>the</strong>r. A number <strong>of</strong>o<strong>the</strong>r remains <strong>of</strong> small <strong>animals</strong> from <strong>the</strong> same formation, previouslysupposed to be allied to <strong>the</strong> Ungulata, are now shown to


chap, vii.] MAMMALIA OF THE NEW WORLD 133belong to <strong>the</strong> Primates ;so that no less than twelve genera <strong>of</strong> <strong>the</strong>se<strong>animals</strong> are recognized by Mr. Marsh, who classes<strong>the</strong>m in tw<strong>of</strong>amilies—Limno<strong>the</strong>ridse, comprising <strong>the</strong> genera Limno<strong>the</strong>rium,(which had larger canine teeth), Thinolestes, Telmatolestes,Mesacodon,Bathrodon, and Antiacodon <strong>of</strong> Marsh, <strong>with</strong> Notharctos,Hipposyus, Microsyops, and Palceacodon previously described byLeidy ;—and Leniuravidse, consisting<strong>of</strong> <strong>the</strong> genera Lemuravus(Marsh) and Hyopsodus (Leidy). <strong>The</strong> <strong>animals</strong> <strong>of</strong> <strong>the</strong> latterfamily were most allied to existing lemurs, but were a moregeneralized form, Lemuravus having forty-four teeth, <strong>the</strong> greatestnumber known in <strong>the</strong> order.<strong>The</strong>se numerous forms ranged from<strong>the</strong> size <strong>of</strong> a small squirrel to that <strong>of</strong> a racoon. It is especiallyinteresting to find <strong>the</strong>se peculiar lemuroid forms in America,just when a lemur has been discovered <strong>of</strong> about <strong>the</strong> same agein Europe ; and as <strong>the</strong> American forms are said to show anaffinity <strong>with</strong> <strong>the</strong> South American marmosets, while <strong>the</strong> Europeananimal is most allied to a West African group, we haveevidently not yet got back far enough to find <strong>the</strong> primevalor ancestral type from which all <strong>the</strong> Primates sprang.About <strong>the</strong> same time, in <strong>the</strong> succeeding Miocene formation,true monkeys were discovered. Mr. Marsh describes Laopi<strong>the</strong>cusas an animal nearly <strong>the</strong> size <strong>of</strong> <strong>the</strong> largest South Americanmonkeys, and allied both to <strong>the</strong> Cebidse and <strong>the</strong> Eocene Limno<strong>the</strong>ridse.Mr. Cope has described Meno<strong>the</strong>rium from <strong>the</strong> Miocene<strong>of</strong> Colorado, as a lemuroid animal, <strong>the</strong> size <strong>of</strong> a cat, andperhaps allied to Limno<strong>the</strong>rium. More Miocene remains will,no doubt, be discovered, by which we shall be enabled to trace<strong>the</strong> origin <strong>of</strong> some <strong>of</strong> <strong>the</strong> existing forms <strong>of</strong> South Americanmonkeys ;and perhaps help to decide <strong>the</strong> question (now in disputeamong anatomists) whe<strong>the</strong>r <strong>the</strong> lemurs are really Primates,or form an altoge<strong>the</strong>r distinct and isolated order <strong>of</strong> mammalia.Insectivora.—This order is represented by comparatively fewforms in <strong>the</strong> tertiary beds, and <strong>the</strong>se areall very different fromexisting types. In <strong>the</strong> Upper Miocene <strong>of</strong> Dakota are foundremains indicating two extinct genera, Lepictis and Lctops. In<strong>the</strong> Miocene <strong>of</strong> Colorado, Pr<strong>of</strong>essor Cope has recently discoveredfour new genera, Isacis ; allied to <strong>the</strong> preceding, but as large as a


—;134 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.Mephitis or skunk ; Herpeto<strong>the</strong>rium, near <strong>the</strong> moles ; Embasis,more allied to <strong>the</strong> shrews ; and Dommina, <strong>of</strong> uncertain affinities.Two o<strong>the</strong>rs have been found in <strong>the</strong> Eocene <strong>of</strong> Wyoming;Amo?nys,having some resemblance to hedgehogs and to <strong>the</strong> Eastern Tupaia;and Washakms, <strong>of</strong> doubtful affinities.Far back in <strong>the</strong> Triassic coal <strong>of</strong> North Carolina has been found<strong>the</strong> jaw <strong>of</strong> a small mammal (Dromothcrium), <strong>the</strong>teeth <strong>of</strong> whichsomewhat resemble those <strong>of</strong> <strong>the</strong> Australian Myrmccobius, andmay belong ei<strong>the</strong>r to <strong>the</strong> Insectivora or Marsupials ; if indeed, atthat early period <strong>the</strong>se orders were differentiated.Carnivora.—<strong>The</strong> most ancient forms <strong>of</strong> this order are someremains found in <strong>the</strong> Middle Eocene <strong>of</strong> Wyoming, and o<strong>the</strong>rsrecently described by Pr<strong>of</strong>essor Cope (1875) from <strong>the</strong> Eocene <strong>of</strong>New Mexico, <strong>of</strong> perhaps earlier date. <strong>The</strong> former consist <strong>of</strong> threegenera, Patri<strong>of</strong>elis,Vintacyon, and Sinopa,—<strong>animals</strong> <strong>of</strong> large sizebut which cannot be classed in any existing family ; and twoo<strong>the</strong>rs, Mcsonyx and Synoplo<strong>the</strong>riurn,allied to Hyccnodon.believed by Mr. Cope to be<strong>The</strong> latter consist <strong>of</strong> four genera,— Oxycena,consisting <strong>of</strong> several species, some as large as a jaguar, wasallied to Hyccnodon and Pterodon ; Pachyccna, allied to <strong>the</strong> lastPrototomus, allied to Amphicyon and <strong>the</strong> Viverridas ; and Limnocyon,a civet-like carnivore <strong>with</strong> resemblances to <strong>the</strong> Canidse.In <strong>the</strong> Miocene formations we find <strong>the</strong> Feline type well developed.<strong>The</strong> wonderful Machairodus, which in Europe lived downto Post-Pliocene times, is found in <strong>the</strong> Upper Miocene <strong>of</strong> Dakotaand perfect crania have been discovered, showing that <strong>the</strong> chinwas leng<strong>the</strong>ned downwards toreceive and protect <strong>the</strong> enormouscanines. Dinyctis was allied both to Machairodus and to <strong>the</strong>weasels.Three new genera have been lately described by Pr<strong>of</strong>essorCope from <strong>the</strong> Miocene <strong>of</strong> Colorado,Buncclurus, <strong>with</strong> characters<strong>of</strong> both cats and weasels ; Baptophilus, allied to Dinyctis ; andHoplophoncus, more allied to Machairodus. <strong>The</strong> Canidas are representedby Amphicyon, which occurs in deposits <strong>of</strong> <strong>the</strong> sameage in Europe ; and by Canis, four species <strong>of</strong> which genus arerecorded by Pr<strong>of</strong>essor Cope from <strong>the</strong> Miocene <strong>of</strong> Colorado, and italso occurs in <strong>the</strong> Pliocene. <strong>The</strong> Hyccnodon is represented bythree species in <strong>the</strong> Miocene <strong>of</strong> Dakota and Colorado.It occurs


chap, vii.] MAMMALIA OF THE NEW WORLD. 135also in <strong>the</strong> European Miocene and Upper Eocene formations, andconstitutes a distinct family Hyaenodontidse, allied,according toDr. Leidy, to wolves, cats, hyaenas and weasels. <strong>The</strong> Ursidaeare represented by only one species <strong>of</strong> an extinct genus, Leptarchus,from <strong>the</strong> Pliocene <strong>of</strong> Nebraska. From <strong>the</strong> Pliocene <strong>of</strong>Colorado, Pr<strong>of</strong>. Cope has recently described Tomarctos, as a" short-faced type <strong>of</strong> dog ;" as well as species <strong>of</strong> Cains andMartes.Ungulata.—<strong>The</strong> <strong>animals</strong> belonging to this order being usually<strong>of</strong> large size and accustomed to feed and travel in herds, areliable to wholesale destruction by floods, bogs, precipices, droughtor hunger. It is for <strong>the</strong>se reasons, probably, that <strong>the</strong>ir remainsare almost always more numerous than those <strong>of</strong> o<strong>the</strong>r orders <strong>of</strong>mammalia. In America <strong>the</strong>y are especially abundant ; and <strong>the</strong>number <strong>of</strong> new and intermediate types about whose position<strong>the</strong>re is much difference <strong>of</strong> opinion among Palaeontologists, rendersit very difficult to give a connected summary <strong>of</strong> <strong>the</strong>m <strong>with</strong>any approach to systematic accuracy.Beginning <strong>with</strong> <strong>the</strong> Perissodactyla,or odd-toed ungulates, wefind <strong>the</strong> Equine <strong>animals</strong> remarkably numerous and interesting.<strong>The</strong> true horses <strong>of</strong> <strong>the</strong> genus Equus, so abundant in <strong>the</strong> Post-Pliocene formations, are represented in <strong>the</strong> Pliocene by severalancestral forms.<strong>The</strong> most nearly allied to Equus is Pliohippus,consisting <strong>of</strong> <strong>animals</strong> about <strong>the</strong> size <strong>of</strong> an ass, <strong>with</strong> <strong>the</strong> lateraltoes not externally developed, but <strong>with</strong> some differences <strong>of</strong> dentition.Next come Protohippus and Hipparion, in which <strong>the</strong>lateral toes are developed but are small and functionless. <strong>The</strong>nwe have <strong>the</strong> allied genera, Anchippus, Merychippus, and Hyohippus,related to <strong>the</strong> European Hippo<strong>the</strong>rium, which were all stillsmaller <strong>animals</strong>, Protohippus being only 2| feet high. In <strong>the</strong>older deposits we come to a series <strong>of</strong> forms, still unmistakablyequine, but <strong>with</strong> three or more toes used for locomotion and <strong>with</strong>numerous differentiations in form, proportions, and dentition.<strong>The</strong>se constitute <strong>the</strong> family Anchi<strong>the</strong>ridae. In <strong>the</strong> MioceDe wehave <strong>the</strong> genera Anchi<strong>the</strong>rium (found also in <strong>the</strong> EuropeanMiocene), Miohippus and Mesohippus, all <strong>with</strong> three toes on eachfoot, and about <strong>the</strong> size <strong>of</strong> a sheep or large goat.In <strong>the</strong> Eocene <strong>of</strong>


—13G DISTRIBUTION OF EXTINCT ANIMALS. [part ii.Utah and Wyoming, we get a step fur<strong>the</strong>r back, severalspecieshaving been discovered about <strong>the</strong> size <strong>of</strong> a fox <strong>with</strong> four toes infront and three behind. <strong>The</strong>se form <strong>the</strong> genus Orohippus, andare <strong>the</strong> oldest ancestral horse known. Pr<strong>of</strong>. Marsh points out <strong>the</strong>remarkably perfect series <strong>of</strong> forms in America, which, beginning<strong>with</strong> this minute ancient type, is gradually modified by gainingincreased size, increased speed by concentration <strong>of</strong> <strong>the</strong> limb-bones,elongation <strong>of</strong> <strong>the</strong> head and neck, <strong>the</strong> canine teeth decreased insize, <strong>the</strong> molars becoming longer and being coated <strong>with</strong> cementtill we at last come to <strong>animals</strong> hardly distinguishable, specifically,from <strong>the</strong> living horse.Allied to <strong>the</strong>se, are a series <strong>of</strong> forms showing a transition to <strong>the</strong>tapirs, and to <strong>the</strong> Palwo<strong>the</strong>rium <strong>of</strong> <strong>the</strong> European Eocene. In <strong>the</strong>Pliocene we have Parahippus ; in <strong>the</strong> Miocene Lophiodon, foundin <strong>the</strong> same formation and in <strong>the</strong> Eocene <strong>of</strong> Europe, and alliedto <strong>the</strong> tapir ; and in <strong>the</strong> Eocene, Palwosyops, as large as a rhinoceros,which had large canines and was allied to <strong>the</strong> tapir andPalcco<strong>the</strong>rium ; Limnohyus, forming <strong>the</strong> type <strong>of</strong> a family Limnohyidse,which included <strong>the</strong> last genus and some o<strong>the</strong>rs mentionedfur<strong>the</strong>r on ; and Hyrachyus, allied to Lophiodon, and to Hyracodonan extinct form <strong>of</strong> rhinoceros. Besides <strong>the</strong>se we have Lophio<strong>the</strong>rium(also from <strong>the</strong> Eocene <strong>of</strong> Europe) ; Diplacodon allied toLimnohyus, hut <strong>with</strong> affinities to modern Perissodactyla and nearlyas large as a rhinoceros ; and Colonoceras, also belonging to <strong>the</strong>Limnohyidse, an animal which was <strong>the</strong> size <strong>of</strong> a sheep, and haddivergent protuberances or horns on its nose. A remarkablegenus, JBathmodon, lately described by Pr<strong>of</strong>essor Cope, and <strong>of</strong>which five species have been found in <strong>the</strong> Eocene <strong>of</strong> New Mexicoand Wyoming, is believed to form <strong>the</strong> type <strong>of</strong> a new family,having some affinity to Palwosyops and to <strong>the</strong> extinct Bronto<strong>the</strong>ridse.It had large canine tusks but no horns.<strong>The</strong> Bhinocerotidae are represented in America by <strong>the</strong> genusRhinoceros in <strong>the</strong> Pliocene and Miocene, and by Acera<strong>the</strong>riumand Hyracodon in <strong>the</strong> Miocene. Both <strong>the</strong> latter were hornless,and Hyracodon was allied to <strong>the</strong> Eocene Hyrachyus, one <strong>of</strong> <strong>the</strong>Lophiodontidse. In <strong>the</strong> Eocene and Miocene deposits <strong>of</strong> Utah,and Oregon, several remarkable extinct rhinoceroses have been


chap, vii.] MAMMALIA OF THE NEW WORLD. 137recently discovered, forming <strong>the</strong> genus Diccra<strong>the</strong>rium.<strong>The</strong>se hada pair <strong>of</strong> nasal horns placed side by side on <strong>the</strong> snout, not behindeach o<strong>the</strong>r as in existing two-horned rhinoceroses, <strong>the</strong> rest <strong>of</strong><strong>the</strong>ir skeleton resembling <strong>the</strong> hornless Acera<strong>the</strong>rium. <strong>The</strong>y were<strong>of</strong> ra<strong>the</strong>r small size.Next to <strong>the</strong>se extinct rhinoceroses come <strong>the</strong> Bronto<strong>the</strong>ridse,an extraordinary family <strong>of</strong> large mammalia, some <strong>of</strong> whichexceeded in bulk <strong>the</strong> largest living rhinoceros. <strong>The</strong>y had fourtoes to <strong>the</strong> front and three to <strong>the</strong> hindfeet, <strong>with</strong> a pair <strong>of</strong> largedivergent horns on <strong>the</strong> front <strong>of</strong> <strong>the</strong> head, in both sexes. Pr<strong>of</strong>essorMarsh and Dr. Leidy have described four genera, Bronto<strong>the</strong>rium,Titano<strong>the</strong>rium, Megacerops, and Anisacodon, distinguished bypeculiarities <strong>of</strong> dentition. Though most nearly allied to <strong>the</strong>rhinoceroses, <strong>the</strong>y show some affinity for <strong>the</strong> gigantic Dinocerata<strong>of</strong> <strong>the</strong> Eocene to be noticed fur<strong>the</strong>r on. Pr<strong>of</strong>essor Copehas since described ano<strong>the</strong>r genus, Symborodon, from <strong>the</strong> Miocene<strong>of</strong> Colorado, <strong>with</strong> no less than seven species, one nearly<strong>the</strong> size <strong>of</strong> an elephant. He thinks <strong>the</strong>y had a short tapir-likeproboscis. <strong>The</strong> species differ greatly in <strong>the</strong> form <strong>of</strong> <strong>the</strong> craniumand development <strong>of</strong> <strong>the</strong> horn -bearing processes.We commence <strong>the</strong> Artiodactyla, or even-toed Ungulates, <strong>with</strong><strong>the</strong> hog tribe. <strong>The</strong>se are represented by species <strong>of</strong> peccaries,{Dicotyles) from <strong>the</strong> Pliocene <strong>of</strong> Nebraska and Oregon ; and byan allied form Thinolujus, very like Dicotyles, but having anadditional premolar tooth and a much smaller brain-cavity.From <strong>the</strong> Miocene are three allied genera, Nanohyus, Leptochcerus,and Perchce-rus. Pr<strong>of</strong>essor Cope, however, thinks Leptochcerusmay be Lemuroid, and allied to Meno<strong>the</strong>rium. <strong>The</strong>Anthraco<strong>the</strong>ridae, a family which connects <strong>the</strong> Hippopotamidseand Euminants, and which occurs in <strong>the</strong> Miocene <strong>of</strong> Europeand India, are represented in America by <strong>the</strong> genus Hyopotamusfrom <strong>the</strong> Miocene <strong>of</strong> Dakota, and Elo<strong>the</strong>rium from <strong>the</strong> Miocene <strong>of</strong>Oregon and <strong>the</strong> Eocene <strong>of</strong> Wyoming; <strong>the</strong> latter genus being sometimesclassed <strong>with</strong> <strong>the</strong> preceding family, and lately placed byPr<strong>of</strong>essor Marsh, in <strong>the</strong> new order, Tillodontia. Pr<strong>of</strong>essor Copehas since described three o<strong>the</strong>r genera from <strong>the</strong> Eocene <strong>of</strong>Vol. L—11New


—138 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.Mexico : Menisco<strong>the</strong>rium, having resemblances to Palccosyops,Hyopotamus, and <strong>the</strong> Limno<strong>the</strong>ridie ;Phenacodus, <strong>the</strong> size <strong>of</strong> ahog, <strong>of</strong> doubtful position, but perhaps near Mo<strong>the</strong>rium ; andAchmnodon, as large as a cow, but more hog-like than <strong>the</strong> preceding.Ano<strong>the</strong>r new genus from <strong>the</strong> Miocene <strong>of</strong> ColoradoPclonax—issaid by Pr<strong>of</strong>essor Cope to come between Elo<strong>the</strong>riurnand Hippopotamus.<strong>The</strong> Camelidse are very abundant, and form one <strong>of</strong> <strong>the</strong> moststriking features <strong>of</strong> <strong>the</strong> ancient fauna <strong>of</strong> America. Procamelus,Homocamelus, and Megalomeryx, are extinct genera found inPliocene formation ; <strong>the</strong> first very closely allied to <strong>the</strong> OldWorld camel, <strong>the</strong> last smaller and more sheep-like. In <strong>the</strong>Miocene two o<strong>the</strong>r genera<strong>the</strong>occur, Poebro<strong>the</strong>rium and Protomeryx,<strong>the</strong> former allied to both <strong>the</strong> camel and <strong>the</strong> llama.Deer are represented by a single species <strong>of</strong> Ccrvus in <strong>the</strong>Pliocene, while two extinct genera, Leptomeryx and Mcrycodus,are found in <strong>the</strong> Miocene deposits, <strong>the</strong> latter indicating a transitionbetween camels and deer. Two o<strong>the</strong>r genera, Hypisodusand Hypertracjulus, <strong>of</strong> very small size, are said by Pr<strong>of</strong>essorCope to be allied to <strong>the</strong> Tragulidse and to Leptomeryx.<strong>The</strong> Bovidse, or hollow-horned ruminants, are only representedin <strong>the</strong> Newer Pliocene by a single species <strong>of</strong> an extinct genus,Casoryx, said toWe now come toan exclusively American family, <strong>the</strong> Oreodontidw,which consisted <strong>of</strong>be intermediate between antelopes and deer.small <strong>animals</strong> termed by Dr. Leidy,"ruminating hogs," and which had some general structuralresemblances to deer and camels. <strong>The</strong>y abounded in NorthAmerica during <strong>the</strong> Pliocene, and especially during <strong>the</strong> Mioceneepoch, no less than six genera and twenty species having beendiscovered. Mcrychus contains <strong>the</strong> Pliocene forms ; whileOreodon, Eporeodon, Merychochcerus, Zeptauchcnia, and Agriochcerusare Miocene. <strong>The</strong> last genus extends back into <strong>the</strong>Eocene period, and shows affinity to <strong>the</strong> European Anoplo<strong>the</strong>ridtB<strong>of</strong> <strong>the</strong> same epoch.Proboscidea.—<strong>The</strong> Elephantidse are only represented inAmerica by one species <strong>of</strong> Mastodon and one <strong>of</strong> Elephas, in<strong>the</strong> Newer Pliocene deposits. In <strong>the</strong> Older Pliocene, Miocene,


;chap, vii.] MAMMALIA OF THE NEW WORLD. 139and Upper Eocene, no remains <strong>of</strong> this order have been foundand in 1869, Dr. Leidy remarked on <strong>the</strong> small average size <strong>of</strong><strong>the</strong> extinct North American mammalia, which were almost allsmaller than <strong>the</strong>ir living analogues. Since <strong>the</strong>n, however, wonderfuldiscoveries have been made in deposits<strong>of</strong> Middle Eoceneage in Wyoming and Colorado, <strong>of</strong> a group <strong>of</strong> huge <strong>animals</strong> notonly rivalling <strong>the</strong> elephants in size, but <strong>of</strong> so remarkable andpeculiar a structure as to require <strong>the</strong> formation <strong>of</strong> a new order <strong>of</strong>mammals—Dinocerata—for <strong>the</strong>ir reception.This order consists <strong>of</strong> <strong>animals</strong> <strong>with</strong> generalisedUngulate andProboscidean affinities. <strong>The</strong> lower jaw resembles that <strong>of</strong> <strong>the</strong>hippopotamus ;<strong>the</strong> posterior ;<strong>the</strong>y had five toes on <strong>the</strong> anterior feet and four onthree pairs <strong>of</strong> horns, <strong>the</strong> first pair on <strong>the</strong> top <strong>of</strong> <strong>the</strong>head, large and perhaps palmated, <strong>the</strong> second pair above <strong>the</strong> eyes,while <strong>the</strong> third and smallest stood out sideways on <strong>the</strong> snout.<strong>The</strong>y had enormous upper canines, <strong>of</strong> which <strong>the</strong> roots entered<strong>the</strong> middle horn cores, no upper incisors, and small molars.Pr<strong>of</strong>essor Marsh believes that <strong>the</strong>y had no trunk. <strong>The</strong> remainsdiscovered indicate four genera, Dinoceras (3 sp.), Tinoceras(2 sp.), Uinta<strong>the</strong>rium (1 sp.), and Eobasilcus (2 sp.). Many o<strong>the</strong>rnames have been given to fragments <strong>of</strong> <strong>the</strong>se <strong>animals</strong>, and eventhose here given may not be all distinct.Ano<strong>the</strong>r new order, Tillodontia, recently established by Pr<strong>of</strong>essorMarsh, is perhaps yet more remarkable in a zoologicalpoint <strong>of</strong> view, since it combines <strong>the</strong> characters <strong>of</strong> Carnivora,Ungulata, and Rodents.<strong>The</strong>se <strong>animals</strong> have been formed intotwo families, Tillo<strong>the</strong>ridse and Stylinodontidae; and three genera,Tillo<strong>the</strong>rium, Anchippodus, and Stylinodontia.Eocene <strong>of</strong> Wyoming and New Jersey.added Elo<strong>the</strong>riumAll are from <strong>the</strong>Perhaps to <strong>the</strong>se must befrom <strong>the</strong> Miocene <strong>of</strong> Dakota, <strong>the</strong> o<strong>the</strong>r formsbeing all Eocene. <strong>The</strong>y were mostly <strong>animals</strong> <strong>of</strong> small size,between that <strong>of</strong> <strong>the</strong> capybara and tapir. <strong>The</strong> skull resembled inform that <strong>of</strong> a bear ;<strong>the</strong> molar teeth were <strong>of</strong> Ungulate type, and<strong>the</strong> incisors like those <strong>of</strong> a Rodent ; but <strong>the</strong> skeleton was morethat <strong>of</strong> <strong>the</strong> Ursidpe, <strong>the</strong> feet being plantigrade. Pr<strong>of</strong>essor Copehas since described three new genera from <strong>the</strong> Eocene <strong>of</strong>New Mexico, Ectoganus, Calamodon, and Estlwnyr, comprising


———140 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.seven species allied to Tillo<strong>the</strong>rium and Anchippodus, and havingalso <strong>relations</strong>, as Pr<strong>of</strong>essor Cope believes, <strong>with</strong> <strong>the</strong> South AmericanToxodontidse.Rodentia.—This order is represented in <strong>the</strong> Pliocene by abeaver, a porcupine, and an American mouse (Hespervmys), allextinct species <strong>of</strong> living genera, <strong>the</strong> Hystrix being an Old Worldtype ; and Pr<strong>of</strong>essor Cope has recently described Panolax, anew genus <strong>of</strong> hares from <strong>the</strong> Pliocene <strong>of</strong> New Mexico. <strong>The</strong>Miocene deposits have furnished an extinct genus allied to <strong>the</strong>haresPalccolagus ; one <strong>of</strong> <strong>the</strong> squirrel familya small extinct form <strong>of</strong> beavermouseIschyromys ;Palceocastor ; and an extinctEumys. <strong>The</strong> Eocene strata <strong>of</strong> Wyoming have latelyfurnished two extinct forms <strong>of</strong> squirrel, Paramys and Sciuravus ;and ano<strong>the</strong>r <strong>of</strong> <strong>the</strong> Muridse (or mouse family), Mysops.Cetacea.—Numerous remains <strong>of</strong> dolphins and whales, belongingto no less than twelve genera, mostly extinct, have beenfound in <strong>the</strong> Miocene deposits <strong>of</strong> <strong>the</strong> Atlantic and Gulf States,from New Jersey to South Carolina and Louisiana ;while sevengenera <strong>of</strong> <strong>the</strong> extinct family, Zeuglodontidse, have been found inMiocene and Eocene beds <strong>of</strong> <strong>the</strong> same districts. Some remainsassociated <strong>with</strong> <strong>the</strong>se are doubtfully referred to <strong>the</strong> Seal family(Phocidae) among <strong>the</strong> Carnivora.Edentata.—Tillquite recently no remains <strong>of</strong> this order haveoccurred in any North American deposits below <strong>the</strong> Post- Pliocene;but in 1874 Pr<strong>of</strong>. Marsh described some remains allied toMegalonyx and Mylodon, from <strong>the</strong> Pliocene beds <strong>of</strong> Californiaand Idaho, and forming a new genus, Moro<strong>the</strong>rinm. As <strong>the</strong>seremains have only occurred to <strong>the</strong> west <strong>of</strong> <strong>the</strong> Eocky Mountains,and in Pliocene deposits whose exact age is not ascertained, <strong>the</strong>yhardly affect <strong>the</strong> remarkable absence <strong>of</strong> this group from <strong>the</strong>whole <strong>of</strong> <strong>the</strong> exceedingly rich Tertiary deposits in all o<strong>the</strong>r parts<strong>of</strong> North America.General Relations <strong>of</strong> <strong>the</strong> extinct Tertiary Mammalia <strong>of</strong> NorthAmerica and Europe.—Having now given a sketch <strong>of</strong> <strong>the</strong> extinctMammalia which inhabited Europe and North Americaduring <strong>the</strong> Tertiary period, we are enabled by comparing <strong>the</strong>m,


chap, vii.] MAMMALIA OF THE NEW WORLD. HIto ascertain <strong>the</strong>ir <strong>relations</strong> to each o<strong>the</strong>r, and to see how tar<strong>the</strong>y elucidate <strong>the</strong> problem <strong>of</strong> <strong>the</strong> birth-place and subsequentmigrations <strong>of</strong> <strong>the</strong> several families and genera. We have alreadypointed out <strong>the</strong> remarkable features <strong>of</strong> <strong>the</strong> Quaternary (or Post-Pliocene) fauna <strong>of</strong> North America, and now proceed to discussthat <strong>of</strong> <strong>the</strong> various Tertiary periods, which isclosely connected<strong>with</strong> <strong>the</strong> extinct fauna <strong>of</strong> Europe.<strong>The</strong> Tertiary Mammalia <strong>of</strong> North America at present describedbelong to from eighty to one hundred genera, whilethose <strong>of</strong> Europe are nearly double that number ;yet onlyeighteen genera are common to <strong>the</strong> two faunas, and <strong>of</strong> <strong>the</strong>seeight are living and belong chiefly to <strong>the</strong> Pliocene period.Taking first, <strong>the</strong> genera which in America do not go back beyond<strong>the</strong> Pliocene period (ten in number), we find that eight <strong>of</strong> <strong>the</strong>min Europe go back to <strong>the</strong> Upper Miocene. <strong>The</strong>se are Felis,Pseudmlurus, Hipparion, Cervus, Mastodon, Elephas (in India),Castor and Hystrix ; while ano<strong>the</strong>r, Canis, goes back to <strong>the</strong>Upper Eocene and <strong>the</strong> tenth, Equus, confined to <strong>the</strong> newerPliocene or perhaps to <strong>the</strong> Post-Pliocene in America, extendsback to <strong>the</strong> older Pliocene in Europe. Of <strong>the</strong> seven Europeangenera which are confined to <strong>the</strong> Miocene period in America,three, Hymnodon, Anchi<strong>the</strong>rium, and Lophiodon go back to <strong>the</strong>Eocene in Europe; three o<strong>the</strong>rs, Machairodus, Rhinoceros, andAcera<strong>the</strong>rium, are also <strong>of</strong> Miocene age in Europe; Amphicyon goesback to <strong>the</strong> Lower Miocene <strong>of</strong> Europe. Lophio<strong>the</strong>rium belongsto <strong>the</strong> Eocene <strong>of</strong> both countries.If we turn now to families instead <strong>of</strong> genera, we find that <strong>the</strong>same general rule prevails.Mustelidae (weasels), Ursidae (bears),true Equidae (horses), and Bovidae (oxen &c), go no fur<strong>the</strong>r backin America than <strong>the</strong> Pliocene, while <strong>the</strong>y all go back to <strong>the</strong>Miocene in Europe.Suidae (swine) and Anoplo<strong>the</strong>ridae (extinct)are found in <strong>the</strong> American Miocene and in <strong>the</strong> European Eocene.Anchi<strong>the</strong>ridae (extinct) reach <strong>the</strong> Upper Eocene in America,while in Europe <strong>the</strong>y range through Upper, Middle, and LowerEocene. Cervidae (deer) alone are Miocene in both countries.<strong>The</strong>re remain two families in which America has <strong>the</strong> preeminence.Camelidae (camels) were •wonderfully developed in


142 DISTRIBUTION OF EXTINCT ANIMALS. [part 11.<strong>the</strong> American Pliocene and Miocene periods, abounding ingenera and species ; whereas in Europe <strong>the</strong> group only exists in<strong>the</strong> Post-Pliocene or Lower Pliocene, <strong>with</strong> one Upper Miocenespecies <strong>of</strong> Camelus in N. India. <strong>The</strong> Anthraco<strong>the</strong>ridae (extinct),found only in <strong>the</strong> Upper Miocene <strong>of</strong> France and India, reacheven <strong>the</strong> Lower Eocene in America.<strong>The</strong>se facts may be due, in part, to a want <strong>of</strong> strictco-ordinationbetween <strong>the</strong> Tertiary deposits <strong>of</strong> Europe and North America,—in part to <strong>the</strong> imperfection <strong>of</strong> <strong>the</strong> record in <strong>the</strong> latter country.Yet it does not seem probable that <strong>the</strong>y are altoge<strong>the</strong>r due to<strong>the</strong>se causes, because <strong>the</strong> Miocene beds, which are by far <strong>the</strong>best know T n in America as in Europe, exhibit deficiencies <strong>of</strong> <strong>the</strong>same kind as <strong>the</strong> less known Eocene deposits. <strong>The</strong> fossil fauna<strong>of</strong> both countries is so rich, that we can hardly impute great andwell marked differences to imperfect knowledge; yet we findsuch important families as <strong>the</strong> Civets, Hyamas, Giraffes, andHippopotami absent from America, <strong>with</strong> <strong>the</strong> Weasels, and Antelopesalmost so ; while America possesses almost all <strong>the</strong> Camelidaa,two peculiar orders, Dinocerata and Tillodontia, and four remarkablypeculiar families, Limno<strong>the</strong>ridse, Lemuravidse, Oreodontidseand Bronto<strong>the</strong>ridae. If <strong>the</strong>n <strong>the</strong> facts at present known representapproximately <strong>the</strong> real time-<strong>relations</strong> <strong>of</strong> <strong>the</strong> groups in questionon <strong>the</strong> two continents, <strong>the</strong>y render it probable that weasels, bears,true horses, swine, oxen, sheep and antelopes, originated on <strong>the</strong>Old World continent, and were transmitted to America duringsome part <strong>of</strong> <strong>the</strong> Miocene period ;while camels originated in <strong>the</strong>New World, and somewhere about <strong>the</strong> same time passed over toEurope. Of <strong>the</strong> extinct families common to <strong>the</strong> two hemispheres,<strong>the</strong> Anthraco<strong>the</strong>ridse alone seem to have had anAmerican origin. Of <strong>the</strong> genera common to <strong>the</strong> two countries,almost all seem to have had a European origin, <strong>the</strong> only genera<strong>of</strong> equal date being <strong>the</strong> two rhinoceroses and three Anchi<strong>the</strong>ridse;but if <strong>the</strong> Bronto<strong>the</strong>ridse are allied to <strong>the</strong> Ehinocerotidae,<strong>the</strong>se latter may have originated in America, althoughnow anexclusively Old World type. <strong>The</strong>se conclusions are not improbablewhen we consider <strong>the</strong> much greater size <strong>of</strong> <strong>the</strong> OldWorld continents, extending far into <strong>the</strong> tropics and probably


chap, vii.] MAMMALIA OF THE NEW WORLD. 143always more or less united to <strong>the</strong> tropical areas ; while <strong>the</strong>evidence <strong>of</strong> <strong>the</strong> extinct mammalia <strong>the</strong>mselves shows, that SouthAmerica has been for <strong>the</strong> most part isolated from <strong>the</strong> nor<strong>the</strong>rncontinent, and did not take part in <strong>the</strong> development <strong>of</strong> itscharacteristic Tertiary fauna.Before speculating fur<strong>the</strong>r on this subject, it will be well tolay before our readers a summary <strong>of</strong> South American palaeontology,after which we shall be in a better position to drawcorrect inferences from <strong>the</strong> whole body <strong>of</strong> <strong>the</strong> evidence.South America.Unfortunately, our knowledge <strong>of</strong> <strong>the</strong> interesting fossil fauna<strong>of</strong> this continent, is almost wholly confined to <strong>the</strong> Post-Plioceneand Pliocene periods. A few remains have been discovered indeposits believed to be <strong>of</strong> Eocene age, but nothing whateverrepresenting <strong>the</strong> vast intervening period, so rich in peculiarforms <strong>of</strong> animal life both in North America and Europe.Fauna <strong>of</strong> <strong>the</strong> Brazilian caves.—What we know <strong>of</strong> <strong>the</strong> Post-Pliocene period is chiefly due to <strong>the</strong> long-continued researches<strong>of</strong> Dr. Lund in <strong>the</strong> caves <strong>of</strong> Central Brazil, mostly situated ina district near <strong>the</strong> head waters <strong>of</strong> <strong>the</strong> San Francisco riverin <strong>the</strong> Province <strong>of</strong> Minas Geraes. <strong>The</strong> caves are formed inlimestone rocks, and are so numerous that Dr. Lund visitedthousands, but only sixty contained bones in any quantity.<strong>The</strong>se caves have a floor <strong>of</strong> reddish earth, <strong>of</strong>ten crowded <strong>with</strong>bones. In one experiment, half a cubic foot <strong>of</strong> this earthcontained jaws <strong>of</strong> 400 opossums, 2,000 mice, besides remains<strong>of</strong> bats, porcupines and small birds. In ano<strong>the</strong>r trial, <strong>the</strong>whole <strong>of</strong> <strong>the</strong> earth in a cavern was carried out for examination,amounting to 6,552 firkins ; and, from a calculationmade by measured samples, it was estimated to containnearly seven millions <strong>of</strong> jaw-bones <strong>of</strong> cavies, opossums, porcupines,and mice, besides small birds, lizards, and frogs. _ Thisimmense accumulation is believed to have been formed from<strong>the</strong> bodies <strong>of</strong> <strong>animals</strong> brought into <strong>the</strong> cavern by owls ;and,as <strong>the</strong>se are unsocial birds, <strong>the</strong> quantity found implies an


144 DISTRIBUTION OF EXTINCT ANIMALS. [part n.immense lapse <strong>of</strong> time, probably some thousands <strong>of</strong> years.More than 100 species <strong>of</strong> Mammalia, in all, were obtainedin <strong>the</strong>se caves. Some were living species or closely alliedto such; but <strong>the</strong> majority were extinct, and a considerablenumber, about one-fourth, belonged to extinct genera, orgenera not now inhabiting South America. Stone implementsand human remains were found in several <strong>of</strong> <strong>the</strong> caves <strong>with</strong>extinct <strong>animals</strong>. <strong>The</strong> following enumeration <strong>of</strong> <strong>the</strong>se remainsis from <strong>the</strong> corrected list <strong>of</strong> M. Gervais.Primates.—Extinct species <strong>of</strong> Cebus, Callithrix, and Jacchus—South American genera <strong>of</strong> monkeys; <strong>with</strong> an extinct genus,Protop>ithccus—axi animal <strong>of</strong> large size but belonging to <strong>the</strong>American family Cebidae.Chiroptera.—Species belonging to <strong>the</strong> South American Phyllostomidre,and to two South American genera <strong>of</strong> o<strong>the</strong>r families.Carnivora.—Five species <strong>of</strong> Felis, some allied to living <strong>animals</strong>,o<strong>the</strong>rs extinct ; a species <strong>of</strong> <strong>the</strong> widespread extinct genusMachairodiis ; and a small species referred to Cynmlurus, <strong>the</strong>genus containing <strong>the</strong> hunting leopard now found only inAfricaand India. Canidse are represented by Canis and Icticyon (aliving Brazilian species <strong>of</strong> <strong>the</strong> latter genus), and <strong>the</strong> extinctgenus Speothos. Mustelidse are represented by extinct species<strong>of</strong> <strong>the</strong> South American genera Mephitis and G-alietis. ProcyonidaB,by a species <strong>of</strong> Nasua. Ursidse, by Arcto<strong>the</strong>rium, agenus closely resembling, if not identical <strong>with</strong>, that containing<strong>the</strong> " spectacled bear " <strong>of</strong> Chili.Ungulata.— Equus, Tapirus, Dicotyles, Auchenia, Cervus,Lepto<strong>the</strong>rium,and Antilope, are <strong>the</strong> cave-genera <strong>of</strong> this order.Equus and Antelope are particularly interesting, asrepresentinggroups forming no part <strong>of</strong> existing South American zoology;while <strong>the</strong> presence also <strong>of</strong> Lepto<strong>the</strong>rium, an extinct genus <strong>of</strong>antelopes, shows that <strong>the</strong> group was fairly represented in SouthAmerica at this comparatively recent period.Proboscidea.—A species <strong>of</strong> Mastodon, found also in <strong>the</strong> Pliocene<strong>of</strong> La Plata, represents this order.Rodentia.—<strong>The</strong>se abound. Dasyprocta, Cmlogemjs, Cavia,Kerodon, all living genera <strong>of</strong> Caviidae, are represented by


;chap, vii.] MAMMALIA OF THE NEW WORLD. 145extinct species. Cercolabes, <strong>the</strong> ' tree porcupine ' (Cercolabidse)has two species, one as large as a peccary ;Myopotamus, Zoncheres,Carterodon, are existing genera <strong>of</strong> spiny rats (Echimyidse)and <strong>the</strong>re are two extinct genera <strong>of</strong> <strong>the</strong> same family, Lonchophorusand Phyllomys. Lagodomus (Chinchillidae), <strong>the</strong> viscacha<strong>of</strong> <strong>the</strong> Pampas, is represented by an extinct species.<strong>The</strong>re is also an extinct species <strong>of</strong> Lepus ; several species <strong>of</strong>Hesperomys and Oxymycterus ;living in South America.and a large Arvicola, a genus notEdentata.—<strong>The</strong>se, which constitute <strong>the</strong> great feature <strong>of</strong> <strong>the</strong>existing South American fauna, were still more abundant andvaried in <strong>the</strong> Cave period, and it is remarkable that most <strong>of</strong><strong>the</strong>m are extinct genera. <strong>The</strong> armadillos are alone representedby living forms, Dasypus, and Xenurus ; Ewrydon and Heterodon,are extinct genera <strong>of</strong> <strong>the</strong> same family, as well as Chlamydo<strong>the</strong>rium—hugearmadillos <strong>the</strong> size <strong>of</strong> a tapir or rhinoceros, andPachy<strong>the</strong>rium, which was nearly as large. <strong>The</strong> ant-eaters arerepresented only by Glosso<strong>the</strong>rium, an extinct form allied toMyrmecophaga and Manis. <strong>The</strong> sloths were more numerous,being represented by <strong>the</strong> extinct genera Ccelodon, Sphenodonand Ochotlierium, <strong>the</strong> last <strong>of</strong> large size. <strong>The</strong> huge terrestrialsloths—Mega<strong>the</strong>ridse, also abounded ; <strong>the</strong>re being species <strong>of</strong>Mega<strong>the</strong>rium and Megalonyx, as well as <strong>the</strong> allied Scclido<strong>the</strong>rium,supposed to have some affinity for <strong>the</strong> African Orycteropus.Marsupials.—No new forms <strong>of</strong> <strong>the</strong>se appear, but numerousspecies <strong>of</strong> Didelphys, all closely allied to opossums still livingin South America.<strong>The</strong> preceding sketch <strong>of</strong> <strong>the</strong> wonderful cave fauna <strong>of</strong> CentralBrazil, is sufficient to show that it represents, in <strong>the</strong> main, aperiod <strong>of</strong> great antiquity. Not only are almost <strong>the</strong> whole <strong>of</strong><strong>the</strong> species extinct, but <strong>the</strong>re are twenty extinct genera, and threeo<strong>the</strong>rs not now inhabitants <strong>of</strong> South America. <strong>The</strong> fact that s<strong>of</strong>ew remains <strong>of</strong> <strong>the</strong> living <strong>animals</strong> <strong>of</strong> <strong>the</strong> country are found in<strong>the</strong>se caves, indicates that some change <strong>of</strong> physical conditionshas occurred since <strong>the</strong>y were <strong>the</strong> receptacles <strong>of</strong> so*many <strong>of</strong> <strong>the</strong>larger <strong>animals</strong>; and <strong>the</strong> presence <strong>of</strong> many extinct genera <strong>of</strong>


146 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.large size, especially among <strong>the</strong> Edentata and American families<strong>of</strong> Kodents, are additional pro<strong>of</strong>s <strong>of</strong> a very high antiquity.Yet many <strong>of</strong> <strong>the</strong>se cave <strong>animals</strong> are closely allied to thosewhich are found in North America in <strong>the</strong> Post-Pliocene depositsonly, so that we have no reason to suppose <strong>the</strong> cave-fauna tobe <strong>of</strong> much earlier date. But <strong>the</strong> great amount <strong>of</strong> organicchange it implies, must give us an enlarged idea <strong>of</strong> <strong>the</strong> vastperiods <strong>of</strong> time, as measured by years, which are included inthis, <strong>the</strong> most recent <strong>of</strong> all geological epochs.Pliocene Period <strong>of</strong> Temperate South America.—We have nowto consider <strong>the</strong> numerous remains <strong>of</strong> extinct <strong>animals</strong> found invarious deposits in <strong>the</strong> Pampas, and in Patagonia, and a few inBolivia. <strong>The</strong> age <strong>of</strong> <strong>the</strong>se is uncertain; but as <strong>the</strong>y are verysimilar to <strong>the</strong> cave-fauna, though containing a somewhat largerproportion <strong>of</strong> extinct genera and some very remarkable newforms, <strong>the</strong>y cannot be very much older, and are perhaps bestreferred at present to <strong>the</strong> newer portion <strong>of</strong> <strong>the</strong> Plioceneformation.Carnivora.—<strong>The</strong> genus Machairodus orsabre-too<strong>the</strong>d tigers,represents <strong>the</strong> Felidae. <strong>The</strong>re are several species <strong>of</strong> wolves(Canis) ; a weasel (Mustela) ;two bears <strong>of</strong> <strong>the</strong> Brazilian cavegenusArcto<strong>the</strong>rium; and <strong>the</strong> extinct European genus Hyamarctos.Ungulata.—<strong>The</strong>re are two species <strong>of</strong> Eqnus, found in <strong>the</strong>Pampas, Chili, and Bolivia; two <strong>of</strong> Macrauchenia, an extraordinaryextinct group allied to <strong>the</strong> tapir and Palceothcrium, but<strong>with</strong> <strong>the</strong> long neck, and general size <strong>of</strong> a camel. A secondspecies found on <strong>the</strong> highlands <strong>of</strong> Bolivia is much smaller.A more recent discovery, in Patagonia, is <strong>the</strong> almost perfectseries <strong>of</strong> teeth <strong>of</strong> a large animal named Homalodonto<strong>the</strong>rium; andwhich is believed by Pr<strong>of</strong>essor Flower, who has described it, tohave been allied to Rhinoceros, and still more to <strong>the</strong> MioceneHyracodon from North America ; and also to present someresemblances to Macrauchenia, and though much more remotely,to <strong>the</strong> curious genus Nesodon mentioned fur<strong>the</strong>r on.<strong>The</strong> Artiodactyla, or even-toed Ungulates, are represented by aspecies <strong>of</strong> Picotyles, or peccary, found in <strong>the</strong> deposits <strong>of</strong> <strong>the</strong>


—;chap, vii.] MAMMALIA OF THE NEW WORLD. 147Pampas ; by Auchenia, or llama, <strong>of</strong> which three extinct speciesinhabited Bolivia, in which country two allied but extinctgenera, Palmolama and Camelo<strong>the</strong>rium, have also been found.Three species<strong>of</strong> deer (Cervus), from <strong>the</strong> Pampas deposits, complete<strong>the</strong> list <strong>of</strong> Pliocene Ungulates.Proboscidea.—<strong>The</strong> cave species <strong>of</strong> Mastodon is found also in<strong>the</strong> Pampas deposits, and ano<strong>the</strong>r in <strong>the</strong> Andes <strong>of</strong> Chili andBolivia.Rodents.—<strong>The</strong>se are not so numerous as in <strong>the</strong> caves. <strong>The</strong>reare species <strong>of</strong> <strong>the</strong> existing genera, Kerodon and Cavia (Caviidse)Zagostomus (Chinchillidse) ; Ctenomys (Octodontidse) ; Lepus(hare) ; Hesperomys and Oxymycterus (Muridre) ; Arvicola, agenus not living in South America ; and an extinct genus, Cardiodus.<strong>The</strong>re is also a remarkable extinct form, Typo<strong>the</strong>rium,larger than <strong>the</strong> capybara, and having affinities to Edentates andUngulates. Three species have been found in <strong>the</strong> Pampas deposits.Edentata.—<strong>The</strong>se are as abundant and remarkable as in <strong>the</strong>cave deposits. Scelido<strong>the</strong>rium, Mega<strong>the</strong>rium, Megalonyx, Glosso<strong>the</strong>riumand Dasypus, have already been noticed as from <strong>the</strong>Brazilian caves. We have here, in addition, <strong>the</strong> huge Mylodonallied to <strong>the</strong> Mega<strong>the</strong>rium, and <strong>the</strong> allied genera Gnathopsisand Lestodon. We <strong>the</strong>n come to <strong>the</strong> huge extinct armadillos,Glyptodon and Schistopleurum, <strong>the</strong> former consisting <strong>of</strong> numerousspecies, some <strong>of</strong> which were as large as an elephant. Ano<strong>the</strong>rgenus, Eutatus, is allied to <strong>the</strong> living three-banded armadillosand a species <strong>of</strong> <strong>the</strong> existing genus Euphractus has been foundin Bolivia.Toxodontidce.—<strong>The</strong>re remain a number <strong>of</strong> huge <strong>animals</strong>rivalling <strong>the</strong> Mega<strong>the</strong>rium in size, and forming <strong>the</strong> generaToxodon and Nesodon, but whose position is doubtful. Severalspecies have been found in <strong>the</strong> deposits <strong>of</strong> <strong>the</strong> Pampas andPatagonia. <strong>The</strong>y are allied at once to Ungulates, Eodents, Edentates,and <strong>the</strong> aquatic Sirenia, in so puzzling a manner that itimpossible to determine to what order <strong>the</strong>y belong, or whe<strong>the</strong>r<strong>the</strong>y require a new order to be formed for <strong>the</strong>ir reception. Someare believed to date back to <strong>the</strong> Miocene period, and <strong>the</strong>yindicate what strange forms may still be discovered, should anyis


—;148 DISTRIBUTION QF EXTINCT ANIMALS. [part 11.productive deposits be found in South America <strong>of</strong> middle Tertiaryage.Pliocene Mammalia <strong>of</strong> <strong>the</strong> Antilles.—<strong>The</strong>se may be noticedhere, as <strong>the</strong>y are <strong>of</strong> special interest, proving <strong>the</strong> connection <strong>of</strong><strong>the</strong> larger West Indian Islands <strong>with</strong> <strong>the</strong> Continent some time in<strong>the</strong> later Tertiary period. <strong>The</strong>y consist <strong>of</strong> remains <strong>of</strong> two large<strong>animals</strong> belonging to <strong>the</strong> South American Chinchillidse, found incave deposits in <strong>the</strong> island <strong>of</strong> Anguilla, and forming two newgenera, AmUyrhiza and Loxomylus ; and remain allied to Megalonyxfrom Cuba, which have been named Megalocnus andMyomorpkus.Eocene fauna <strong>of</strong> South America.—<strong>The</strong> few remains yet discoveredin <strong>the</strong> Tertiary deposits <strong>of</strong> <strong>the</strong> Pampas which are believedto be <strong>of</strong> Eocene age, are exceedingly interesting, because <strong>the</strong>yshow us ano<strong>the</strong>r change in <strong>the</strong> scenery <strong>of</strong> <strong>the</strong> great drama <strong>of</strong>life ; <strong>the</strong>re being apparently a considerable resemblance, at thisepoch, between South America and Europe. <strong>The</strong>y consist <strong>of</strong> alarge extinct feline animal, Eutemnodus ; <strong>of</strong> Palceo<strong>the</strong>rium andAnoplo<strong>the</strong>rium, <strong>the</strong> well-known extinct Ungulates <strong>of</strong> <strong>the</strong> EuropeanTertiaries, and which have never been found in NorthAmerica; and <strong>of</strong> three genera <strong>of</strong> Eodents,<strong>The</strong>ridromys,alliedto Echimys, and found also in <strong>the</strong> Eocene and Miocene <strong>of</strong> FranceMcgamys, allied to <strong>the</strong> living Capromys <strong>of</strong> <strong>the</strong> Antilles, andalso to Palaiomys, an extinct form <strong>of</strong> <strong>the</strong> French Miocene ;anda very large animal referred to Arvicola, a genus found also in<strong>the</strong> Pliocene deposits <strong>of</strong> South America, and abundant in <strong>the</strong>nor<strong>the</strong>rn hemisphere. No Edentates have been found.<strong>The</strong> resemblances <strong>of</strong> this fauna to that <strong>of</strong> Europe ra<strong>the</strong>r thanto any part <strong>of</strong> America, are so strong, that <strong>the</strong>y can hardly beaccidental. We greatly want, however, more information on thispoint, as well as some corresponding evidences as to <strong>the</strong> condition<strong>of</strong> West and South Africa about <strong>the</strong>same epoch, before we canventure to speculate on <strong>the</strong>ir bearing as regards <strong>the</strong> early migrations<strong>of</strong> organic forms.G 1 "" 1""'' "* "*O lGeneral Remarks on <strong>the</strong> Extinct Mammalian Fauna <strong>of</strong> <strong>the</strong> Old


chap, vii.] MAMMALIA OF THE NEW WORLD. 149and New Worlds.—Leaving <strong>the</strong> more special applications <strong>of</strong>palseontological evidence to be made after discussing <strong>the</strong> <strong>relations</strong><strong>of</strong> <strong>the</strong> existing fauna <strong>of</strong> <strong>the</strong> several regions, we propose here toindicate briefly, some <strong>of</strong> <strong>the</strong> more general deductions from <strong>the</strong>evidence which has now been laid before our readers.<strong>The</strong> first, and perhaps <strong>the</strong> most startling fact brought out byour systematic review, is <strong>the</strong> very recent and almost universalchange that has taken place in <strong>the</strong> character <strong>of</strong> <strong>the</strong> fauna,all <strong>the</strong> areas we have been considering ;overa change which seemsto be altoge<strong>the</strong>r unprecedented in <strong>the</strong> past history <strong>of</strong> <strong>the</strong> samecountries as revealed by <strong>the</strong> geological record. In Europe, inNorth America, and in South America, we have evidence that avery similar change occurred about <strong>the</strong> same time. In all threewe find, in <strong>the</strong> most recent deposits—cave-earths, peat-bogs, andgravels—<strong>the</strong> remains <strong>of</strong> a whole series <strong>of</strong> large <strong>animals</strong>, whichhave since become wholly extinct or only survive in far-distantlands. In Europe, <strong>the</strong> great Irish elk, <strong>the</strong> Machairodus andcave-lion, <strong>the</strong> rhinoceros, hippopotamus, and elephant ;—in NorthAmerica, equally large felines, horses and tapirs larger than anynow living, a llama as large as a camel, great mastodons andelephants, and abimdance <strong>of</strong> huge mega<strong>the</strong>roid <strong>animals</strong> <strong>of</strong> almostequal size ;—in South America <strong>the</strong>se same mega<strong>the</strong>roids in greatervariety, numerous huge armadillos, a mastodon, large horses andtapirs, large porcupines, two forms <strong>of</strong> antelope, numerous bears andfelines, including a Machairodus, and a large monkey,—have all becomeextinct since <strong>the</strong> deposition <strong>of</strong> <strong>the</strong> most recent <strong>of</strong> <strong>the</strong> fossilbearingstrata. This is certainly not a great while ago, geologically; and it is almost certain that this great organic revolution,implying physical changes <strong>of</strong> such vast proportions that <strong>the</strong>ymust have been due to causes <strong>of</strong> adequate intensity and proportionaterange, has taken place since man lived on <strong>the</strong> earth.This is proved to have been <strong>the</strong> case in Europe, and is supportedby much evidence both as regards North and South America.It is clear that so complete and sudden a change in <strong>the</strong> higherforms <strong>of</strong> life, does not represent <strong>the</strong> normal state <strong>of</strong> things.Species and genera have not, at all times, become so rapidlyextinct<strong>The</strong> time occupied by <strong>the</strong> " Eecent period," that is <strong>the</strong>


150 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.time since <strong>the</strong>se changes took place is, geologically, minute.<strong>The</strong> time <strong>of</strong> <strong>the</strong> whole <strong>of</strong> <strong>the</strong> Post-Pliocene period, as measuredby <strong>the</strong> amount <strong>of</strong> physical and general organic change known tohave taken place, is exceedingly small when compared <strong>with</strong> <strong>the</strong>duration <strong>of</strong> <strong>the</strong> Pliocene period, and still smaller, probably, ascompared <strong>with</strong> <strong>the</strong> Miocene. Yet during <strong>the</strong>se two periods wemeet <strong>with</strong> no such break in <strong>the</strong> continuity <strong>of</strong> <strong>the</strong> forms <strong>of</strong> life, nosuch radical change in <strong>the</strong> character <strong>of</strong> <strong>the</strong> fauna (though <strong>the</strong>number <strong>of</strong> specific and generic changes may be as great) as wefind in passing from <strong>the</strong> Post-Pliocene to recent times. Forexample, in Central Europe numerous hysenas, rhinoceroses, andantelopes, <strong>with</strong> <strong>the</strong> great Machairodus, continued from Mioceneall through Pliocene into Post-Pliocene times ; while hippopotamiand elephants continued to live through a good part <strong>of</strong><strong>the</strong> Pliocene and Post-Pliocene periods,—and <strong>the</strong>n allsuddenlybecame extinct or left <strong>the</strong> country. In North America <strong>the</strong>re hasbeen more movement <strong>of</strong> <strong>the</strong> fauna in all <strong>the</strong> periods ; but wehave similar great felines, horses, mastodons, and elephants, in<strong>the</strong> Pliocene and Post-Pliocene periods, while Rhinoceros is commonto <strong>the</strong> Miocene and Pliocene, and camels range continuouslyfrom Miocene, through Pliocene, toall alike became extinct. Even in South America <strong>the</strong> evidence is,as far as it goes, all <strong>the</strong> same way. We find Machairodus, Equus,Mastodon, Mega<strong>the</strong>rium, Scelido<strong>the</strong>rium, Megalonyx, and numerousPost-Pliocene times ;—whengigantic armadillos, alike in <strong>the</strong> caves and in <strong>the</strong> stratifiedtertiary deposits <strong>of</strong> <strong>the</strong> Pampas ;— yet all have since passed away.It is clear, <strong>the</strong>refore, that we are now in an altoge<strong>the</strong>rexceptional period <strong>of</strong> <strong>the</strong> earth's history. We live in a zoologicallyimpoverished world, from which all <strong>the</strong> hugest, and fiercest,and strangest forms have recently disappeared ;and it is, nodoubt, a much better world for us now <strong>the</strong>y have gone.Yet itis surely a marvellous fact, and one that has hardly been sufficientlydwelt upon, this sudden dying out <strong>of</strong> so many largemammalia, not in one place only but over half <strong>the</strong> land surface<strong>of</strong> <strong>the</strong> globe. We cannot but believe that <strong>the</strong>re must have beensome physical cause for this great change ; and it must havebeen a cause capable <strong>of</strong> acting almost simultaneously over large


chap, vii.] MAMMALIA OF THE NEW WORLD. 151portions <strong>of</strong> <strong>the</strong> earth's surface, and one which, as far as <strong>the</strong>Tertiary period at least isconcerned, was <strong>of</strong> an exceptional character.Such a cause exists in <strong>the</strong> great and recent physical changeknown as " <strong>the</strong> Glacial epoch." We have pro<strong>of</strong> in both Europeand North America, that just about <strong>the</strong> time <strong>the</strong>se large <strong>animals</strong>were disappearing, all <strong>the</strong> nor<strong>the</strong>rn parts <strong>of</strong> <strong>the</strong>se continentswere wrapped in a mantle <strong>of</strong> ice ;believe that <strong>the</strong> presence <strong>of</strong> thisand we have every reason tolarge quantity <strong>of</strong> ice (known tohave been thousands <strong>of</strong> feet if not some miles in thickness) musthave acted in various ways to have produced alterations <strong>of</strong>level <strong>of</strong> <strong>the</strong> ocean as well as vast local floods, which would havecombined <strong>with</strong> <strong>the</strong> excessive cold to destroy animal life.<strong>The</strong>reis great difference <strong>of</strong> opinion among geologists and physicists asto <strong>the</strong> extent, nature, and duration <strong>of</strong> <strong>the</strong> Glacial epoch. Somebelieve it to have prevailed alternately in <strong>the</strong> nor<strong>the</strong>rn andsou<strong>the</strong>rn hemispheres ;o<strong>the</strong>rs that it was simultaneous in both.Some think <strong>the</strong>re was a succession <strong>of</strong> cold periods, each lastingmany thousands <strong>of</strong> years, but <strong>with</strong> intercalated warm periods <strong>of</strong>equal duration ; o<strong>the</strong>rs deny that <strong>the</strong>re is any evidence <strong>of</strong> suchchanges, and maintain that <strong>the</strong> Glacial epoch was one continuousperiod <strong>of</strong> arctic conditions in <strong>the</strong> temperate zones, <strong>with</strong> somefluctuations perhaps but <strong>with</strong> no regular alternations <strong>of</strong> warmperiods. Some believe in a huge ice-cap covering <strong>the</strong> wholenor<strong>the</strong>rn hemisphere from <strong>the</strong> pole to near 50° north latitudein <strong>the</strong> eastern, and 40° in <strong>the</strong> western hemisphere ;impute <strong>the</strong> observed effects ei<strong>the</strong>r to glacierswhile o<strong>the</strong>rsfrom local centres,or to floating icebergs <strong>of</strong> vast size passing over <strong>the</strong> surface duringa period <strong>of</strong> submersion.Without venturing to decide which <strong>of</strong> <strong>the</strong>se various <strong>the</strong>orieswill be ultimately proved to be correct, we may state, that<strong>the</strong>re is an increasing belief among geologists in <strong>the</strong> long duration<strong>of</strong> this ice-period, and <strong>the</strong> vast extent and great thicknessattained by <strong>the</strong> ice-sheet. One <strong>of</strong> <strong>the</strong> most recent, andnot <strong>the</strong> least able, <strong>of</strong> <strong>the</strong> writers on this question (Mr. Belt) showsstrong reasons for adopting <strong>the</strong> view that <strong>the</strong> ice-period wassimultaneous in both hemispheres ; and he calculates that <strong>the</strong>vast amount <strong>of</strong> water abstracted from <strong>the</strong> ocean aid locked up


152 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.in mountains <strong>of</strong> ice around <strong>the</strong> two poles, would lower <strong>the</strong> generallevel <strong>of</strong> <strong>the</strong> ocean about 2,000 feet. This would be equivalentto a general elevation <strong>of</strong> <strong>the</strong> land to <strong>the</strong> same amount, and wouldthus tend to intensify <strong>the</strong> cold ; and this elevation may enableus to understand <strong>the</strong> recent discoveries <strong>of</strong> signs <strong>of</strong> glacialaction at moderate elevations in Central America and Brazil, far<strong>with</strong>in <strong>the</strong> tropics. At <strong>the</strong> same time, <strong>the</strong> weight <strong>of</strong> ice piled upin <strong>the</strong> north would cause <strong>the</strong> land surface to sink <strong>the</strong>re, perhapsunequally, according to <strong>the</strong> varying nature <strong>of</strong> <strong>the</strong> interior crust<strong>of</strong> <strong>the</strong> earth ;and since <strong>the</strong> weight has been removed land wouldand thus <strong>the</strong> phenomenarise again, still somewhat irregularly ;<strong>of</strong> raised beds <strong>of</strong> arctic shells in temperate latitudes, are explained.Now, it is evident, that <strong>the</strong> phenomena we have been considering—<strong>of</strong><strong>the</strong> recent changes <strong>of</strong> <strong>the</strong> mammalian fauna inEurope, North America, South Temperate America, and <strong>the</strong>highlands <strong>of</strong> Brazil—are such as might be explained by <strong>the</strong> mostextreme views as to <strong>the</strong> extent and vastness <strong>of</strong> <strong>the</strong> ice-sheet,and especially as toand sou<strong>the</strong>rn hemispheres ;and where two such completely independentsets <strong>of</strong> facts areits simultaneous occurrence in <strong>the</strong> nor<strong>the</strong>rnfound to combine harmoniously, andsupplement each o<strong>the</strong>r on a particular hypo<strong>the</strong>sis, <strong>the</strong> evidencein favour <strong>of</strong> that hypo<strong>the</strong>sis is greatly streng<strong>the</strong>ned. An objectionthat will occur to zoologists, may here be noticed. If<strong>the</strong> Glacial epoch extended over so much <strong>of</strong><strong>the</strong> temperate andeven parts <strong>of</strong> <strong>the</strong> tropical zone, and led to <strong>the</strong> extinction <strong>of</strong> somany forms <strong>of</strong> life even <strong>with</strong>in <strong>the</strong> tropics, how is it that somuch <strong>of</strong> <strong>the</strong> purely tropical fauna <strong>of</strong> South America has maintaineditself, and that <strong>the</strong>re are still such a vast number <strong>of</strong>forms, both <strong>of</strong> mammalia, birds, reptiles, and insects, that seemorganized for an exclusive existence in tropical forests ? NowMr. Belt's <strong>the</strong>ory, <strong>of</strong> <strong>the</strong> subsidence <strong>of</strong> <strong>the</strong> ocean to <strong>the</strong> extent <strong>of</strong>about 2,000 feet, supplies an answer to this objection ; for weshould thus have a tract <strong>of</strong> lowland <strong>of</strong> an average width <strong>of</strong>some hundreds <strong>of</strong> miles, added to <strong>the</strong> whole east coast <strong>of</strong> Centraland South America.This tract would, no doubt, become covered<strong>with</strong> forests as it was slowly formed, would enjoy a perfectly


chap, vil] MAMMALIA OF THE NEW WORLD. 153tropical climate, and would thus afford an ample area for <strong>the</strong>continued existence and development <strong>of</strong> <strong>the</strong> typical SouthAmerican fauna ; even had glaciers descended in places so lowas what is now <strong>the</strong> level <strong>of</strong> <strong>the</strong> sea, which, however, <strong>the</strong>re is noreason to believe <strong>the</strong>y ever did. It is probable too, that tinslow tract, winch all round <strong>the</strong> Gulf <strong>of</strong> Mexico would be <strong>of</strong> considerablewidth, <strong>of</strong>fered that passage for intermigration betweenNorth and South America, which led to <strong>the</strong> sudden appearancein <strong>the</strong> former country in Post-Pliocene times, <strong>of</strong> <strong>the</strong> huge Mega<strong>the</strong>roidsfrom <strong>the</strong> latter ; a migration which took place in oppositedirections as we shall presently show.<strong>The</strong> birthplaceand migrations <strong>of</strong> some mammalian familiesand genera.—We have now to consider a few <strong>of</strong> those casesin which <strong>the</strong> evidence already at our command, is sufficientlydefinite and complete, to enable us to pronounce <strong>with</strong> some confidenceas to<strong>the</strong> last movements <strong>of</strong> several important groups <strong>of</strong>mammalia.Primates.—<strong>The</strong> occurrence in North America <strong>of</strong> numerousforms <strong>of</strong> Lemuroidea, forming two extinct families, which arebelieved by American paleontologists to present generalizedfeatures <strong>of</strong> both Lemuridae and Hapalida?, while in Europe onlyLemurine forms allied to those <strong>of</strong> Africa have occurred indeposits <strong>of</strong> <strong>the</strong> same age (Eocene), renders it possible that <strong>the</strong>Primates may have originated in America, and sent one branchto South America to form <strong>the</strong> Hapalida? and Cebidas, andano<strong>the</strong>r to <strong>the</strong> Old "World, giving rise to <strong>the</strong> lemurs and trueapes. But <strong>the</strong> fact that apes <strong>of</strong> a high degree <strong>of</strong> organizationoccur in <strong>the</strong> European Miocene, while in <strong>the</strong> Eocene, a monkeybelieved to have <strong>relations</strong> to <strong>the</strong> Lemuroids and Cebidse has alsobeen discovered, make it more probable that <strong>the</strong> ancestral forms<strong>of</strong> this order originated in <strong>the</strong> Old World at a still earlier period.<strong>The</strong> absence <strong>of</strong> any early tertiary remains from <strong>the</strong> tropical parts<strong>of</strong> <strong>the</strong> two hemispheres, renders it impossible to arrive at anydefinite conclusions as to <strong>the</strong> origin <strong>of</strong> groups which were, nodoubt, always best developed in tropical regions.Carnivora.—This is a very ancient and wide-spread group, <strong>the</strong>families and genera <strong>of</strong> which had an extensive range in veryVol. I.—12


—154 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.early times. <strong>The</strong> true bears (Ursus) are almost <strong>the</strong> onlyimportant genus that seems to have recently migrated. InEurope it dates back to <strong>the</strong> Older Pliocene, while in NorthAmerica it is Post-Pliocene only. Bears, <strong>the</strong>refore, seem tohave passed into America from <strong>the</strong> Palaearctic region in <strong>the</strong> latterpart <strong>of</strong> <strong>the</strong> Pliocene period. <strong>The</strong>y probably came in on <strong>the</strong>north-west, and passed down <strong>the</strong> Andes into South America,where one isolated species still exists.Ungulata.—Horses are very interesting. In Europe <strong>the</strong>y dateback under various forms to <strong>the</strong> Miocene period, and true Equusto <strong>the</strong> Older Pliocene. In North America <strong>the</strong>y are chieflyPliocene, true Equus being Post- Pliocene, <strong>with</strong> perhaps one ortwo species Newer Pliocene ;but numerous ancestral forms dateback to <strong>the</strong> Miocene and Eocene, giving a more perfect " pedigree<strong>of</strong> <strong>the</strong> horse " than <strong>the</strong> European forms, and going back to amore primitivetypeOrohippus.In South America, Equus is<strong>the</strong> only genus, and is Post-Pliocene or at most Newer Pliocene.While, <strong>the</strong>refore, <strong>the</strong> ancient progenitors <strong>of</strong> <strong>the</strong> Equidae werecommon to North America and Europe, in Miocene and evenEocene times, true horses appear to have arisen in <strong>the</strong> Palasarcticregion, to have passed into North America in <strong>the</strong> latter part <strong>of</strong><strong>the</strong> Pliocene period, and <strong>the</strong>nce tohave spread over all suitabledistricts in South America. <strong>The</strong>y w T ere not, however, able tomaintain <strong>the</strong>mselves permanently in <strong>the</strong>ir new territory, and allbecame extinct ;while in <strong>the</strong>ir birth-place, <strong>the</strong> Old World, <strong>the</strong>ycontinue to exist under several varied forms.True tapirs are an Old World group. <strong>The</strong>y go back to <strong>the</strong>Lower Miocene in Europe, while in both North and SouthAmerica <strong>the</strong>y are exclusively Post-Pliocene. <strong>The</strong>y occur inFrance down to <strong>the</strong> Newer Pliocene, and must, about thattime, have entered America. <strong>The</strong> land connection by whichthis and so many o<strong>the</strong>r <strong>animals</strong> passed between <strong>the</strong> Old andNew Worlds in late Tertiary times, was almost certainly in <strong>the</strong>North Pacific, south <strong>of</strong> Behring's Straits, where, as will be seenby our geueral map, <strong>the</strong>re is a large expanse <strong>of</strong> shallow water,which a moderate elevation would convert into dry land, in asufficiently temperate latitude.


Jchap. vii.MAMMALIA OF THE NEW WORLD. 155<strong>The</strong> peccary (Dicotyles), now a characteristic South Americangenus, is a recent immigrant from North America, where itappears to have.been developed from ancestral forms <strong>of</strong> swinedating back to <strong>the</strong> Miocene period.Antelopes are an Old World type, but a few <strong>of</strong> <strong>the</strong>m appear tohave entered North, and reached South America in late Pliocenetimes.Camels, strange to say, are a special North American type,since <strong>the</strong>y abounded in that continent under various ancientforms in <strong>the</strong> Miocene period. Towards <strong>the</strong> end <strong>of</strong> that period<strong>the</strong>y appear to have entered eastern Asia, and developed into <strong>the</strong>Siberian Meryco<strong>the</strong>riurn and <strong>the</strong> North Indian Camelus, whilein <strong>the</strong> Pliocene age <strong>the</strong> ancestral llamas entered SouthAmerica.Cervidce are a wide-spread nor<strong>the</strong>rn type in <strong>the</strong>irgeneralizedform, but true deer (Cervus) are Palsearctic. <strong>The</strong>y abounded inEurope in Miocene times, but only appear inNorth and SouthAmerica in <strong>the</strong> later Pliocene and Post-Pliocene periods.True oxen (Bovince) seem to be an Oriental type (Miocene),while <strong>the</strong>y appear in Europe only late in <strong>the</strong> Pliocene period,and in America are confined to <strong>the</strong> Post-Pliocene.Elephants (Mephantidce) are an Old World type, aboundingin <strong>the</strong> Miocene period in Europe and India, and first appearingin America in Post-Pliocene or later Pliocene times.Ancestralforms, doubtfully Proboscidean (Dinocerata), existed in NorthAmerica in <strong>the</strong> Eocene period, but <strong>the</strong>se became extinct <strong>with</strong>outleaving any direct descendants, unless <strong>the</strong> Bronto<strong>the</strong>ridce andrhinoceroses may be so considered.Marsupials are almost certainly a recent introduction intoSouth and North America from Asia. <strong>The</strong>y existed in Europein Eocene and Miocene times,and presumably over a considerablepart <strong>of</strong> <strong>the</strong> Old World ; but no trace <strong>of</strong> <strong>the</strong>m appears inNorth or South America before <strong>the</strong> Post-Pliocene period.Edentata.— <strong>The</strong>se <strong>of</strong>fer a most curious and difficult problem.In South America <strong>the</strong>y abound, and were so much more numerousand varied in <strong>the</strong> Post-Pliocene and Pliocene, that wemay be sure <strong>the</strong>y lived also in <strong>the</strong> preceding Miocene period. Afew living Edentates are scattered over Africa and Asia, and.


156 DISTRIBUTION OF EXTINCT ANIMALS. [part 11.<strong>the</strong>y flourished in Europe during <strong>the</strong> Miocene age—<strong>animals</strong> aslarge(in some species) as a rhinoceros, and most allied to livingAfrican forms. In North America no trace <strong>of</strong> Edentata has beenfound earlier than <strong>the</strong> Post-Pliocene period, or perhaps <strong>the</strong> NewerPliocene on <strong>the</strong> west coast. Nei<strong>the</strong>r is <strong>the</strong>re any trace <strong>of</strong> <strong>the</strong>min South America in <strong>the</strong> Eocene formations ;but this may wellbe owing to our very imperfect knowledge <strong>of</strong> <strong>the</strong> forms <strong>of</strong>that epoch. <strong>The</strong>ir absence from North America is, however,probably real ; and we have to account for <strong>the</strong>ir presence in <strong>the</strong>Old World and in South America. <strong>The</strong>ir antiquity is no doubtvery great, and <strong>the</strong> point <strong>of</strong> divergence <strong>of</strong> <strong>the</strong> Old World andSouth American groups, may take us back to early Eocene, oreven to Pre-Eocene times. <strong>The</strong> <strong>distribution</strong> <strong>of</strong> land and sea may<strong>the</strong>n have been very different from what it is now ; and to thosewho would create a continent to account for <strong>the</strong> migrations <strong>of</strong>a beetle, nothing would seem more probable than that a SouthAtlantic continent, <strong>the</strong>n united parts <strong>of</strong> what are now Africaand South America. <strong>The</strong>re is, however, so much evidence for<strong>the</strong> general permanence <strong>of</strong> what are now <strong>the</strong> great continentsand deep oceans, that Pr<strong>of</strong>essor Huxley's supposition <strong>of</strong> a considerableextension <strong>of</strong> land round <strong>the</strong> borders <strong>of</strong> <strong>the</strong> North PacificOcean in Mesozoic times, best indicates <strong>the</strong> probable area inwhich <strong>the</strong> Edentate type originated, and <strong>the</strong>nce spread over much<strong>of</strong> <strong>the</strong> Old World and South America. But while in <strong>the</strong> lattercountry it flourished and increased <strong>with</strong> little check, in <strong>the</strong>o<strong>the</strong>r great continents it was soon overcome by <strong>the</strong> competition<strong>of</strong> higher forms, only leaving a few small-sized representativesin Africa and Asia.


CHAPTER VIII.VARIOUS EXTINCT ANIMALS;—AND ON THE ANTIQUITYGENEEA OF INSECTS AND LAND MOLLUSCA.OF THEEXTINCT MAMMALIA OF AUSTRALIA.<strong>The</strong>se have all been obtained from caves and late Tertiary orPost-Tertiary deposits, and consist <strong>of</strong> a large number <strong>of</strong> extinctforms, some <strong>of</strong> gigantic size, but all marsupials and allied to <strong>the</strong>existing fauna. <strong>The</strong>re are numerous forms <strong>of</strong> kangaroos, somelarger than any living species ;and among <strong>the</strong>se are two genera,Protemnodon and S<strong>the</strong>nurus, which Pr<strong>of</strong>essor Garrod has latelyshown to have been allied, not to any Australian forms, but to<strong>the</strong> Dmdrolagi or tree-kangaroos <strong>of</strong> New Guinea. We havealso remains <strong>of</strong> Thylacinus and Dasyurus, which now only existin Tasmania ; and extinct species <strong>of</strong> Hypsiprymnus and Phasco-Among <strong>the</strong> more remarkablelomys, <strong>the</strong> latter as large as a tapir.extinct genera are Diprotodon, a huge thick-limbed animalallied to <strong>the</strong> kangaroos, but nearly as large as an elephant;Noto<strong>the</strong>rivm, having characters <strong>of</strong> Macropus and Phascolarctoscombined, and as large as a rhinoceros ; and Thylacoho, a phaianger-likemarsupial nearly as large as a lion, and supposed byPr<strong>of</strong>essor Owen to have been <strong>of</strong>opinion isnot held by o<strong>the</strong>r naturalists.Here <strong>the</strong>n we find <strong>the</strong> same phenomena as incarnivorous habits, though this<strong>the</strong> o<strong>the</strong>r countrieswe have already discussed,—<strong>the</strong> very recent disappearance<strong>of</strong> a large number <strong>of</strong> peculiar forms, many <strong>of</strong> <strong>the</strong>m far surpassingin size any that continue to exist. It hardly seems probablethat in this case <strong>the</strong>ir disappearance can have been due to <strong>the</strong>direct effects <strong>of</strong> <strong>the</strong> Glacial epoch, since no very extensive glacia-


158 EXTINCT ANIMALS OF AUSTRALIA. [part ii.tion could have occurred in a country like Australia ; but if <strong>the</strong>ocean sank 2,000 feet, <strong>the</strong> great eastern mountain range mighthave given rise to local glaciers. It is, however, almost certainthat during late Tertiary times Australia must have been muchmore extensive than it is now. This is necessary to allow <strong>of</strong> <strong>the</strong>development <strong>of</strong> its peculiar and extensive fauna, especially as wesee that that fauna comprised <strong>animals</strong> rivalling in bulk those <strong>of</strong><strong>the</strong> great continents. It is fur<strong>the</strong>r indicated by <strong>the</strong> <strong>relations</strong><strong>with</strong> New Guinea, already alluded to, and by <strong>the</strong> general character<strong>of</strong> <strong>the</strong> various faunas which compose <strong>the</strong> Australian region, details<strong>of</strong> which will be found in <strong>the</strong> succeeding part <strong>of</strong> this work.<strong>The</strong> lowering <strong>of</strong> <strong>the</strong> ocean during <strong>the</strong> Glacial period would befavourable to <strong>the</strong> stillfur<strong>the</strong>r development <strong>of</strong> <strong>the</strong> fauna <strong>of</strong> sucha country ; and it is to <strong>the</strong> unfavourable conditions produced byits subsequent rising—equivalent to a depression <strong>of</strong> <strong>the</strong> land to<strong>the</strong> amount <strong>of</strong> two thousand feet—that we must impute <strong>the</strong>extinction <strong>of</strong> so many remarkable groups <strong>of</strong> <strong>animals</strong>. It is notimprobable, that <strong>the</strong> disappearance <strong>of</strong> <strong>the</strong> ice and <strong>the</strong> consequent(apparent) subsidence <strong>of</strong> <strong>the</strong> land, might have been rapid ascompared <strong>with</strong> <strong>the</strong> rate at which large <strong>animals</strong> can becomemodified to meet new conditions. Extensive tracts <strong>of</strong> fertileland might have been submerged, and <strong>the</strong>consequent crowding<strong>of</strong> large numbers <strong>of</strong> species and individuals on limited areaswould have led to a struggle for existence in which <strong>the</strong> lessadapted and less easily modifiable, not <strong>the</strong> physically weaker,would succumb.<strong>The</strong>re is, however, ano<strong>the</strong>r cause for <strong>the</strong> extinction <strong>of</strong> largera<strong>the</strong>r than small <strong>animals</strong> whenever an important change <strong>of</strong>conditions occurs, which has been suggested to me by a correspondent,1 but which has not, I believe, been adduced by Mr.Darwin or by any o<strong>the</strong>r writer on <strong>the</strong> subject. It is dependenton <strong>the</strong> fact, that large <strong>animals</strong> as compared <strong>with</strong> small ones arealmost invariably slow breeders, and as <strong>the</strong>y also necessarilyexist in much smaller numbers in a given area, <strong>the</strong>y <strong>of</strong>fer farless materials forIn such an extreme case as thatfavourable variations than do smaller <strong>animals</strong>.1Mr. John Hiekman <strong>of</strong> Desborongh.<strong>of</strong> <strong>the</strong> rabbit and elephant, <strong>the</strong>


chap, viii.] MESOZOIC MAMMALIA. 159young born each year in <strong>the</strong> world are probably as some millionsto one ; and it is very easily conceivable that in a thousandyears <strong>the</strong> former might, under pressure <strong>of</strong> rapidly changing conditions,become modified into a distinct species, while <strong>the</strong> latter,not <strong>of</strong>fering enough favourable variations to effect a suitableadaptation, would become extinct. We must also remember<strong>the</strong> extreme specialization <strong>of</strong> many <strong>of</strong> <strong>the</strong> large <strong>animals</strong> thathave become extinct—a specializationwhich would necessarilyrender modification in any new direction difficult, since <strong>the</strong> inheritedtendency <strong>of</strong> variation would probably be to increase <strong>the</strong>specialization in <strong>the</strong> same directionswhich had heret<strong>of</strong>ore beenbeneficial. If to <strong>the</strong>se two causes we add <strong>the</strong> difficulty <strong>of</strong>obtaining sufficient food for such large <strong>animals</strong>, and perhaps <strong>the</strong>injurious effects <strong>of</strong> changes <strong>of</strong> climate, we shall not find it difficultto understand how such a vast physical revolution as <strong>the</strong>Glacial epoch, <strong>with</strong> its attendant phenomena <strong>of</strong> elevations andsubsidences, icy winds, and sudden floods by <strong>the</strong> bursting <strong>of</strong> lakebarriers, might have led to <strong>the</strong> total extinction <strong>of</strong> a vast number<strong>of</strong> <strong>the</strong> most bulky forms <strong>of</strong> mammalia, while <strong>the</strong> less bulkywere able to survive, ei<strong>the</strong>rby greater hardiness <strong>of</strong> constitutionor bv becoming more or less modified. <strong>The</strong> result is apparent intrie comparatively small or moderate size <strong>of</strong> <strong>the</strong> species constituting<strong>the</strong> temperate fauna, in all parts <strong>of</strong> <strong>the</strong> globe.It is much to be regretted that no mammalian remains <strong>of</strong> earlierdate have been found in Australia, as we should <strong>the</strong>n see ifit isreally <strong>the</strong> case that marsupials have always formed its highesttype <strong>of</strong> mammalian life. At present its fossil fauna is chieflyinteresting to <strong>the</strong> zoologist, but throws little light on <strong>the</strong> past<strong>relations</strong> <strong>of</strong> this isolated country <strong>with</strong> o<strong>the</strong>r parts <strong>of</strong> <strong>the</strong> globe.Mammalian remains in<strong>the</strong> Secondary Formations.In <strong>the</strong> oldest Tertiary beds <strong>of</strong> Europe and North America, wehave (even <strong>with</strong> our present imperfect record) a rich and variedmammalian fauna. As compared <strong>with</strong> our living or recent highlyspecialized forms, it may be said to consist <strong>of</strong> generalised types ;but as compared <strong>with</strong> any primeval mammalian type, it must bepronounced highly specialized. Not only are such diversified


;160 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.groups as Carnivora, Perrissodactyle and Artiodactyle Ungulates,Primates, Chiroptera, Rodents, and Marsupials already wellmarked, but in many <strong>of</strong> <strong>the</strong>se <strong>the</strong>re is a differentiation intonumerous families and genera <strong>of</strong> diverse character.It is impossible<strong>the</strong>refore to doubt, that many peculiar forms <strong>of</strong> mammaliamust have lived long anterior to <strong>the</strong> Eocene period ;but <strong>the</strong>reis unfortunately a great gap in <strong>the</strong> record between <strong>the</strong> Eoceneand Cretaceous beds, and <strong>the</strong>se latter being for <strong>the</strong> most partmarine continue <strong>the</strong> gap as regards mammals over an enormouslapse <strong>of</strong> time. Yet far beyond both <strong>the</strong>se chasms in <strong>the</strong> UpperOolitic strata, remains <strong>of</strong> small mammalia have been foundagain, in <strong>the</strong> Stonesfield slate, a member <strong>of</strong> <strong>the</strong> Lower Oolite,o<strong>the</strong>r forms appear. <strong>The</strong>n comes <strong>the</strong> marine Lias formation<strong>with</strong> ano<strong>the</strong>r huge gap ; but beyond this again in <strong>the</strong> UpperTrias, <strong>the</strong> oldest <strong>of</strong> <strong>the</strong> secondary formations, mammalian teethhave been discovered inboth England and Germany, and <strong>the</strong>seare, as nearly as can be ascertained, <strong>of</strong> <strong>the</strong> same age as <strong>the</strong>Dromatlurium already noticed, from North America. <strong>The</strong>yhave bee:i named Microlestes, and show some resemblance tothose <strong>of</strong> <strong>the</strong> West Australian Myrmecobius. In <strong>the</strong> Ooliticstrata numerous small jawbones have been found, which haveserved to characterise eight genera, all <strong>of</strong> which are believed tohave been Marsupials, and in some <strong>of</strong> <strong>the</strong>m a resemblance can betraced to some <strong>of</strong> <strong>the</strong> smaller living Australian species.however, are mere indications<strong>The</strong>se,<strong>of</strong> <strong>the</strong> number <strong>of</strong> mammalia thatmust have lived in <strong>the</strong> secondary period, so long thought to beexclusively " <strong>the</strong> age <strong>of</strong> reptiles ;" and <strong>the</strong> fact that <strong>the</strong> few yetfound are at all comparable <strong>with</strong> such specialised forms as stillexist, must convince us, that we shall have to seek far beyondeven <strong>the</strong> earliest <strong>of</strong> <strong>the</strong>se remains, for <strong>the</strong> first appearance <strong>of</strong> <strong>the</strong>mammalian type <strong>of</strong> vertebrata.Extinct Birds.Compared <strong>with</strong> those <strong>of</strong> mammalia, <strong>the</strong> remains <strong>of</strong> birds areexceedingly scarce in Europe and America ;and from <strong>the</strong> wanderin"habits <strong>of</strong> so many <strong>of</strong> this class, <strong>the</strong>y are <strong>of</strong> much less value


;chap, viii.] BIRDS. . 161as indications <strong>of</strong> past changes in physical geography. A largeproportion <strong>of</strong> <strong>the</strong> remains belong to aquatic or wading types,and as <strong>the</strong>se have now <strong>of</strong>ten a world-wide range, <strong>the</strong> occurrence<strong>of</strong> extinct forms can have little bearing on our present inquiry.<strong>The</strong>re are, however, a few interesting cases<strong>of</strong> extinct land-birdsbelonging to groups now quite strangers to <strong>the</strong> country in which<strong>the</strong>y are found ; and o<strong>the</strong>rs scarcely less interesting, in whichgroups now peculiar to certain areas are shown to have beenpreceded by allied species or genera <strong>of</strong> gigantic size.Palcearctic Region and N. India.—In <strong>the</strong> caves and o<strong>the</strong>rPost-Pliocene deposits <strong>of</strong> <strong>the</strong>se countries, <strong>the</strong> remains <strong>of</strong> birdsalmost all belong to genera now inhabiting <strong>the</strong> same districts.Almost <strong>the</strong> only exceptions are, <strong>the</strong> great auk and <strong>the</strong>capercailzie, already mentioned as being found in <strong>the</strong> Danishmounds ; <strong>the</strong> latter bird, <strong>with</strong> Tetrao albus, in Italian cavernsand a species <strong>of</strong> pheasant (Phasianus) said to have occurred in<strong>the</strong> Post- Pliocene <strong>of</strong> France, considerably west <strong>of</strong> <strong>the</strong> existingrange <strong>of</strong> <strong>the</strong> genus in a wild state.In <strong>the</strong> preceding Pliocene deposits, but few remains havebeen found, and all <strong>of</strong> existing genera but one, a gallinaceousbird (Gallus bravardi) allied to <strong>the</strong> domestic fowl and peacock.<strong>The</strong> Miocene beds <strong>of</strong> France and Central Europe have producedmany more remains <strong>of</strong>birds, but <strong>the</strong>se, too, are mostly <strong>of</strong>existing European genera, though <strong>the</strong>re are some notable exceptions.Along <strong>with</strong> forms undistinguishable from crows (Corvus),shrikes (Lanius),wagtails (Motacilla), and woodpeckers (Picus),are found remains allied to <strong>the</strong> Oriental edible-nest swift (Collocalia)and Trogon ; a parrot resembling <strong>the</strong> African genusPsittacus ; an extinct form Necrornis, perhaps allied to <strong>the</strong>plantain-eaters (Musophaga) ;Homalophus, doubtfully allied towoodpeckers, and Limnatornis to <strong>the</strong> hoopoes. <strong>The</strong> gallinaceousbirds arerepresented by three species <strong>of</strong> pheasants, somevery close to <strong>the</strong> domesticated species ; Palccoperdix allied to <strong>the</strong>partridges ; and Palceortyx, small birds allied to <strong>the</strong> Americangenus Ortyx, but <strong>with</strong> larger wings. <strong>The</strong>re are also species <strong>of</strong>Pterocles allied to living birds, and a small pigeon. <strong>The</strong>re arenumerous living genera <strong>of</strong> Accipitres ; such as eagle {Aquila),


;162 . DISTRIBUTION OF EXTINCT ANIMALS. [part ii.kite (Milvus), eagle-owl {Bubo), and screech-owl (Stria) ; <strong>with</strong><strong>the</strong> African secretary-bird (Serpentarius), and some extinct forms,as Palceocercus, Palceohierix and Palceetus.Aquatic and wading birds were abundant, including numerousrails, bustards, herons, sandpipers, gulls, divers, and pelicans.<strong>The</strong>re were also many ducks, some allied to <strong>the</strong> genus Denclrocygna;<strong>the</strong> Oriental genus <strong>of</strong> storks, Lcptoptilus ; lbidipodia, aremarkable form allied to Ibis and Ciconia ; Elornis, nearLimosa; Pelagornis, a large bird allied to gannets and pelicans ;Hydrornis, allied to <strong>the</strong> ducks and petrels ; DolicJwpterus, alliedto plovers. Perhaps <strong>the</strong> most interesting <strong>of</strong> <strong>the</strong>se extinct birdsare, however, <strong>the</strong> flamingoes, represented by forms hardly distinguishablefrom living species, and by one extinct genus Palcelodus,which had very long toes, and probably walked on aquaticplants like <strong>the</strong> tropical jacanas.<strong>The</strong> Miocene beds <strong>of</strong> North India have furnished few birds<strong>the</strong> only one <strong>of</strong> <strong>geographical</strong> interest being an extinct species<strong>of</strong>ostrich, not very different from that now inhabiting Arabia.On <strong>the</strong> whole, <strong>the</strong> birds <strong>of</strong> Europe at this period were verylike those now living, <strong>with</strong> <strong>the</strong> addition <strong>of</strong> a few tropicalforms.<strong>The</strong>se latter were, however, perhaps more numerous and importantthan <strong>the</strong>y appear to be, as <strong>the</strong>y belong to inland and foresthauntingtypes, which would not be so frequently preserved as<strong>the</strong> marsh and lake-dwelling species. Taking this into consideration,<strong>the</strong> assemblage <strong>of</strong> Miocene birds accords well <strong>with</strong>what we know <strong>of</strong> <strong>the</strong> mammalian fauna. We have <strong>the</strong> sameindications <strong>of</strong> a luxuriant vegetation and subtropical climate,and <strong>the</strong> same appearance <strong>of</strong> Oriental and especially <strong>of</strong> Africantypes. Trogon is perhaps <strong>the</strong> most interesting <strong>of</strong> all <strong>the</strong> formsyet discovered, since itfurnishes us <strong>with</strong> a central point whence<strong>the</strong> living trogons <strong>of</strong> Asia, Africa, and South America mighthave diverged.In <strong>the</strong> Eocene we find ourselves almost wholly among extinctforms <strong>of</strong> birds. <strong>The</strong> earliest known Passerine bird is heremet <strong>with</strong>, in Protornis, somewhat similar to a lark, found in<strong>the</strong> Lower Eocene <strong>of</strong> Switzerland ;while ano<strong>the</strong>r Passerine form,Palwgithalus, and one allied to <strong>the</strong> nuthatch {Sitta), have been


chap, vm.] BIRDS. 163discovered in <strong>the</strong> Upper Eocene <strong>of</strong> Paris.Picariae <strong>of</strong> equal antiquityare found. Cryptomis, from <strong>the</strong> Paris Eocene, and Halcyornisfrom <strong>the</strong> Lower Eocene <strong>of</strong> <strong>the</strong> Isle <strong>of</strong> Sheppey, wereboth allied to kingfishers ; while a form allied to Centropus agenus <strong>of</strong> cuckoos, or,as Milne -Edwards thinks, to <strong>the</strong> MadagascarLeptosomus, has been found in <strong>the</strong> Upper Eocene <strong>of</strong>.France.Several Accipitrcs <strong>of</strong> somewhat doubtful affinities have beenfound in <strong>the</strong> same country; while Lithomis, from <strong>the</strong> LowerEocene <strong>of</strong> <strong>the</strong> Isle <strong>of</strong> Sheppey, was a small vulturine bird supposedto be allied to <strong>the</strong> American group, Catliartcs. Among<strong>the</strong> waders, some extinct forms <strong>of</strong> plovers have been found, and agenus (Agnoptems), allied to <strong>the</strong> flamingoes ; while <strong>the</strong>re aremany swimming birds, such as pelicans, divers, and severalextinct types <strong>of</strong> doubtful affinities. Most intersting <strong>of</strong> all is aportion <strong>of</strong> a cranium discovered in <strong>the</strong> Lower Eocene <strong>of</strong> Sheppey,and lately pronounced by Pr<strong>of</strong>essor Owen to belong to alarge Struthious bird, allied to <strong>the</strong> New T Zealand Dinornis andalso perhaps to <strong>the</strong> ostrich. Ano<strong>the</strong>r gigantic bird is <strong>the</strong> Gastornis,from <strong>the</strong> Lower Eocene <strong>of</strong> Paris, which was as large as anostrich, but which is believed to have been a generalised type,allied to wading and swimming birds as well as to <strong>the</strong> Struthiones.Beyond this epoch we have no remains <strong>of</strong> birds in Europeanstrata till we come to <strong>the</strong> wonderful Archceoptcryx from <strong>the</strong>Upper Oolite <strong>of</strong> Bavaria ; a bird <strong>of</strong> a totally new type, <strong>with</strong> abony tail, longer than <strong>the</strong> body, each vertebra <strong>of</strong> which carrieda pair <strong>of</strong> diverging fea<strong>the</strong>rs.North America.—A number <strong>of</strong>bird-remains have lately beenfound in <strong>the</strong> rich Tertiary and Cretaceous deposits <strong>of</strong> <strong>the</strong> UnitedStates ; but here, too, comparatively few are terrestrial forms.No Passerine bird has yet been found. <strong>The</strong> Picariae are representedby Uintornis, an extinct form allied to woodpeckers, from<strong>the</strong> Eocene <strong>of</strong> Wyoming. Species <strong>of</strong> turkey {Melcagris) occurin <strong>the</strong> Post-Pliocene and as far back as <strong>the</strong> Miocene strata,showing that this interesting type isa true denizen <strong>of</strong> temperateNorth America. <strong>The</strong> o<strong>the</strong>r birds are, Accipitrcs; waders andaquatics <strong>of</strong> existing genera ;and a number <strong>of</strong> extinct forms <strong>of</strong><strong>the</strong> two latter orders—such as, Alctornis an Eocene wader;


;164 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.Palceotringa, allied to <strong>the</strong> sandpipers, and Telmatobms to <strong>the</strong>rails, both Cretaceous ;<strong>with</strong> Graculavus, allied to GraculusLaornis allied to <strong>the</strong> swans ; Hcspcrornis a gigantic diver ; andIcthyornis a very low form, <strong>with</strong> biconcave vertebra,such as areonly found in fishes and some reptiles—also from Cretaceousdeposits.South America.—<strong>The</strong> caverns <strong>of</strong> Brazil produced thirty-fourspecies <strong>of</strong> birds, most <strong>of</strong> <strong>the</strong>m referable to Brazilian genera, andmany to still existing species. <strong>The</strong> most interesting were twospecies <strong>of</strong> American ostrich {Rhea), one larger than ei<strong>the</strong>r <strong>of</strong><strong>the</strong> living species ; a large turkey-buzzard (Cathartes) ; a newspecies <strong>of</strong> <strong>the</strong> very isolated South American genus Opisthocomus ;and a Cariama, or allied new genus. *Madagascar and <strong>the</strong>Mascarene Islands.—We have here onlyevidence <strong>of</strong> birds that have become extinct in <strong>the</strong> historicalperiod or very little earlier.First we have a group <strong>of</strong> birds incapable<strong>of</strong> flight, allied to pigeons, but forming a separate family,Dididoz ;and which, so far as we yet know, inhabited Mauritius,Eodriguez, and probably Bourbon. Aphanapteryx, an extinctgenus <strong>of</strong> rails, inhabited Mauritius ; and ano<strong>the</strong>r genus, {Erythromachus),Rodriguez. A large parrot, said by Pr<strong>of</strong>. Milne Edwardsto be allied to Ara and Microglossia, also inhabited Mauritiusand ano<strong>the</strong>r allied to Eclectus, <strong>the</strong> island <strong>of</strong> Rodriguez. None<strong>of</strong> <strong>the</strong>se have been found in Madagascar ;but a gigantic Struthiousbird, JEpyomis, forming a peculiar family distinct both from<strong>the</strong> ostriches <strong>of</strong> Africa and <strong>the</strong> Dinornis <strong>of</strong> New Zealandinhabited that island ; and <strong>the</strong>re is reason to believe thatthis may have lived less than 200 years ago.New Zealand,—A number <strong>of</strong> extinct Struthious birds, formingtwo families, Dinomithidce and Palapterygidoe, have beenfound in New Zealand. Some were <strong>of</strong> gigantic size. <strong>The</strong>y seemallied both to <strong>the</strong> living Apteryx <strong>of</strong> New Zealand and <strong>the</strong> emu<strong>of</strong> Australia. <strong>The</strong>y are quite recent, and some <strong>of</strong> <strong>the</strong>m haveprobably lived <strong>with</strong>in <strong>the</strong> last few centuries. Remains <strong>of</strong>Dinornis have also been found in a Post-Pliocene deposit inQueensland, N. E. Australia x—a very important discovery, asit1Trans. Zool. Soc. <strong>of</strong> London, vol. viii. p. 381.


chap, viii.] TERTIARY REPTILES. 1G5gives support to <strong>the</strong> <strong>the</strong>ory <strong>of</strong> a great eastward extension <strong>of</strong>Australia in Tertiary times.Extinct Teetiaey Eepttles.<strong>The</strong>se will not occupy us long, as no very great number areknown, and most <strong>of</strong> <strong>the</strong>m belong to a few principal forms <strong>of</strong>comparatively little <strong>geographical</strong> interest.Tortoises are perhaps <strong>the</strong> most abundant <strong>of</strong> <strong>the</strong> Tertiaryreptiles. <strong>The</strong>y are numerous in <strong>the</strong> Eocene and Mioceneformations both in Europe and North America. <strong>The</strong> 'generaEmys and Trionyx abound in both countries, as well as in <strong>the</strong>Miocene <strong>of</strong> India. Land tortoises occur in <strong>the</strong> Eocene <strong>of</strong> NorthAmerica and in <strong>the</strong> Miocene <strong>of</strong> Europe and India, where <strong>the</strong>huge Colossochelys, twelve feet long, has been found. In <strong>the</strong>Pliocene deposits <strong>of</strong> Switzerland <strong>the</strong> living American genusChelydra has been met <strong>with</strong>. <strong>The</strong>se facts, toge<strong>the</strong>r <strong>with</strong> <strong>the</strong>occurrence <strong>of</strong> a living species in <strong>the</strong> Miocene <strong>of</strong> India, showthat this order <strong>of</strong> reptiles is <strong>of</strong> great antiquity, and that most<strong>of</strong> <strong>the</strong> genera once had a wider range than now.Crocodiles, allied to <strong>the</strong> three forms now characteristic <strong>of</strong>India, Africa, and America, have been found in <strong>the</strong> Eocene <strong>of</strong>our own country, and several species <strong>of</strong> Crocodihis have occurredin beds <strong>of</strong> <strong>the</strong> same age in North America.Lizards are very ancient, many small terrestrial formsoccurring in all <strong>the</strong> Tertiary deposits. A species <strong>of</strong> <strong>the</strong> genusChamceleo is recorded from <strong>the</strong> Eocene <strong>of</strong> North America, toge<strong>the</strong>r<strong>with</strong> several extinct genera.Snakes were well developed in <strong>the</strong> Eocene period, whereremains <strong>of</strong> several have been found which must have been fromtwelve to twenty feet long. An extinct species <strong>of</strong> true viper hasoccurred in <strong>the</strong> Miocene <strong>of</strong> France, and one <strong>of</strong> <strong>the</strong> Pythonideein <strong>the</strong> Miocene brown coal <strong>of</strong> Germany.Batrachia occur but sparingly in a fossil state in <strong>the</strong> Tertiarydeposits. <strong>The</strong> most remarkable is <strong>the</strong> large Salamander{Andreas) from <strong>the</strong> Upper Miocene <strong>of</strong> Switzerland, which


1GG DISTRIBUTION OF EXTINCT ANIMALS. [part ii.is allied to <strong>the</strong> Menopoma living in North America. Species<strong>of</strong> frog (Bana), and Palceophryus an extinct genus <strong>of</strong> toads,have been found in <strong>the</strong> Miocene deposits <strong>of</strong> Germany andSwitzerland.Fresh water fish are almost unknown in <strong>the</strong> Tertiarydeposits <strong>of</strong> Europe, although most <strong>of</strong> <strong>the</strong> families and somegenera <strong>of</strong> living marine fish are represented from <strong>the</strong> Eocenedownwards.Antiquity <strong>of</strong> <strong>the</strong> Geneka <strong>of</strong> Insects.Fossil insects are far too rarely found, to aid us in our determination<strong>of</strong> difficult questions <strong>of</strong> <strong>geographical</strong> <strong>distribution</strong> ; but indiscussing <strong>the</strong>se questions it will be important to know, whe<strong>the</strong>rwe are to look upon <strong>the</strong> existing generic forms <strong>of</strong> insects as <strong>of</strong>gieat or small antiquity, compared <strong>with</strong> <strong>the</strong> higher vertebrates ;and to decide this question <strong>the</strong> materials at our command areample.<strong>The</strong> conditions requisite for <strong>the</strong> preservation <strong>of</strong> insects in afossil state are no doubt very local and peculiar ;<strong>the</strong> result being,that it is only at long intervals in <strong>the</strong> geological record that wemeet <strong>with</strong> remains <strong>of</strong> insects in a recognisable condition. Noneappear to have been found in <strong>the</strong> Pliocene formation ; but in <strong>the</strong>Upper Miocene <strong>of</strong> (Eninghen in Switzerland, associated <strong>with</strong> <strong>the</strong>wonderfully rich fossil flora, are found immense quantities <strong>of</strong>insects. Pr<strong>of</strong>. Heer examined more than 5,000 specimens belongingto over 800 species, and many have been found in o<strong>the</strong>rlocalities in Switzerland ;so that more than 1,300 species <strong>of</strong>Miocene insects have now been determined.Most <strong>of</strong> <strong>the</strong> ordersare represented, but <strong>the</strong> beetles (Coleoptera) are far <strong>the</strong> mostabundant.Almost all belong to existing genera, and <strong>the</strong> majority<strong>of</strong> <strong>the</strong>se genera now inhabit Europe, only three or four beingexclusively Indian, African, or American.In <strong>the</strong> Lower Miocene <strong>of</strong> Croatia <strong>the</strong>re is ano<strong>the</strong>r rich deposit<strong>of</strong> insects, somewhat more tropical in character, comprisinglarge white-ants and dragon-flies differently marked from any


——;chap, vm.] ANTIQUITY OF INSECTS. 167now inhabiting Europe. A butterfly is also well preserved, <strong>with</strong>all <strong>the</strong> markings <strong>of</strong> <strong>the</strong> wings ;and it seems to be a Junonia, atropical genus, though it may be a Vanessa, which is European,but <strong>the</strong> fossil most resembles Indian species <strong>of</strong> Junonia.<strong>The</strong> Eocene formations seem to have produced no insectremains ;but <strong>the</strong>y occur again in <strong>the</strong> Upper Cretaceousat Aix-la-Chapelle, where two butterflies have been found,Cyllo sepulta and Satyrites Reynesii, both belonging to <strong>the</strong> Satyridse,and <strong>the</strong> former to a genus now spread over Africa, India,and Australia.A little earlier, in <strong>the</strong> Wealden formation <strong>of</strong> our own country,numerous insects have been found, principally dragon flies (Libellula,JEshna) ; aquatic Hemiptera (Velia Hydrometrci) ; cricketscockroaches, and cicadas, <strong>of</strong> familiar types.Fur<strong>the</strong>r back in <strong>the</strong> Upper Oolite <strong>of</strong> Bavaria—which produced<strong>the</strong> wonderful long-tailed bird, Archceopteryx—insects <strong>of</strong> all ordershave been found, including a moth referred to <strong>the</strong> existing genusSphinx.In <strong>the</strong> Lower Oolite <strong>of</strong> Oxfordshire many fossil beetles havebeen found whose affinities are shown by <strong>the</strong>ir names :Buprestidium,Curculionidium, Blapsidium, Mclolonthidium, and Prionidium; a wing <strong>of</strong> a butterfly has also been found, allied to <strong>the</strong>Brassolidse now confined to tropical America, and named Palmontinaoolitica.Still more remote are <strong>the</strong> insects <strong>of</strong> <strong>the</strong> Lias <strong>of</strong> Gloucestershire,yet <strong>the</strong>y too can be referred to well-known family typesCarabidse, Melolonthidse, Telephoridae,Elateridse, and Curculionidse,among beetles;Gryllidse and Blattidae among Orthoptera;<strong>with</strong> Libelhda, Agrion, JEshna, Ephemera, and some extinctgenera. When we consider that almost <strong>the</strong> only vertebruta <strong>of</strong>this period were huge Saurian repliles like <strong>the</strong> Icthyosaurus,Plesiosaurtts, and Dinosaurus, <strong>with</strong> <strong>the</strong> flying Pterodactylesand that <strong>the</strong> great mass <strong>of</strong> our existing genera,and even families,<strong>of</strong> fish and reptiles had almost certainly not come into existence,we see at once that types <strong>of</strong> insect-form are, proportionately,far more ancient. At this remote epoch we find <strong>the</strong> chief familytypes (<strong>the</strong> genera <strong>of</strong> <strong>the</strong> time <strong>of</strong> Linnaeus) perfectly differentiated


168 DISTRIBUTION OF EXTINCT ANIMALS. [part II.and recognisable.It is only when we go fur<strong>the</strong>r back still, into<strong>the</strong> Palseozoic formations, that <strong>the</strong> insect forms begin to show thatgeneralization <strong>of</strong> type which renders it impossible to classify<strong>the</strong>m in any existing groups. Yet even in <strong>the</strong> coal formation <strong>of</strong>Nova Scotia and Durham, <strong>the</strong> fossil insects are said by competententomologists to be " allied to Ephemera" " near Blatta,"Phasmidce ; "" nearand in deposits <strong>of</strong> <strong>the</strong> same age at Saarbruck nearTreves, a well-preserved wing <strong>of</strong> a grasshopper or locust has beenfound, as well as a beetle referred to <strong>the</strong> Scarabeidse. Moreremarkable, however, is <strong>the</strong> recent discovery in <strong>the</strong> carboniferousshales <strong>of</strong> Belgium, <strong>of</strong> <strong>the</strong> clearly-defined wing <strong>of</strong> a large moth(Breyeria borinensis), closely resembling some <strong>of</strong> <strong>the</strong> Saturniidae ;so that we have now all <strong>the</strong> chief orders <strong>of</strong> Insects—includingthose supposed to be <strong>the</strong> most highly developed and <strong>the</strong> mostrecent—well represented at this very remote epoch. Even <strong>the</strong>oldest insects, from <strong>the</strong> Devonian rocks <strong>of</strong> North America, canmostly be classed as Neuroptera or Myriapoda, but appear t<strong>of</strong>orm new families.We may consider it, <strong>the</strong>refore, as proved, that many <strong>of</strong> <strong>the</strong>larger and more important genera <strong>of</strong> insects date back to <strong>the</strong>beginning <strong>of</strong> <strong>the</strong> Tertiary period, or perhaps beyond it ;but <strong>the</strong>family types are far older, and must have been differentiated veryearly in <strong>the</strong> Secondary period, while some <strong>of</strong> <strong>the</strong>m perhaps goback to Palseozoic times. <strong>The</strong> great comparative antiquity <strong>of</strong><strong>the</strong> genera is however <strong>the</strong> important fact for us, and we shallhave occasion <strong>of</strong>ten to refer to it, in endeavouring to ascertain<strong>the</strong> true bearing <strong>of</strong> <strong>the</strong> facts <strong>of</strong> insect <strong>distribution</strong>, as elucidatingor invalidating <strong>the</strong> conclusions arrived at from a <strong>study</strong> <strong>of</strong> <strong>the</strong><strong>distribution</strong> <strong>of</strong> <strong>the</strong> higher <strong>animals</strong>.Antiquity <strong>of</strong> <strong>the</strong> Genera <strong>of</strong> Land and Fresh-WaterShells.<strong>The</strong> remains <strong>of</strong> land and fresh-water shells are not much morefrequent than those <strong>of</strong> insects. Like <strong>the</strong>m, too, <strong>the</strong>ir forms arevery stable, continuing unchanged through several geological


—chap, viii.] ANTIQUITY OF LAND SHELLS. 169periods. In <strong>the</strong> Pliocene and Miocene formations, most <strong>of</strong> <strong>the</strong>shells are very similar to living species, and some are quite identical.In <strong>the</strong> Eocene we meet <strong>with</strong> ordinary forms <strong>of</strong> <strong>the</strong> generaHelix, Clausilia, Pupa, Bulimus, Glandina, Cyclostoma, Megalostoma,Planorbis, Paludina and Limncea, some resembling Europeanspecies, o<strong>the</strong>rs more like tropical forms. A British Eocenespecies <strong>of</strong> Helix is still living in Texas ;and in <strong>the</strong> South <strong>of</strong> Franceare found species <strong>of</strong> <strong>the</strong> Brazilian sub-genera Megaspira andAnastoma. In <strong>the</strong> secondary formation no true land shells havebeen found, but fresh water shells are tolerably abundant, andalmost all are still <strong>of</strong> living forms. In <strong>the</strong> Wealden (LowerCretaceous) and Purbeck (Upper Oolite) are found Unio, Melania,Paludina, Planorbis, and Limncea ; while <strong>the</strong> last named genusoccurs even in <strong>the</strong> Lias.<strong>The</strong> notion that land shells were really not in existence during<strong>the</strong> secondary period is, however, proved to be erroneous by <strong>the</strong>startling discovery, in <strong>the</strong> Palaeozoic coal measures <strong>of</strong> Nova Scotia,<strong>of</strong> two species <strong>of</strong> Helicidae, both <strong>of</strong> living genera Pupa vetusta,and Zonites priscus. <strong>The</strong>y have been found in <strong>the</strong> hollow trunk<strong>of</strong> a Sigillaria, and in great quantities in a bed full <strong>of</strong> Stigmarianrootlets. <strong>The</strong> most minute examination detects no importantdifferences <strong>of</strong> form or <strong>of</strong> microscopic structure, between <strong>the</strong>seshells and living species <strong>of</strong> <strong>the</strong> same genera !<strong>The</strong>se mollusca were<strong>the</strong> contemporaries <strong>of</strong> Labyrinthodonts and strange Ganoid fishes,which formed almost <strong>the</strong> whole vertebrate fauna. This unexpecteddiscovery renders it almost certain, that numbers <strong>of</strong> o<strong>the</strong>rexisting genera, <strong>of</strong> which we have found no traces, lived <strong>with</strong><strong>the</strong>se two through <strong>the</strong> whole secondary period ; and we are thusobliged to assume as a probability, that any particular genus haslived through a long succession <strong>of</strong> geological ages. In estimating<strong>the</strong> importance <strong>of</strong> any peculiarities or anomalies in <strong>the</strong><strong>geographical</strong> <strong>distribution</strong> <strong>of</strong> land shells as compared <strong>with</strong> <strong>the</strong>higher vertebrates, we shall, <strong>the</strong>refore, have to keep this possible,and even probable high antiquity, constantly in mind.We have now concluded our sketch <strong>of</strong> Tertiary Palaeontologyas a preparation for <strong>the</strong> intelligent <strong>study</strong> <strong>of</strong> <strong>the</strong> GeographicalVol. I.—13


170 DISTRIBUTION OF EXTINCT ANIMALS. [part ii.Distribution <strong>of</strong> Land Animals ; and however imperfectly <strong>the</strong> taskhas been performed, <strong>the</strong> reader will at all events have been convincedthat some such preliminary investigation is an essentialand most important part <strong>of</strong> our work.So much <strong>of</strong> palaeontologyis at present tentative and conjectural, that in combining <strong>the</strong>information derived from numerous writers, many errors <strong>of</strong> detailmust have been made. <strong>The</strong> main conclusions have, however, beendrawn from as large a basis <strong>of</strong> facts as possible ;and althoughfresh discoveries may show thatour views as to <strong>the</strong> past history<strong>of</strong> some <strong>of</strong> <strong>the</strong> less important genera or families are erroneous,<strong>the</strong>y can hardly invalidate our results to any important degree,ei<strong>the</strong>r as regards <strong>the</strong> intercommunications between separateregions in <strong>the</strong> various geological epochs, or as to <strong>the</strong> centresfrom which some <strong>of</strong> <strong>the</strong> more important groups have been dispersed.


PART III.ZOOLOGICALGEOGRAPHY:AREVIEW OF TEE CHIEF FORMS OF ANIMAL LIFE IN TEESEVERAL REGIONS AND SUB-REGIONS, WITH THE INDICA-TIONS THEY AFFORD OF GEOGRAPHICAL MUTATIONS.


CHAPTER IX.THE ORDER OF SUCCESSION OF THE REGIONS.— COSMOPOLITANGROUPS OF ANIMALS.—TABLES OF DISTRIBUTION.Having discussed, in our First Part, such general and preliminarymatters as are necessary to a proper comprehension <strong>of</strong> oursubject ; and having made ourselves acquainted, in our SecondPart, <strong>with</strong> <strong>the</strong> most important results <strong>of</strong> Palaeontology, we nowcome to our more immediate subject, which we propose to treatfirst under its <strong>geographical</strong> aspect. Taking each <strong>of</strong> our sixregions in succession, we shall point out in some detail <strong>the</strong> chiefzoological features <strong>the</strong>y present, as influenced by climate, vegetation,and o<strong>the</strong>r physical features. We shall <strong>the</strong>n treat each<strong>of</strong> <strong>the</strong> sub-regions by itself, as well as such <strong>of</strong> <strong>the</strong> islands oro<strong>the</strong>r sub-divisions as present features <strong>of</strong> special interest; endeavouringto ascertain <strong>the</strong>ir true <strong>relations</strong> to each o<strong>the</strong>r, and <strong>the</strong>more important changes <strong>of</strong> physical geography that seem necessaryto account for <strong>the</strong>ir present zoological condition.Order <strong>of</strong> Succession <strong>of</strong> <strong>the</strong> Regions.— We may here explain<strong>the</strong> reason for taking <strong>the</strong> several regions in a different successionfrom that in which <strong>the</strong>y appear in <strong>the</strong> tabular or diagrammaticheadings to each family, in <strong>the</strong> Fourth, and concluding part <strong>of</strong>this work. It will have been seen, by our examination <strong>of</strong> extinct<strong>animals</strong> (and it will be made still clearer during our <strong>study</strong> <strong>of</strong><strong>the</strong> several regions) that all <strong>the</strong> chief types <strong>of</strong> animal life appearto have originated in <strong>the</strong> great north temperate or nor<strong>the</strong>rncontinents ; while <strong>the</strong> sou<strong>the</strong>rn continents—now represented by


—174 ZOOLOGICAL GEOGRAPHY. |>art in.South America, Australia, and South Africa <strong>with</strong> Madagascarhave been more or less completely isolated, during long periods,both from <strong>the</strong> nor<strong>the</strong>rn continent and from each o<strong>the</strong>r. <strong>The</strong>selatter countries have, however, been subject to more or less immigrationfrom <strong>the</strong> north during rare epochs <strong>of</strong> approximationto, or partial union <strong>with</strong> it. In <strong>the</strong> nor<strong>the</strong>rn, more extensive, andprobably more ancient land, <strong>the</strong> process <strong>of</strong> development hasbeen more rapid, and has resulted in more varied and highertypes ; while <strong>the</strong> sou<strong>the</strong>rn lands, for <strong>the</strong> most part, seem to haveproduced numerous diverging modifications <strong>of</strong> <strong>the</strong> lower grades<strong>of</strong> organization, <strong>the</strong> original types <strong>of</strong> which <strong>the</strong>y derived ei<strong>the</strong>rfrom <strong>the</strong> north, or from some <strong>of</strong> <strong>the</strong> ancient continents in Mesozoicor Palaeozoic times. Hence those curious resemblances in<strong>the</strong> fauna <strong>of</strong> South America, Australia, and, to a less extent,Madagascar, which have led to a somewhat general belief that<strong>the</strong>se distant countries must at one time or o<strong>the</strong>r have beenunited ; a belief which, after a careful examination <strong>of</strong> all <strong>the</strong>facts, does not seem to <strong>the</strong> author <strong>of</strong> this work to be wellfounded. Oh <strong>the</strong> o<strong>the</strong>r hand, <strong>the</strong>re is <strong>the</strong> most satisfactoryevidence that each sou<strong>the</strong>rn region has been more or lessclosely united (during <strong>the</strong> tertiary or later secondary epoch)<strong>with</strong> <strong>the</strong> great nor<strong>the</strong>rn continents, leading to numerous resemblancesand affinities in <strong>the</strong>ir productions.In endeavouring to present at a glance in <strong>the</strong> most convenientmanner, <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> families in <strong>the</strong> several regionsand sub-regions, it was necessary to arrange <strong>the</strong>m, so that thosewhose <strong>relations</strong> to each o<strong>the</strong>r were closest should stand side byside ;<strong>the</strong> first and last being those between which <strong>the</strong> <strong>relations</strong>were least numerous and least important. Influenced by <strong>the</strong>usual opinions as to <strong>the</strong> <strong>relations</strong> between Australia and SouthAmerica, <strong>the</strong> series was at first begun <strong>with</strong> <strong>the</strong> Nearetic, andterminated <strong>with</strong> <strong>the</strong> Australian and Neotropical regions ; and itwas not till <strong>the</strong> whole <strong>of</strong> <strong>the</strong> vertebrate families had been gonethrough, and <strong>the</strong>ir <strong>distribution</strong> carefully studied, that <strong>the</strong>se lasttwo regions were seen to be really wider apart than any o<strong>the</strong>rs<strong>of</strong> <strong>the</strong> series. It was <strong>the</strong>refore decided to alter <strong>the</strong> arrangement,beginning <strong>with</strong> <strong>the</strong> Neotropical, and ending <strong>with</strong> <strong>the</strong> Australian


;chap, ix.] ORDER OF SUCCESSION OF THE REGIONS. 175regions ; and a careful inspection <strong>of</strong> <strong>the</strong> diagrams <strong>the</strong>mselves,taken in <strong>the</strong>ir entirety, will, it is believed, show that this is<strong>the</strong> most natural plan, and most truly exhibits <strong>the</strong> <strong>relations</strong> <strong>of</strong><strong>the</strong> several regions.In <strong>the</strong> portion <strong>of</strong> our work now commencing, we are not,however, by any means bound to begin at ei<strong>the</strong>r end <strong>of</strong> thisseries. Each region is studied by itself, but reference will <strong>of</strong>tenhave to be made to all <strong>the</strong> o<strong>the</strong>r regions ;and wherever webegin, we must occasionally refer to facts which will be givenfur<strong>the</strong>r on. As, however, <strong>the</strong> great nor<strong>the</strong>rn continents form<strong>the</strong> central mass from which <strong>the</strong> sou<strong>the</strong>rn regions, as it were,diverge, and as <strong>the</strong> Pala?arctic region isboth more extensive andmuch better known than any o<strong>the</strong>r, it undoubtedly forms <strong>the</strong>most convenient starting-point for our proposed survey <strong>of</strong><strong>the</strong> zoological history <strong>of</strong> <strong>the</strong> earth. We thus pass from <strong>the</strong>better known to <strong>the</strong> less known—from Europe to Africa andtropical Asia, and <strong>the</strong>nce to Australia, completing <strong>the</strong> series <strong>of</strong>regions <strong>of</strong> <strong>the</strong> Eastern Hemisphere.Beginning again <strong>with</strong> <strong>the</strong>Neotropical region, we pass to <strong>the</strong> Nearctic, which has suchstriking <strong>relations</strong> <strong>with</strong> <strong>the</strong> preceding and <strong>with</strong> <strong>the</strong> Palsearcticregion, that it can only be properly understood by constantreference to both. We thus keep separate <strong>the</strong> Eastern and Westernhemispheres, which form, from our point <strong>of</strong> view, <strong>the</strong>most radical and most suggestivedivision <strong>of</strong> terrestrial faunasand as we are able to make this also <strong>the</strong> dividing point <strong>of</strong> ourtwo volumes, reference to <strong>the</strong> work will be <strong>the</strong>reby facilitated.Cosmopolitan Groups.— Before proceeding to sketch <strong>the</strong> zoologicalfeatures <strong>of</strong> <strong>the</strong> several Begions it will be well to noticethose family groups which belong to <strong>the</strong> earth as a whole, andwhich are so widely and universally distributed over itthat itwill be unnecessary, in some cases, to do more than refer to<strong>the</strong>m under <strong>the</strong> separate <strong>geographical</strong> divisions.<strong>The</strong> only absolutely cosmopolitan families <strong>of</strong> Mammalia arethose which are aerial or marine; and this is one <strong>of</strong> <strong>the</strong> strikingpro<strong>of</strong>s that <strong>the</strong>ir <strong>distribution</strong> has been effected by natural causes,and that <strong>the</strong> permanence <strong>of</strong> barriers is one <strong>of</strong> <strong>the</strong> chief


and176 ZOOLOGICAL GEOGRAPHY. [part hi.agencies in <strong>the</strong> limitation <strong>of</strong> <strong>the</strong>ir range. Even among <strong>the</strong>aerial bats, however, only one family-—<strong>the</strong> Vespertilionidae— istruly cosmopolitan, <strong>the</strong> o<strong>the</strong>rs having a more or less restrictedrange.Nei<strong>the</strong>r are <strong>the</strong> Cetacea necessarily cosmopolitan, most <strong>of</strong><strong>the</strong> families being restricted ei<strong>the</strong>r to warm or to cold seas ; butone family, <strong>the</strong> dolphins (Delphinidae), is truly so. This orderhowever will not require fur<strong>the</strong>r notice, as, being exclusivelymarine <strong>the</strong> groups do not enter into any <strong>of</strong> our terrestrialregions. <strong>The</strong> only o<strong>the</strong>r family <strong>of</strong> mammals that may be consideredto be cosmopolitan, is <strong>the</strong> Muridae (rats .mice) : yet<strong>the</strong>se are not entirely so, since none are known to be trulyindigenous in any part <strong>of</strong> <strong>the</strong> Australian region except Australiaitself.In <strong>the</strong> class <strong>of</strong> Birds, a number <strong>of</strong> families are cosmopolites,if we reckon as such all which are found in each region andsub-region ;but several <strong>of</strong> <strong>the</strong>se are so abundant in some parts,while <strong>the</strong>y are so sparingly represented in o<strong>the</strong>rs, that <strong>the</strong>ycannot fairly be considered so. We shall confine that term<strong>the</strong>refore, to such as, <strong>the</strong>re is reason to believe, inhabit everyimportant sub-division <strong>of</strong> each region. Such are, among <strong>the</strong>Passerine birds <strong>the</strong> crows (Corvidae), and swallows (Hiruudinidae);.among <strong>the</strong> Picariae <strong>the</strong> kingfishers (Alcedinidae) ; amongo<strong>the</strong>r Land birds <strong>the</strong> pigeons (Columbidae), grouse and partridges(Tetraonidae), hawks (Falconidae), and owls (Strigidae) ; among<strong>the</strong> Waders <strong>the</strong> rails (Kallidae), snipes (Scolopacidae), plovers(Charadriadae), and herons (Ardeidae) ;and among <strong>the</strong> Swimmers<strong>the</strong> ducks (Anatidae), gulls (Laridae), petrels (Procellariidae),pelicans (Pelecanidae), and grebes (Podicipidae).In <strong>the</strong> class <strong>of</strong> Reptiles <strong>the</strong>re are few absolutely cosmopolitanfamilies, owing to <strong>the</strong> scarcity <strong>of</strong> members <strong>of</strong> this group in someinsular sub-regions, such as New Zealand and <strong>the</strong> Pacific Islands.Those which are most nearly so are <strong>the</strong> Colubridae among snakes,and <strong>the</strong> Scincidae among lizards.<strong>The</strong>re isno cosmopolitan family <strong>of</strong> Amphibia, <strong>the</strong> true frogs(Eanidae) being <strong>the</strong> most widely distributed.Nei<strong>the</strong>r is any family <strong>of</strong> Freshwater Fishes cosmopolitan,<strong>the</strong> Siluridae, which have <strong>the</strong> widest range, being confined


—;chap, ix.] TABLES OF DISTRIBUTION. 177to warm regions, and becoming very scarce in <strong>the</strong> temperatezones.Among <strong>the</strong> Diurnal and Crepuscular Lepidoptera(butterfliesand sphinges) <strong>the</strong> following families are cosmopolitan :—Satyridse,Nymphalidse, Zygaenidse, Pieridse, Papilionidae, Hesperidae,Lycaenidee, and Sphingidse.Of <strong>the</strong> Coleoptera almost all, except some <strong>of</strong> <strong>the</strong> small andobscure families, are cosmopolitan.Of <strong>the</strong> terrestrial Mollusca, <strong>the</strong> Helicidae alone are true cosmopolites.Tables <strong>of</strong> Distribution <strong>of</strong> Families and Genera.—Having beenobliged to construct numerous tables <strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong>various groups for <strong>the</strong> purposes <strong>of</strong> <strong>the</strong> descriptive part <strong>of</strong> <strong>the</strong>work, I have thought it well to append <strong>the</strong> most important <strong>of</strong><strong>the</strong>m, in a convenient form, to <strong>the</strong> chapter on each region ; asmuch information will <strong>the</strong>reby be given, which can only beobtained from existing works at <strong>the</strong> cost <strong>of</strong> great labour. All<strong>the</strong>se tables are drawn up on a uniform plan, <strong>the</strong> same genericand family names being used in each ; and all are arranged in<strong>the</strong> same systematic order, so as to be readily comparable<strong>with</strong> each o<strong>the</strong>r. This, although it seems a simple and naturalthing to do, has involved a very great amount <strong>of</strong> labour, becausehardly two authors use <strong>the</strong> same names or follow <strong>the</strong> samearrangement. Hence comparison between <strong>the</strong>m is impossible, tillall <strong>the</strong>ir work has been picked to pieces, <strong>the</strong>ir synonymy unravelled,<strong>the</strong>ir differences accounted for, and <strong>the</strong> materials recastand this has to be done, not for two or three authors only, butfor <strong>the</strong> majority <strong>of</strong> those whose works have been consulted on<strong>the</strong> zoology <strong>of</strong> any part <strong>of</strong> <strong>the</strong> globe.Except in <strong>the</strong> two higher orders—Mammalia and Birdsmaterials do not exist for complete tables<strong>of</strong> <strong>the</strong> genera broughtdown to <strong>the</strong> present time. We have given <strong>the</strong>refore, first, acomplete table <strong>of</strong> all <strong>the</strong> families <strong>of</strong> Vertebrata and DiurnalLepidoptera found in each region, showing <strong>the</strong> sub-regions inwhich <strong>the</strong>y occur, and <strong>the</strong>ir range beyond <strong>the</strong> limits <strong>of</strong> <strong>the</strong>region. Families which are wholly peculiar to <strong>the</strong> region, or


;178 ZOOLOGICAL GEOGRAPHY. [part hi.very characteristic and almost exclusively confined to it, are initalics. <strong>The</strong> number prefixed to each family corresponds tothat <strong>of</strong> <strong>the</strong> series <strong>of</strong> families in <strong>the</strong> Fourth Part <strong>of</strong> this work,so that if fur<strong>the</strong>r information is required it can be readilyreferred to <strong>with</strong>out consulting <strong>the</strong> index. Names inclosed inparen<strong>the</strong>ses—( . . . ) thus—indicate families which only justenter a region from an adjacent one, to which <strong>the</strong>y properlybelong. <strong>The</strong> eye is thus directed to <strong>the</strong> more, and <strong>the</strong> less importantfamilies ;and a considerable amount <strong>of</strong> information asto <strong>the</strong> general features <strong>of</strong> <strong>the</strong> zoology <strong>of</strong> <strong>the</strong> region, isconveyedin <strong>the</strong> easiest manner.<strong>The</strong> tables <strong>of</strong> genera <strong>of</strong> Mammalia and Birds, are arranged ona somewhat different plan. Each genus is given under itsFamily and Order, and <strong>the</strong>y follow in <strong>the</strong>same succession in all<strong>the</strong> tables. <strong>The</strong> number <strong>of</strong> species <strong>of</strong> each genus, inhabitingbut in many<strong>the</strong> region, is given as nearly as can be ascertained ;cases this can only be a general approximation.<strong>of</strong> <strong>the</strong> genera <strong>with</strong>in <strong>the</strong> region, is<strong>The</strong> <strong>distribution</strong><strong>the</strong>n given <strong>with</strong> some detailand, lastly, <strong>the</strong> range <strong>of</strong> <strong>the</strong> genus beyond <strong>the</strong> region is givenin general terms, <strong>the</strong> words " Oriental," " Ethiopian," &c, beingused for brevity, to indicate that <strong>the</strong> genus occurs over a considerablepart <strong>of</strong> such regions. Genera which are restricted to<strong>the</strong> region (or which are very characteristic <strong>of</strong> it though justtransgressing its limits) are given in italics ;while those whichonly just enter <strong>the</strong> region from ano<strong>the</strong>r to which <strong>the</strong>y reallybelong, are enclosed in paren<strong>the</strong>ses— ( . . . ) thus. <strong>The</strong> generaare here numbered consecutively, in order that <strong>the</strong> number <strong>of</strong>genera in each family or each order, in <strong>the</strong> region, may be readilyascertained (by one process <strong>of</strong> subtraction), and thus comparisonsmade <strong>with</strong> o<strong>the</strong>r regions or <strong>with</strong> any o<strong>the</strong>r area. As <strong>the</strong>tables <strong>of</strong> birds would be swelled to an inconvenient length by<strong>the</strong> insertion in each region <strong>of</strong> all <strong>the</strong> genera <strong>of</strong> Waders andAquatics, most <strong>of</strong> which have a very wide range and wouldhave to be repeated in several or all <strong>the</strong> regions, <strong>the</strong>se havebeen omitted ; but a list has been given <strong>of</strong> such <strong>of</strong> <strong>the</strong> generaas are peculiar to, or highly characteristic <strong>of</strong> each region.As this is <strong>the</strong> first time that any such extensive tables <strong>of</strong>


chap, ix.] TABLES OF DISTRIBUTION. 179<strong>distribution</strong> have been constructed for <strong>the</strong> whole <strong>of</strong> <strong>the</strong> Mammaliaand Birds, <strong>the</strong>y must necessarily contain many errors <strong>of</strong>detail ; but <strong>with</strong> all <strong>the</strong>ir imperfections it is believed <strong>the</strong>y willprove very useful to naturalists, to teachers, and to all who takean intelligent interest in <strong>the</strong> wider problems <strong>of</strong> geography andnatural history.


;CHAPTER X.THE PALiEAECTICREGION.This region is<strong>of</strong> immense extent, comprising all <strong>the</strong> temperateportions <strong>of</strong> <strong>the</strong> great eastern continents. It thus extends from<strong>the</strong> Azores and Canary Islands on <strong>the</strong> west to Japan on <strong>the</strong>east, a distance not far short <strong>of</strong> half <strong>the</strong> circumference <strong>of</strong> <strong>the</strong>globe. Yet so great is <strong>the</strong> zoological unity <strong>of</strong> this vast tract,that <strong>the</strong> majority <strong>of</strong> <strong>the</strong> genera <strong>of</strong> <strong>animals</strong> in countries so farremoved as Great Britain and Nor<strong>the</strong>rn Japan are identical.Throughout its nor<strong>the</strong>rn half <strong>the</strong> animal productions <strong>of</strong> <strong>the</strong>Palaearctic region are very uniform, except that <strong>the</strong> vast elevateddesert-regions <strong>of</strong> Central Asia possess some characteristic formsbut in its sou<strong>the</strong>rn portion,we find a warm district at each ex-On <strong>the</strong> west we havetremity <strong>with</strong> somewhat contrasted features.<strong>the</strong> rich and luxuriant Mediterranean sub-region, possessing manypeculiar forms <strong>of</strong> life,characteristic <strong>of</strong> <strong>the</strong> Ethiopian region.as well as a few which are more especiallyOn <strong>the</strong> east we have <strong>the</strong>fertile plains <strong>of</strong> Nor<strong>the</strong>rn China and <strong>the</strong> rich and varied islands<strong>of</strong> Japan, possessing a very distinct set <strong>of</strong> peculiar forms, <strong>with</strong>o<strong>the</strong>rs belonging to <strong>the</strong> Oriental region, into which this part <strong>of</strong><strong>the</strong> Palaearctic region merges gradually as we approach <strong>the</strong>Tropic <strong>of</strong> Cancer.Thus, <strong>the</strong> countries roughly indicated by <strong>the</strong>names—Nor<strong>the</strong>rn Europe, <strong>the</strong> Mediterranean district, Centraland Nor<strong>the</strong>rn Asia, and China <strong>with</strong> Japan—have each wellmarkedminor characteristics which entitle <strong>the</strong>m to <strong>the</strong> rank <strong>of</strong>sub-regions. <strong>The</strong>ir boundaries are <strong>of</strong>ten indefinable ; and thosehere adopted have been fixed upon to some extent by considera-


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;chap, x.] THE PALiEARCTIC REGION. 181tions <strong>of</strong> convenience, dependent on custom and on <strong>the</strong> more orless perfect knowledge we possess <strong>of</strong> some <strong>of</strong> <strong>the</strong> interveningcountries.Zoological Characteristics <strong>of</strong> <strong>the</strong> Palmarctic Region.—<strong>The</strong> Palaearcticregion has representatives <strong>of</strong> thirty-five families <strong>of</strong>mammalia, fifty-five <strong>of</strong> birds, twenty-five <strong>of</strong> reptiles, nine <strong>of</strong>amphibia, and thirteen <strong>of</strong> freshwater fishes. Comparing it <strong>with</strong><strong>the</strong> only o<strong>the</strong>r wholly temperate region, <strong>the</strong> Nearctic, we findmuch greater variety <strong>of</strong> types <strong>of</strong> mammalia and birds. Thismay be due in part to its greater area, but more, probably,to itssou<strong>the</strong>rn boundary being conterminous for an enormousdistance <strong>with</strong> two tropical regions, <strong>the</strong>Ethiopean and Orientalwhereas <strong>the</strong> Nearctic has a comparatively short sou<strong>the</strong>rn boundaryconterminous <strong>with</strong> <strong>the</strong> Neotropical region only. This is sovery important a difference, that it is ra<strong>the</strong>r a matter <strong>of</strong> surprisethat <strong>the</strong> two north temperate regions should not be more unequalin <strong>the</strong> number <strong>of</strong> <strong>the</strong>ir higher vertebrate forms, than <strong>the</strong>yactually are.It is also to <strong>the</strong> interblending<strong>of</strong> <strong>the</strong> Palsearctic <strong>with</strong> <strong>the</strong> twoadjacent tropical regions, that we must attribute its possession<strong>of</strong> so few peculiar family groups. <strong>The</strong>se are only three ; two<strong>of</strong> reptiles, Trogonophidcc and Ophiomoridce, and one <strong>of</strong> fishes,Comephoridm. <strong>The</strong> number <strong>of</strong> peculiar genera is, however, considerable,as <strong>the</strong> following enumeration will show.Mammalia.—<strong>The</strong> monkey <strong>of</strong> Gibraltar and North Africa, andan allied species found in Japan, are now considered to belongto <strong>the</strong> extensive eastern genus Macacus. <strong>The</strong> former, however,is peculiar in <strong>the</strong> entire absence <strong>of</strong> <strong>the</strong> tail, and has by manynaturalists, been held to form a distinct genus, Inuus, confinedto<strong>the</strong> Palsearctic region.Of bats <strong>the</strong>re are one or two genera (Barbastelhis, Plecotus)which seem to be mainly or wholly Palsearctic, but <strong>the</strong> classification<strong>of</strong> <strong>the</strong>se <strong>animals</strong> is in such an unsettled state that <strong>the</strong><strong>distribution</strong> <strong>of</strong> <strong>the</strong> genera is <strong>of</strong> little importance.In <strong>the</strong> next order, Insectivora, we have almost <strong>the</strong> entirefamily <strong>of</strong> <strong>the</strong> Moles confined to <strong>the</strong> region. Talpa just entersNor<strong>the</strong>rn India ;and Urotrichus is common to Japan and North-a


182 ZOOLOGICAL GEOGRAPHY. [part hi.Western America, but <strong>the</strong> remaining genera,six in number, areall exclusively Palaearctic.Among Carnivora we have Nyctereutes, <strong>the</strong> curious racoon-dog<strong>of</strong> Japan and North-Eastern Asia ;Lutronectes, an otter peculiarto Japan ; and <strong>the</strong> badger (Meles), which ranges over <strong>the</strong> wholeregion, and just enters <strong>the</strong> Oriental region as far as Hongkong;JEliLropus, a curious form <strong>of</strong> <strong>the</strong> Himalayan panda, inhabiting<strong>the</strong> high mountains <strong>of</strong> Eastern Thibet ;and Pclagius, a genus <strong>of</strong>seals, ranging from <strong>the</strong> shores <strong>of</strong> Madeira to <strong>the</strong> Black Sea.<strong>The</strong> Ungulata, or ho<strong>of</strong>ed <strong>animals</strong>, are stillmore productive <strong>of</strong>forms peculiar to this region. First we have <strong>the</strong> Camels, whosenative home is <strong>the</strong> desert region <strong>of</strong> Central and Western Asia andNor<strong>the</strong>rn Africa, and which, even in <strong>the</strong>ir domesticated condition,are confined almost wholly <strong>with</strong>in <strong>the</strong> limits <strong>of</strong> <strong>the</strong> Palaearcticregion.Of Deer we have six peculiar genera, Dama and Capreolusfound in Europe, <strong>with</strong> Elaphodus, Lophotragus, Hydropotes, andMoschus, confined to Nor<strong>the</strong>rn China and Mongolia.<strong>The</strong> greatfamily Bovidse—comprising <strong>the</strong> oxen, sheep, goats and antelopes—furnishes no less than seven peculiar Palaearctic genera.<strong>The</strong>se are Poephagus, <strong>the</strong> yak <strong>of</strong> Thibet; Addax, a well-knownantelope <strong>of</strong> Nor<strong>the</strong>rn Africa and Syria ; Procapra, Pantholopsand Budorcas, antelopine genera peculiar to Thibet andMongolia ; <strong>with</strong> Rupicapra (<strong>the</strong> chamois), and <strong>the</strong> extraordinary•large-nosed antelope Saiga, confined to Europe and Western Asia.Besides <strong>the</strong>se we have Capra (<strong>the</strong> wild sheep and goats), all <strong>the</strong>numerous species <strong>of</strong> which, except two, are exclusively Pala?arctic.Coming to <strong>the</strong> Eodents, we have again many peculiar forms.Of Murida3 (<strong>the</strong> mouse and rat tribe), we have six peculiargenera, <strong>the</strong> more important being Cricetus, Phombomys Sminthus,and Myospalax. Of Spalacidae (mole-rats) both <strong>the</strong> Palaearcticgenera, Ellobius and Spalax, are peculiar. Ctenodadylus,a genus <strong>of</strong> <strong>the</strong> South American family Octodontidae, is foundonly in North Africa. To <strong>the</strong>se we may add Myoxus (<strong>the</strong>dormice) and Lagomys (<strong>the</strong> pikas or tail-less hares) as essentiallyPalaearctic, since but one species <strong>of</strong> each genus is found beyond<strong>the</strong> limits <strong>of</strong> <strong>the</strong> region.Birds.—It appears to have been <strong>the</strong> opinion <strong>of</strong> many natural-


——chap, x.] THE PAL.EARCTIC REGION. 183ists that <strong>the</strong> Palsearctic region could not be well characterised byits peculiar genera <strong>of</strong> birds. In Mr. Sclater's celebrated paperalready referred to, he remarks, " It cannot be denied that <strong>the</strong>ornithology <strong>of</strong> <strong>the</strong> Palsearctic region isby what it has not than by what itmore easily characterisedhas," and this has been quiterecently quoted by Mr. Allen, in his essay on <strong>the</strong> <strong>distribution</strong><strong>of</strong> North American birds, as ifit represented our present know•ledge <strong>of</strong> <strong>the</strong> subject. But, thanks to <strong>the</strong> labours <strong>of</strong> Dr.Jerdon, Mr. Swinhoe, Pere David and o<strong>the</strong>rs, we have nowlearnt that a large number <strong>of</strong> birds included in <strong>the</strong> Indianlist, are ei<strong>the</strong>r mere winter emigrants from Central Asia, or onlyinhabit <strong>the</strong> higher ranges <strong>of</strong> <strong>the</strong> Himalayas, and thus reallybelong to <strong>the</strong> Pakearctic region. <strong>The</strong> result is, that a host <strong>of</strong>genera are now seen to be ei<strong>the</strong>r exclusively or characteristicallyPalsearctic, and we have no fur<strong>the</strong>rornithological characters to <strong>the</strong> region.difficulty in giving positiveIn <strong>the</strong> tables appendedto this chapter, all <strong>the</strong>se truly Palsearctic genera willbe foundprinted in italics, <strong>with</strong> an indication <strong>of</strong> <strong>the</strong>ir <strong>distribution</strong>, whichwill sometimes be found more fully given under <strong>the</strong> respectivefamilies in <strong>the</strong> fourth part <strong>of</strong> this work. Eeferring to this tablefor details we shall here summarise <strong>the</strong> results.Of <strong>the</strong> Sylviidse or warblers, no less than fourteen genera areei<strong>the</strong>r exclusively or characteristically Palsearctic, <strong>of</strong> whichLocustella, Sylvia, Curruca and Erithacus are good examples.Of <strong>the</strong> oriental family Timaliidse, <strong>the</strong> genus Pterorhinus isPalsearctic.Of Panuridse, or reedlings, <strong>the</strong>re are four peculiar genera(comprising almost <strong>the</strong> whole family) ;one<strong>of</strong> Certhiidse, or creepers,Tichodroma—which extends southward to <strong>the</strong> Abyssinianhighlands. Of Paridse, or tits, oneAcredula ;<strong>of</strong> Corvidse, orcrows, four—Pica (containing our magpie) being a good example; <strong>of</strong>Fringillidse, or finches and buntings,twelve,amongwhichvl canthis,Pyrrhula and Emberiza are good illustrations ;<strong>of</strong> Alaudidse, orlarks, <strong>the</strong>re are two peculiar genera.Leaving <strong>the</strong> Passeres we nextSyrrhaptescome to peculiar forms among <strong>the</strong> gallinaceous birds :among <strong>the</strong> Pteroclidse or sand grouse; four genera <strong>of</strong> Tetraonidse orgrouse and partridges, and five <strong>of</strong> Phasianidse or pheasants, comprisingsome <strong>of</strong> <strong>the</strong> most magnificent birds in <strong>the</strong> world.Lastly


—;184 ZOOLOGICAL GEOGRAPHY. [part hi.among <strong>the</strong> far-wandering aquatic birds we have no less than fivegenera which are more especially Palsearctic,corn-crake, and Otis, <strong>the</strong> great bustard, being typicalOrtygometra,<strong>the</strong>examples.We may add to <strong>the</strong>se, several genera almost confined to thisregion, such as Garrulus (jays), Fringilla (true finches), Yunx(wrynecks) and some o<strong>the</strong>rs ; so that in proportion to its totalgeneric forms a very large number are found to be peculiar orcharacteristic.This view, <strong>of</strong> <strong>the</strong> high degree <strong>of</strong> speciality <strong>of</strong> <strong>the</strong> Palsearcticregion, will no doubt be objected toby some naturalists, on <strong>the</strong>ground that many <strong>of</strong> <strong>the</strong> genera reckoned as exclusivelyPalsearctic are not so, but extend more or less into o<strong>the</strong>r regions.It is well, <strong>the</strong>refore, to consider what principles should guide usin a matter <strong>of</strong> this kind, especially as we shall have to apply<strong>the</strong> same rules to each <strong>of</strong> <strong>the</strong> o<strong>the</strong>r regions. We may remarkfirst, that <strong>the</strong> limits <strong>of</strong> <strong>the</strong> regions <strong>the</strong>mselves are, when notformed by <strong>the</strong> ocean, somewhat arbitrary, depending on <strong>the</strong>average <strong>distribution</strong> <strong>of</strong> a number <strong>of</strong> characteristic forms; andthat slight local peculiarities <strong>of</strong> soil, elevation, or climate, maycause <strong>the</strong> species <strong>of</strong> one region to penetrate more or lessdeeplyinto ano<strong>the</strong>r. <strong>The</strong> land boundary between two regions will be,not a defined line but a neutral territory <strong>of</strong> greater or lesswidth, <strong>with</strong>in which <strong>the</strong> forms <strong>of</strong>both regions will intermingleand this neutral territory itself will merge imperceptibly intoboth regions. So long <strong>the</strong>refore as a species or genus does notpermanently reside considerably beyond <strong>the</strong> possible limits <strong>of</strong>this neutral territory, we should not claim it as an inhabitant<strong>of</strong> <strong>the</strong> adjacent region. A consideration <strong>of</strong> perhaps more importancearises, from <strong>the</strong> varying extent <strong>of</strong> <strong>the</strong> range <strong>of</strong> a genus,over <strong>the</strong> area occupied by <strong>the</strong> region. Some genera are representedby single species existing only in a very limited areao<strong>the</strong>rs by numerous species which occupy, entirely or verynearly, <strong>the</strong> whole extent <strong>of</strong> <strong>the</strong> region ;and <strong>the</strong>re is every intermediategrade between <strong>the</strong>se extremes. Now, <strong>the</strong> small localisedgenera, are always reckoned as among <strong>the</strong> best examples<strong>of</strong> types peculiar to a region ;while <strong>the</strong> more wide-spread groupsare <strong>of</strong>ten denied that character if <strong>the</strong>y extend a little beyond


;chap. x.J THE PAL^ARCTIC REGION. 185<strong>the</strong> supposed regional limits, or send one or two, out <strong>of</strong> a largenumber <strong>of</strong> species, into adjacent regions ;yet <strong>the</strong>re is somereason to believe that <strong>the</strong> latter are really more important ascharacterising a zoological region than <strong>the</strong> former.In <strong>the</strong> case<strong>of</strong> a single isolated species or genus we have a dying-out groupand we have so many cases <strong>of</strong> discontinuous species <strong>of</strong> suchgroups (<strong>of</strong> which Urotrichus in Japan and British Columbia,Eupetes in Sumatra and New Guinea are examples), that it isquite as probable as not, that any such isolated specieshas onlybecome peculiar to <strong>the</strong> region by <strong>the</strong> recent extinction <strong>of</strong> anallied form or forms in some o<strong>the</strong>r region. On <strong>the</strong> o<strong>the</strong>r hand,a genus consisting <strong>of</strong> numerous speciesranging over an entireregion or <strong>the</strong> greater part <strong>of</strong> one, is a dominant group, whichhas most likely been for some time extending its range, andwhose origin dates back to a remote period. <strong>The</strong> slight extension<strong>of</strong> such a group beyond <strong>the</strong> limits <strong>of</strong> <strong>the</strong> region to whichit mainly belongs, isprobably a recent phenomenon, and in thatcase cannot be held in any degree to detract from its value asone <strong>of</strong> <strong>the</strong> peculiar forms <strong>of</strong> that region.<strong>The</strong> most numerous examples <strong>of</strong> this class, are those birds <strong>of</strong><strong>the</strong> temperate regions which in winter migrate, ei<strong>the</strong>rpartially, into adjacent warmer countries.wholly orThis migration mostlikely began subsequent to <strong>the</strong> Miocene period, during thatgradual refrigeration <strong>of</strong> <strong>the</strong> temperate zones which culminatedin <strong>the</strong> glacial epoch, and which still continues in a mitigatedform. Most <strong>of</strong> <strong>the</strong> genera, and many even <strong>of</strong> <strong>the</strong> species <strong>of</strong>birds which migrate southwards in winter, have <strong>the</strong>refore, mostlikely, always been inhabitants <strong>of</strong> our present Palsearctic andNearctic regions ;permanent residents during warm epochs, butonly able now to maintain <strong>the</strong>ir existence by migration inwinter. Such groups belong truly to <strong>the</strong> temperate zones, and<strong>the</strong> test <strong>of</strong> this is<strong>the</strong> fact <strong>of</strong> <strong>the</strong>ir not having any, or very few,representatives, which are permanent residents in <strong>the</strong> adjacenttropical regions.When <strong>the</strong>re are such representative species, wedo not claim <strong>the</strong>m as peculiar to <strong>the</strong> Nor<strong>the</strong>rn regions.Bearingin mind <strong>the</strong>se various considerations, it will be found that we havebeen very moderate in our estimate <strong>of</strong> <strong>the</strong> number <strong>of</strong> generaYol. I.—14


186 ZOOLOGICAL GEOGRAPHY. [part hi.that may fairly be considered as exclusively or characteristicallyPalsearctic.Reptiles and Amphibia.—<strong>The</strong> Palsearctic region possesses,proportion to its limited reptilian fauna, a full proportion <strong>of</strong>peculiar types. We have for instance two genera <strong>of</strong> snakes,Bhinechic and Halys ; seven <strong>of</strong> lizards, Trigonophis, Bsammodromus,Hyalosaurus, Scincus, Ophiomorus, Mcgalochilus, andBhrynocephalus ;eight <strong>of</strong> tailed batrachians, Proteus, Salamandra,Seiranota, Chioglossa, Hynobius, Onychodactylus, Geotriton,and Sieboldia ; and eight <strong>of</strong> tail-less batrachians, Bombinator,Pelobates, Didocus, Alytes, Pelodytes, Discoglossus, Laprissa, andLatonia.<strong>The</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong>se and o<strong>the</strong>r Palsearctic generawill be found in our second vol.chap. xix.Freshwater Fish.—About twenty genera <strong>of</strong> freshwater fishesare wholly confined to this region, and constitute a feature whichought not to be overlooked in estimating its claim to <strong>the</strong> rank<strong>of</strong> a separate primary division <strong>of</strong> <strong>the</strong> earth. <strong>The</strong>y belong to <strong>the</strong>following families :—Percidre (three genera), Acerina, Percarina,Aspro ; Comephoridie (one genus), Comephorus, found only inLake Baikal ; Salmonidse (three genera), Brachymystax, Luciotrutta,and Plecoglossus ; Cyprinodontidse (one genus), Tellia,found only in Alpine pools on <strong>the</strong> Atlas Mountains ;inCyprinidse(thirteen genera), Cyprinus, Carassus, Paraphoxinus, Tinea,Aehilognathus, Bhodeus, Chondrostoma, Pseialoperilampus, Ochetebius,Aspius, Alburnus, Misgurnus, and Nemachilus.Summary <strong>of</strong> Balccarctic Vertebrata.—Summarising <strong>the</strong>se details,we find that <strong>the</strong> Palsearctic region possesses thirty-fivepeculiar genera <strong>of</strong> mammalia, fifty-seven <strong>of</strong> birds, nine <strong>of</strong>reptiles, sixteen <strong>of</strong> amphibia, and twenty-one <strong>of</strong> freshwaterfishes; or a total <strong>of</strong> 138 peculiar generic types <strong>of</strong> vertebrata.Of <strong>the</strong>se, 87 are mammalia and land-birds out <strong>of</strong> a total<strong>of</strong> 274 genera <strong>of</strong> <strong>the</strong>se groups ; or ra<strong>the</strong>r less than one-thirdpeculiar, a number which will serve usefully to compare <strong>with</strong><strong>the</strong> results obtained in o<strong>the</strong>r regions.In our chapter on Zoological Ptegions we have already pointedout <strong>the</strong> main features which distinguish <strong>the</strong> Pakearctic from <strong>the</strong>Oriental and Ethiopian regions. <strong>The</strong> details now given will


—chap. x.J THE PALiEARCTIC REGION. 187streng<strong>the</strong>n our view <strong>of</strong> <strong>the</strong>ir radical distinctness, by showing tohow considerable an extent <strong>the</strong> former is inhabited by peculiar,and <strong>of</strong>ten very remarkable generic types.Insects :Lepidoptera.—<strong>The</strong> Diurnal Lepidoptera, or butterflies,are not very abundant in species, <strong>the</strong>ir number being probablysomewhat over 500, and <strong>the</strong>se belong to not more than fifty genera.But no less than fifteen <strong>of</strong> <strong>the</strong>se genera are wholly confined to<strong>the</strong> region. Nine <strong>of</strong> <strong>the</strong> families are represented, as follows :1. Danaidce; having only a single species in South Europe.2. Satyridce ; well represented, <strong>the</strong>re being more than 100 speciesin Europe, and three peculiar genera. 3. Nymphalidce ; ra<strong>the</strong>rpoorly represented, Europe having only about sixty species, but<strong>the</strong>re is one peculiar genus. 4. Liby<strong>the</strong>idm ; a very small family,represented by a single species occurring in South Europe.5. Nemeobiidce ; a ra<strong>the</strong>r small family, also having only onespecies in Europe, but which constitutes a peculiar genera.6. Lyccenidw ; an extensive family, fairly represented, havingabout eighty European species ; <strong>the</strong>re are two peculiar generain <strong>the</strong> Palasarctic region. 7. Pieridm ; ra<strong>the</strong>r poorly represented<strong>with</strong> thirty-two European species ;two <strong>of</strong> <strong>the</strong> genera are, however,peculiar. 8. Papilionidce ; very poorly represented inEurope <strong>with</strong> only twelve species, but <strong>the</strong>re are many more inSiberia and Japan. No less than five <strong>of</strong> <strong>the</strong> small number <strong>of</strong>genera in this family are wholly confined to <strong>the</strong> region, afact <strong>of</strong> much importance, and which to a great extent redeems<strong>the</strong> character <strong>of</strong> <strong>the</strong> Palaearctic region as regard this order <strong>of</strong>insects. <strong>The</strong>ir names are Mesapia, Hypermnestra, Doritis,Sericinus, and Thais ; and besides <strong>the</strong>se we have Parnassius—<strong>the</strong> " Apollo " butterflies—highly characteristic,and only foundelsewhere in <strong>the</strong> mountains <strong>of</strong> <strong>the</strong> Nearctic region. 9. Hesperidcc; poorly represented <strong>with</strong> about thirty European species,and one peculiar genus.Four families <strong>of</strong> Sphingina occur in <strong>the</strong> Palsearctic region,and <strong>the</strong>re are several peculiar genera.In <strong>the</strong> Zygmnidm <strong>the</strong>re are two exclusively European genera,and <strong>the</strong> extensive genus Zygoma is itself mainly Palaearctic.<strong>The</strong> small family Stygiidas has two out <strong>of</strong> its three genera


188 ZOOLOGICAL GEOGRAPHY. [part hi.confined to <strong>the</strong> Palsearctic region. In <strong>the</strong> JEgeriidce <strong>the</strong> genus^Egeria is mainly Palsearctic. <strong>The</strong> Sphingidce have a widergeneral range, and none <strong>of</strong> <strong>the</strong> larger genera are peculiar to anyone region.Colecptera.—<strong>The</strong> Palaearctic region is <strong>the</strong> richest portion <strong>of</strong><strong>the</strong> globe in <strong>the</strong> great family <strong>of</strong> Carabidce, or predacious groundbeetles,about 50 <strong>of</strong> <strong>the</strong> genera being confined to it, while manyo<strong>the</strong>rs, including <strong>the</strong> magnificent genus Carabus, have here <strong>the</strong>irhighest development. While several <strong>of</strong> <strong>the</strong> smaller genera areconfined to <strong>the</strong> eastern or western sub-regions, most <strong>of</strong> <strong>the</strong> largerones extend over <strong>the</strong> whole area, and give it an unmistakableaspect ; while in passing from east to west or vice-versa, alliedspecies and genera replace each o<strong>the</strong>r <strong>with</strong> considerable regularity,except in <strong>the</strong> extreme south-east, where, in China andJapan, some Oriental forms appear, as do a few Ethiopian typesin <strong>the</strong> south-west.Cicindelida?, or tiger-beetles, are but poorly represented byabout 70 species <strong>of</strong> <strong>the</strong> genus Cicindela, and a single Tetrachain South Europe.Lucanidss, or stag-beetles, are also poor, <strong>the</strong>re being representatives<strong>of</strong> 8 genera. One <strong>of</strong> <strong>the</strong>se, yEsalus (a single species),is peculiar to South Europe, and two o<strong>the</strong>rs, Cladognathus andGyclopthalmus, are only represented in Japan, China, and Thibet.Cetoniidse, or rose-chafers, are represented by 13 genera,two <strong>of</strong> which are peculiar to South Europe (Tropinota andHeterocncmis), while Stalagmosoma, ranging from Persia toNubia, and <strong>the</strong> fine Dicranocephahis inhabiting North China,Corea, and Nipal, may also be considered to belong to it. <strong>The</strong>genera Trichius, Gnorimus, and Osmoderma are confined to <strong>the</strong>two north temperate regions.Buprestidee, or metallic beetles, are ra<strong>the</strong>r abundant in <strong>the</strong>warmer parts <strong>of</strong> <strong>the</strong> region, 27 genera being represented, nine<strong>of</strong> which are peculiar. By far <strong>the</strong> larger portion <strong>of</strong> <strong>the</strong>se areconfined to <strong>the</strong> Mediterranean sub-region. A considerablenumber also inhabit Japan and China.<strong>The</strong> Longicorns, or long-horned beetles, are represented byno less than 196 genera, 51 <strong>of</strong> which are peculiar. <strong>The</strong>y are


Ghap. x.] THE PAL^ARCTIC REGION. 189much more abundant in <strong>the</strong> sou<strong>the</strong>rn than <strong>the</strong> nor<strong>the</strong>rn half<strong>of</strong> <strong>the</strong> region. Several Oriental genera extend to Japan andNorth China, and a few Ethiopian genera to North Africa.Thirteen genera are confinedto <strong>the</strong> two north temperate regions.Several large genera, such as Dorcadion (154 species), Phytcecia(85 species), Pogonochmrus (22 species), Agapanthia (22 species),and Vesperus (7 species), are altoge<strong>the</strong>r peculiar to <strong>the</strong> Patearcticregion ; and <strong>with</strong> a preponderance <strong>of</strong> Leptura, Grammoptera,Stenocorus, and several o<strong>the</strong>rs, strongly characterise it asdistinct from <strong>the</strong> Nearctic and Oriental regions.<strong>The</strong> o<strong>the</strong>r familieswhich are well developed in <strong>the</strong> Palsearcticregions, are, <strong>the</strong> Staphylinidre or rove-beetles, Silphidse orburying-beetles, Histeridae or mimic-beetles, Nitidulidse, Aphodiidse,Copridse (especially in South Europe), Geotrupidse ordung-beetles, Melolonthidse or chafers, Elateridee or clickbeetles,<strong>the</strong> various families <strong>of</strong> Malacoderms and Heteromera,especially Pimeliida? in <strong>the</strong> Mediterranean sub-region, Curculionidseor weevils, <strong>the</strong> Phytophaga or leaf-eaters, and Cocinellidaeor lady-birds.<strong>The</strong> number <strong>of</strong> species <strong>of</strong> Coleoptera in <strong>the</strong> western part <strong>of</strong><strong>the</strong> Palrearctic region is about 15,000, and <strong>the</strong>re are probablynot more than 2,000 to add to this number from Siberia.Japan, and North China; but were <strong>the</strong>se countries as wellexplored as Europe, we may expect that <strong>the</strong>y would add atleast 5,000 to <strong>the</strong> number above given, raising <strong>the</strong> PalaearcticColeopterous fauna to 20,000 species. As <strong>the</strong> total number <strong>of</strong>species at present known to exist in collections isestimated (andperhaps somewhat over-estimated) at 70,000 species, we maybe sure that were <strong>the</strong> whole earth as thoroughly investigatedas Europe, <strong>the</strong> number would be at least doubled, since wecannot suppose that Europe, <strong>with</strong> <strong>the</strong> Mediterranean basin, cancontain more than one-fifth <strong>of</strong> <strong>the</strong> whole <strong>of</strong> <strong>the</strong> Coleoptera <strong>of</strong><strong>the</strong> globe.Of <strong>the</strong> o<strong>the</strong>r orders <strong>of</strong> insects we here say nothing, because in<strong>the</strong>ir case much more than in that <strong>of</strong> <strong>the</strong> Coleopteraand Lepidoptera,is <strong>the</strong> disproportion enormous between our knowledge <strong>of</strong><strong>the</strong> European fauna and that <strong>of</strong> almost all <strong>the</strong> rest <strong>of</strong> <strong>the</strong> globe.


;190 ZOOLOGICAL GEOGRAPHY. [part in.<strong>The</strong>y are, <strong>the</strong>refore, at present <strong>of</strong> comparatively little use forpurposes <strong>of</strong> <strong>geographical</strong> <strong>distribution</strong>, even were it advisableto enter into <strong>the</strong> subject in a work which will, perhaps, be toomuch overburdened <strong>with</strong> details only <strong>of</strong> interest to specialists.Land Shells.—<strong>The</strong>se are very numerous in <strong>the</strong> warmer parts<strong>of</strong> <strong>the</strong> region, but comparatively scarce towards <strong>the</strong> North.South Europe alone possesses over 600 species, wmereas <strong>the</strong>re areonly 200 in all Nor<strong>the</strong>rn Europe and Asia. <strong>The</strong> total number<strong>of</strong> species in <strong>the</strong> whole region is probably about 1,250, <strong>of</strong> which<strong>the</strong> great majority are Helicidse;<strong>the</strong> Operculated families beingvery poorly represented. Several small genera or sub-generaare peculiar to <strong>the</strong> region, as Testacclla (West Europe andCanaries); Leucochroa (Mediterranean district) ;Acicula (Europe);Craspedopoma (Atlantic Islands) ; Lconia (Algeria and Spain)Pamatias (Europe and Canaries) ; Cecina (Mongolia). <strong>The</strong> largestgenera are Helix and Clausilia, which toge<strong>the</strong>r comprise morethan half <strong>the</strong> species ; Pupa, very numerous ; Bidimus andAchatina in moderate numbers, and all <strong>the</strong> rest smallHelix is <strong>the</strong> only genus which contains large and handsomespecies ;Bulimus and Achatina, so magnificent in tropical countries,being here represented by small and obscure forms only.Daudehardia is confined to Central and South Europe and NewZealand ; Glandina is chiefly South American ; Hyalina is onlyAmerican and European ; Buliminus ranges over all <strong>the</strong> worldexcept America ;and <strong>the</strong> o<strong>the</strong>r European genera <strong>of</strong> Helicidae arewidely distributed. Of <strong>the</strong> Operculata, Cyclotus, Cyclophorus,and Pupina extend from <strong>the</strong> Oriental region into Japan andNorth China ;Tudoria is found in Algeria and <strong>the</strong> West IndiesHydrocena is widely scattered, and occurs in South Europe andJapan. <strong>The</strong> genera <strong>of</strong> freshwater shells are all widely distributed.<strong>The</strong> Pal^earctic Sub-regions.<strong>The</strong> four sub-regions which are here adopted, have beenfixed upon as those which are, in <strong>the</strong> present state <strong>of</strong> our knowledge,at once <strong>the</strong> most natural and <strong>the</strong> only practicable ones.


;chap, x.] THE PAL^IARCTIC REGION. 191No doubt all <strong>of</strong> <strong>the</strong>m could be advantageously again subdivided,in a detailed <strong>study</strong> <strong>of</strong> <strong>the</strong> <strong>geographical</strong> <strong>distribution</strong> <strong>of</strong>species.But in a general work, which aims at treating all parts<strong>of</strong> <strong>the</strong> world <strong>with</strong> equal fulness, and which <strong>the</strong>refore is confinedalmost wholly to <strong>the</strong> <strong>distribution</strong> <strong>of</strong> families and genera, suchfur<strong>the</strong>r subdivision would be out <strong>of</strong> place. It is even difficult,in some <strong>of</strong> <strong>the</strong> classes <strong>of</strong> <strong>animals</strong>, to find peculiar or evencharacteristic genera for <strong>the</strong> present sub-regions ; but <strong>the</strong>y allhave well marked climatic and physical differences, and thissuffi-leads to an assemblage <strong>of</strong> species and <strong>of</strong> groups which areciently distinctive./. Central and Nor<strong>the</strong>rn Europe.This sub-region, which may perhaps be termed <strong>the</strong> " European,"is zoologically and botanically <strong>the</strong> best known on <strong>the</strong> globe. Itcan be pretty accurately defined, as bounded on <strong>the</strong> south by<strong>the</strong> Pyrenees, <strong>the</strong> Alps, <strong>the</strong> Balkans, <strong>the</strong> Black Sea, and <strong>the</strong>Caucasus range ; and by <strong>the</strong> Ural Mountains, or perhaps morecorrectly <strong>the</strong> valley <strong>of</strong> <strong>the</strong> Irtish and Caspian Sea, on <strong>the</strong> eastwhile Ireland and Iceland are its fur<strong>the</strong>st outliers in <strong>the</strong> west.To <strong>the</strong> north, it merges so gradually into <strong>the</strong> Arctic zone thatno demarcation is possible. <strong>The</strong> great extent to which thissub-region is interpenetrated by <strong>the</strong> sea, and <strong>the</strong> prevalence<strong>of</strong> westerly winds bringing warmth and moisture from an oceaninfluenced by <strong>the</strong> gulf-stream, give it a climate for <strong>the</strong> mostpart genial, and free from extremes <strong>of</strong> heat and cold. Itis thus broadly distinguished from Siberia and Nor<strong>the</strong>rnAsia generally, where a more extreme and rigorous climateprevails.<strong>The</strong> whole <strong>of</strong> this sub-region is well watered, being penetratedby rivers in every direction ; and it consists mainly <strong>of</strong>plains and undulating country <strong>of</strong> moderate elevation, <strong>the</strong> chiefmountain ranges being those <strong>of</strong> Scandinavia in <strong>the</strong> north-west,and <strong>the</strong> extensive alpine system <strong>of</strong> Central Europe.But <strong>the</strong>seare both <strong>of</strong> moderate height, and a very small portion <strong>of</strong> <strong>the</strong>nsurfaceis occupied ei<strong>the</strong>r by permanent snow-fields, or bybarren uplands inimical to vegetable and animal life. It is, in


192 ZOOLOGICAL GEOGRAPHY. [part hi.fact, to <strong>the</strong>se, and <strong>the</strong> numerous lesser mountains and hillswhich everywhere diversify <strong>the</strong> surface <strong>of</strong> Europe, that <strong>the</strong>variety and abundance <strong>of</strong> its animal life is greatly due. <strong>The</strong>yafford <strong>the</strong> perennial supplies to rivers, and furnish in <strong>the</strong>irvalleys and ever varying slopes, stations suited to every form<strong>of</strong> existence. A considerable area <strong>of</strong> Central Europe is occupiedby uplands <strong>of</strong> moderate elevation, a comparativelysmall portion being flat and marshy plains.Most <strong>of</strong> <strong>the</strong> nor<strong>the</strong>rn and much <strong>of</strong> <strong>the</strong> central portions <strong>of</strong>Europe are covered <strong>with</strong> vast forests <strong>of</strong> coniferous trees ;and<strong>the</strong>se, occupying as <strong>the</strong>y do those tracts where <strong>the</strong> winter ismost severe, supply food and shelter to many <strong>animals</strong> who couldnot o<strong>the</strong>rwise maintain <strong>the</strong>ir existence. It is probable that<strong>the</strong> original condition <strong>of</strong> <strong>the</strong> greater part, if not <strong>the</strong> whole, <strong>of</strong>temperate Europe, except <strong>the</strong> flat marshes <strong>of</strong> <strong>the</strong> river valleysand <strong>the</strong> sandy downs <strong>of</strong> <strong>the</strong> coast, was that <strong>of</strong> woodland andforest, mostly <strong>of</strong> deciduous trees, but <strong>with</strong> a plentiful admixture<strong>of</strong> such hardy evergreens as holly, ivy, privet, and yew. Asufficient proportion <strong>of</strong> <strong>the</strong>se primeval woods, and <strong>of</strong> artificialplantations which have replaced <strong>the</strong>m, fortunately remain, topreserve for us most <strong>of</strong> <strong>the</strong> interesting forms <strong>of</strong> life, whichwere developed before man had so greatly modified <strong>the</strong> surface<strong>of</strong> <strong>the</strong> earth, and so nearly exterminated many <strong>of</strong> its originaltenants. Almost exactly in proportion to <strong>the</strong> amount <strong>of</strong>woodland that still remains in any part <strong>of</strong> Europe, do wefind (o<strong>the</strong>r things being equal) <strong>the</strong> abundance and variety<strong>of</strong> wild <strong>animals</strong>; a pretty clear indication that <strong>the</strong> originalcondition <strong>of</strong> <strong>the</strong> country was essentially that <strong>of</strong> a forest, acondition which only<strong>of</strong> <strong>the</strong> north.now exists in <strong>the</strong> thinly inhabited regionsAlthough <strong>the</strong> sub-region we are considering is, for its extentand latitude, richly peopled <strong>with</strong> animal life, <strong>the</strong> number <strong>of</strong>genera altoge<strong>the</strong>r peculiar to it is not great.<strong>The</strong>re are, however,several which are very characteristic, and many species, both<strong>of</strong> <strong>the</strong> smaller mammalia and <strong>of</strong> birds, are wholly restrictedto it.Mammalia.—<strong>The</strong> genera wholly confined to this sub-region are


chap, x.] THE PAL^ARCTIC REGION. 193only two. Myogalc, <strong>the</strong> desman, is a curious long-snoutedInsectivorous animal somewhat resembling <strong>the</strong> water-rat in itshabits. <strong>The</strong>re are two species, one found only on <strong>the</strong> banks <strong>of</strong>streams in <strong>the</strong> French Pyrenees, <strong>the</strong> o<strong>the</strong>r on <strong>the</strong> great rivers <strong>of</strong>Sou<strong>the</strong>rn Russia. <strong>The</strong> o<strong>the</strong>r peculiar genus, Eupicapra (<strong>the</strong>chamois <strong>of</strong> <strong>the</strong> Alps), is found on all <strong>the</strong> high mountains <strong>of</strong>Central Europe. Almost peculiar are Spalax (<strong>the</strong> mole-rat)found only in Eastern Europe and Western Siberia ; and Saiga,an extraordinary large-nosed antelope which has a nearlysimilar <strong>distribution</strong>.Highly characteristic forms, which inhabitnearly every part <strong>of</strong> <strong>the</strong> sub-region, are, Talpa (<strong>the</strong> mole),Erinaceus, (<strong>the</strong> hedgehog), Sorex (<strong>the</strong> shrew), Meles (<strong>the</strong> badger),Ursus (<strong>the</strong> bear), Canis (<strong>the</strong> wolf and fox), Mustela (<strong>the</strong> weasel),Lutra (<strong>the</strong> otter), Arvicola (<strong>the</strong> vole), Myoxus (<strong>the</strong> dormouse),and Lepits (<strong>the</strong> hare and rabbit) ; while Bos (<strong>the</strong> wild bull) was,until exterminated by man, no doubt equally characteristic.O<strong>the</strong>r genera inhabiting <strong>the</strong> sub-region will be found ingiven at <strong>the</strong> end <strong>of</strong> this chapter.<strong>the</strong> listBirds.—It is difficult to name <strong>the</strong> birds that are most characteristic<strong>of</strong> this sub-region, because so many <strong>of</strong> <strong>the</strong> most familiarand abundant are emigrants from <strong>the</strong> south, and belong togroups that have a different range. <strong>The</strong>re is peril aps not asingle genus wholly confined to it, and very few that have notequal claims to be placed elsewhere.Among <strong>the</strong> more characteristicwe may name Turdus (<strong>the</strong> thrushes), Sylvia (<strong>the</strong> warblers),Panurus (<strong>the</strong> reedling) Parus (<strong>the</strong> tits), Anthus (<strong>the</strong>pipits), Motacilla (<strong>the</strong> wagtails), which are perhaps more abundan<strong>the</strong>re than in any o<strong>the</strong>r part <strong>of</strong> <strong>the</strong> world, Emberiza (<strong>the</strong>buntings), Plcrtrophanes (<strong>the</strong> snow buntings), Passer (<strong>the</strong> housesparrows), Loxia (<strong>the</strong> crossbills), Linota (<strong>the</strong> linnets), Pica (<strong>the</strong>magpies), Tetrao (grouse), Zagopus (ptarmigan) and many o<strong>the</strong>rs.I am indebted to Mr. H. E. Dresser, who is personally acquainted<strong>with</strong> <strong>the</strong> ornithology <strong>of</strong> much <strong>of</strong> <strong>the</strong> North <strong>of</strong> Europe,for some valuable notes on <strong>the</strong> nor<strong>the</strong>rn range <strong>of</strong> many Europeanbirds. Those which are characteristic <strong>of</strong> <strong>the</strong> extremeArctic zone, extending beyond 70° north latitude, and tolerablyabundant, are two falcons (Ealco gyrfalco and F. peregrinus) ;


194 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>the</strong> rough-legged buzzard (Archibuteo lagopus) ; <strong>the</strong> snowy owl(Nyctea scandiaca) ; <strong>the</strong> raven (Corvus corax) ; three buntings(Emberiza schceniculus, Plectrophanes nivalis and P. calcarata) ;lark (Otocorys alpestris) ;several pipits, <strong>the</strong> most nor<strong>the</strong>rn beingAnthus cervinus ; a wagtail (Budytes cinereocapilla) ; a dipper(Cinclus melanogaster) ; a warbler (Cyanecula suecica) ;<strong>the</strong>wheatear (Saxicola cenan<strong>the</strong>) ; and two ptarmigans (Lagopusalius and L. salicetus). Most <strong>of</strong> <strong>the</strong>se birds are, <strong>of</strong> course, onlysummer visitors to <strong>the</strong> Arctic regions, <strong>the</strong> only species noted asa permanent resident in East Finmark (north <strong>of</strong> latitude 70°)being <strong>the</strong> snow-bunting (Plectrophanes nivalis).<strong>The</strong> birds that are characteristic <strong>of</strong> <strong>the</strong> zone <strong>of</strong> pine forests,or from about 61° to 70° north latitude, are very numerous, andit will be sufficient to note <strong>the</strong> genera and <strong>the</strong> number <strong>of</strong>species (where more than one) to give an idea <strong>of</strong> <strong>the</strong> ornithology<strong>of</strong> this part <strong>of</strong> Europe.<strong>The</strong> birds <strong>of</strong> prey are, Falco (threespecies), Astur (two species), Butco, Pandion, Sicrnia, Bubo,Syrnium, Asio, Nyctala. <strong>The</strong> chief Passerine birds are, Corvus(two species), Pica, Garrulus (two species), Nucifraga, Bombycilla,Hirundo (two species), Muscicapa (two species), Lanius,Sturnus, Passer (two species), Pyrrhula,Carpodacus, Loxia (twospecies), Pinicola, Fringilla (eight species), Emberiza (fivespecies), Alauda, Anthus, Turdus (fivespecies), Ruticilla, Pratincola,Accentor, Sylvia (four species), Hypolais, Regulus,Phylloscopus(two species), Acrocephalus, Troglodytes, and Parus (six species).Woodpeckers are abundant, Picas (four species), Gecinus, andYunx. <strong>The</strong> kingfisher (Alcedo), goatsucker (Caprimulgus), andswift (Cypselus) are also common. <strong>The</strong> wood-pigeon (Columba)is plentiful.<strong>The</strong> gallinaceous birds are three grouse, Tetrao (twospecies) and Bonasa, and <strong>the</strong> common quail (Coturnix).<strong>The</strong> remaining genera and species <strong>of</strong> temperate or north-European birds, do not usually range beyond <strong>the</strong> region <strong>of</strong>deciduous trees, roughly indicated by <strong>the</strong> parallel <strong>of</strong> 60° northlatitude.Plate I. — Illustrating <strong>the</strong> Zoology <strong>of</strong> Central Europe.—Before considering <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> o<strong>the</strong>r classes <strong>of</strong>vertebrata, it will be convenient to introduce our first illustra-a


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chap, x.] THE PALiEARCTIC REGION. 195tion, which represents a scene in <strong>the</strong> Alps <strong>of</strong> Central Europe,<strong>with</strong> figures <strong>of</strong> some <strong>of</strong> <strong>the</strong> most characteristic Mammaliaand Birds <strong>of</strong> this sub-region. On <strong>the</strong> left is <strong>the</strong> badger(Meles Taxus) one <strong>of</strong> <strong>the</strong> weasel family, and belonging to agenus which is strictly Palaearctic. It abounds in Central andNor<strong>the</strong>rn Europe and also extends into North Asia, but is representedby ano<strong>the</strong>r species in Thibet and by a third in Japan.<strong>The</strong> elegantly-formed creatures on <strong>the</strong> right are chamois [Rupicapratragus), almost <strong>the</strong> only European antelopes, and whollyconfined to <strong>the</strong> higher mountains, from <strong>the</strong> Pyrenees to <strong>the</strong>Carpathians and <strong>the</strong> Caucasus. <strong>The</strong> chamois is <strong>the</strong> onlyspecies <strong>of</strong> <strong>the</strong> genus, and is thus perhaps <strong>the</strong> most characteristicEuropean mammal. <strong>The</strong> bird on <strong>the</strong> left, above <strong>the</strong> badgers, is<strong>the</strong> Alpine chough, (Fregilus pyrrhocorax). It is found in <strong>the</strong>high mountains from <strong>the</strong> Alps to <strong>the</strong> Himalayas, and is alliedto <strong>the</strong> Cornish chough, which is still found on our southwesterncoasts, and which ranges to Abyssinia and NorthChina. <strong>The</strong> Alpine chough differs in having a shorter bill <strong>of</strong>an orange colour, and vermilion red feet as in <strong>the</strong> o<strong>the</strong>rspecies. In <strong>the</strong> foreground are a pair <strong>of</strong> ruffs {Machetes pugnax)belonging to <strong>the</strong> Scolopacidae or snipe family, and most nearlyallied to <strong>the</strong> genus Tringa or sandpiper. This bird is remarkablefor <strong>the</strong> fine collar <strong>of</strong> plumes which adorns <strong>the</strong> males inbreeding season, when <strong>the</strong>y are excessively pugnacious. It is<strong>the</strong> only species <strong>of</strong> its genus, and ranges over all Europe andmuch <strong>of</strong> Nor<strong>the</strong>rn Asia, migrating in <strong>the</strong> winter to <strong>the</strong> plains <strong>of</strong>India, and even down <strong>the</strong> east coast <strong>of</strong> Africa as far as <strong>the</strong>Cape <strong>of</strong> Good Hope ; but it only breeds in <strong>the</strong> Palsearcticregion, over <strong>the</strong> greater part <strong>of</strong> which itranges.<strong>the</strong>Reptiles and Amphibia.—<strong>The</strong>re are no genera <strong>of</strong> reptilespeculiar to this sub-region. Both snakes and lizards are comparativelyscarce, <strong>the</strong>re being about fourteen species <strong>of</strong> <strong>the</strong> formerand twelve <strong>of</strong> <strong>the</strong> latter. Our common snake {Troyidonotusnatrix) extends into Sweden and North Eussia, but <strong>the</strong> viper(Viperus berus) goes fur<strong>the</strong>r north, as far as Archangel (64° N.),and in Scandinavia (67° N.), and is<strong>the</strong> most Arctic <strong>of</strong> all known


—;,196 ZOOLOGICAL GEOGRAPHY. [part hi.snakes. Of <strong>the</strong> lizards, Lacerta stirpium (<strong>the</strong> sand lizard) has<strong>the</strong> most nor<strong>the</strong>rly range, extending into Poland and Nor<strong>the</strong>rnRussia; and Anguis fragilis (<strong>the</strong> blind or slow-worm) has almostan equal range.Amphibia, being more adapted to a nor<strong>the</strong>rn climate, haveacquired a more special development, and thus several formsare peculiar to <strong>the</strong> North European sub-region. Most remarkableis Proteus, a singular eel -like aquatic creature <strong>with</strong> small legs,found only in <strong>the</strong> subterranean lakes in Carniola and CarinthiaAlytes, a curioustoad, <strong>the</strong> male <strong>of</strong> which carries about <strong>the</strong> eggstill <strong>the</strong>y are hatched, found only in Central Europe fromFrance to <strong>the</strong> east <strong>of</strong> Hungary ; and Pelodytcs, a frog found onlyin France.Frogs and toads are very abundant all over Europe, <strong>the</strong>common frog {Rana temporaria) extending to <strong>the</strong> extreme north.<strong>The</strong> newts {Triton) are also very abundant and widely spread,though not ranging so far north as <strong>the</strong> frogs. <strong>The</strong> genera Bombinator(a toad-like frog), and Hyla (<strong>the</strong> tree frog) are also commonin Central Europe.Freshvjater Fish,—Two genera <strong>of</strong> <strong>the</strong> perch family (Percidse) arepeculiar to thissub-region,Percarina, a fishfound only in <strong>the</strong>river Dniester, and Aspro, confined to <strong>the</strong> rivers <strong>of</strong> CentralEurope. Of <strong>the</strong> very characteristic forms are, Gastcrosteus(stickle-back), which alone forms a peculiar family—Gasterosteidse; Perca, Acerina and Luciopcrca, genera <strong>of</strong> <strong>the</strong> perch family;Silurus, a large fish found in <strong>the</strong> rivers <strong>of</strong> Cenrtal Europe, <strong>of</strong><strong>the</strong> family Siluridae ; Esox (<strong>the</strong> pike), <strong>of</strong> <strong>the</strong> family EsocidteCyprinus (carp), Gobio (gudgeon), Leuciscus (roach, chub, dace,&c), Tinea (tench), Abramus (bream), Alburnus (bleak), Cobitis(loach), all genera <strong>of</strong> <strong>the</strong> family Cyprinidoe.Insects— Lepidoptera, — No genera <strong>of</strong> butterflies are actuallyconfined to this sub-region, but many are characteristic <strong>of</strong> it.Parnassius, Aporia, Leucophasia, Colias, Melitcea, Argynnis,Vanessa, Limenitis, and Chionobas, are all very abundant andwidespread, and give a feature tocountries included in it.<strong>the</strong> entomology <strong>of</strong> most <strong>of</strong> <strong>the</strong>Coleoptera.—This sub-region is very rich in Carabidae ; <strong>the</strong>genera Elaphrus, Nebria, Carabtis, Cychrus, Pterostichits, Amara,


chap, x.] THE PAL^ARCTIC REGION. 197Trechus and Peryphus being especially characteristic. Staphylinidaeabound. Among Lamellicorns <strong>the</strong> genus Aphodius ismost characteristic. Buprestidse are scarce ; Elateridse moreabundant. Among Malacoderms Telephones and Malachius arecharacteristic. Curculionidse abound : Otiovhyuchus, Omias,Erirhinus, Bagous, Rhynchites and Ceutorhynchus being verycharacteristic genera. Of Longicorns Callidium, Dorcadion,Pogonocficerus, Pachyta and Leptura are perhaps <strong>the</strong> best representatives.Donacia, Crioceris, Chrysomela, and Attica, aretypical Phytophaga; while Coccinella is <strong>the</strong> best representative<strong>of</strong> <strong>the</strong>Securipalpes.Forth European Islands.—<strong>The</strong> British Islands are known tohave been recently connected <strong>with</strong> <strong>the</strong> Continent, and <strong>the</strong>iranimal productions areso uniformly identical <strong>with</strong> continentalspecies as to require no special note. <strong>The</strong> only general fact <strong>of</strong>importance is, that <strong>the</strong> number <strong>of</strong> .species in all groups is muchless than in continental districts <strong>of</strong> equal extent, and that thisnumber is still far<strong>the</strong>r diminished in Ireland. This may beaccounted for by <strong>the</strong> smaller areaand less varied surface <strong>of</strong> <strong>the</strong>latter island ; and it may also be partly due to <strong>the</strong> great extent<strong>of</strong> low land, so that a very small depression would reduce itto <strong>the</strong> condition <strong>of</strong> a cluster <strong>of</strong> small islands capable <strong>of</strong> supportinga very limited amount <strong>of</strong> animal life. Yet fur<strong>the</strong>r, ifafter such a submergence had destroyed much <strong>of</strong> <strong>the</strong> higherforms <strong>of</strong> life in Great Britain and Ireland, both were elevated soas to again form part <strong>of</strong> <strong>the</strong> Continent, a migration would commenceby which <strong>the</strong>y would be stocked afresh ; but this migrationwould be a work <strong>of</strong> time, and it is to be expected thatmany species would never reach Ireland or would find its excessivelymoist climate unsuited to <strong>the</strong>m.Some few British species differ slightly from <strong>the</strong>ir continentalallies, and are considered by many naturalists to be distinct.This is <strong>the</strong> case <strong>with</strong> <strong>the</strong> red grouse (Lagopus scoticus) amongbirds ;and a few <strong>of</strong> <strong>the</strong> smaller Passeres have also been foundto vary somewhat from <strong>the</strong> allied forms on <strong>the</strong> Continent, showingthat <strong>the</strong> comparatively short interval since <strong>the</strong> glacial period,and <strong>the</strong> slightly different physical conditions dependent on


—198 ZOOLOGICAL GEOGRAPHY. [part hi.insularity, have sufficed to commence <strong>the</strong> work <strong>of</strong> specificmodification.<strong>The</strong>re are also a few small land-shells and severalinsects not yet found elsewhere than in Britain ;<strong>of</strong> <strong>the</strong> smaller Mammalia—a shrew (Sorcx rusticus).and even one<strong>The</strong>se factsare all readily explained by <strong>the</strong> former union <strong>of</strong> <strong>the</strong>se islands<strong>with</strong> <strong>the</strong> Continent, and <strong>the</strong> alternate depressions and elevationswhich are proved by geological evidence to have occurred, bywhich <strong>the</strong>y have been more than once separated and unitedagain in recent times. For <strong>the</strong> evidence <strong>of</strong> this elevation anddepression, <strong>the</strong> reader may consult Sir Charles Lyell's Antiquity<strong>of</strong> Man.Iceland is <strong>the</strong> only o<strong>the</strong>r island <strong>of</strong> importance belonging tothis sub-region, and it contrasts strongly <strong>with</strong> Great Britain,both in its Arctic climate and oceanic position. It is situatedjust south <strong>of</strong> <strong>the</strong> Arctic circle and considerably nearer Greenlandthan Europe, yet its productions are almost wholly European.<strong>The</strong> only indigenous land mammalia are <strong>the</strong> Arctic fox (Canislagopus), and <strong>the</strong> polar bear as an occasional visitant, <strong>with</strong> amouse {Mus islandicus), said to be <strong>of</strong> a peculiar species. Fourspecies <strong>of</strong> seals visit its shores. <strong>The</strong> birds are more interesting.According to Pr<strong>of</strong>essor Newton, ninety-five species have beenobserved ; but many <strong>of</strong> <strong>the</strong>se are mere stragglers. <strong>The</strong>re aretwenty-three land, and seventy-two aquatic birds and waders.Four or five are peculiar species, though very closely related too<strong>the</strong>rs inhabiting Scandinavia or Greenland. Only two orthree species are more nearly related to Greenland birds than tothose <strong>of</strong> Nor<strong>the</strong>rn Europe, so that <strong>the</strong> Palrearctic character <strong>of</strong><strong>the</strong> fauna is unmistakable. <strong>The</strong> following lists, compiled froma paper by Pr<strong>of</strong>essor Newton, may be interesting as showingmore exactly <strong>the</strong> character <strong>of</strong> Icelandic ornithology.1. Peculiar species. — Troglodytes borealis (closely alliedto <strong>the</strong> common wren, found also in <strong>the</strong> Faroe Islands) ; Falcoislandicus (closely allied to F. gyrfalco) ;Lagopus islandorum(closely allied to L. rupestris <strong>of</strong> Greenland).2. European species resident in Iceland.Emberiza nivalis,Corvus corax, Haliazctus albicilla, Rallus aquaticus, Hamiatopusostralegus, Cygnus fcrus, Mcrgus (two species), Phalacocorax (two


———chap, x.] THE PAL^ARCTIC REGION. 199species), Sula bassana, Lams (two species), Stercorarius catarractes,Pujrfinus anglorum, Mergulus alle, Una (three species),Alca torda.3. American species resident in Iceland. Clangula islandica,Histrionicus torquatus.4. Annual visitants from Europe.titliys,Turdus iliacus, ButicillaSaxicola mnan<strong>the</strong>, Motacilla alba, Anthus pratensis, Linotalinaria, Chelidon urbica, Hirundo rustica, Falco msalon, Surnianyctea, Otus brachyotus, Charadrius pluvialis, AZgialites hiaticida,Strepsilas interpres, Phalaropus fulicarius, Totanus calidris,Limosa (species), Tringa (three species), Calidris arenaria,Gallinago media, Numenius plimopus, Ardea cinerea, Anser (twospecies), Bernicla (two species), Anas (four species), Fuligulamarila, Harelda glacialis, Somateria mollissima, (Edemia nigra,Sterna macrura, Bissa tridactyla, Larus luecopterus, Stercorarius(two species), Fratercula artica, Colymbus (two species), Podicepscornutus.5. Annual visitant from Greenland. Falco candicans.6.—Former resident, now extinct. Alca impennis (<strong>the</strong>great auk).II.— Mediterranean Sub-region.This is by far <strong>the</strong> richest' portion <strong>of</strong> <strong>the</strong> Paloearctic region,for although <strong>of</strong> moderate extent much <strong>of</strong> it enjoys a climate inwhich <strong>the</strong> rigours <strong>of</strong> winter are almost unknown. It includesall <strong>the</strong> countries south <strong>of</strong> <strong>the</strong> Pyrenees, Alps, Balkans, andCaucasus mountains ;all <strong>the</strong> sou<strong>the</strong>rn shores <strong>of</strong> <strong>the</strong> Mediterraneanto <strong>the</strong> Atlas range, and even beyond it to include <strong>the</strong>extra-tropical portion <strong>of</strong> <strong>the</strong> Sahara ; and in <strong>the</strong> Nile valleyas far as <strong>the</strong> second cataract. Fur<strong>the</strong>r east it includes <strong>the</strong>nor<strong>the</strong>rn half <strong>of</strong> Arabia and <strong>the</strong> whole <strong>of</strong> Persia, as well asBeluchistan, and perhaps Afghanistan up to <strong>the</strong> banks <strong>of</strong> <strong>the</strong>Indus. This extensive district is almost wholly a region <strong>of</strong>mountains and elevated plateaus. On <strong>the</strong> west, Spain ismainly a table-land <strong>of</strong> more than 2000 feet elevation, deeplypenetrated by extensive valleys and rising into l<strong>of</strong>ty mountainchains. Italy, Corsica, Sardinia, and Sicily, are all very


200 ZOOLOGICAL GEOGRAPHY. [part in.mountainous, and much <strong>of</strong> <strong>the</strong>ir surface considerably elevated.Fur<strong>the</strong>r east we have all European Turkey and Greece, amountain region <strong>with</strong> a comparatively small extent <strong>of</strong> levelplain. Id Asia <strong>the</strong> whole country, from Smyrna throughArmenia and Persia to <strong>the</strong> fur<strong>the</strong>r borders <strong>of</strong> ,Affghanistan, is avast mountainous plateau, almost all above 2000, and extensivedistricts above 5000 feet in elevation. <strong>The</strong> only large tract <strong>of</strong>low-land is <strong>the</strong> valley <strong>of</strong> <strong>the</strong> Euphrates. <strong>The</strong>re is also somelow-land south <strong>of</strong> <strong>the</strong> Caucasus, and in Syria <strong>the</strong> valley <strong>of</strong> <strong>the</strong>Jordan. In North Africa <strong>the</strong> valley <strong>of</strong> <strong>the</strong> Nile and <strong>the</strong> coastplains <strong>of</strong> Tripoli and Algiers are almost <strong>the</strong> only exceptions to<strong>the</strong> more or less mountainous and plateau-like character <strong>of</strong> <strong>the</strong>country. Much <strong>of</strong> this extensive area is now bare and arid,and <strong>of</strong>ten even <strong>of</strong> a desert character ; a fact no doubt due, ingreat part, to <strong>the</strong> destruction <strong>of</strong> aboriginal forests. This lossis rendered permanent by <strong>the</strong> absence <strong>of</strong> irrigation, and, it isalso thought, by <strong>the</strong> abundance <strong>of</strong> camels and goats, <strong>animals</strong>which are exceedingly injurious to woody vegetation, and areable to keep down <strong>the</strong> natural growth <strong>of</strong> forests. Mr. Marshwhose valuable work Man and Nature gives much informationon this subject) believes that even large portions <strong>of</strong> <strong>the</strong> Africanand Asiatic deserts would become covered <strong>with</strong> woods, and <strong>the</strong>climate <strong>the</strong>reby greatly improved, were <strong>the</strong>y protected from<strong>the</strong>se destructive domestic <strong>animals</strong>, which are probably notindigenous to <strong>the</strong> country. Spain, in proportion to its extent,is very barren ;Italy and European Turkey are more woody andluxuriant ; but it is perhaps in Asia Minor, on <strong>the</strong> range <strong>of</strong> <strong>the</strong>Taurus, along <strong>the</strong> shores <strong>of</strong> <strong>the</strong> Black Sea, and to <strong>the</strong> south <strong>of</strong><strong>the</strong> Caucasus range, that this sub-region attains its maximum <strong>of</strong>luxuriance in vegetation and in animal life. From <strong>the</strong> Caspianeastward extends a region <strong>of</strong> arid plains and barren deserts,diversified by a few more fertile valleys, in which <strong>the</strong> characteristicflora and fauna <strong>of</strong> this portion <strong>of</strong> <strong>the</strong> Palsearctic regionabounds. Fur<strong>the</strong>r east we come to <strong>the</strong> forests <strong>of</strong> <strong>the</strong> HindooKoosh, which probably form <strong>the</strong> limit <strong>of</strong> <strong>the</strong> sub-region.Beyond <strong>the</strong>se we enter on <strong>the</strong> Siberian sub-region to <strong>the</strong> north,and on <strong>the</strong> outlying portion <strong>of</strong> <strong>the</strong> Oriental region on <strong>the</strong> south.


chai\ x.] THE PAL^IARCTIC REGION. 201In addition to <strong>the</strong> territories now indicated as forming part<strong>of</strong> <strong>the</strong> Mediterranean sub-region, we must add <strong>the</strong> group <strong>of</strong>Canary Islands <strong>of</strong>f <strong>the</strong> west coast <strong>of</strong> Africa which seem to bean extension <strong>of</strong> <strong>the</strong> Atlas mountains, and <strong>the</strong> oceanic groups <strong>of</strong>Madeira and <strong>the</strong> Azores ; <strong>the</strong> latter about 1,000 miles from <strong>the</strong>continent <strong>of</strong> Europe, yet still unmistakably allied to it both in<strong>the</strong>ir vegetable and animal productions. <strong>The</strong> peculiarities <strong>of</strong> <strong>the</strong>faunas <strong>of</strong> <strong>the</strong>se islands will be subsequently referred to.It seems at first sight very extraordinary, that so large andwide a sea as <strong>the</strong> Mediterranean should not separate distinctfaimas, and this is <strong>the</strong> more remarkable when we find how verydeep <strong>the</strong> Mediterranean is, and <strong>the</strong>refore how ancient we maywell suppose it to be. Its eastern portion reaches a depth <strong>of</strong>2,100 fathoms or 12,600 feet, while its western basin is about1,600 fathoms or 9,600 feet in greatest depth, and a considerablearea <strong>of</strong> both basins is more than 1,000 fathoms deep. But afur<strong>the</strong>r examination shows, that a comparatively shallow sea orsubmerged bank incloses Malta and Sicily, and that on <strong>the</strong>opposite coast a similar bank stretches out from <strong>the</strong> coast <strong>of</strong>Tripolileaving a narrow channel <strong>the</strong> greatest depth <strong>of</strong> which is240 fathoms. Here <strong>the</strong>refore is a broad plateau, which anelevation <strong>of</strong> about 1,500 feet<strong>of</strong> land connecting Italy <strong>with</strong> Africa ;would alsowould convert into a wide extentwhile <strong>the</strong> same elevationconnect Morocco <strong>with</strong> Spain, leaving two extensivelakes to represent what is now <strong>the</strong> Mediterranean Sea, and affordingfree communication for laud <strong>animals</strong> between Europe andNorth Africa. That such a state <strong>of</strong> things existed at a comparativelyrecent period, is almost certain ; not only because aconsiderable number <strong>of</strong> identical species <strong>of</strong> mammalia inhabit <strong>the</strong>opposite shores <strong>of</strong> <strong>the</strong> Mediterranean, but also because numerousremains <strong>of</strong> three species <strong>of</strong> elephants have been found in cavesin Malta,—now a small rocky island in which it would be impossiblefor such <strong>animals</strong> to live even if <strong>the</strong>y could reach it.Remains <strong>of</strong> hippopotami are also found atGibraltar, and manyo<strong>the</strong>r <strong>animals</strong> <strong>of</strong> African types in Greece ; all indicating means<strong>of</strong> communication between South Europe and North Africa whichno longer exist. (See Chapter VI. pp. 113—115.)Vol. I.—15


202 ZOOLOGICAL GEOGRAPHY. [part hi.Mammalia.—<strong>The</strong>re are a few groups <strong>of</strong> Paleearctic Mammaliathat are peculiar to this sub-region. Such are, Dama, <strong>the</strong>fallow deer, which is now found only in South Europe and NorthAfrica ; Psammomys, a peculiar genus <strong>of</strong> Muridse, found only inEgypt and Palestine ; while Ctenodactylus, a rat-like animalclassed in <strong>the</strong> South American family Octodontidse, inhabitsTripoli. Among characteristic genera not found in o<strong>the</strong>r subregions,are, Dysopes, a bat <strong>of</strong> <strong>the</strong> family Noctilionidoe; Macroscelides,<strong>the</strong> elephant shrew, in North Africa ; Genetta, <strong>the</strong>civet, in South Europe ; Herpestes, <strong>the</strong> ichneumon, in NorthAfrica and (?) Spain ; Hyaena, in South Europe ; Gaizclla, Oryx,Alcephalus, and Addax, genera <strong>of</strong> antelopes in North Africaand Palestine ; Hyrax, in Syria : and Hystrix, <strong>the</strong> porcupine,in South Europe. Besides <strong>the</strong>se, <strong>the</strong> camel and <strong>the</strong> horsewere perhaps once indigenous in <strong>the</strong> eastern parts <strong>of</strong> <strong>the</strong> subregion; and a wild sheep (Ovis musmon) still inhabits Sardinia,Corsica, and <strong>the</strong> mountains <strong>of</strong> <strong>the</strong> south-east <strong>of</strong> Spain. <strong>The</strong>presence <strong>of</strong> <strong>the</strong> large feline <strong>animals</strong>—such as <strong>the</strong> lion, <strong>the</strong>leopard, <strong>the</strong> serval, and <strong>the</strong> hunting leopard—in North Africa,toge<strong>the</strong>r <strong>with</strong> several o<strong>the</strong>r quadrupeds not found in Europe,have been thought by some naturalists to prove, that this districtshould not form part <strong>of</strong> <strong>the</strong> Palrearctic region. No doubtseveral Ethiopian groups and species have entered it from <strong>the</strong>south, but <strong>the</strong> bulk <strong>of</strong> its Mammalia still remains Palsearctic,although several <strong>of</strong> <strong>the</strong> species have Asiatic ra<strong>the</strong>r than Europeanaffinities. <strong>The</strong> Macacus innuus is allied to an Asiaticra<strong>the</strong>r than an African group <strong>of</strong> monkeys, and thus denotes anOriental affinity. Ethiopian affinity is apparently shown by <strong>the</strong>three genera <strong>of</strong> antelopes, by Herpestes, and by Macroscelides ; butour examination <strong>of</strong> <strong>the</strong> Miocene fauna has shown that <strong>the</strong>se wereprobably derived from Europe originally, and do not form anypart <strong>of</strong> <strong>the</strong> truly indigenous or ancient Ethiopian fauna. Against<strong>the</strong>se, however, we have <strong>the</strong> occurrence in North Africa <strong>of</strong>such purely Palaearctic and non-Ethiopian genera as Vrsus, Males,Putorius, Sus, Cervvs, Dama, Capra, Alactaga ; toge<strong>the</strong>r <strong>with</strong>actual European or West Asiatic species <strong>of</strong> Canis, Genetta, Fclis,Putorius, Lutra, many bats, Sorex, Crocidura, Crossopus, Hystrix,


chap, x.] THE PAL^ARCTIC REGION. 203Dipus, Lepus, and Mus. It is admitted that, as regards everyo<strong>the</strong>r group <strong>of</strong> <strong>animals</strong>, North Africa is Palaearctic, and <strong>the</strong>above enumeration shows that even in Mammalia, <strong>the</strong> intermixture<strong>of</strong> what are now true Ethiopian types is altoge<strong>the</strong>rinsignificant. It must be remembered, also, that <strong>the</strong> lioninhabited Greece even in historic times, while large carnivorawere contemporary <strong>with</strong> man all over Central Europe.Birds.—So many <strong>of</strong> <strong>the</strong> European birds migrate over largeportions <strong>of</strong> <strong>the</strong> region, and so many o<strong>the</strong>rs have a wide permanentrange, that we cannot expect to find more than a fewgenera, consisting <strong>of</strong> one or two species, each, confined to a subregion;and such appear to be, Lusciniola and Pyrophthalma,genera <strong>of</strong> Sylviidae. But many are characteristic <strong>of</strong> this, ascompared <strong>with</strong> o<strong>the</strong>r Palsearctic sub-regions ; such as, Bradyptetus,Aedon, Dromolcea, and Cercomela, among Sylviidae; Crateropusand Malacocercus, among Timaliidse ; Teloplionus amongLaniidse ; Certhilauda and Mirafra among larks ; Pastor amongstarlings ; Upupa, <strong>the</strong> hoopoe ; Halycon and Ceryle amongkingfishers; Turnix and Caccabis among Gallinae, and <strong>the</strong>pheasant as an indigenous bird ;toge<strong>the</strong>r <strong>with</strong> Gyps, Vultur andNeophron, genera <strong>of</strong> vultures. In addition to <strong>the</strong>se, almost allour summer migrants spend <strong>the</strong>ir winter in some part <strong>of</strong> thisfavoured land, mostly in North Africa, toge<strong>the</strong>r <strong>with</strong> manyspecies <strong>of</strong> Central Europe that rarely or never visit us. Itfollows, that a large proportion <strong>of</strong> all <strong>the</strong> birds <strong>of</strong> Europe andWestern Asia are to be found in this sub-region, as will be seenby referring to <strong>the</strong> list <strong>of</strong> <strong>the</strong> genera <strong>of</strong> <strong>the</strong> region. Palestineis one <strong>of</strong> <strong>the</strong> remote portions <strong>of</strong> this region which has beenwell explored by Canon Tristram, and it may be interesting togive his summary <strong>of</strong> <strong>the</strong> range <strong>of</strong> <strong>the</strong> birds. We must bearin mind that <strong>the</strong> great depression <strong>of</strong> <strong>the</strong> Dead Sea has a tropicalclimate, which accounts for <strong>the</strong> presence here only, <strong>of</strong> such atropical form as <strong>the</strong> sun- bird (Nectarinea osea).<strong>The</strong> total number <strong>of</strong> <strong>the</strong> birds <strong>of</strong> Palestine is 322, and <strong>of</strong><strong>the</strong>se no less than 260 are European, at once settling <strong>the</strong> question<strong>of</strong> <strong>the</strong> general affinities <strong>of</strong> <strong>the</strong> fauna.Of <strong>the</strong> remainder elevenbelong to North and East Asia, four to <strong>the</strong> Eed Sea, and thirty-


—;204 ZOOLOGICAL GEOGRAPHY. [part in.one to East Africa, while twenty-seven are peculiar to Palestine.It is evident <strong>the</strong>refore that an unusual number <strong>of</strong> East Africanbirds have extended <strong>the</strong>ir range to this congenial district, butmost <strong>of</strong> <strong>the</strong>se are desert species and hardly true Ethiopians,and do not much interfere <strong>with</strong> <strong>the</strong> general Palsearctic character<strong>of</strong> <strong>the</strong> whole assemblage.As an illustration <strong>of</strong> how wide-spreadare many <strong>of</strong> <strong>the</strong> Palsearctic forms, we may add, that seventyninespecies <strong>of</strong> land birds and fifty-five <strong>of</strong> water birds, are commonto Palestine and Britain. <strong>The</strong> Oriental and Ethiopiangenera Pycnonotus and Ncctarinea are found here, while Bcssornisand Dromolcca are characteristically Ethiopian. Almost all <strong>the</strong>o<strong>the</strong>r genera are Palsearctic.Persia is ano<strong>the</strong>r remote regiongenerally associated <strong>with</strong> <strong>the</strong>idea <strong>of</strong> Oriental and almost tropical forms, but which yet undoubtedlybelongs to <strong>the</strong> Palsearctic region. Mr. Blanford's recentcollections in this country, <strong>with</strong> o<strong>the</strong>r interestinginformation, issummarised in Mr. Elwes's paper on <strong>the</strong> " Geographical Distribution<strong>of</strong> Asiatic Birds " (Proc. Zool. Soc. 1873, p. 647). No lessthan 127 species are found also in Europe, and thirty-seveno<strong>the</strong>rs belong to European genera ; seven are allied to birds <strong>of</strong>Central Asia or Siberia, and fifteen to those <strong>of</strong> North-East Africa,while only three are purely <strong>of</strong> Indian affinities. This shows apreponderance <strong>of</strong> nearly nine-tenths <strong>of</strong> Palsearcticforms, whichis fully as much as can be expected in any country near <strong>the</strong>limits <strong>of</strong> a great region.Reptiles and Amphibia.—<strong>The</strong> climatal conditions being heremore favourable to <strong>the</strong>se groups, and <strong>the</strong> genera being <strong>of</strong>ten <strong>of</strong>limited range, we find some peculiar, and several very interestingforms.Fvhinechis, a genus <strong>of</strong> Colubrine snakes, is found only inSouth Europe ; Trogonophis, one <strong>of</strong> <strong>the</strong> Amphisbsenianscurious snake-like lizards—is known only from North AfricaPsammosaurus, belonging to <strong>the</strong> water lizards (Varanidse) isfound in North Africa and North-West India ;a genus <strong>of</strong> Lacertidse, is peculiar to South Europe ;Psammodromus,Hyalosaurus,belonging to <strong>the</strong> family Zonuridre, is a lizard <strong>of</strong> especial interest,as it inhabits North Africa while its nearest ally is <strong>the</strong>Opliisawrus or " glass snake " <strong>of</strong> North America ; <strong>the</strong> family <strong>of</strong>


—chap, x.] THE PAL^ARCTIC REGION. 205<strong>the</strong> scinks is represented by Scincus found inArabia.North Africa andBesides <strong>the</strong>se Seps, a genus <strong>of</strong> sand lizards (Sepidae) andAgarna, a genus <strong>of</strong> Agamidae, are abundant and characteristic.Of Amphibia we have Seiranota, a genus <strong>of</strong> salamandersfound only in Italy and Dalmatia ;Chioglossa, in Portugal, andGeotriton, in Italy, belonging to <strong>the</strong> same family, are equallypeculiar to <strong>the</strong> sub-region.Freshtvater Fish,—One <strong>of</strong> <strong>the</strong> most interesting is Tellia, agenus <strong>of</strong> Cyprinodontidae found only in alpine pools in <strong>the</strong>Atlas mountains. Paraphoxinius, found in South-East Europe,and Chondrostoma, in Europe and Western Asia, genera <strong>of</strong> Cyprimidae,seem almost peculiar to this sub-region.Insects— Lepidoptera.—Two genera <strong>of</strong> butterflies, Thais andDoritis, are wholly confined to this sub-region, <strong>the</strong> formerranging over all Sou<strong>the</strong>rn Europe, <strong>the</strong>latter confined to EasternEurope and Asia Minor. Anthocharis and Zegris are very characteristic<strong>of</strong> it, <strong>the</strong> latter only extending into South Eussia,while Danais, Charaxes, and Libythca are tropical genera unknownin o<strong>the</strong>r parts <strong>of</strong> Europe.Coleoptera.—This sub-region is very rich in many groups <strong>of</strong>Coleoptera, <strong>of</strong> which a few only can be noticed here. AmongCarabidse it possesses Procerus and Procrustes, almost exclusively,while Brachinus,Cymindis, Lcbia, Graphipterus, Scarites, Chlcenius,Calathus, and many o<strong>the</strong>rs, are abundant and characteristic.Among Lamellicorns—Copridae, G-laphyridae, Melolonthidae, andCetoniidae abound.Buprestidae are plentiful, <strong>the</strong> genera Julodis,Acmccodera, Buprestis, and Sphenoptera being characteristic.Among Malacoderms—Cebrionidae, Lampyridae, and Malachiidaeabound.<strong>The</strong> Tenebrioid Heteromera are very varied and abundant,and give a character to <strong>the</strong> sub-region. <strong>The</strong> Mylabridse,Cantharidae, and OZdemeridae are also characteristic Of <strong>the</strong>immense number <strong>of</strong> CurculionidseBrachycerus,Lixus, and Acalles may be mentioned asThylacites,among <strong>the</strong> most prominent.Of Longicorns <strong>the</strong>re are few genera especially characteristic,but perhaps Prinobms, Purpuricenus, Hesperophanes,and Parmena are most so.Of <strong>the</strong> remaining families, we maymention Clythridae, Hispidae, and Cassididae as being abundant.


206 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>The</strong> Mediterranean and Atlantic Islands.—<strong>The</strong> various islands<strong>of</strong> <strong>the</strong> Mediterranean are interesting to <strong>the</strong> student <strong>of</strong> <strong>geographical</strong><strong>distribution</strong> as affording a few examples <strong>of</strong> local species<strong>of</strong> very restricted range, but as a rule <strong>the</strong>y present us <strong>with</strong>exactly <strong>the</strong> same forms as those <strong>of</strong> <strong>the</strong> adjacent mainland. 1<strong>The</strong>ir peculiarities do not, <strong>the</strong>refore, properly come <strong>with</strong>in <strong>the</strong>scope <strong>of</strong> this work. <strong>The</strong> islands <strong>of</strong> <strong>the</strong> Atlantic Ocean belongingto this sub-region are, from <strong>the</strong>ir isolated position and <strong>the</strong>various problems <strong>the</strong>y suggest, <strong>of</strong> much more interest, and <strong>the</strong>irnatural history has been carefully studied. We shall <strong>the</strong>reforegive a short account <strong>of</strong> <strong>the</strong>ir peculiar features.Of <strong>the</strong> three groups <strong>of</strong> Atlantic islands belonging to this subregion,<strong>the</strong> Canaries are nearest to <strong>the</strong> Continent, some <strong>of</strong> <strong>the</strong>islands being only about fifty miles from <strong>the</strong> coast <strong>of</strong> Africa.<strong>The</strong>y are, however, separated from <strong>the</strong> mainland by a very deepchannel (more than 5,000 feet), as shown on our general map.<strong>The</strong> islands extend over a length <strong>of</strong> 300 miles ;<strong>the</strong>y are verymountainous and wholly volcanic, and <strong>the</strong> celebrated peak <strong>of</strong>Teneriffe rises to a height <strong>of</strong> more than 12,000 feet. <strong>The</strong> smallMadeira group is about 400 miles from <strong>the</strong> coast <strong>of</strong> Moroccoand 600 from <strong>the</strong> sou<strong>the</strong>rn extremity <strong>of</strong> Portugal ; and <strong>the</strong>re isa depth <strong>of</strong> more than 12,000 feet between it and <strong>the</strong> continent.<strong>The</strong> Azores are nearly 1,000 miles west <strong>of</strong> Lisbon. <strong>The</strong>y arequite alone in mid-Atlantic, <strong>the</strong> most westerly islands beingnearer Newfoundland than Europe, and are surrounded by oceandepths <strong>of</strong> from 12,000 to 18,000 feet. It will be convenient totake <strong>the</strong>se islands firstin order.Azores.—Considering <strong>the</strong> remoteness <strong>of</strong> this group from everyo<strong>the</strong>r land, it is surprising to find as many as fifty-three species<strong>of</strong> birds inhabiting or visiting <strong>the</strong> Azores ; and still more to1Malta is interesting as forming a resting-place for migratory birds, whilecrossing <strong>the</strong> Mediterranean. It has only eight land and three aquatic birdswhich are permanent residents ;yet no less than 278 species have beenrecorded by Mr. E. A. Wright as visiting or passing over it, comprising alarge proportion <strong>of</strong> <strong>the</strong> European migratory birds. <strong>The</strong> following are <strong>the</strong>permanent residents : Cerchneis tinnunculus, Strix flammed, Passer salicicola,Emberiza miliaria, Corvu* monedula, Monticola cyanea, Sylvia conspicillata,Columba livia, Puffinus cinerevs, P. anglorum, Thalassidroma pelagica.


chap, x.] THE PAL^EARCTIC REGION. 207find that <strong>the</strong>y are <strong>of</strong> Palsearctic genera and, <strong>with</strong> one exception,all <strong>of</strong> species found ei<strong>the</strong>r in Europe, North Africa, Madeira,or <strong>the</strong> Canaries. <strong>The</strong> exception is a bullfinch peculiar to <strong>the</strong>islands, but closely allied to a European species. Of land birds<strong>the</strong>re are twenty-two, belonging to twenty-one genera, all European.<strong>The</strong>se genera are Cerchneis, Buteo, Asio, Strix, Turdus,Oriolus, Erithacus, Sylvia, Regulus, Saxicola, Motacilla, Plectrophanes,Fringilla, Pyrrhula, Serinus, Stumus, Picus,Upwpa, Columba, Caccabis, and Coturnix. Besides <strong>the</strong> bullfinch{Pyrrhula) o<strong>the</strong>r species show slight differences from <strong>the</strong>irEuropean allies, but not such as to render <strong>the</strong>m more thanvarieties. <strong>The</strong> only truly indigenous mammal is a bat <strong>of</strong> aEuropean species. Nine butterflies inhabit <strong>the</strong> Azores ; eight<strong>of</strong> <strong>the</strong>m are European species, one North American. Of beetles212 have been collected, <strong>of</strong> which no less than 175 are Europeanspecies; <strong>of</strong> <strong>the</strong> remainder, nineteen are found in <strong>the</strong>Canaries or Madeira, three in South America, while fourteenare peculiar to <strong>the</strong> islands.Now <strong>the</strong>se facts (for which we are indebted to Mr. Godman'sNatural History <strong>of</strong> <strong>the</strong> Azores) are both unexpected and exceedinglyinstructive. In most o<strong>the</strong>r cases <strong>of</strong> remote Oceanicislands, a much larger proportion <strong>of</strong> <strong>the</strong> fauna is endemic, orconsists <strong>of</strong> peculiar species and <strong>of</strong>ten <strong>of</strong> peculiar genera ; as iswell shown by <strong>the</strong> case<strong>of</strong> <strong>the</strong> Galapagos and Juan Fernandez,both much nearer to a continent and both containing peculiargenera and species <strong>of</strong> birds. Now we know that <strong>the</strong> cause andmeaning <strong>of</strong> this difference is, that in <strong>the</strong> one case <strong>the</strong> originalimmigration isvery remote and has never or very rarely beenrepeated, so that under <strong>the</strong> unchecked influence <strong>of</strong> new conditions<strong>of</strong> life <strong>the</strong> species have become modified ;in <strong>the</strong> o<strong>the</strong>rcase, ei<strong>the</strong>r <strong>the</strong> original immigration has been recent, or if remotehas been so frequently repeated that <strong>the</strong> new comers have keptup <strong>the</strong> purity <strong>of</strong> <strong>the</strong> stock, and have not allowed time for <strong>the</strong>new conditions to produce <strong>the</strong> effect we are sure <strong>the</strong>y would intime produce if not counteracted. For Mr. Godman tells usthat many <strong>of</strong> <strong>the</strong> birds are modified—instancing <strong>the</strong> gold-crestedwren, blackcap, and rockdove—and he adds, that <strong>the</strong> modifica-


208 ZOOLOGICAL GEOGRAPHY. [part hi.tion all tends in one direction—to produce a more sombreand legs, and a more robustplumage, a greater strength <strong>of</strong> feetbill. We fur<strong>the</strong>r find, that four <strong>of</strong> <strong>the</strong> land- birds, including <strong>the</strong>oriole, snow-bunting, and hoopoe, are not resident birds, butstraggle accidentally to <strong>the</strong> islands by stress <strong>of</strong> wea<strong>the</strong>r; andwe are told that every year some fresh birds are seen afterviolent storms. Add to this <strong>the</strong> fact, that <strong>the</strong> number <strong>of</strong>species diminishes in <strong>the</strong> group as we go from east towest, andthat <strong>the</strong> islands are subject to fierce and frequent stormsblowing from every point <strong>of</strong> <strong>the</strong> compass,—and we have all <strong>the</strong>facts requisite to enable us to understand how this remotearchipelago has become stocked <strong>with</strong> animal life<strong>with</strong>out everprobably being much nearer to Europe than it is now. For<strong>the</strong> islands are all volcanic, <strong>the</strong> only stratified rock that occursbeing believed to be <strong>of</strong> Miocene date.Madeira and <strong>the</strong> Canaries.—Coming next to Madeira, we find<strong>the</strong> number <strong>of</strong> genera <strong>of</strong> land birds has increased to twenty-eight,<strong>of</strong> which seventeen are identical <strong>with</strong> those <strong>of</strong> <strong>the</strong> Azores.Some<strong>of</strong> <strong>the</strong> commonest European birds—swallows, larks, sparrows,linnets, goldfinches, ravens, and partridges, are among <strong>the</strong> additions.A gold-crested warbler, Begulus Maderensis, and a pigeon,Columba Trocaz, are peculiar to Madeira.In <strong>the</strong> Canaries we find that<strong>the</strong> birds have again very muchincreased, <strong>the</strong>re being more than fifty genera <strong>of</strong> land birds ;but<strong>the</strong> additions are wholly European in character, and almost allcommon European species. We find a few more peculiar species(five), while some o<strong>the</strong>rs, including <strong>the</strong> wild canary, arecommon to all <strong>the</strong> Atlantic Islands or to <strong>the</strong> Canaries andMadeira. Here, too, <strong>the</strong> only indigenous mammalia are twoEuropean species <strong>of</strong> bats.Land SMls.-^Ihe land shells<strong>of</strong> Madeira <strong>of</strong>fer us an instructivecontrast to <strong>the</strong> birds <strong>of</strong> <strong>the</strong> Atlantic Islands. About fiftysixspecies have been found in Madeira, and forty-two in <strong>the</strong> smalladjacent island<strong>of</strong> Porto Santo, but only twelve are common toboth, and all or almost all are distinct from <strong>the</strong>ir nearest alliesin Europe and North Africa. Great numbers <strong>of</strong> fossil shellsare also found in deposits <strong>of</strong> <strong>the</strong> Newer Pliocene period ; and


chap, x.] THE PAL^EARCTIC REGION. 209although <strong>the</strong>se comprise many freshspecies, <strong>the</strong> two faunas andthat <strong>of</strong> <strong>the</strong> continent still remain almost as distinct from eacho<strong>the</strong>r as before. It has been already stated (p. 31) that <strong>the</strong>means by which landmollusca have been carried across arms<strong>of</strong> <strong>the</strong> sea are unknown, although several modes may besuggested ; but it is evidently a rare event, requiring someconcurrence <strong>of</strong> favourable conditions not always present. <strong>The</strong>diversity and specialization <strong>of</strong> <strong>the</strong> forms <strong>of</strong> <strong>the</strong>se <strong>animals</strong> is,<strong>the</strong>refore, easily explained by <strong>the</strong> fact, that, once introduced<strong>the</strong>y have been left to multiply under <strong>the</strong> influence <strong>of</strong> a variety<strong>of</strong> local conditions, which inevitably lead, in <strong>the</strong> course <strong>of</strong> ages,to <strong>the</strong> formation <strong>of</strong> new varieties and new species.Colcoptera.—<strong>The</strong> beetles <strong>of</strong> Madeira and <strong>the</strong> Canaries havebeen so carefully collected and examined by Mr. T. V. Wollaston,and those <strong>of</strong> <strong>the</strong> Azores described and compared by Mr. Crotch,and <strong>the</strong>y illustrate so many curious points in <strong>geographical</strong> <strong>distribution</strong>,that it is necessary to give some account <strong>of</strong> <strong>the</strong>m.less than 1,480 species <strong>of</strong> beetles have been obtained from <strong>the</strong>Canaries and Madeira, only 360 <strong>of</strong> which are European, <strong>the</strong>remainder being peculiar to <strong>the</strong> islands. <strong>The</strong> Canaries -areinhabited by a little over 1,000 species, Madeira by about 700,while 240 are common to both ;but it is believed that many ol<strong>the</strong>se have been introduced by man.have been obtained, <strong>of</strong> which 1 75 are European ;NoIn <strong>the</strong> Azores, 212 speciesshowing, as in<strong>the</strong> birds, a closer resemblance to <strong>the</strong> European fauna than in<strong>the</strong> o<strong>the</strong>r islands which, although nearer to <strong>the</strong> continent, <strong>of</strong>fermore shelter and are situated in a less tempestuous zone. Of<strong>the</strong> non-European species in <strong>the</strong> Azores, 19 are found also in<strong>the</strong> o<strong>the</strong>r groups <strong>of</strong> islands, 14 are peculiar, while 3 are American.Of <strong>the</strong> European species, 132 are found also in <strong>the</strong> o<strong>the</strong>r Atlanticislands, while 43 have reached <strong>the</strong> Azores only. This is interestingas showing to how great an extent <strong>the</strong> same insectsreach all <strong>the</strong> islands, not<strong>with</strong>standing <strong>the</strong> difference <strong>of</strong> latitudeand position ; and it becomes <strong>of</strong> great <strong>the</strong>oretical importance,when we find how many extensive families and genera are altoge<strong>the</strong>rabsent.<strong>The</strong> Madeira group has been more thoroughly explored than


210 ZOOLOGICAL GEOGRAPHY. [part hi.any o<strong>the</strong>r, and its comparatively remote situation, combined <strong>with</strong>its luxuriant vegetation, have been favourable to <strong>the</strong> developmentand increase <strong>of</strong> <strong>the</strong> peculiar forms which characterize all<strong>the</strong> Atlantic islands in a more or less marked degree. A consideration<strong>of</strong> some <strong>of</strong> its peculiarities will, <strong>the</strong>refore, best serveto show <strong>the</strong> bearing <strong>of</strong> <strong>the</strong> facts presented by <strong>the</strong> insect fauna <strong>of</strong><strong>the</strong> Atlantic islands, on <strong>the</strong> general laws <strong>of</strong> <strong>distribution</strong>. <strong>The</strong>711 species <strong>of</strong> beetles now known from <strong>the</strong> Madeira group,belong to 236 genera ; and no less than 44 <strong>of</strong> <strong>the</strong>se genera arenot European but are peculiar to <strong>the</strong> Atlantic islands. Most <strong>of</strong><strong>the</strong>m are, however, closely allied to European genera, <strong>of</strong> which<strong>the</strong>y are evidently modifications. A most curious general featurepresented by <strong>the</strong> Madeiran beetles, is <strong>the</strong> total absence <strong>of</strong> manywhole families and large genera abundant in South Europe.Such are <strong>the</strong> Cicindelidse, or tiger beetles ;<strong>the</strong> Melolonthidae, orchafers; <strong>the</strong> Cetoniidae, or rose-chafers; <strong>the</strong> Eumolpidse andGalerucidte, large families <strong>of</strong> Phytophagous, or leaf-eating beetles;and also <strong>the</strong> extensive groups <strong>of</strong> Elateridre and Buprestida?,which are each represented by but one minute species. Ofextensive genera abundant in South Europe, but wholly absentin Madeira, are Carabus, Bhizotrogus, Lampyris, and o<strong>the</strong>rgenera <strong>of</strong> Malacoderms; Otiorhynchus, Brachycerus, and 20o<strong>the</strong>r genera <strong>of</strong> Curculionidse, comprising more than 300 SouthEuropean and North African species ;Pimelia, Tentyra, Blaps,and 18 o<strong>the</strong>r genera <strong>of</strong> Heteromera, comprising about 550 speciesin South Europe and North Africa ; and Timarcha, containing44 South European and North African species.Ano<strong>the</strong>r most remarkable feature <strong>of</strong> <strong>the</strong> Madeiran Coleopterais <strong>the</strong> unusual prevalence <strong>of</strong> apterous or wingless insects.Thisis especially <strong>the</strong> case <strong>with</strong> groups which are confined to <strong>the</strong>Atlantic islands, many <strong>of</strong> which consist wholly <strong>of</strong> winglessspecies ; but it also affects <strong>the</strong> o<strong>the</strong>rs, no less than twenty-twogenera which are usually or sometimes winged in Europe, havingonly wingless species in Madeira ; and even <strong>the</strong> same specieswhich is winged in Europe becomes, in at least three cases,wingless in Madeira, <strong>with</strong>out any o<strong>the</strong>r perceptible change havingtaken place. But <strong>the</strong>re is ano<strong>the</strong>r most curious fact noticed by


'chap, x.] THE PALMARCTIC REGION. 211Mr. Wollaston ; that those species which possess wings inMadeira, <strong>of</strong>ten have <strong>the</strong>m ra<strong>the</strong>r larger than <strong>the</strong>ir allies inEurope. <strong>The</strong>se two facts were connected by Mr. Darwin, whosuggested that flying insects are much more exposed to beblown out to sea and lost, than those which do not fly(and Mr-Wollaston had himself supposed that <strong>the</strong> " stormy atmosphere<strong>of</strong> Madeira had something to do <strong>with</strong> <strong>the</strong> matter) ; so that <strong>the</strong>most frequent fliers would be continually weeded out, while <strong>the</strong>more sluggish individuals, who ei<strong>the</strong>r could not or would notfly, remained to continue <strong>the</strong> race ;and this process going on fromgeneration to generation, would, on <strong>the</strong> well-ascertained principles<strong>of</strong> selection and abortion by disuse, in time lead to <strong>the</strong>entire loss <strong>of</strong> wings by those insects to whom wings were not anecessity. But those whose wings were essential to <strong>the</strong>ir existencewould be acted upon in ano<strong>the</strong>r way.All <strong>the</strong>se must flyto obtain <strong>the</strong>ir food or provide for <strong>the</strong>ir <strong>of</strong>fspring, and those thatflew best would be best able to battle <strong>with</strong> <strong>the</strong>storms, and keep<strong>the</strong>mselves safe, and thus those <strong>with</strong> <strong>the</strong> longest and mostpowerful wings would be preserved. If however all <strong>the</strong> individuals<strong>of</strong> <strong>the</strong> species were too weak on <strong>the</strong> wing to resist <strong>the</strong>storms, <strong>the</strong>y would soon become extinct. 1Now this explanation <strong>of</strong> <strong>the</strong> facts is not only simple andprobable in itself, but it also serves to explain in a remarkablemanner some <strong>of</strong> <strong>the</strong> peculiarities and deficiencies <strong>of</strong> <strong>the</strong> Madeiraninsect fauna, in harmony <strong>with</strong> <strong>the</strong> view (supported by <strong>the</strong> <strong>distribution</strong><strong>of</strong> <strong>the</strong> birds and land shells, and in particular by <strong>the</strong>immigrant birds and insects <strong>of</strong> <strong>the</strong> Azores) that all <strong>the</strong> insectshave been derived from <strong>the</strong>continent or from o<strong>the</strong>r islands, by1A remarkable confirmation <strong>of</strong> this <strong>the</strong>ory, is furnished in <strong>the</strong> Report to<strong>the</strong> Royal Society <strong>of</strong> <strong>the</strong> naturalist to <strong>the</strong> Kerguelen Island, " Transit Expedition—<strong>the</strong> Rev. A. E. Eaton. Insects were assiduously collected, and itwas found that almost all were ei<strong>the</strong>r completely apterous, or had greatlyabbreviated wings. <strong>The</strong> only moth found, several flies, and numerous beetles,were alike incapable <strong>of</strong> flight. As this island is subject to violent, andalmost perpetual gales, even in <strong>the</strong> finest season, <strong>the</strong> meaning <strong>of</strong> <strong>the</strong> extraordinaryloss <strong>of</strong> wings in almost all <strong>the</strong> insects, can, in this case, hardly bemisunderstood.


212 ZOOLOGICAL GEOGRAPHY. [part hi.immigration across <strong>the</strong> ocean, in various ways and during a longperiod. <strong>The</strong>se deficiencies are, on <strong>the</strong> o<strong>the</strong>r hand, quite inconsistent<strong>with</strong> <strong>the</strong> <strong>the</strong>ory (still held by some entomologists) thata land communication is absolutely necessary to account for<strong>the</strong> origin <strong>of</strong> <strong>the</strong> Madeiran fauna.First, <strong>the</strong>n, we can understand how <strong>the</strong> tiger-beetles (Cicindelidae)are absent ; since <strong>the</strong>y are insects which have a short weakflight, but yet to whom flight is necessary. If a few had beenblown over to Madeira, <strong>the</strong>y would soon have become exterminated.<strong>The</strong> same thing applies to <strong>the</strong> Melolonthidse, Cetoniidse,Eumolpidae, and Galerucidse,—all flower and foliagehauntinginsects, yet bulky and <strong>of</strong> comparatively feeble powers<strong>of</strong> flight.Again, all <strong>the</strong> large genera abundant in South Europe,which have been mentioned above as absent from Madeira,wholly apterous (or <strong>with</strong>out wings), and thus <strong>the</strong>ir absence is amost significant fact ; for it proves that in <strong>the</strong> case <strong>of</strong> all insects<strong>of</strong> moderate size, flight was essential to <strong>the</strong>ir reaching <strong>the</strong> island,which could not have been <strong>the</strong> case had <strong>the</strong>re been a land connection.<strong>The</strong>re are, however, one or two curious exceptions to<strong>the</strong> absence <strong>of</strong> <strong>the</strong>se wholly apterous European genera in Madeira,and as in each case <strong>the</strong> reason <strong>of</strong> <strong>the</strong>ir being exceptions canbe pointed out, <strong>the</strong>y are eminently exceptions that prove <strong>the</strong>rule. Two <strong>of</strong> <strong>the</strong> apterous species common to Europe andMadeira are found always in ants' nests ; and as ants, whenwinged, fly in great swarms and are carried by <strong>the</strong> wind togreat distances, <strong>the</strong>y may have conveyed <strong>the</strong> minute eggs <strong>of</strong><strong>the</strong>se very small beetles. Two European species <strong>of</strong> Blapsoccur in Madeira, but <strong>the</strong>se are house beetles, and are admittedto have been introduced by man. <strong>The</strong>re are also three species<strong>of</strong> Meloe, <strong>of</strong> which two are European and one peculiar.<strong>The</strong>se are large, sluggish, wingless insects, but <strong>the</strong>y have amost extraordinary and exceptional metamorphosis, <strong>the</strong> larvaein <strong>the</strong> first state being minute active insects parasitic on bees,and thus easily conveyed across <strong>the</strong> ocean. This case is mostsuggestive, as itareaccounts for what would be o<strong>the</strong>rwise a difficultanomaly. Ano<strong>the</strong>r case, not quite so easily explained, is that<strong>of</strong> <strong>the</strong>genus Acalles, which is very abundant in all <strong>the</strong> Atlantic


chap, x.] THE PAL^ARCTIC REGION. 213islands and also occurs in South Europe, but isalways apterous.It is however closely allied to ano<strong>the</strong>r genus, Cryptorhynchus,which is apterous in some species, winged in o<strong>the</strong>rs. We may<strong>the</strong>refore well suppose that <strong>the</strong> ancestors <strong>of</strong> A calks were once in<strong>the</strong> same condition, and that some <strong>of</strong> <strong>the</strong> winged forms reachedMadeira, <strong>the</strong> genus having since become wholly apterous.We may look at this curious subject in ano<strong>the</strong>r way. <strong>The</strong>Coleoptera <strong>of</strong> Madeira may be divided into those which arefound also in Europe or <strong>the</strong> o<strong>the</strong>r islands, and those which arepeculiar to it. On <strong>the</strong> <strong>the</strong>ory <strong>of</strong> introduction by accidentalimmigration across <strong>the</strong> sea, <strong>the</strong> latter must be <strong>the</strong> more ancient,since <strong>the</strong>y have had time to become modified ;while <strong>the</strong> formerare comparatively recent, and <strong>the</strong>ir introduction may be supposedto be now going on. <strong>The</strong> peculiar influence <strong>of</strong> Madeira inaborting <strong>the</strong> wings should, <strong>the</strong>refore, have actedon <strong>the</strong> ancientand changed forms much more powerfully than on <strong>the</strong> recentand unchanged forms. On carefully comparing <strong>the</strong> two sets <strong>of</strong>insects (omitting those which have almost certainly beenintroduced by man) we find, that out <strong>of</strong> 263 species whichhave a wide range, only 14 are apterous ;while <strong>the</strong> o<strong>the</strong>r class,consisting <strong>of</strong> 393 species, has no less than 178 apterous ; orabout 5 per cent in <strong>the</strong> one case, and 45 per cent in <strong>the</strong> o<strong>the</strong>r. 1On <strong>the</strong> <strong>the</strong>ory <strong>of</strong> a land connection as <strong>the</strong> main agent in introducing<strong>the</strong> fauna, both groups must have been introduced at orabout <strong>the</strong> same time, and why one set should have lost <strong>the</strong>irwings and <strong>the</strong> o<strong>the</strong>r not, isquite inexplicable.TakiDg all <strong>the</strong>se singular facts, in connection <strong>with</strong> <strong>the</strong> totalabsence <strong>of</strong> all truly indigenous terrestrial mammalia and reptilesfrom <strong>the</strong>se islands—even from <strong>the</strong> extensive group <strong>of</strong> <strong>the</strong> Canariesso comparatively near to <strong>the</strong> continent, we are forced toreject <strong>the</strong> <strong>the</strong>ory <strong>of</strong> a land connection as quite untenable ;andthis view becomes almost demonstrated by <strong>the</strong> case <strong>of</strong> <strong>the</strong>Azores, which being so much fur<strong>the</strong>r <strong>of</strong>f, and surrounded bysuch avast expanse <strong>of</strong> deep ocean, could only have been con-1<strong>The</strong> facts on which <strong>the</strong>se statements rest, will be found more fullydetailed in <strong>the</strong> Author's Presidential Address to <strong>the</strong> Entomological Society<strong>of</strong> London for <strong>the</strong> year 1871.


214 ZOOLOGICAL GEOGRAPHY. [part hi.nected <strong>with</strong> Europe at a farremoter epoch, and ought <strong>the</strong>reforeto exhibit to us. a fauna composed almost entirelyforms both <strong>of</strong> birds and insects.<strong>of</strong> peculiarYet, so far from this being <strong>the</strong>case, <strong>the</strong> facts are exactly <strong>the</strong> reverse. Far more <strong>of</strong> <strong>the</strong> birdsand insects are identical <strong>with</strong> those <strong>of</strong> Europe than in <strong>the</strong>o<strong>the</strong>r islands, and this difference is clearly traced to <strong>the</strong> moretempestuous atmosphere, which is shown to be even nowannually bringing fresh immigrants (both birds and insects) toits shores. We here see nature actually at work ; and if <strong>the</strong>case <strong>of</strong> Madeira rendered her mode <strong>of</strong> action probable, that <strong>of</strong><strong>the</strong> Azores may be saidto demonstrate it.Mr. Wollaston has objected to this view that " storms andhurricanes " are somewhat rare in <strong>the</strong> latitude <strong>of</strong> Madeira and<strong>the</strong> Canaries ; but this little affects <strong>the</strong> question, since <strong>the</strong> timeallowed for such operations is so ample. If but one veryviolent storm happened in a century, and ten such stormsrecurred before a single species <strong>of</strong> insect was introduced intoMadeira, that would be more than sufficient to people it, as wenow find it, <strong>with</strong> a varied fauna. But he also adds <strong>the</strong> importantinformation that <strong>the</strong> ordinary winds blow almost uninterruptedlyfrom <strong>the</strong> north-east, so that <strong>the</strong>re would be always achance <strong>of</strong> a little stronger wind than usual bringing insect, orlarva, or egg, attached to leaves or twigs. Nei<strong>the</strong>r Mr. Wollaston,Mr. Crotch, Mr. A. Murray, nor any o<strong>the</strong>r naturalist whoupholds <strong>the</strong> land-connection <strong>the</strong>ory, has attempted to accountfor <strong>the</strong> fact <strong>of</strong> <strong>the</strong> absence <strong>of</strong> so many extensive groups <strong>of</strong>insects that ought to be present, as well as <strong>of</strong> all smallmammalia and reptiles.Cape Vera 1 Islands.—<strong>The</strong>re is yet ano<strong>the</strong>r group <strong>of</strong> Atlanticislands which is very little known, and which is usually consideredto be altoge<strong>the</strong>r African—<strong>the</strong> Cape Verd Islands, situatedbetween 300 and 400 miles west <strong>of</strong> Senegal, and a little to <strong>the</strong>south <strong>of</strong> <strong>the</strong> termination <strong>of</strong> <strong>the</strong> Sahara. <strong>The</strong> evidence that wepossess as to <strong>the</strong> productions <strong>of</strong> <strong>the</strong>se islands, shows that, like<strong>the</strong> preceding groups, <strong>the</strong>y are truly oceanic, and have probablyderived <strong>the</strong>ir fauna from <strong>the</strong> desert and <strong>the</strong> Canaries to <strong>the</strong>north-east <strong>of</strong> <strong>the</strong>m ra<strong>the</strong>r than from <strong>the</strong> fertile and more truly


——chap, x.] THE PAL^ARCTIC REGION. 215Ethiopian districts <strong>of</strong> Senegal and Gambia to <strong>the</strong> east.a mingling <strong>of</strong> <strong>the</strong><strong>The</strong>re istwo faunas, but <strong>the</strong> preponderance seems tobe undoubtedly <strong>with</strong> <strong>the</strong> Palasarctic ra<strong>the</strong>r than <strong>with</strong> <strong>the</strong>Ethiopian. I owe to Mr. E. B. Sharpe <strong>of</strong> <strong>the</strong> British Museum,a MS. list <strong>of</strong> <strong>the</strong> birds <strong>of</strong> <strong>the</strong>se islands, twenty-three species inall. Of <strong>the</strong>se eight are <strong>of</strong> wide <strong>distribution</strong> and may be neglected.Seven are undoubted Palasarctic species, viz. :Milvusictinus, Sylvia atricapilla, S. conspicillata, Corvus corone, Passersalicarius, Certhilauda desertorum, Columba livia. Three arepeculiar species, but <strong>of</strong> Palasarctic genera and affinities, viz. :Calamoherpe brevipennis, Ammomanes cinctura, and Passer jagoensis.Against this we have to set two West African species,Estrilda cinerea and Namida meleagris, both <strong>of</strong> which wereprobably introduced by man ; and three which are <strong>of</strong> Ethiopiangenera and affinities, viz.:—Halcyon erythrorhyncha, closelyallied to H. semAcceruka <strong>of</strong> Arabia and North-east Africa, and<strong>the</strong>refore almost Palasarctic; Accipiter melanoleucus ; an&Pyrrhulaudanigriceps, an Ethiopian form ;but <strong>the</strong> same species occursin <strong>the</strong> Canaries.<strong>The</strong> Coleoptera <strong>of</strong> <strong>the</strong>se islands have been also collected byMr. Wollaston, and he finds that <strong>the</strong>y have generally <strong>the</strong> sameEuropean character as those <strong>of</strong> <strong>the</strong> Canaries and Madeira, several<strong>of</strong> <strong>the</strong> peculiar Atlantic genera, such as Acalles and Hegdcr,occurring, while o<strong>the</strong>rs are represented by new but closely alliedgenera.Out <strong>of</strong> 275 species 91 were found also in <strong>the</strong> Canariesand 81 in <strong>the</strong> Madeiran group ; a wonderful amount <strong>of</strong> similaritywhen we consider <strong>the</strong> distance and isolation <strong>of</strong> <strong>the</strong>se islandsand <strong>the</strong>ir great diversity <strong>of</strong> climate and vegetation.This connection <strong>of</strong> <strong>the</strong> four groups <strong>of</strong> Atlantic islands nowreferred to, receives fur<strong>the</strong>r support from <strong>the</strong> occurrence <strong>of</strong> landshells<strong>of</strong> <strong>the</strong> subgenus Leptaxis in all <strong>the</strong> groups, as well as inMajorca ;and by ano<strong>the</strong>r subgenus, Hemicycla, being common to<strong>the</strong> Canaries and Cape Verd islands. Combining <strong>the</strong>se severalclasses <strong>of</strong> facts, we seem justified in extending <strong>the</strong> Mediterraneansub-region to include <strong>the</strong> Cape Verd Islands.


216 ZOOLOGICAL GEOGRAPHY. [part hi.Ill— <strong>The</strong> Siberian Sub-region, or Nor<strong>the</strong>rn Asia.This large and comparatively little-known subdivision <strong>of</strong> <strong>the</strong>Palrearctic region, extends from <strong>the</strong> Caspian Sea to Kamschatkaand Behring's Straits, a distance <strong>of</strong> about 4,000 miles ; andfrom <strong>the</strong> shores <strong>of</strong> <strong>the</strong> Arctic Ocean to<strong>the</strong> high Himalayas <strong>of</strong>Sikhim in North Latitude 29°, on <strong>the</strong> same parallel as Delhi.To <strong>the</strong> east <strong>of</strong> <strong>the</strong> Caspian Sea and <strong>the</strong> Ural Mountains is a greatextent <strong>of</strong> lowland which is continued round <strong>the</strong> nor<strong>the</strong>rn coast,becoming narrower as it approaches <strong>the</strong> East Cape. Beyondthis, in a general E.KE. direction, rise hills and uplands, soonbecoming l<strong>of</strong>ty mountains, which extend in an unbroken linefrom <strong>the</strong> Hindu Koosh, through <strong>the</strong> Thian Shan, Altai andYablonoi Mountains, to <strong>the</strong>extremity <strong>of</strong> Asia.Stanovoi range in <strong>the</strong> north-easternSouth <strong>of</strong> this region is a great central basin,which is almost wholly desert ; beyond which again is <strong>the</strong> vastplateau <strong>of</strong> Thibet, <strong>with</strong> <strong>the</strong> Kuenlun, Karakorum, and Himalayansnow-capped ranges, formingdistrict on <strong>the</strong> globe.<strong>the</strong> most extensive elevated<strong>The</strong> superficial aspects <strong>of</strong> this vast territory, as determined byits vegetable covering, are very striking and well contrasted. Abroad tract on <strong>the</strong> nor<strong>the</strong>rn coast, varying from 150 to 300 andeven 500 miles wide, is occupied by <strong>the</strong> Tundras or barrens,where nothing grows but mosses and <strong>the</strong> dwarfest Arctic plants,and where <strong>the</strong> ground is permanently frozen to a great depth.This tract has its greatest sou<strong>the</strong>rn extension between <strong>the</strong> riversObi and Yenesi, where it reaches <strong>the</strong> parallel <strong>of</strong> 60° north latitude.Next to this comes a vast extent <strong>of</strong> nor<strong>the</strong>rn forests,mostly <strong>of</strong> conifers in <strong>the</strong> more nor<strong>the</strong>rn and l<strong>of</strong>ty situations,while deciduous trees preponderate in <strong>the</strong> sou<strong>the</strong>rn portions andin <strong>the</strong> more sheltered valleys. <strong>The</strong> greatest extension <strong>of</strong> thisforest region is north <strong>of</strong> Lake Baikal, where it is more than1,200 miles wide. <strong>The</strong>se forests extend along <strong>the</strong> mountainranges to join those <strong>of</strong> <strong>the</strong> Hindu Koosh. South <strong>of</strong> <strong>the</strong> forests<strong>the</strong> remainder <strong>of</strong> <strong>the</strong> sub-region consists <strong>of</strong> open pasture-landsand vast intervening deserts, <strong>of</strong> which <strong>the</strong> Gobi, and those <strong>of</strong>Turkestan between <strong>the</strong> Aral and Balkash lakes, are <strong>the</strong> most


chap, x.] THE PAL^ARCTIC REGION. 217extensive.<strong>The</strong> former is nearly 1,000 miles long, <strong>with</strong> a width<strong>of</strong> from 200 to 350 miles, and isalmost as complete a desert as<strong>the</strong> Sahara.With very few exceptions, this vast territory is exposed toan extreme climate, inimical to animal life.All <strong>the</strong> lower partsbeing situated to <strong>the</strong> north, have an excessively cold winter, sothat <strong>the</strong> limit <strong>of</strong> constantly frozen ground descends below <strong>the</strong>parallel <strong>of</strong> 60° north latitude. To <strong>the</strong> south, <strong>the</strong> land is greatlyelevated, and <strong>the</strong> climate extremely dry. In summer <strong>the</strong> heatis excessive, while <strong>the</strong> winter is almost as severe as fur<strong>the</strong>rnorth. <strong>The</strong> whole country, too, is subject to violent storms, bothin summer and winter ; and <strong>the</strong> rich vegetation that clo<strong>the</strong>s <strong>the</strong>steppes in spring, is soon parched up and replaced by dustyplains. Under <strong>the</strong>se adverse influences we cannot expectanimal life to be so abundant as in those sub-regions subject tomore favourable physical conditions ;yet <strong>the</strong> country is so extensiveand so varied, that it does actually, as we shall see, possessa very considerable and interesting *auna.Mammalia.—Four genera seem to be absolutely confined tothis sub-region, Nectogak, a peculiar form <strong>of</strong> <strong>the</strong> mole family(TalpidaB) ; Poephagus, <strong>the</strong> yak, or hairy bison <strong>of</strong> Thibet ;<strong>with</strong>Procapra and Pantholops, Thibetan antelopes. Some o<strong>the</strong>rsmore especially belong here, although <strong>the</strong>y justenter Europe, asSaiga, <strong>the</strong> Tartarian antelope ;Sminthus, a desert rat ;andEllobius, a burrowing mole-rat ; while Myospalax, a curiousrodent alliedand North China ;to <strong>the</strong> voles, is found only in <strong>the</strong> Altai mountainsand Moschus, <strong>the</strong> musk-deer, is almost confinedto this sub-region. Among <strong>the</strong> characteristic <strong>animals</strong> <strong>of</strong> <strong>the</strong>extreme north, are Mustela, and Martes, including <strong>the</strong> ermineand sable; Gulo, <strong>the</strong> glutton ; Tarandus, <strong>the</strong> reindeer; Myodcs,<strong>the</strong> lemming; <strong>with</strong> <strong>the</strong> lynx, arctic fox, and polar bear; andhere, in <strong>the</strong> Post-pliocene epoch, ranged <strong>the</strong> hairy rhinocerosand Siberian mammoth, whose entire bodies still remain preservedin <strong>the</strong> ice-cliffs near <strong>the</strong> mouths <strong>of</strong> <strong>the</strong> great rivers. Far<strong>the</strong>rsouth, species <strong>of</strong> wild cat, bear, wolf, deer, and pika (Lagomys)abound ; while in <strong>the</strong> mountains we find wild goats and sheep<strong>of</strong> several species, and in <strong>the</strong> plains and deserts wild horsesVol. L—16


218 ZOOLOGICAL GEOGRAPHY. [part m.and asses, gazelles, two species <strong>of</strong> antelopes, flying squirrels(Pteromys), ground squirrels (Tamias), marmots, <strong>of</strong> <strong>the</strong> genusSpermophihis, <strong>with</strong> camels and dromedaries, probably natives<strong>of</strong> <strong>the</strong> south-western part <strong>of</strong> this sub-region. <strong>The</strong> mostabundant and conspicuous <strong>of</strong> <strong>the</strong> mammalia are <strong>the</strong> grea<strong>the</strong>rds <strong>of</strong> reindeer in <strong>the</strong> north, <strong>the</strong> wolves <strong>of</strong> <strong>the</strong> steppes, <strong>with</strong><strong>the</strong> wild horses,goats, sheep, and antelopes <strong>of</strong> <strong>the</strong> plateaus andmountains.Among <strong>the</strong> curiosities <strong>of</strong> this sub-region we must notice <strong>the</strong>seal, found in <strong>the</strong> inland and freshwater lake Baikal, at an elevation<strong>of</strong> about 2,000 feet above <strong>the</strong> sea. It is a species <strong>of</strong> Callocephalus,closely allied to, if not identical <strong>with</strong>, one inhabitingnor<strong>the</strong>rn seas as well as <strong>the</strong> Caspian and Lake Aral. Thiswould indicate that almost all nor<strong>the</strong>rn Asia was depressedbeneath <strong>the</strong> sea very recently ; and Mr. Belt's view, <strong>of</strong> <strong>the</strong> iceduring <strong>the</strong> glacial epoch having dammed up <strong>the</strong> rivers and convertedmuch <strong>of</strong> Siberia into a vast freshwater or brackish lake,perhaps <strong>of</strong>fers <strong>the</strong> best solution <strong>of</strong> <strong>the</strong> difficulty. 1Plate II.— Characteristic Mammalia <strong>of</strong> Western Tartary.—Several <strong>of</strong> <strong>the</strong> most remarkable <strong>animals</strong> <strong>of</strong>inhabit Western Tartary, and are<strong>the</strong> Palrearctic regioncommon to <strong>the</strong> European andSiberian sub-regions. We <strong>the</strong>refore choose this district for one<strong>of</strong> our illustrative plates.<strong>The</strong> large <strong>animals</strong> in <strong>the</strong> centre are <strong>the</strong>remarkable saiga antelopes (Saiga Tartarica), distinguished fromall o<strong>the</strong>rs by a large and fleshy proboscis-like nose, which gives<strong>the</strong>m a singular appearance.<strong>The</strong>y differ so much from all o<strong>the</strong>rantelopes that <strong>the</strong>y have been formed into a distinct family bysome naturalists, but are here referred to <strong>the</strong> great family Bovidse.<strong>The</strong>y inhabit <strong>the</strong> open plains from Poland to <strong>the</strong> Irtish RiverOn <strong>the</strong> left is <strong>the</strong> mole-rat, or sand-rat (Spalax murinus). Thisanimal burrows under ground like a mole, feeding on bulbousroots. It inhabits <strong>the</strong> same country as <strong>the</strong> saiga, but extendsfar<strong>the</strong>r south in Europe. On <strong>the</strong> right is a still more curiousanimal, <strong>the</strong> desman (Myogalc Muscovitica), a long-snoutedwater-mole. This creature is fifteen inches long, including <strong>the</strong>tail ; it burrows in <strong>the</strong> banks <strong>of</strong> streams, feeding on insects,1Quarterly Journal <strong>of</strong> <strong>the</strong> Geological Society, 1874, p. 494.


chap, x.] THE PAL.EARCTIC REGION. 219worms, and leeches ; it swims well, and remains long underwater, raising <strong>the</strong> tip <strong>of</strong> <strong>the</strong> snout, where <strong>the</strong> nostrils aresituated, to <strong>the</strong> surface when it wants to brea<strong>the</strong>. It is thuswell concealed ; and this may be one use <strong>of</strong> <strong>the</strong> development<strong>of</strong> <strong>the</strong> long snout, as well as serving to follow worms into<strong>the</strong>ir holes in <strong>the</strong> s<strong>of</strong>t earth. This species is confined to <strong>the</strong>rivers Volga and Don in Sou<strong>the</strong>rn Eussia, and <strong>the</strong> only o<strong>the</strong>rspecies known inhabits some <strong>of</strong> <strong>the</strong> valleys on <strong>the</strong> north side<strong>of</strong> <strong>the</strong> Pyrenees. In <strong>the</strong> distance are wolves, a characteristicfeature <strong>of</strong> <strong>the</strong>se wastes.Birds.—But few genera <strong>of</strong> birds are absolutely restricted tothis sub-region. Podoces, a curious form <strong>of</strong> starling, is <strong>the</strong> mostdecidedly so ; Mycerobas and Pyrrhospiza are genera <strong>of</strong> finchesconfined to Thibet and <strong>the</strong> snowy Himalayas ;Leucosticte, ano<strong>the</strong>rgenus <strong>of</strong> finches, is confined to <strong>the</strong> eastern half <strong>of</strong> <strong>the</strong> subregionand North America ; Tetraogallus, a large kind <strong>of</strong>partridge, ranges west to <strong>the</strong> Caucasus; Syrrhaptcs, a form <strong>of</strong>sand-grouse, and Leriva (snow-partridge), are almost confinedhere, only extending into <strong>the</strong> next sub-region ; as do Grandalo,and Calliope, genera <strong>of</strong> warblers, Uragus, a finch allied to <strong>the</strong>North American cardinals, and Crossoptilon, a remarkable group<strong>of</strong> pheasants.Almost all <strong>the</strong> genera <strong>of</strong> central and nor<strong>the</strong>rn Europe arefound here, and give quite a European character to <strong>the</strong> ornithology,though a considerable number <strong>of</strong> <strong>the</strong>species are different.<strong>The</strong>re are a few Oriental forms, such as Abrornis and Larvivora(warblers) ; <strong>with</strong> Ceriornis and Ithaginis, genera <strong>of</strong> pheasants,which reach <strong>the</strong> snow-line in <strong>the</strong> Himalayas and thus just enterthis sub-region, but as <strong>the</strong>y do not penetrate far<strong>the</strong>r north, <strong>the</strong>yhardly serve to modify <strong>the</strong> exclusively Palsearctic character <strong>of</strong>its ornithology.According to Middendorf, <strong>the</strong> extreme nor<strong>the</strong>rn Asiatic birdsare <strong>the</strong> Alpine ptarmigan [Lagopus mutus); <strong>the</strong> snow-bunting(Plectrophanes nivalis) ; <strong>the</strong> raven, <strong>the</strong> gyrfalcon and <strong>the</strong> snowyowl.Those which are characteristic <strong>of</strong> <strong>the</strong> barren " tundras,"but which do not range so far north as <strong>the</strong> preceding are,—<strong>the</strong>willow-grouse (Lagopus albus); <strong>the</strong> Lapland-bunting (Plectrophanes


;220 ZOOLOGICAL GEOGRAPHY. [part hi.lapponica) ; <strong>the</strong> shore-lark (Gtocorys alpestris) ; <strong>the</strong> sand-martin(Cotyle riparia), and <strong>the</strong> sea-eagle {Haliceetus albicilla).Those which are more characteristic <strong>of</strong> <strong>the</strong> nor<strong>the</strong>rn forests,and which do not pass beyond <strong>the</strong>m, are—<strong>the</strong> linnet ;two crossbills(Loxia Zeucoptera and L. Curvirostra) ; <strong>the</strong> pine grosbeak[Pinicola enucleator) ; <strong>the</strong> waxwing ; <strong>the</strong> common magpie ; <strong>the</strong>common swallow ; <strong>the</strong> peregrine falcon ; <strong>the</strong> rough-legged buzzard ;and three species <strong>of</strong> owls.Fully one-half <strong>of</strong> <strong>the</strong> land-birds <strong>of</strong> Siberia are identical <strong>with</strong>those <strong>of</strong> Europe, <strong>the</strong> remainder being mostly representativespecies peculiar to Nor<strong>the</strong>rn Asia, <strong>with</strong> a few stragglers andimmigrants from China and Japan or <strong>the</strong> Himalayas.A muchlarger proportion <strong>of</strong> <strong>the</strong> wading and aquatic families are Europeanor Arctic, <strong>the</strong>se groups having always aland birds.Reptiles and Amphibia.—From <strong>the</strong> nature <strong>of</strong>wider range than<strong>the</strong> country andclimate <strong>the</strong>se are comparatively few, but in <strong>the</strong> more temperatedistricts snakes and lizards seem to be not uncommon. Halys,a genus <strong>of</strong> Crotaline snakes, and Phryotocephalus, lizards <strong>of</strong> <strong>the</strong>family Agamidse, are characteristic <strong>of</strong> <strong>the</strong>se parts. Simotes, asnake <strong>of</strong> <strong>the</strong> family Oligodontidae, reaches an elevation <strong>of</strong> 16,000feet in <strong>the</strong> Himalayas, and <strong>the</strong>refore enters this sub-region.Insects.— Mesapia and Hypermnestra, genera <strong>of</strong> Papilionidse,are butterflies peculiar to this sub-region ; and Parnassius is ascharacteristic as it is <strong>of</strong> our European mountains. Carabidseare also abundant, as will be seen by referring to <strong>the</strong> Chapteron <strong>the</strong> Distribution <strong>of</strong> Insects in <strong>the</strong> succeeding part <strong>of</strong> thiswork. <strong>The</strong> insects, on <strong>the</strong> whole, have a strictly Europeancharacter, although a large proportion <strong>of</strong> <strong>the</strong> species are peculiar,and several new genera appear.IV.— Japan and North China, or <strong>the</strong> Manchurian Sub-region.Tliis is an interesting and very productive district, correspondingin <strong>the</strong> east to <strong>the</strong> Mediterranean sub-region in <strong>the</strong> west, orra<strong>the</strong>r perhaps to all western temperate Europe. Its limits arenot very well defined, but it probably includes all Japan<strong>the</strong> Corea and Manchuria to <strong>the</strong> Amour river and to <strong>the</strong> lower


chap x.] THE PAL^AECTIC REGION. 221slopes <strong>of</strong> <strong>the</strong> Khingan and Peling mountains ; and China to<strong>the</strong> Nanlirt mountains south <strong>of</strong> <strong>the</strong> Yang-tse-kiang. On <strong>the</strong>coast <strong>of</strong> China <strong>the</strong> dividing line between it and <strong>the</strong> Orientalregion seems to be somewhere about Foo-chow, but as <strong>the</strong>re ishere no natural barrier, a great intermingling <strong>of</strong> nor<strong>the</strong>rn andsou<strong>the</strong>rn forms takes place.Japan is volcanic and mountainous, <strong>with</strong> a fine climate and amost luxuriant and varied vegetation. Manchuria is hilly, <strong>with</strong>a high range <strong>of</strong> mountains on <strong>the</strong> coast, and some desert tractsin <strong>the</strong> interior, but fairly wooded in many parts. Much <strong>of</strong>nor<strong>the</strong>rn China is a vast alluvial plain, backed by hills andmountains <strong>with</strong> belts <strong>of</strong> forest, above which are <strong>the</strong> dry andbarren uplands <strong>of</strong> Mongolia. We have a tolerable knowledge<strong>of</strong> China, <strong>of</strong> Japan, and <strong>of</strong> <strong>the</strong> Amoor valley, but very little<strong>of</strong>Corea and Manchuria. <strong>The</strong> recent researches <strong>of</strong> Pere David inMoupin, in east Thibet, said to be between 31° and 32° northlatitude, show, that <strong>the</strong> fauna <strong>of</strong> <strong>the</strong> Oriental region here advancesnorthward along <strong>the</strong> flanks <strong>of</strong> <strong>the</strong> Yun-ling mountains (acontinuation <strong>of</strong> <strong>the</strong> Himalayas) ; since he found at differentaltitudes representatives <strong>of</strong> <strong>the</strong> Indo-Chinese, Manchurian, andSiberian faunas. On <strong>the</strong> higher slopes <strong>of</strong> <strong>the</strong> Himalayas, <strong>the</strong>remust be a narrow strip from about 8,000 to 11,000 feet elevationintervening between <strong>the</strong> tropical fauna <strong>of</strong> <strong>the</strong> Indo-Chinese subregionand <strong>the</strong> almost arctic fauna <strong>of</strong> Thibet ; and <strong>the</strong> <strong>animals</strong><strong>of</strong> this zone will for <strong>the</strong> most part belong to <strong>the</strong> fauna <strong>of</strong>temperate China and Manchuria, except in <strong>the</strong> extreme westtowards Cashmere, where <strong>the</strong> Mediterranean fauna will in likemanner intervene. On a map <strong>of</strong> sufficiently large scale, <strong>the</strong>refore,it would be necessary to extend our present sub-regionwestward along <strong>the</strong> Himalayas, in a narrow strip just below<strong>the</strong> upper limits <strong>of</strong> forests. It is evident that <strong>the</strong> large number<strong>of</strong> Fringillidae, Corvidae, Troglodytidae, and Paridse, <strong>of</strong>ten <strong>of</strong> southPalsearctic forms, that abound in <strong>the</strong> higher Himalayas, are somewhatout <strong>of</strong> place as members <strong>of</strong> <strong>the</strong> Oriental fauna, and areequally so in that <strong>of</strong> Thibet and Siberia; but <strong>the</strong>y form anatural portion <strong>of</strong> that <strong>of</strong> North China on <strong>the</strong> one side, or <strong>of</strong>South Europe on <strong>the</strong> o<strong>the</strong>r.


—;222 ZOOLOGICAL GEOGRAPHY. [part in.Mammalia.—This sub-region contains a number <strong>of</strong> peculiarand very interesting forms, most <strong>of</strong> which have been recentlydiscovered by Pere David in North and West China and EastThibet. <strong>The</strong> following are <strong>the</strong> peculiar genera Rhinopithccus,:a sub-genus <strong>of</strong> monkeys, here classed under Semnopi<strong>the</strong>ousAnurosorex, Scaptochirus,Uropsilus and Scaptonyx, new forms <strong>of</strong>Talpiche or moles ; JEluropus (iEluridte) ; Nyctereutes (Canidae)Lutroncctes (Mustelida?) ; Cricetulus (Muridse) ; Hydropotes, Moschus,and Elaphodus (Cervidse). <strong>The</strong> Rhinopi<strong>the</strong>cus appears tobe a permanent inhabitant <strong>of</strong> <strong>the</strong> highest forests <strong>of</strong> Moupin,in a cold climate. It has a very thick fur, as has also a newspecies <strong>of</strong> Macacus found in <strong>the</strong> same district.East Thibet seem to be very rich in Insectivora.North China andScaptochirus islike a mole ; Uropsilus between <strong>the</strong> Japanese Urotrichus andSorex ; Scaptonyx between Urotrichus and Talpa. JEluropusseems to be <strong>the</strong>most remarkable mammal discovered by PereDavid. It is allied to <strong>the</strong> singular panda {JElurus fulgens) <strong>of</strong>Nepal, but is as large as a bear, <strong>the</strong> body wholly white, <strong>with</strong><strong>the</strong> feet, ears, and tip <strong>of</strong> <strong>the</strong> tail black. It inhabits <strong>the</strong> highestforests, and is <strong>the</strong>refore a true Palsearctic animal, as most likelyis <strong>the</strong> JElurus. Nyctereutes, a curious racoon-like dog, rangesfrom Canton to North China, <strong>the</strong> Amoor and Japan,and <strong>the</strong>reforeseems to come best in this sub-region ; Hydropotes andLophotragus are small hornless deer confined to North China ;Elaphodus, from East Thibet, isano<strong>the</strong>r peculiar form <strong>of</strong> deer;while <strong>the</strong> musk deer (MoscJms) is confined to this sub-region and<strong>the</strong> last. Besides <strong>the</strong> above, <strong>the</strong> following Palaearctic generawere found by Pere David in this sub-region : Macacus ; fivegenera or sub-genera <strong>of</strong> bats (Vespertilio, Vcsperus, Vesperugo,Rhinolophus, and Murina) ; Erinaceus, Nectogale, Talpa, Crociduraand Sorcx, among Insectivora; Mustela, Putorius, Martes,Lutra, Viverra, Mcles, JElurus, Ursus, Fclis, and Canis, amongCarnivora ; Hystrix, Arctomys, Myospalax, Spcrmophilus, Gerbillus,Dipus, Lagomys, Lepus, Sciurus, Ptcromys, Arvicola, andMus, among Eodentia; Budorcas, Ncmorhcdus, Antilope, Ovis,Moschus, Cervulus and Cervus among Kuminants ;and <strong>the</strong> widespreadSus or wild boar.<strong>The</strong> following Oriental genera are also


—chap, x.] THE PAL^ARCTIC REGION. 223included in Pere David's list, but no doubt occur only in <strong>the</strong>lowlands and warm valleys, and can hardly be considered tobelong to <strong>the</strong> Paloearctic region : Paguma, Helictis, Arctonyx,Rhizomys, Manis. <strong>The</strong> Rhizomys from Moupin is a peculiarspecies <strong>of</strong> this tropical genus, but all <strong>the</strong> o<strong>the</strong>rs inhabit Sou<strong>the</strong>rnChina.A few additional forms occur in Japan :Urotrichus, a peculiarMole, which is found also in north-west America ; Enhydra, <strong>the</strong>sea otter <strong>of</strong> California; and <strong>the</strong> dormouse (Myoxus). Japan alsopossesses peculiar species <strong>of</strong> Macacus, Talpa, Meles, Canis, andSciuropterus.It will be seen that this sub-region is remarkably richin Insectivora, <strong>of</strong> which it possesses ten genera ; and thatit has also several peculiar forms <strong>of</strong> Carnivora, Ptodentia, andRuminants.Birds.—To give an accurate idea <strong>of</strong> <strong>the</strong> ornithology <strong>of</strong> thissub-region is very difficult,both on account <strong>of</strong> its extreme richnessand <strong>the</strong> impossibility <strong>of</strong> defining <strong>the</strong> limits between it and<strong>the</strong> Oriental region. A considerable number <strong>of</strong> genera whicharewell developed in <strong>the</strong> high Himalayas, and some which arepeculiar to that district, have hi<strong>the</strong>rto always been classed asIndian, and <strong>the</strong>refore Oriental groups ; but <strong>the</strong>y more properlybelong to this sub-region. Many <strong>of</strong> <strong>the</strong>m frequent <strong>the</strong> highestforests, or descend into <strong>the</strong> Himalayan temperate zone only inwinter ; and o<strong>the</strong>rs are so intimately connected <strong>with</strong> Paleearcticspecies, that <strong>the</strong>y can only be considered as stragglers into <strong>the</strong>border land <strong>of</strong> <strong>the</strong> Oriental region. On <strong>the</strong>se principles weconsider <strong>the</strong> following genera to be confined to this sub-region :Grandala, Nemura (Sylviidee) ; Pterorhinus (Timaliidie) ;Cholomis, Conostoma, Heteromorpha (Panuridre) ; Cyanoptila(Muscicapidae) ; Eophona (Fringillidae) ; Dendrotreron (Columbidae);Lophoylioms, Tetraophasis, Crossoptilon, Pucrasia, Thauinnlea, and Ithaginis (Phasianidse). This may be called <strong>the</strong>sub-region <strong>of</strong> Pheasants ; for <strong>the</strong> above six genera, comprisingsixteen species <strong>of</strong> <strong>the</strong> most magnificent birds in <strong>the</strong> world, areall confined to <strong>the</strong> temperate or cold mountainous regions <strong>of</strong><strong>the</strong> Himalayas, Thibet, and China ; and in addition we have


224 ZOOLOGICAL GEOGRAPHY. [PART III.most <strong>of</strong> <strong>the</strong> species <strong>of</strong> tragopan (Ceriornis), and some <strong>of</strong> <strong>the</strong>true pheasants (Phasianus).<strong>The</strong> most abundant and characteristic <strong>of</strong> <strong>the</strong> smaller birds arewarblers, tits, and finches, <strong>of</strong> Palsearctic types ;but <strong>the</strong>re arealso a considerable number <strong>of</strong> Oriental forms which penetratefar into <strong>the</strong> country, and mingling <strong>with</strong> <strong>the</strong> nor<strong>the</strong>rn birdsgive a character to <strong>the</strong> Ornithology <strong>of</strong> this sub-region verydifferent from that <strong>of</strong> <strong>the</strong> Mediterranean district at <strong>the</strong> westernend <strong>of</strong> <strong>the</strong> region. Leaving out a large number <strong>of</strong> wide-ranginggivinggroups, this mixture <strong>of</strong> types may be best exhibited bylists <strong>of</strong> <strong>the</strong> more striking Palsearctic and Oriental genera whichare here found intermingled.Pal^earcticGenera.Sylviid^e.Erithacus.Ruticilla.Locustella.Cyanecula.Sylvia.Potamodus.Reguloides.Regulus.Accentor.ClNCLID^E.Cinclus.TROGLODYTIDiE.Troglodytidas.CeRTHIIDjE.Certhia.Sitta.Tichodroma.PaRIDjE.Parus.Lophophanes.Acredula.CoRVID^E.


CHAP. X.] THE PAL^ARCTIC REGION. 225Omental Genera— continued.TimaliiDjE— (continued).Pomatorhinus.Suthora.Panurid^:.Paradoxornis.ClNCLID^.Enicurus.Myiophonus.TrOGLODYTID^E.Pnoepyga.Liotrichid^e.Liothrix.Yuhina.Pteruthius.Pycnonotid^e.Microscelis.Pycnonotus.Hypsipetes.Campephagid.e.Pericrocotus.Dicruridje.Dicrurus.Chibia.Buchanga.Muscicapid^e.Xanthopygia.Niltava.Tchitrea.CoRyiD^;.Urocissa.Nectarineid^;.^Ethopyga.MOTACILLID.E.Nemoricola.DicjEID.e.Zosterops.Fringillid^e.Melophus.Pyrgilauda.Ploceid.e.Munia.Sturnid^e.Acrido<strong>the</strong>res.Sturnia.Pittid.e.Pitta.PlCID^E.Vivia.Yungipicus.Gecinus.CoRACIlDiF.Eurystomus.AlCEDINIDjE.Halcyon.Ceryle.Ufupid^e.Upupa.PsiTTACID^E.Palseornis.CoLUMBID-E.Treron.Ianthsenas.Macropygia.Phasianid^e.Phasianus.Ceriornis.Strigid^e.Scops.In <strong>the</strong> above lists <strong>the</strong>re are ra<strong>the</strong>r more Oriental than Palaearcticgenera ; but it must be remembered that most <strong>of</strong> <strong>the</strong>former are summer migrants only, or stragglers just entering <strong>the</strong>sub-region; whereas <strong>the</strong> great majority <strong>of</strong> <strong>the</strong> latter are permanentresidents, and a large proportion <strong>of</strong> <strong>the</strong>m range over <strong>the</strong>greater part <strong>of</strong> <strong>the</strong> Manchurian district. Many <strong>of</strong> those in <strong>the</strong>Oriental column should perhaps be omitted, as we have no exactdetermination <strong>of</strong> <strong>the</strong>ir range, and <strong>the</strong> limits <strong>of</strong> <strong>the</strong> regions arevery uncertain.It must be remembered, too, that <strong>the</strong> Palsearcticgenera <strong>of</strong> Sylviida?, Paridoe, and Fringillidae, are <strong>of</strong>ten representedby numerous species, whereas <strong>the</strong> corresponding Oriental generahave for <strong>the</strong> most part only single species ; and we shall <strong>the</strong>nfind that, except towards <strong>the</strong> borders <strong>of</strong> <strong>the</strong> Oriental region <strong>the</strong>Palaearctic element is strongly predominant. Four <strong>of</strong> <strong>the</strong> moreespecially Oriental groups are confined to Japan, <strong>the</strong> sou<strong>the</strong>rn


226 ZOOLOGICAL GEOGRAPHY [part hi.extremity <strong>of</strong> which should perhaps come in <strong>the</strong> Oriental region.<strong>The</strong> great richness <strong>of</strong> this sub-region compared <strong>with</strong> that <strong>of</strong>Siberia is well shown by <strong>the</strong> fact, that a list <strong>of</strong> all <strong>the</strong> know nland-birds<strong>of</strong> East Siberia, including Dahuria and <strong>the</strong> comparativelyfertile Amoor Valley, contains only 190 species ; whereasPere David's catalogue <strong>of</strong> <strong>the</strong> birds <strong>of</strong> Nor<strong>the</strong>rn China <strong>with</strong>adjacent parts <strong>of</strong> East Thibet and Mongolia (a very muchsmaller area) contains for <strong>the</strong> same families 366 species. Of <strong>the</strong>Siberian birds more than 50 per cent, are European species, whilethose <strong>of</strong> <strong>the</strong> Manchurian sub-region comprise about half thatproportion <strong>of</strong> land-birds which are identical <strong>with</strong> those <strong>of</strong>Europe.Japan is no doubt very imperfectly known, as only 134 landbirdsare recorded from it. Of <strong>the</strong>se twenty- two are peculiarspecies, a number that would probably bediminished were <strong>the</strong>Corea to be explored. Of <strong>the</strong> genera, only nine are Indo-Malayan, while forty-three are Palsearctic.Plate III.— Scene on <strong>the</strong> Borders <strong>of</strong> North- West China andMongolia <strong>with</strong> Characteristic Mammalia and Birds. — <strong>The</strong>mountainous districts <strong>of</strong> Nor<strong>the</strong>rn China, <strong>with</strong> <strong>the</strong> adjacentportions <strong>of</strong> Thibet and Mongolia, are <strong>the</strong> head-quarters <strong>of</strong> <strong>the</strong>pheasant tribe, many <strong>of</strong> <strong>the</strong> most beautiful and remarkablespecies being found <strong>the</strong>re only.In <strong>the</strong> north-western provinces<strong>of</strong> China and <strong>the</strong> sou<strong>the</strong>rn parts <strong>of</strong> Mongolia may be found <strong>the</strong>species figured. That in <strong>the</strong> foreground is <strong>the</strong> superb goldenpheasant (Thaumalea picta), a bird that can hardly be surpassedfor splendour <strong>of</strong> plumage by any denizen <strong>of</strong> <strong>the</strong> tropics. <strong>The</strong>large bird perched above is <strong>the</strong> eared pheasant {Crossoptilonauriium), a species <strong>of</strong> comparatively sober plumage but <strong>of</strong>remarkable and elegant form.In <strong>the</strong> middle distance is Pallas'ssand grouse {Syrrhaptes paradoxus), a curious bird, whose nativecountry seems to be <strong>the</strong> high plains <strong>of</strong> Nor<strong>the</strong>rn Asia, but which<strong>of</strong>ten abounds near Pekin, and in 1863 astonished Europeanornithologists by appearing inconsiderable numbers in Centraland Western Europe, in every part <strong>of</strong> Great Britain, and evenm Ireland.<strong>The</strong> quadruped figured is <strong>the</strong> curious racoon dog (Nyctermtes


PLATE III.CHARACTERISTIC ANIMALS OF NORTH CHINA.


;chap, x.] THE PALiEARCTIC REGION. 227procyonoides), an animal confined to North China, Japan, and<strong>the</strong>Amoor Valley, and having no close allies in any o<strong>the</strong>r part<strong>of</strong> <strong>the</strong> globe.In <strong>the</strong> distance are some deer, a group <strong>of</strong> <strong>animals</strong>very abundant and varied in this part <strong>of</strong> <strong>the</strong> Palaearctic region.Reptiles and Amphibia.—Reptiles are scarce in North China,only four or five species <strong>of</strong> snakes, a lizard and one <strong>of</strong> <strong>the</strong> Geckotidaeoccurring in <strong>the</strong> country round Pekin. <strong>The</strong> genus Halysis <strong>the</strong> most characteristic form <strong>of</strong> snake, while Callophis, anoriental genus, extends to Japan. Among lizards, Plestiodon,Maybouya, Tachydromus, and Gecko reach Japan, <strong>the</strong> two latterbeing very characteristic <strong>of</strong> <strong>the</strong> Oriental region.Amphibia are more abundant and interesting ; Hynobius,Onychodactylus, and Sieboldtia (Salamandridae) being peculiarto it,while most <strong>of</strong> <strong>the</strong> European genera are also represented.Fresh-water Fish.—Of <strong>the</strong>se <strong>the</strong>re are a few peculiar genera;as Plecoglossvs (Salmonidae) from Japan; Achilognathus, Pseudoperilampus,Ochetobius, and Opsariichthys (Cyprinidae) ; and<strong>the</strong>re are many o<strong>the</strong>r Chinese Cyprinidae belonging to <strong>the</strong> borderland <strong>of</strong> <strong>the</strong> Palaearctic and Oriental regions.Insects—<strong>The</strong> butterflies <strong>of</strong> this sub-region exhibit <strong>the</strong> samemixture <strong>of</strong> tropical and temperate forms as <strong>the</strong> birds. Most <strong>of</strong><strong>the</strong> common European genera are represented, and <strong>the</strong>re arespecies <strong>of</strong> Parnassius in Japan and <strong>the</strong> Amoor. Isodema, apeculiar genus <strong>of</strong> Nymphalidae isfound near Ningpo, just <strong>with</strong>inour limits ; and Sericinus, one <strong>of</strong> <strong>the</strong> most beautiful genera <strong>of</strong>Papilionidae is peculiar to North China, where four species occur,thus balancing <strong>the</strong> Thais and Doritis <strong>of</strong> Europe. <strong>The</strong> genusZephyrus (Lycaenidae) is well represented by six species in Japanand <strong>the</strong> Amoor, against two in Europe. Papilio parts andP. Manor, magnificent insects <strong>of</strong> wholly tropical appearance,abound near Pekin, and allied forms inhabit Japan and <strong>the</strong>Amoor, as well as P. demetrius and P. alcinous belonging to<strong>the</strong> " Protenor " group <strong>of</strong> <strong>the</strong> Himalayas.O<strong>the</strong>r tropical generaoccurring in Japan, <strong>the</strong> Amoor, or North China are, Debis,Neope, Mycalesis, Ypthimia (Satyridae) ;Thaumantis (Morphidae),at Shanghae ;Euripus, Neptis, Athyma (Nymphalidae)Terias (Pieridae) ;and <strong>the</strong> above-mentioned Papilionidae.


;228 ZOOLOGICAL GEOGRAPHY. [part hi.Coleoptera.—<strong>The</strong> beetles <strong>of</strong> Japan decidedly exhibit a mixture<strong>of</strong> tropical forms <strong>with</strong> o<strong>the</strong>rs truly Palsearctic, and it has been<strong>with</strong> some naturalists a matter <strong>of</strong> doubt whe<strong>the</strong>r <strong>the</strong> sou<strong>the</strong>rn andbest known portion <strong>of</strong> <strong>the</strong> islands should not be joined to <strong>the</strong>Oriental region. An important addition to our knowledge <strong>of</strong><strong>the</strong> insects <strong>of</strong> this country has recently been made by Mr. GeorgeLewis, and a portion <strong>of</strong> his collections have been described byvarious entomologists in <strong>the</strong> Transactions <strong>of</strong> <strong>the</strong> EntomologicalSociety <strong>of</strong> London. As <strong>the</strong> question is one <strong>of</strong> considerable interestwe shall give a summary <strong>of</strong> <strong>the</strong> results fairly deduciblefrom what is now known <strong>of</strong> <strong>the</strong> entomology <strong>of</strong> Japan ;and itmust be remembered that almost all our collections come from<strong>the</strong> sou<strong>the</strong>rn districts, in what is almost a sub-tropical climateso that if we find a considerable proportion <strong>of</strong> Palsearctic forms,we may be pretty sure that <strong>the</strong> preponderance will be muchgreater a little fur<strong>the</strong>r north.Of Carabidae Mr. Bates enumerates 244 speciesbelonging to84 genera, and by comparing <strong>the</strong>se <strong>with</strong> <strong>the</strong> Coleoptera <strong>of</strong> atract <strong>of</strong> about equal extent in western Europe, he concludes that<strong>the</strong>re is little similarity, and that <strong>the</strong> cases <strong>of</strong> affinity to <strong>the</strong> forms<strong>of</strong> eastern tropical Asia preponderate. By comparing his genera<strong>with</strong> <strong>the</strong> <strong>distribution</strong>s as given in Gemminger and Harold'sCatalogue, a somewhat different result is arrived at. Leavingout <strong>the</strong> generic types altoge<strong>the</strong>r peculiar to Japan, and also thoseo-enera <strong>of</strong> such world-wide <strong>distribution</strong> that <strong>the</strong>y afford no clearindications for our purpose, it appears that no less than twentytwogenera, containing seventy-four <strong>of</strong> <strong>the</strong> Japanese species, areei<strong>the</strong>r exclusively Palsearctic, Palsearctic and Nearctic, or highlycharacteristic <strong>of</strong> <strong>the</strong> Palsearctic region ; <strong>the</strong>n come thirteen generacontaining eighty-seven <strong>of</strong> <strong>the</strong> species which have a very wide<strong>distribution</strong>, but are also Palsearctic : we next have seventeen^enera containing twenty-four <strong>of</strong> <strong>the</strong> Japanese species which aredecidedly Oriental and tropical. Here <strong>the</strong>n <strong>the</strong> fair comparisonis between <strong>the</strong> twenty-two genera and seventy-four species whoseaffinities are clearly Palsearctic or at least north temperate, andseventeen genera <strong>with</strong> twenty-four species which are Asiaticand tropical; and this seems to prove that, although South


chap, x.] THE PAL^ARCTIC REGION. 229Japan (like North China) has a considerable infusion <strong>of</strong> tropicalforms, <strong>the</strong>re is a preponderating substratum <strong>of</strong> Palrearctic forms,which clearly indicate <strong>the</strong> true position <strong>of</strong> <strong>the</strong> islandsin zoologicalgeography. <strong>The</strong>re are also a few cases <strong>of</strong> what may bewhich show that Japan, like manycalled eccentric <strong>distribution</strong> ;o<strong>the</strong>r island-groups, has served as a kind <strong>of</strong> refuge in whichdying-out forms continue to maintain <strong>the</strong>mselves.<strong>The</strong>se, whichare worthy <strong>of</strong> notice, are as follows : Orthotrichus (1 sp.) has<strong>the</strong> only o<strong>the</strong>r species in Egypt ; Trechichus (1 sp.) has twoo<strong>the</strong>r species, <strong>of</strong> which one inhabits Madeira, <strong>the</strong> o<strong>the</strong>r <strong>the</strong>Sou<strong>the</strong>rn United States ;Perileptus (1 sp.) has two o<strong>the</strong>r species,<strong>of</strong> which one inhabits Bourbon, <strong>the</strong> o<strong>the</strong>r West Europe ; andlastly, Crepidogaster (1 sp.) has <strong>the</strong> o<strong>the</strong>r known species inSouth Africa.<strong>The</strong>se cases diminish <strong>the</strong> value <strong>of</strong> <strong>the</strong> indicationsafforded by some <strong>of</strong> <strong>the</strong> Japanese forms, whose only allies aresingle species in various remote parts <strong>of</strong> <strong>the</strong> Oriental region.<strong>The</strong> Staphylinidse have been describedby Dr. Sharp, and hislist exhibits a great preponderance <strong>of</strong> north temperate, or cosmopolitanforms, <strong>with</strong> a few which are decidedly tropical. <strong>The</strong>Pselaphidae and Scydmenidre, also described by Dr. Sharp,exhibit, according to that gentleman, " even a greater resemblanceto those <strong>of</strong> North America than to those <strong>of</strong> Europe," but he saysnothing <strong>of</strong> any tropical affinities. <strong>The</strong> water-beetles are allei<strong>the</strong>r Paleearctic or <strong>of</strong> wide <strong>distribution</strong>.<strong>The</strong> Lucanidse{Gemm. and Har. Cat, 1868) exhibit an intermingling<strong>of</strong> Palaearctic and Oriental genera.<strong>The</strong> Cetoniidse {Gemm. and Har. Gat. 1869) show, for NorthChina and Japan, three Oriental to two Palaearctic genera.<strong>The</strong> Buprestidse collected by Mr. Lewis have been describedby Mr. Edward Saunders in <strong>the</strong> Journal <strong>of</strong> <strong>the</strong> Linnman Society,vol. xi. p. 509. <strong>The</strong> collection consisted <strong>of</strong> thirty-six speciesbelonging to fourteen genera. No less than thirteen <strong>of</strong> <strong>the</strong>seare known also from India and <strong>the</strong> Malay Islands ; nine fromEurope ; seven from Africa ; six from America, and four fromChina.In six <strong>of</strong> <strong>the</strong> genera <strong>the</strong> Japanese species are said to beallied to those <strong>of</strong> <strong>the</strong> Oriental region ;while in three <strong>the</strong>y areallied to European forms, and in two to American.Considering


;230 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>the</strong> sou<strong>the</strong>rn latitude and warm climate in which <strong>the</strong>seinsectswere mostly collected, and <strong>the</strong> proximity to Formosa and <strong>the</strong>Malay Islands compared <strong>with</strong> <strong>the</strong> enormous distance fromEurope, this shows as much Palaearctic affinity as can beexpected. In <strong>the</strong> Palaearctic region <strong>the</strong> group is only plentifulin <strong>the</strong> sou<strong>the</strong>rn parts <strong>of</strong> Europe, which is cut<strong>of</strong>f by <strong>the</strong> coldplateau <strong>of</strong> Thibet from all direct communication <strong>with</strong> Japan;while in <strong>the</strong> Oriental region it everywhere abounds and is, infact, one <strong>of</strong> <strong>the</strong> most conspicuous and dominant families <strong>of</strong>Coleoptera.<strong>The</strong> Longicorns collected by Mr. Lewis have been describedby Mr. Bates in <strong>the</strong> Annals <strong>of</strong> Natural History for 1873.number <strong>of</strong> species now known from Japan is<strong>The</strong>107, belonging tosixty-four genera. <strong>The</strong> most important genera are Lcptura,'Clytanthus, Monohammus, Praonctha, Exoccntrus, Glenea, andOberea. <strong>The</strong>re are twenty-one tropical genera, and sevenpeculiar to Japan, leaving thirty-six ei<strong>the</strong>r Palaearctic or <strong>of</strong> verywide range. A number <strong>of</strong> <strong>the</strong> genera are Oriental and Malayan,and many characteristic European genera seem to be absentbut it is certaiu that not half <strong>the</strong> Japanese Longicorns are yetknown, and many <strong>of</strong> <strong>the</strong>se gaps will doubtless be filled up when<strong>the</strong> more nor<strong>the</strong>rn islands are explored.<strong>The</strong> Phytophaga, described by Mr. Baly, appear to have aconsiderable preponderance <strong>of</strong> tropical Oriental forms.A considerable collection <strong>of</strong> Hymenoptera formed by Mr.Lewis have been described by Mr. Frederick Smith ;and exhibit<strong>the</strong> interesting result, that while <strong>the</strong> bees and wasps are decidedly<strong>of</strong> tropical and Oriental forms, <strong>the</strong> Tenthredinidae and Ichneumonidaeare as decidedly Pala?arctic, " <strong>the</strong> generalaspect <strong>of</strong> <strong>the</strong>collection being that <strong>of</strong> a European one, only a single exoticform being found among <strong>the</strong>m."Remarks on <strong>the</strong> General Character <strong>of</strong> <strong>the</strong> Fauna <strong>of</strong> Japan —From a general view <strong>of</strong> <strong>the</strong> phenomena <strong>of</strong> <strong>distribution</strong> we feeljustified in placing Japan in <strong>the</strong> Palaearctic region ; althoughsome tropical groups, especially <strong>of</strong> reptiles and insects, havelargely occupied its sou<strong>the</strong>rn portions ;and <strong>the</strong>se same groupshave in many cases spread into Nor<strong>the</strong>rn China, beyond <strong>the</strong>


chap, x.] THE PAL^EARCTIC REGION 231usual dividing line <strong>of</strong> <strong>the</strong> Palsearctic and Oriental regions. <strong>The</strong>causes <strong>of</strong> such a phenomenon are not difficult to conceive. Evennow, that portion <strong>of</strong> <strong>the</strong> Palsearctic region between WesternAsia and Japan is, for <strong>the</strong> most part, a bleak and inhospitableregion, abounding in desert plateaus, and <strong>with</strong> a rigorous climateeven in its most favoured districts, and can, <strong>the</strong>refore, supportbut a scanty population <strong>of</strong> snakes, and <strong>of</strong> such groups <strong>of</strong>insects as require flowers, forests, or a considerable period <strong>of</strong>warm summer wea<strong>the</strong>r ; and it is precisely <strong>the</strong>se which arerepresented in Japan and North China by tropical forms.Wemust also consider, that during <strong>the</strong> Glacial epoch this wholeregion would have become stillless productive, and that, as <strong>the</strong>sou<strong>the</strong>rn limit <strong>of</strong> <strong>the</strong> ice retired northward, it would be followedup by many tropical forms along <strong>with</strong> such as had been drivensouth by its advance, and had survived to return to <strong>the</strong>irnor<strong>the</strong>rn homes.It isalso evident that Japan has a more equable and probablymoister climate than <strong>the</strong> opposite shores <strong>of</strong> China, and has alsoa very different geological character, being rocky and broken,<strong>of</strong>ten volcanic, and supporting a rich,varied, and peculiar vegetation.It would thus be well adapted to support all <strong>the</strong> morehardy denizens <strong>of</strong> <strong>the</strong> tropics which might at various timesreach it, while it might not be so well adapted for <strong>the</strong> moreboreal forms from Mongolia or Siberia. <strong>The</strong> fact that a mixture<strong>of</strong> such forms occurs <strong>the</strong>re, is <strong>the</strong>n, little to be wondered at, butwe may ra<strong>the</strong>r marvel that <strong>the</strong>y are not more predominant, andthat even in <strong>the</strong> extreme south, <strong>the</strong> most abundant forms <strong>of</strong>mammal, bird, and insect, are modifications <strong>of</strong> familiar Palaearctictypes. <strong>The</strong> fact clearly indicates that <strong>the</strong> former land connections<strong>of</strong> Japan <strong>with</strong> <strong>the</strong> continent have been in a nor<strong>the</strong>rlyra<strong>the</strong>r than in a sou<strong>the</strong>rly direction, and that <strong>the</strong> tropical immigrantshave had difficulties to contend <strong>with</strong>, and have found <strong>the</strong>land already fairly stocked <strong>with</strong> nor<strong>the</strong>rn aborigines inevery class and order <strong>of</strong> <strong>animals</strong>.almostGeneral Conclusions as to <strong>the</strong> Fauna <strong>of</strong> <strong>the</strong> Palccarctic Region.—From<strong>the</strong> account that has now been given <strong>of</strong> <strong>the</strong> fauna


232 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>of</strong> <strong>the</strong> Palaearctic region, it is evident that it owes many <strong>of</strong> itsdeficiencies and some <strong>of</strong> its peculiarities to <strong>the</strong> influence <strong>of</strong> <strong>the</strong>Glacial epoch, combined <strong>with</strong> those important changes <strong>of</strong> physicalgeography which accompanied or preceded it. <strong>The</strong> elevation <strong>of</strong><strong>the</strong> old Sarahan sea and <strong>the</strong> complete formation <strong>of</strong> <strong>the</strong> Mediterranean,are <strong>the</strong> most important <strong>of</strong> <strong>the</strong>se changes in <strong>the</strong> westernportion <strong>of</strong> <strong>the</strong> region. In <strong>the</strong> centre, a wide arm <strong>of</strong> <strong>the</strong> ArcticOcean extended southward from <strong>the</strong> Gulf <strong>of</strong> Obi to <strong>the</strong> Aral and<strong>the</strong> Caspian, dividing nor<strong>the</strong>rn Europe and Asia. At this timeour European and Siberian sub-regions were probably moredistinct than <strong>the</strong>y are now, <strong>the</strong>ir complete fusion having beeneffected since <strong>the</strong> Glacial epoch. As we know that <strong>the</strong> Himalayashave greatly increased in altitude during <strong>the</strong> Tertiary period, it isnot impossible that during <strong>the</strong> Miocene and Pliocene epochs <strong>the</strong>vast plateau <strong>of</strong> Central Asia was much less elevated and lesscompletely cut <strong>of</strong>f from <strong>the</strong> influence <strong>of</strong> rain- bearing winds. Itmight <strong>the</strong>n have been far more fertile, and have supported a richand varied animal population, a few relics <strong>of</strong> which we see in<strong>the</strong> Thibetan antelopes, yaks, and wild horses. <strong>The</strong> influence<strong>of</strong> yet earlier changes <strong>of</strong> physical geography, and <strong>the</strong> <strong>relations</strong> <strong>of</strong><strong>the</strong> Palsearctic to <strong>the</strong> tropical regions immediately south <strong>of</strong> it,will be better understood when we have examined and discussed<strong>the</strong> faunas <strong>of</strong> <strong>the</strong> Ethiopian and Oriental regions.


;chap, x.] THE PAL^ARCTIC REGION. 233TABLES OF DISTRIBUTION.In constructing <strong>the</strong>se tablesshowing <strong>the</strong> <strong>distribution</strong> <strong>of</strong> variousclasses <strong>of</strong> <strong>animals</strong> in <strong>the</strong> Palaearctic region, <strong>the</strong> followingsources <strong>of</strong> information have been chiefly relied on, in additionto <strong>the</strong> general treatises, monographs, and catalogues used incompiling <strong>the</strong> fourth part <strong>of</strong> this work.Mammalia.—Lord Clement's Mammalia and Pieptiles <strong>of</strong>Europe ;Siebold's Eauna Japonica ; Pere David's List <strong>of</strong>Mammalia <strong>of</strong> North China and Thibet ; Swinhoe's ChineseMammalia ;Eadde's List <strong>of</strong> Mammalia <strong>of</strong> South-Eastern SiberiaCanon Tristram's, Lists for Sahara and Palestine ; Papers byPr<strong>of</strong>essor Milne-Edwards, Mr. Blanford, Mr. Sclater, and <strong>the</strong>local lists given by Mr, A. Murray in <strong>the</strong> Appendix to hisGeographical Distribution <strong>of</strong> Mammalia.Birds.—Blasius' List <strong>of</strong> Birds <strong>of</strong> Europe; Godman, OnBirds <strong>of</strong> Azores, Madeira, and Canaries ; Middendorf, forSiberia ;Pere David and Mr, Swinhoe, for China and MongoliaHomeyer, for East Siberia; Mr. Blanford, for Persia and <strong>the</strong>high Himalayas ; Mr. Elwes's paper on <strong>the</strong> Distribution <strong>of</strong>Asiatic Birds ; Canon Tristram, for <strong>the</strong> Sahara and PalestinePr<strong>of</strong>essor Newton, for Iceland and Greenland ; Mr. Dresser,for Scandinavia ; and numerous papers and notes in <strong>the</strong> IbisJournal fiir Ornithologie ; Annals and Mag. <strong>of</strong> Nat. History ; andProceedings <strong>of</strong> <strong>the</strong> Zoological Society.Reptiles and Amphibia.— Schreiber's European Herpetology.Vol. I.—17


.234 ZOOLOGICAL GEOGRAPHY. [part in.TABLE I.FAMILIES OF ANIMALS INHABITING THE PAL^EARCTIC REGION.Explanation.Names in italics show families peculiar to <strong>the</strong> region.Names inclosed thus ( ) barely enter <strong>the</strong> region, and are not considered properlyto belong to it.Numbers are not consecutive, but correspond to those in Part IV.(Order aud Family.OSub-regions.is ~ sRange beyond <strong>the</strong> Region.MAMMALIA.PPvIMATES.3. Cynopi<strong>the</strong>cidseChiroptera.9. (Pteropidse^ ...11. Rhinolophidae12. Vespertilionidae13. NoctilionidaB . .Insectivora.15. Macroscelididte17. Erinaceidse ...21. Talpidae22. SoricidseCarnivora.23. Felidfe ...25. Viverridae27. Hyamidse28. Canidte ...29. Mustelidae31. jEluridae32. Ursidse ...33. Otariidae...34. Trichechidse35. PhocidseCetacea.36 to 41.Sirenia.42. ManatidseUngulata43. Equidse ...47. Suidae ...48. Camelidae50. Cervidse...52. Bovidae ...Ethiopian, OrientalTropics <strong>of</strong> E. HemisphereWarmer parts <strong>of</strong> E. Hemis.CosmopoliteTropical regionsEthiopianOriental, S. AfricaNearctic, OrientalCosmopolite, excl. Australia and S.All regions but AustralianEthiopian, OrientalEthiopian, OrientalAll regions but AustralianAll regions but AustralianOrientalNearctic, Oriental, AndesN. and S. temperate zonesArctic regionsN. and S. temperate zonesOceanicTropics, from Brazil to N. AustraliaAmericaEthiopianCosmopolite, excl. Nearctic reg. and AustraliaAndesAll regions but Ethiopian and AustralianAll regions but Neotropical and Australian


CHAP. X.] THE PAL^ARCTIC REGION. 235Sub-regions.Order and Family.PiO2SS•sS3aP.csRange beyond <strong>the</strong> Region.Hyracoidae.54. (Hyracidae)Ethiopian familyRODENTIA.55. Muridae ...56. Spalacidse57. Dipodidae58. Myoxidse60. Castoridae61. Sciuridae...64. Octodontidae67. Hystricidae69. Lagomyidse70. LeporidaeAlmost CosmopoliteEthiopian, OrientalEthiopian, NearcticEthiopianNearcticAll regions but AustralianAbyssinia, NeotropicalEthiopian, OrientalNearcticAll regions but AustralianBIRDS.Passeres.1. Turdidag...2. Sylviidae...3. Timaliida?4. Panuridae5. Cinclidae6. Troglodytidae8. Certhiidae9. Sittidae ...10. Paridae ...13. Pycnonotidae14. Oriolidae...17. Muscicapidae19. Laniidae ...20. Corridae...23. (Nectariniidae)24. (Dicseidse)29. Ampelidae30. Hirundinidae33. Fringillidae35. Sturnidae37. Alaudidae38. Motacillidae47. (Pittidae)CosmopoliteCosmopoliteEthiopian, Oriental, AustralianNearctic, OrientalOrientalAmerican, OrientalOriental, NearcticNearctic, Oriental, Australian, MadagascarNearctic, Oriental, Australian [?]Oriental, EthiopianEthiopian, Oriental, AustralianEastern HemisphereEastern Hemisphere and N. AmericaCosmopoliteEthiopian, Oriental, AustralianEthiopian, Oriental, AustralianNearcticCosmopoliteAll regions but AustralianEastern HemisphereAll regions but NeotropicalCosmopoliteOriental, Australian, EthiopianPlCARIiE.51. Picidae ...52. Yungidae58. Cuculidae62. Coraciidae63. Meropidae67. Alcedinidae69. Upupidae73. Caprimulgidae74. CypselidaeAll regions but AustralianN. W. India, N. E. Africa, S. AfricaAlmost CosmopoliteEthiopian, Oriental, AustralianEthiopian, Oriental, AustralianCosmopoliteEthiopian, OrientalCosmopoliteAlmost Cosmopolite


23G ZOOLOGICAL GEOGRAPHY. [PART III.Sub-regions.Order and Family.o3s 5T3 a33p.Range beyond <strong>the</strong> Region.Columb-s:.84. Columbidae ...CosmopoliteGalling.86. Pteroclidse ...87. Tctraonidae ...88. Phasiauidse ...89. Turnicidse ..Ethiopian, IndianNearctic, Ethiopian, OrientalOriental, Ethiopian, NearcticEthiopian, Oriental, AustralianAccipitres.94. Vulturidae ...96. Falconidae ...97. Pandionidse...98. Strigidae ...All regions but AustralianCosmopoliteCosmopoliteCosmopoliteGralije.99. Rallidce100. Scolopacidse...104. Glareolidffi ...105. Charadriidse...106. Otididee107. Gruidse113. Ardeidoe114. Plataleidse ...115. Ciconiidce ...117. PhsenicopteridseCosmopoliteCosmopoliteEthiopian, Oriental, AustralianCosmopoliteEthiopian, Oriental. AustralianEastern Hemisphere, and N. AmericaCosmopoliteAlmost CosmopoliteNearly CosmopoliteNeotropical, Ethiopian, IndianAnseres.118. Anatidse119. Laridse120. Procellariidte121. Pelecanidae ...123. Colymbidse ...124. Podicipidse ..125. AlcidaeCosmopoliteCosmopoliteCosmopoliteCosmopoliteArctic and N. TemperateCosmopoliteN. Temperate zoneREPT1LIA.Ophidia.1. Typhlopidae...5. Calamariidse...6. Oligodontidse7. Colubridse ...8. Homalopsidae9. Psammophidse18. Erycidse20. Elapidae24. Orotalidae ...25. ViperidaeAll regions but NearcticAll o<strong>the</strong>r regionsOriental and NeotropicalAlmost CosmopoliteOriental, and all o<strong>the</strong>r regionsEthiopian and OrientalOriental and EthiopianAustralian and all o<strong>the</strong>r regionsNearctic, Neotropical, OrientalEthiopian, Oriental


CHAP. X.J


238 ZOOLOGICAL GEOGRAPHY. [PART III.


.CHAP. X.] THE PAL.EARCTIC REGION. 239TABLELIST OF THE GENERA OF TERRESTIAL MAMMALIA AND BIRDSINHABITING THE PAL.EARCTIO REGION.II.Explanation.Names in italics show genera peculiar to <strong>the</strong> region.Names inclosed thus (...)show genera which just enter <strong>the</strong> region, hut are not consideredproperly to helong to it.Genera which undoubtedly helong to <strong>the</strong> region are numbered consecutively.MAMMALIA.Order, Family, audGenus.O o>02Range <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.PRIMATES.SEMNOPITHECIDiE.(Semnopi<strong>the</strong>cusEastern Thibet)Oriental genusCynopi<strong>the</strong>cid^;.1. MacacusCHIROPTERA.Gibraltar, N. Africa, E. Thibetto JapanOrientalPteropid^;.(Pteropus(Xantharpyia ...Egypt, Japan)N. Africa, Palestine)Tropics <strong>of</strong> <strong>the</strong> E. Hemis.Oriental, Austro-MalayanRhINOLOPHIDjE.2. Rhinolphus(Asellia(Rhinopoma .(Nycteris.Temperate & Sou<strong>the</strong>rn partsRegionEgypt) „., .Egypt, Palestine)Egypt)<strong>of</strong>Warmer parts E. HemisphereEthiopian, Java[?] IndiaNubia, HimalayaVESPERTILIONIDiE.3. Vesperugo4. Otonycteris5. Vespertilio(Kerivoula6. Miniopteris ...7. Plecotus8. Barbastellus ...11351112Siberia, AmoorlandEgypt<strong>The</strong> whole regionN. China)S. Europe, N. Africa, JapanS. EuropeMid. and S. Europe, PalestinePI[?]CosmopoliteOriental, S. AfricaS. Afric. Malaya, Austral.HimalayasDarjeeling, TimorNoCTILIONIDiE.9 Molossus(NSECTIVORA.ERINACEID.E.10. ErinaceusS. Europe, N. Africa<strong>The</strong> whole region ;excl. JapanEthiop., Neotrop., AustralianOriental, Africa.


.240 ZOOLOGICAL GEOGRAPHY. [PART 111.Order, Family, andGenus.O


—CHAP. X.] THE PAL^ARCTIC REGION. 241Order,Family, andGenus.'Z, p,03Range <strong>with</strong>in <strong>the</strong>Region.Range beyond <strong>the</strong> RegionOTARIID.E.40. Callorhinus41. Zalophus42. Eunietopias .Tkichechid^:.43. TrickechusPhOCIDvE.44. Callocephalus45. Pagomys ..46. Pagophilus47. Phoca48. Halichserus49. Pelagius ...50. CystophoraSIRENIACETACEAUNGULATA.Equid^e.51. EquusKamschatka andBekring's StraitsJapanJapan, Bekring's StraitsPolar SeasNorth Sea, Caspian, Lake BaikalNortli Sea, JapanNor<strong>the</strong>rn SeasNor<strong>the</strong>rn SeasNorth Sea and BalticMadeira to Black SeaN. AtlanticCent. & and W. Asia & N. AfricaCaliforniaCaliforniaArctic AmericaGreenlandN. PacificN. PacificN. PacificGreenlandN. AtlanticTropics &Behnng'sStrts.OceanicEthiopianSuid^;.52. Sus<strong>The</strong> whole regionOriental, Austro- MalayanCamelid^e.53. Camelus ...Cervid^e.54. Alces55. Tarandus56. Cervus57. Dama58. Elaphodus59. Lophotragus60. Caprcoins61. Moschus ...62. HydropotesDeserts <strong>of</strong> Cent, and W. Asia andN. AfricaNorth Europe and AsiaArctic Europe and Asia<strong>The</strong> whole regionMediterranean districtN. W. ChinaN. ChinaTemp. Europe and W. Asia andN. ChinaAmoor R., N. China, to HimalayasN. CkinaN. AmericaArctic AmericaAll regions but Austral.Bovim:.63. ( Bos64. < Bison65. ( Pocpliagus66. Addax67. Oryx68. ( Gazella ...69. ( Procapra1111112Europe, (not wild)Poland and CaucasusThibetN. Africa to SyriaN. Africa to SyriaN. Africa to Persia, and BeloochistanW. Thibet and MongoliaOrientalNearcticEthiopian desertsS. Africa, India


242 ZOOLOGICAL GEOGRAPHY. [part in.Order, Family, andGenus.


CHAP. X.J THE PALiEARCTIC REGION. 243Order,Family, andGenus.


.244 ZOOLOGICAL GEOGRAPHY. [PART III.Order, Family, andGenus.27. I Ruticilla28. j Larvivora29. Dromolaea30. Saxicola—31. Cercomela32. Pratincola33. Accentor ...Timaliim;.34. Ptcrorhinus(Malacocercus . .(Crateropus —(Trochalopteron(Ianthocincla —Panueid^;.(Paradoxoruis35. Conostoma36. Suthora37. Pannrus38. Hcteromorpha .39. Cholomis


..)CHAP. X.] THE PAL.EARCTIC REGION. 245Order,Family, andGenus.o £"A ftRange <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.PYCNONOTID.fi.49. Microseelis50. Pycnonotus ...JapanPalestine,N. China, JapanOriental genusOriental and EthiopianORIOLID.E.51. OriolusS. Europe, ChinaEthiopian and OrientalMUSCICAFIDfi.52. Muscicapa53. Butalis54. Erythrosterna...(Xanthopygia .(Eumyias— .(Cyanoptila(Siphia55. Tchitrea.."W. and Central Europe"W. Europe to Japan and ChinaCentral Europe to N. China andJapanJapan)E. Thibet)Japan and Arnoor)Moupin, E. Thibet)N. China and JapanEthiopian.E. and S. Africa, MoluccasOriental & MadagascarOriental genusOriental genusOriental genusOriental genusEthiopian and OrientalLANiiD.fi.56. Lanius(Telephouus ..111<strong>The</strong> whole region (excl. AtlanticIslands)N. Africa)Nearctic, Ethiopian,OrientalEthiopian genusCoitviD.£.57. Garrulus58. Perisoreus(Urocissa59. Nucifraga60. Pica61. Cyanopica62. Corvus63. Frcgilus ...NECTARINIID.E.(Arachnecthra2123"W. Europe, N. Africa, to JapanN. Europe and SiberiaCashmere, Japan)W. Europe to Japan, and HimalayasW. Europe to China and JapanSpain, N. E. Asia and Japan<strong>The</strong> whole region\V. Europe to N. China, HimalayasPalestine)Himalayas, FormosaN. AmericaOriental genusHimalayan pine forestsS. China and Formosamigrants [?]Cosmopolite(excl.S. Am.Abyssinian mountainsOriental genusDic.fiiD.fi.(Zosterops1 Amoor and Japan)Ethiop., Orien., Austral.AMPELIDjE.64. Ampelis ...HlRUNDIXID.fi.65. Hirundo66. Cotyle67. ChelidonNor<strong>the</strong>rn half <strong>of</strong> region<strong>The</strong> whole region<strong>The</strong> whole region (excl. Atlan. Is.<strong>The</strong> whole regionNorth AmericaCosmopoliteNearctic, Ethiop., Orien.OrientalFRINGILLID.fi.68. Fringilla<strong>The</strong> whole regionAfrica


246 ZOOLOGICAL GEOGRAPHY. [part hi.Order, Family, andGenus.


CHAP. X.] THE PAL^ARCTIC REGION. 247Order,Family, andGenus.


—248 ZOOLOGICAL GEOGRAPHY. [part hi.Order, F amily, andGenus.Range <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong>Region.COLUMB^E.COLUMBID^!.125. Columba126. Turtur ... .(Alsascomus .GALLING.Pteroclidjo.127. Pterocles128. Syrrhaptes .<strong>The</strong> whole regionW. Europe to JapanE. Thibet)S. Europe, N. Africa, to W. IndiaCentral Asia, N. ChinaAfrica, Asia, AmericaEthiopian and OrientalOriental genusEthiopian genusTetraonid^:.129. Francolinus .130. Perdix ...'131. Coturnix132. Lerwa ...133. Caccabis ...134. Tctraogallus .135. Tetrao ...136. Bonasa ...137. Lagopus...Phasianid^e.138. Crossojitilon .139. Lophophorus140. Tetraophasis141. Ceriornis142. Pucrasia143. PJmsianus144. Thaumalea145. Ithaginis431131032Borders <strong>of</strong> MediterraneanEthiopian, OrientalEurope to MongoliaCentral and S. Europe to Japan Ethiop., Orien., Austral.Snowy Himalayas to E. ThibetCen. Europe and N. Africa to N. Abyssinia, ArabiaW. HimalayasCaucasus to E. Thibet and AltaiMountainsEurope and N. AsiaEurope and N. AsiaIceland, W. Europe to JapanN. AmericaN. AmericaN. America, GreenlandThibet, Mongolia, N. ChinaCashmere to E. Thibet (highestwoods)E. Thibet E. Thibet (?)N. W. Himalayas (high) Himalayas to W. ChinaN. W. Himalayas to N. W. China HimalayasWestern Asia to JapanE. Thibet to Amoor, N. ChinaNepaul to E. Thibet (high)W. Himalayas, FormosaWest ChinaTURNICID.E146. Turnix ...Spain and N. Africa,N. ChinaEthiop., Orien., Austral.ACCIPITRES.VULTURID^!.147. Vultur ...148. Gyps ...149. Otogyps150. NeophronFalconid^e.151. Circus ...152. Astur ...153. Aecipiter154. Buteo ...Spain and N. Africa to N. ChinaS. Europe, Palestine, Cen. AsiaS. Europe, N. AfricaAtlantic Isds. to PalestineEurope to JapanEurope to N. ChinaEurope to JapanEurope to JapanE. Africa, IndiaS. Africa, IndiaAfrica, IndiaAlmost CosmopoliteAlmost CosmopoliteAlmost CosmopoliteCosmopolite (excl. Australia)


CHAP. X.] THE PAL^IARCTIC REGION. 249Order, Family, andGenus.


250 ZOOLOGICAL GEOGRAPHY. [PART III.Order, Family, andGenus.


5cpqa


CHAPTER XI.THE ETHIOPIAN REGION.This is one <strong>of</strong> <strong>the</strong> best defined <strong>of</strong> <strong>the</strong> great zoological regions,consisting <strong>of</strong> tropical and South Africa, to which must be addedtropical Arabia, Madagascar, and a few o<strong>the</strong>r islands, all popularlyknown as African. Some naturalists would extend <strong>the</strong>region northwards to <strong>the</strong> Atlas Mountains and include <strong>the</strong> whole<strong>of</strong> <strong>the</strong> Sahara ; but <strong>the</strong> animal life <strong>of</strong> <strong>the</strong> nor<strong>the</strong>rn part <strong>of</strong> thatgreat desert seems more akin to <strong>the</strong> Palsearctic fauna <strong>of</strong> NorthAfrica. <strong>The</strong> Sahara is really a debatable land which has beenpeopled from both regions;and until we know more <strong>of</strong> <strong>the</strong> naturalhistory <strong>of</strong> <strong>the</strong> great plateaus which rise like islands in <strong>the</strong> waste<strong>of</strong> sand, it will be safer to make <strong>the</strong> provisional boundary line ator near <strong>the</strong> tropic, thus giving <strong>the</strong> nor<strong>the</strong>rn half to <strong>the</strong> Palsearctic,<strong>the</strong> sou<strong>the</strong>rn to <strong>the</strong> Ethiopian region. <strong>The</strong> same line may becontinued across ArabiaWith our present imperfect knowledge <strong>of</strong> <strong>the</strong> interior <strong>of</strong>Africa, only three great continental sub-regions can be well defined.<strong>The</strong> open pasture lands <strong>of</strong> interior tropical Africa arewonderfully uniform in <strong>the</strong>ir productions ; a great number <strong>of</strong>species ranging from Senegal to Abyssinia and <strong>the</strong>nce to <strong>the</strong>Zambesi, while almost all<strong>the</strong> commoner African genera extendover <strong>the</strong> whole <strong>of</strong> this area. Almost all this extensive tract <strong>of</strong>country is a moderately elevated plateau, <strong>with</strong> a hot and dryclimate, and characterised by a grassy vegetation interspersed<strong>with</strong> patches <strong>of</strong> forest. This forms our first or East Africansub-region. <strong>The</strong> whole <strong>of</strong> <strong>the</strong> west coast from <strong>the</strong> south side <strong>of</strong><strong>the</strong> Gambia Paver to about 10° or 12° south latitude, is a very


252 ZOOLOGICAL GEOGRAPHY. [part hi.different kind <strong>of</strong> country ; being almost wholly dense forestswhere not cleared by man, and having <strong>the</strong> hot moist uniformclimate, and perennial luxuriance <strong>of</strong> vegetation, which characterise<strong>the</strong> great equatorial belt <strong>of</strong> forest all round <strong>the</strong> globe.Thisforest country extends to an unknown distance inland, but it wasfound, <strong>with</strong> its features well marked, by Dr. Schweinfurth directlyhe crossed <strong>the</strong> south-western watershed <strong>of</strong> <strong>the</strong> Nile ; and far to<strong>the</strong> south we find itagain unmistakably indicated, in <strong>the</strong> excessivelymoist forest country about <strong>the</strong> head waters <strong>of</strong> <strong>the</strong> Congo,where <strong>the</strong> heroic Livingstone met his death. In this forestdistrict many <strong>of</strong> <strong>the</strong> more remarkable African types are alonefound, and its productions occasionally present us <strong>with</strong> curioussimilarities to those <strong>of</strong> <strong>the</strong> far removed South American orMalayan forests. This is our second or West African subregion.Extra-tropical South Africa possesses features <strong>of</strong> its own, quitedistinct from those <strong>of</strong> both <strong>the</strong> preceding regions (although it hasalso much in common <strong>with</strong> <strong>the</strong> first). Its vegetation is knownto be one <strong>of</strong> <strong>the</strong> richest,most peculiar, and most remarkable on<strong>the</strong> globe ; and in its zoology it has a speciality, similar in kindbut less in degree, which renders it both natural and convenientto separate it as our third, or South African sub-region. Itslimits are not very clearly ascertained, but it is probably boundedby <strong>the</strong> Kalahari desert on <strong>the</strong> north-west, and by <strong>the</strong> LimpopoValley, or <strong>the</strong> mountain range beyond, on <strong>the</strong> north-east, althoughsome <strong>of</strong> its peculiar forms extend to Mozambique. <strong>The</strong>reremains <strong>the</strong> great Island <strong>of</strong> Madagascar, one <strong>of</strong> <strong>the</strong> most isolatedand most interesting on <strong>the</strong> globe, asregards its animal productions;and to this must be added, <strong>the</strong> smaller islands <strong>of</strong> Bourbon,Mauritius and Rodriguez, <strong>the</strong> Seychelles and <strong>the</strong> Comoro Islands,forming toge<strong>the</strong>r <strong>the</strong> Mascarene Islands,—<strong>the</strong> whole constitutingour fourth sub-region.Zoological Characteristics <strong>of</strong> <strong>the</strong>Ethiopian Region.—We havenow to consider briefly, what are <strong>the</strong> peculiarities and characteristics<strong>of</strong> <strong>the</strong> Ethiopian Region asit its distinctive features and broadly separate itprimary zoological regions.a whole,—those which givefrom <strong>the</strong> o<strong>the</strong>r


;chap, xi.] THE ETHIOPIAN REGION. 253Mammalia.—This region has 9 peculiar families <strong>of</strong> mammalia.Chiromyidae (containing <strong>the</strong> aye-aye) ; Potamogalidae and Chrysochloridae(Insectivora) ; Cryptoproctidae and Protelidse (Carnivora); Hippopotamidee and Camelopardalidae (Ungulata) ; andOrycteropodidae (Edentata). Besides <strong>the</strong>se it possesses 7 peculiargenera <strong>of</strong> apes, Troglodytes, Colobus, Myiopitliecus Cercopi<strong>the</strong>cus,Cercocefais, <strong>The</strong>ropitliecus, and Cynocephalus ; 2 subfamilies<strong>of</strong> lemurs containing 6 genera, confined to Madagascar,<strong>with</strong> 3 genera <strong>of</strong> two o<strong>the</strong>r sub-families confined to <strong>the</strong> continent; <strong>of</strong> Insectivora a family, Centetidae, <strong>with</strong> 5 genera,peculiar to Madagascar, and <strong>the</strong> genera Petrodromus and Rhynchocyonbelonging to <strong>the</strong> Macroscelididae, or elephant-shrews,restricted to <strong>the</strong> continent ; numerous peculiar genera or subgenera<strong>of</strong> civets ; Lycaon and Megalotis, remarkable genera <strong>of</strong>Canidae ; Ictonyx, <strong>the</strong> zorilla, a genus allied to <strong>the</strong> weasels ;13 peculiar genera <strong>of</strong> Muridae ; Pectinator, a genus <strong>of</strong> <strong>the</strong> SouthAmerican family Octodontidae ; and 2 genera <strong>of</strong> <strong>the</strong> SouthAmerican Eehimyidse or spiny rats. Of abundant and characteristicgroups it possesses Macroscelides, Felis, Hyaena, Hyrax,Rhinoceros, and Elephas, as well as several species <strong>of</strong> zebra anda great variety <strong>of</strong> antelopes.<strong>The</strong> great speciality indicated by <strong>the</strong>se numerous peculiarfamilies and genera, is still far<strong>the</strong>r increased by <strong>the</strong> absence<strong>of</strong> certain groups dominant in <strong>the</strong> Old-World continent,an absence which we can only account for by <strong>the</strong> persistence,through long epochs, <strong>of</strong> barriers isolating <strong>the</strong> greater part <strong>of</strong> Africafrom <strong>the</strong> rest <strong>of</strong> <strong>the</strong> world. <strong>The</strong>se groups are, Ursidas, <strong>the</strong> bears ;Talpidae <strong>the</strong> moles ; Camelidae, <strong>the</strong> camels ; Cervidae, <strong>the</strong> deerCaprinae, <strong>the</strong> goats and sheep ; and <strong>the</strong> genera Bos (wild ox) ; andSus (wild boar). Combining <strong>the</strong>se striking deficiencies, <strong>with</strong><strong>the</strong> no less striking peculiarities above enumerated, it seemshardly possible to have a region more sharply divided from<strong>the</strong> rest <strong>of</strong> <strong>the</strong> globe than this is, by its whole assemblage <strong>of</strong>mammalia.Birds.—In birds <strong>the</strong> Ethiopian region is by no means sostrikingly peculiar, many <strong>of</strong> <strong>the</strong>se having been able to pass <strong>the</strong>ancient barriers which so long limited <strong>the</strong> range <strong>of</strong> mammalia.


—;254 ZOOLOGICAL GEOGRAPHY. [part in.It is, however, sufficiently rich, possessing 54 families <strong>of</strong> landbirds, besides a few genera whose position is not well ascertained,and which may' constitute distinct families. Of <strong>the</strong>se 6 arepeculiar, Musophagidae (<strong>the</strong> plantain eaters); Coliidse (<strong>the</strong> colies) ;Leptosomidse, allied to <strong>the</strong> cuckoos ;Irrisoridee, allied to <strong>the</strong>hoopoes ; and Serpentaridse, allied to <strong>the</strong> hawks. Only onePasserine family is peculiar—Paictidee, while most <strong>of</strong> <strong>the</strong> o<strong>the</strong>rtropical regions possess several ; but Euryceros and Buphaga,here classed <strong>with</strong> <strong>the</strong> Sturnidse, ought, perhaps, to form twomore.It has, however, many peculiar genera, especially among <strong>the</strong>fruit-thrushes, Pycnonotidse ; flycatchers, Muscicapidae ; shrikes,Lanidse ; crows, Corvidse ;starlings, Sturnidse ;and weaver-birds,Ploceidae ; <strong>the</strong> latter family being very characteristic <strong>of</strong> <strong>the</strong> region.It is also rich in barbets, Megalaemidae (7 peculiar genera)cuckoos, Cuculidas ; rollers, Coraciidse ; bee-eaters, Meropidsehornbills, Bucerotidre ; and goat-suckers, Caprimulgidae. It ispoor in parrots and ra<strong>the</strong>r so in pigeons ; but it abounds inPterocles and Francolinus, genera <strong>of</strong> Gallinae, and possesses 4genera <strong>of</strong> <strong>the</strong> peculiar group <strong>of</strong> <strong>the</strong> guinea-fowls, forming part <strong>of</strong><strong>the</strong> pheasant family. It abounds in vultures, eagles, and o<strong>the</strong>rbirds <strong>of</strong> prey, among which is <strong>the</strong> anomalous genus Seiyentarius,<strong>the</strong> secretary-bird, constituting a distinct family. Many <strong>of</strong> <strong>the</strong>most remarkable forms are confined to Madagascar and <strong>the</strong>adjacent islands, and will be noticed in our account <strong>of</strong> that subregion.Reptiles.—Of <strong>the</strong> reptiles <strong>the</strong>re are 4 peculiar Ethiopianfamilies ;— 3 <strong>of</strong> snakes, Rachiodontidse, Dendraspidae, and Atractaspidseand 1 <strong>of</strong> lizards, Chamsesauridse.Psammophidse (desert snakes) are abuudant, as are Lycodontidse(fanged ground-snakes), and Viperidae (vipers). <strong>The</strong> followinggenera <strong>of</strong> snakes are peculiar or highly characteristic :Lcptorhynclms,Bhamnophis, Herpetetliiops and Grayia (Colubridae)Hopsidrophis and Bucephalus (Dendrophidae) ; Langalia (Dryophidie); Pythonodipsas (Dipsadidse) ; Boedon, Lycophidion, Holuropholis,Simocephahis and Lamprophis (Lycodontiche) ;Hortidiaand Sanzinia (Pythonidae) ; Cyrptopliis, Elapsoidea and Pcecilophis(Elapidae) ; and A<strong>the</strong>ris (Viperidse). <strong>The</strong> following genera


—chap, xl] THE ETHIOPIAN REGION. 255<strong>of</strong> lizards are <strong>the</strong> most characteristic:Monotrophis(Lepidosternidse);Cordylus, Pseudocordylus, Platysaurus, Cordylosaurus,Pleurostichus, Saurophis and Zonurus (Zormridse) ; Sphamops,Scelotes, Sphcenocephalus and Sepsina (Sepidaj) ; Pachydactylu's(Geckotidas) ;Agama (Agamidae); and Chameleon (Chaineleonidae).Of tortoises, Cynyxis, Pyxis and Chersina (Testudinidse), andCycloderma (Trionychidae) are <strong>the</strong> most characteristic.Amphibia.—Of <strong>the</strong> 9 families <strong>of</strong> amphibia <strong>the</strong>re is only1 peculiar, <strong>the</strong> Dactylethridae, a group <strong>of</strong> toads; but <strong>the</strong>Alytidae, a family <strong>of</strong> frogs, are abundant.Fresh-water Fish.—Oi <strong>the</strong> 14 families <strong>of</strong> fr-^-water fishes3 are peculiar: Mormyridae and Gynmarchidse, small groupsnot far removed from <strong>the</strong> pikes ; and Polypteridaa, a small group<strong>of</strong> ganoid fishes allied to <strong>the</strong> gar-pikes (Lepidosteidte) <strong>of</strong> NorthAmerica.Summary <strong>of</strong> Ethiopian Vertebrates.—Combining <strong>the</strong> resultshere indicated and set forth in greater detail in <strong>the</strong> tables <strong>of</strong><strong>distribution</strong>, we find that <strong>the</strong> Ethiopian region possesses examples<strong>of</strong> 44 families<strong>of</strong> mammalia, 72 <strong>of</strong> birds, 35 <strong>of</strong> reptiles,9 <strong>of</strong> amphibia, and 15 <strong>of</strong> fresh-water fishes. It has 23 (orperhaps 25) families <strong>of</strong> Vertebrata altoge<strong>the</strong>r peculiar to it out<strong>of</strong> a total <strong>of</strong> 175 families, or almost exactly one-eighth <strong>of</strong> <strong>the</strong>whole. Out <strong>of</strong> 142 genera <strong>of</strong> mammalia found <strong>with</strong>in <strong>the</strong>region, 90 are peculiar to it ; a proportion not much short <strong>of</strong>two-thirds. Of land birds <strong>the</strong>re are 294 genera, <strong>of</strong> which179 are peculiar; giving a proportion <strong>of</strong> a little less thanthree-fifths.Compared <strong>with</strong> <strong>the</strong> Oriental region this shows a considerablylarger amount <strong>of</strong> speciality under all <strong>the</strong> heads;but <strong>the</strong> superiority is mainly due to <strong>the</strong> wonderful and isolatedfauna <strong>of</strong> Madagascar, to which <strong>the</strong> Oriental region hasnothing comparable. Without this <strong>the</strong> regions would be nearlyequal.Insects: Lepidoptera,— 11 out <strong>of</strong> <strong>the</strong> 16 families <strong>of</strong> butterflieshave representatives in Africa, but none are peculiar.Acrseidae is one <strong>of</strong> <strong>the</strong> most characteristic families, and <strong>the</strong>re


;256 ZOOLOGICAL GEOGRAPHY. [part hi.are many interesting forms <strong>of</strong> Nymphalida?, Lyceenidse, andPapilionidse. <strong>The</strong> peculiar or characteristic forms are Amauris(Danaidse); Gnophodes, Leptoneura, Bicyclus, Heteropsis andCcenyra (Satyridse) ; Acrcea (Acraeidae) ; LacJmoptcra, Precis,Salamis, Crenis, Godartia, Amphidcma, Pscudacrcea, Catuna,Euryphcnc, Romalceosoma, Hamanumida, Aterica, Harma,Mencris, Charaxes, and Philognoma (Nymphalidae) ; Pentila,Liptena, Durbania, Zcritis, Capys, Phytala, Epitola, Heivitsoniaand Deloneura (Lycamidffi) ;Pscudopontia, Idmais, Tcracolus,Callosune (Pieridse) ; Abantis, Ceratrichia and Caprona (Hesperidse).<strong>The</strong> total number <strong>of</strong> species known is about 750which is very poor for an extensive tropical region, but thisis not to be wondered at when <strong>the</strong> nature <strong>of</strong> much <strong>of</strong> <strong>the</strong>country is considered. It is also, no doubt, partly due to ourcomparative ignorance <strong>of</strong> <strong>the</strong> great equatorial forest district,which is <strong>the</strong> only part likely to be very productive in thisorder <strong>of</strong> insects.Coleoptcra.— In our first representative family, Cicindelidseor tiger-beetles, <strong>the</strong> Ethiopian region is ra<strong>the</strong>r rich, having 13genera, 11 <strong>of</strong> which are peculiar to it; and among <strong>the</strong>se aresuch remarkable forms as Manticora, Myrmecoptcra and Dromica<strong>with</strong> Mcgacephala, a genus only found elsewhere in Australiaand South America.In Carabidae or carnivorous ground beetles, <strong>the</strong>re areabout 75 peculiar genera. Among <strong>the</strong> most characteristicare Anthia, Polyrhina, Graphipierus and Piczia, which arealmost all peculiar ; while Orthogonius, Hexagonia, Macrochilus,Thyreopterus, Eudcma, and Abacctus are common to this and<strong>the</strong> Oriental region ;and Hypolithus to <strong>the</strong> Neotropical.Out <strong>of</strong> 27 genera <strong>of</strong> Buprestidse, or metallic beetles, only 6 arepeculiar to <strong>the</strong> region, one <strong>of</strong> <strong>the</strong> most remarkable being Polybothrus,confined to Madagascar.Sternocera and Chrysochroa arecharacteristic <strong>of</strong> this region and <strong>the</strong> Oriental ; it has Julodis incommon <strong>with</strong> <strong>the</strong> Mediterranean sub-region, and Bclionota<strong>with</strong> <strong>the</strong> Malayan.<strong>The</strong> region is not rich in Lucanidae, or stag-beetles, possessingonly 10 genera, 7 <strong>of</strong> which are peculiar, but most <strong>of</strong> <strong>the</strong>m con-


chap, xi.] THE ETHIOPIAN REGION. 257sist <strong>of</strong> single species. <strong>The</strong> o<strong>the</strong>r three genera, Cladognathus,Nigidius, and Figulus, are <strong>the</strong> most characteristic, though a]lhave a tolerably wide range in <strong>the</strong> Old World.In <strong>the</strong> elegant Cetoniida?, or rose-chafers, this region standspreeminent, possessing 76 genera, 64 <strong>of</strong> which are peculiar toit. <strong>The</strong> o<strong>the</strong>rs are chiefly Oriental, except Oxythrcea which isEuropean, and Stethodesma which is Neotropical. Preeminentin size and beauty is Goliathus, comprising perhaps <strong>the</strong> mostbulky <strong>of</strong> all highly-coloured beetles. O<strong>the</strong>r large and characteristicgenera are Ceratorhina, Ischnostoma, Anochilia,Diplognatha, Agenius, and many o<strong>the</strong>rs <strong>of</strong> less extent.In <strong>the</strong> enormous tribe <strong>of</strong> Longicorns, or long-horned beetles,<strong>the</strong> Ethiopian is not so rich as <strong>the</strong> o<strong>the</strong>r three tropical regions ;but this may be, in great part, owing to its more productivedistricts having never been explored by any competent entomologists.It never<strong>the</strong>less possesses 262 genera, 216 <strong>of</strong> which arepeculiar, <strong>the</strong> o<strong>the</strong>rs being mostly groups <strong>of</strong> very wide range.Out <strong>of</strong> such a large number it is difficult to select a few as mostcharacteristic, but some <strong>of</strong> <strong>the</strong> peculiarities <strong>of</strong> <strong>distribution</strong> asregards o<strong>the</strong>r regions may be named.Among Prionidae, Tithocsis a characteristic Ethiopian genus. A few species <strong>of</strong> <strong>the</strong>American genera Parandra and Mallodon occur here, while <strong>the</strong>North Temperate genus Prionus is only found in Madagascar.Among Cerambycidae, Promeces is <strong>the</strong> most characteristic. <strong>The</strong>American genera Oeme and Cyrtomerus occur ; while Homalachnusand Philaga<strong>the</strong>s are Malayan, and Leptocera occurs only inMadagascar, Ceylon, Austro-Malaya, and Australia.<strong>The</strong> Lamiidaeare very fine; Sternotomis, Tragocephala. Ceroplesis, Phryneta,Volumnia, and Nitocris, being very abundant and characteristic.Most <strong>of</strong> <strong>the</strong> non-peculiar genera <strong>of</strong> this family are Oriental,but Spalacopsis and Acanthoderes are American, while Tetruglenesand Schcenionta have been found only in East and South Africaand in Malaya.Terrestrial Mollusca—In <strong>the</strong> extensive family <strong>of</strong> <strong>the</strong> Helicidaeor snails, 13 genera are represented, only one <strong>of</strong> which,Columna, is peculiar. This region is however <strong>the</strong> metropolis <strong>of</strong>Achatina, some <strong>of</strong> <strong>the</strong> species being <strong>the</strong> largest land-shells


258 ZOOLOGICAL GEOGRAPHY. [part hi.known. Buliminus, Stenogyra, and Pupa are characteristicgenera. Bulimus is absent, though one species inhabits St.Helena. <strong>The</strong> operculated shells are not very well represented,<strong>the</strong> great family <strong>of</strong> Cyclostomidse having here only nine genera,With but one peculiar, Zithidion, found inMadagascar, Socotra,and Arabia. None <strong>of</strong> <strong>the</strong> genera appear to be well representedthroughout <strong>the</strong> region, and <strong>the</strong>y are almost or quite absent fromWest Africa.According to Woodward's Manual (1868) West Africa hasabout 200 species <strong>of</strong> land-shells, South Africa about 100,Madagascar nearly 100, Mauritius about 50. All <strong>the</strong> islandshave <strong>the</strong>ir peculiar species; and are, in proportion to <strong>the</strong>irextent, much richer than <strong>the</strong> continent ;as is usually <strong>the</strong> case.<strong>The</strong> Ethiopian Sub-eegions.It has been already explained that <strong>the</strong>se are to some extentprovisional ;yet it is believed that <strong>the</strong>y represent generally <strong>the</strong>primary natural divisions <strong>of</strong> <strong>the</strong> region, however <strong>the</strong>y may besubdivided when our knowledge <strong>of</strong> <strong>the</strong>ir productions becomesmore accurate.I. <strong>The</strong> East African Sub-region, or Central and East Africa.This division includes all <strong>the</strong> open country <strong>of</strong> tropical Africasouth <strong>of</strong><strong>the</strong> Sahara, as well as an undefined sou<strong>the</strong>rn margin <strong>of</strong>that great desert. With <strong>the</strong> exception <strong>of</strong> a narrow strip along<strong>the</strong> east coast and <strong>the</strong> valleys <strong>of</strong> <strong>the</strong> Niger and Nile, it is a vastelevated plateau from 1,000 to 4,000 feet high, hilly ra<strong>the</strong>rthanmountainous, except <strong>the</strong> l<strong>of</strong>ty table land <strong>of</strong> Abyssinia, <strong>with</strong>mountains rising to 16,000 feet and extending south to <strong>the</strong>equator, where it terminates in <strong>the</strong> peaks <strong>of</strong> Kenia and Kilimandjaro,18,000 and 20,000 feet high. <strong>The</strong> nor<strong>the</strong>rn portion<strong>of</strong> this sub-region is a belt about 300 miles wide between <strong>the</strong>Sahara on <strong>the</strong> north and <strong>the</strong> great equatorial forest on <strong>the</strong> south,extending from Cape Verd, <strong>the</strong> extreme western point <strong>of</strong> Africa,across <strong>the</strong> nor<strong>the</strong>rn bend <strong>of</strong> <strong>the</strong> Niger and Lake Tchad to <strong>the</strong>mountains <strong>of</strong> Abyssinia. <strong>The</strong> greater part <strong>of</strong> this tract has a


chap, xi.] THE ETHIOPIAN REGION. 259moderate elevation.<strong>The</strong> eastern portion reaches from about <strong>the</strong>second cataract <strong>of</strong> <strong>the</strong> Nile, or perhaps from about <strong>the</strong> parallel<strong>of</strong> 20° N. Latitude, down to about 20° S.Latitude, and from <strong>the</strong>east coast to where <strong>the</strong> great forest region commences, or to LakeTanganyika and about <strong>the</strong> meridian <strong>of</strong> 28° to 30° E. Longitude.<strong>The</strong> greater part <strong>of</strong> this tract is a l<strong>of</strong>ty plateau.<strong>The</strong> surface <strong>of</strong> all this sub-region is generally open, covered<strong>with</strong> a vegetation <strong>of</strong> high grasses or thorny shrubs, <strong>with</strong> scatteredtrees and isolated patches <strong>of</strong> forest in favourable situations.<strong>The</strong> only parts where extensive continuous forests occur, are on<strong>the</strong> eastern and western slopes<strong>of</strong> <strong>the</strong> great Abyssiuian plateau,and on <strong>the</strong> Mozambique coast from Zanzibar to S<strong>of</strong>ala. <strong>The</strong>whole <strong>of</strong> this great district has one general zoological character.Many species range from Senegal to Abyssinia, o<strong>the</strong>rs fromAbyssinia to <strong>the</strong> Zambesi, and a few, asMungos fasciatus andPhacochcerus cethiopicus, range over <strong>the</strong> entire sub-region.Fennecus,Ictonyx, and several genera <strong>of</strong> antelopes, characterise everypart <strong>of</strong> it, as do many genera <strong>of</strong> birds. Coracias ncevia, Corytlwrniscyanostigma, Tockus nasutus, T. erythrorhynchus, Parusleucopterus, Buphaga africana, Vidua paradisea, are examples<strong>of</strong> species, which are found in <strong>the</strong> Gambia, Abyssinia and SouthEast Africa, but not in <strong>the</strong> West African sub-region ; and consideringhow very little is known <strong>of</strong> <strong>the</strong> natural history<strong>of</strong> <strong>the</strong> country immediately south <strong>of</strong> <strong>the</strong> Sahara, it maywell be supposed that <strong>the</strong>se are only a small portion <strong>of</strong> <strong>the</strong>species really common to <strong>the</strong> whole area inprove itsfundamental unity.question, and whichAlthough this sub-region is so extensive and so generallyuniform in physical features, it is by far <strong>the</strong> least peculiar part<strong>of</strong> Africa.typesIt possesses, <strong>of</strong> course, all those wide-spread Ethiopianwhich inhabit every part <strong>of</strong> <strong>the</strong> region, but it has hardlyany special features <strong>of</strong> its own. <strong>The</strong> few genera which arepeculiar to it have generally a limited range, and for <strong>the</strong> mostpart belong, ei<strong>the</strong>r to <strong>the</strong> isolated mountain-plateau <strong>of</strong> Abyssiniawhich is almost as much Palaearctic as Ethiopian, or to <strong>the</strong> woodydistricts <strong>of</strong> Mozambique where <strong>the</strong> fauna has more <strong>of</strong> a Westor South African character.


—;260 ZOOLOGICAL GEOGRAPHY. [part in.Mammalia.—<strong>The</strong> only forms <strong>of</strong> Mammalia peculiar to thissub-region are Tlieropithccus, one <strong>of</strong> <strong>the</strong> Cynopi<strong>the</strong>cidae confinedto Abyssinia ; Petrodromus and Rhynchocyon, belonging to <strong>the</strong>insectivorous Macroscelididse, have only been found in Mozambique; <strong>the</strong> Antelopine genus Neotragus, from Abyssinia southward;Saccostomus and Pclomys genera <strong>of</strong> Muridas inhabitingMozambique ; Rcteroccphalus from Abyssinia, and Heliophobiusfrom Mozambique, belonging to <strong>the</strong> Spalacidre ; and Pectinatorfrom Abyssinia, belonging to <strong>the</strong> Octodontidse. Cynocephalus,Rhinoceros, Camclopardalis, and antelopes <strong>of</strong> <strong>the</strong> genera Oryx,Cervicapra, Kobus, Nanotragus, Cephalophus,Hippotragus, Alcephalus,and Catoblepas, are characteristic; as well as Felis,Hyama, and numerous civets and ichneumons.Birds.—Peculiar forms <strong>of</strong> birds are hardly to be found herewe only meet <strong>with</strong> two Hypocolius, a genus <strong>of</strong> shrikes in Abyssinia;and Balmniccps, <strong>the</strong> great boat-billed heron <strong>of</strong> <strong>the</strong> UpperNile.Yet throughout <strong>the</strong> country birds are abundant, and most<strong>of</strong> <strong>the</strong> typical Ethiopian forms are well represented.Reptiles.—Of reptiles, <strong>the</strong> only peculiar forms recorded areXenocalamus, a genus <strong>of</strong> snakes, belonging to<strong>the</strong> Calamariid*and Pythonodipsas, one <strong>of</strong> <strong>the</strong> Dipsadidae, both from <strong>the</strong> Zambesiand among lizards, Pisturus, one <strong>of</strong> <strong>the</strong> Geckotidse,Amphibia and Fishes.—-<strong>The</strong>re are no peculiar forms <strong>of</strong> amphibiaor <strong>of</strong> fresh-water fishes.Insects.—Insects are almost equallyforms.unproductive <strong>of</strong> peculiarAmong butterflies we have Abantis, one <strong>of</strong> <strong>the</strong> Hesperidae,from Mozambique ; and in Coleoptera, 2 genera <strong>of</strong> Cicindelidre,8 <strong>of</strong> Carabidaa, 1 or 2 <strong>of</strong> Cetoniidre, and about half-a-dozen <strong>of</strong>Longicorns :a mere nothing, as we shall see, compared <strong>with</strong> <strong>the</strong>hosts <strong>of</strong> peculiar genera that characterise each <strong>of</strong> <strong>the</strong> o<strong>the</strong>rfrom Abyssinia.subregions.Nei<strong>the</strong>r do land-shells appear to present any peculiarforms.<strong>The</strong> fact that so very few special types characterise <strong>the</strong> extensivearea now under consideration is very noteworthy.It justifiesus in uniting this large and widespread tract <strong>of</strong> country asforming essentially but one sub-division <strong>of</strong> <strong>the</strong> great Ethiopian


PLATE IV.CHARACTERISTIC ANIMALS OF EAST AFRICA.


chap, xi.] THE ETHIOPIAN REGION. 261region, and it suggests some curious speculations as to <strong>the</strong> formerhistory <strong>of</strong> that region, a subject which must be deferred to <strong>the</strong>latter part <strong>of</strong> this chapter. In none <strong>of</strong> <strong>the</strong> o<strong>the</strong>r great tropicalregions does it occur, that <strong>the</strong> largest portion <strong>of</strong> <strong>the</strong>ir area,although swarming <strong>with</strong> life, yet possesses hardly any distinctivefeatures except <strong>the</strong> absence <strong>of</strong> numerous types characteristic <strong>of</strong><strong>the</strong> o<strong>the</strong>r sub-regions.Plate IV.— Illustrating <strong>the</strong> Zoology <strong>of</strong> East Africa.—Althoughthis sub-region has so little speciality, it is that which aboundsmost in large <strong>animals</strong>, and is, perhaps, <strong>the</strong> best representative <strong>of</strong>Africa as regards zoology. Some <strong>of</strong> <strong>the</strong> most distinctive <strong>of</strong> African<strong>animals</strong> range over <strong>the</strong> whole <strong>of</strong> it, and as, from recent explorations,many parts <strong>of</strong> this wide area have been made known to <strong>the</strong>reading public, we devote one <strong>of</strong> our plates to illustrate <strong>the</strong>especially African forms <strong>of</strong> life that here abound. <strong>The</strong> antelopesrepresented are <strong>the</strong> koodoo (Tragclaphus strepsiceros) one <strong>of</strong> <strong>the</strong>handsomest <strong>of</strong> <strong>the</strong> family, which ranges over all <strong>the</strong> highlands<strong>of</strong> Africa from Abyssinia to <strong>the</strong> sou<strong>the</strong>rn districts. To <strong>the</strong> leftis <strong>the</strong> aardvark, or earth pig, <strong>of</strong> North Eastern Africa {Orycteropuscethiopicus) which, to <strong>the</strong> north <strong>of</strong> <strong>the</strong> equator in East Africa,represents <strong>the</strong> allied species <strong>of</strong> <strong>the</strong> Cape <strong>of</strong> Good Hope, <strong>The</strong>seEdentata are probably remnants <strong>of</strong> <strong>the</strong> ancient fauna <strong>of</strong> Africa,when it was completely isolated from <strong>the</strong> nor<strong>the</strong>rn continentsand few <strong>of</strong> <strong>the</strong> higher types had been introduced. <strong>The</strong> largebird in <strong>the</strong> foreground is <strong>the</strong> secretary-bird, or serpent-killer(Serpentarius rcptilivorus), which has affinities both for <strong>the</strong> birds<strong>of</strong>-preyand <strong>the</strong> waders. It is common over almost all <strong>the</strong> opencountry <strong>of</strong> Africa, destroying and feeding on <strong>the</strong> most venomousserpents. <strong>The</strong> bird on <strong>the</strong> wing is <strong>the</strong> red-billed promerops(Irrisor erythrorhynchus), a handsome bird <strong>with</strong> glossy plumageand coral-red bill. It is allied to <strong>the</strong> hoopoes, and feeds oninsects which it hunts for among <strong>the</strong> branches <strong>of</strong> trees. Thisspecies also ranges over a large part <strong>of</strong> east and central Africa tonear <strong>the</strong> Cape <strong>of</strong> Good Hope. O<strong>the</strong>r species are found in <strong>the</strong>west ; and <strong>the</strong> genus, which forms a distinct family, Irrisoridm, isone <strong>of</strong> <strong>the</strong> best marked Ethiopian types <strong>of</strong> birds. In <strong>the</strong> distanceisa rhinoceros, now one <strong>of</strong> <strong>the</strong> characteristic features <strong>of</strong> African


262 ZOOLOGICAL GEOGRAPHY. [part hi.zoology, though <strong>the</strong>re is reason to believe that it is a comparativelyrecent intruder into <strong>the</strong> country.II.<strong>The</strong> West-African Sub-region.This may be defined as <strong>the</strong> equatorial-forest sub-region, since itcomprises all that portion <strong>of</strong> Africa, from <strong>the</strong> west coast inland,over which <strong>the</strong> great equatorial forests prevail more or less uninterruptedly.<strong>The</strong>se commence to <strong>the</strong> south <strong>of</strong> <strong>the</strong> Gambia Eiver,and extend eastwards in a line roughly parallel to <strong>the</strong> sou<strong>the</strong>rnmargin <strong>of</strong> <strong>the</strong> great desert, as far as <strong>the</strong> sources <strong>of</strong> <strong>the</strong> upperNile and <strong>the</strong> mountains forming <strong>the</strong> western boundary <strong>of</strong> <strong>the</strong>basin <strong>of</strong> <strong>the</strong> great lakes ;and southward to that high but marshyforest-country in which Livingstone was travelling at <strong>the</strong> time<strong>of</strong> his death. Its sou<strong>the</strong>rn limits are undetermined, but are probablysomewhere about <strong>the</strong> parallel <strong>of</strong> 11° S. Latitude.1This extensive and luxuriant district has only been exploredzoologically in <strong>the</strong> neighbourhood <strong>of</strong> <strong>the</strong> West coast. Much, nodoubt, remains to be done in <strong>the</strong> interior, yet its main featuresare sufficiently well known, and most <strong>of</strong> itscharacteristic types<strong>of</strong> animal life have, no doubt, been discovered,Mammalia.—Several very important groups <strong>of</strong> mammals arepeculiar to this sub-region. Most prominent are <strong>the</strong> greatanthropoid apes—<strong>the</strong> gorilla and <strong>the</strong> chimpanzee—forming <strong>the</strong>genus Troglodytes ; and monkeys <strong>of</strong> <strong>the</strong> genera Myiopi<strong>the</strong>cusand Cercocebus. Two remarkable forms <strong>of</strong> lemurs, Perodicticusand Arctocebus, are also peculiar to West Africa. Among <strong>the</strong>Insectivora is Potamogale, a semi-aquatic animal, forming adistinct family ; and three peculiar genera <strong>of</strong> civets ( Viverridaa)have been described. Hyomoschus, a small, deer-like animal,belongs to <strong>the</strong> Tragulidse, or chevrotains, a family o<strong>the</strong>rwise1Dr. Schweinfurth has accurately determined <strong>the</strong> limits <strong>of</strong> <strong>the</strong> suh-regionat <strong>the</strong> point where he crossed <strong>the</strong> watershed between <strong>the</strong> Nile tributaries andthose <strong>of</strong> <strong>the</strong> Shari, in 4^° N. Lat. and 28£° E. Long. He describes a suddenchange in <strong>the</strong> character <strong>of</strong> <strong>the</strong> vegetation, which to <strong>the</strong> southward <strong>of</strong> thispoint assumes a West-African character. Here also <strong>the</strong> chimpanzee andgrey parrot first appear, and certain species <strong>of</strong> plants only known elsewherein Western Africa.


;CHAP. XI.] THE ETHIOPIAN REGION. 2G3confined to <strong>the</strong> Oriental region ; and in <strong>the</strong> squirrel family is acurious genus, Anomalurus, which resembles <strong>the</strong> flying squirrels<strong>of</strong> o<strong>the</strong>r parts <strong>of</strong> <strong>the</strong> world, <strong>with</strong>out be.ing directly allied to <strong>the</strong>m.Birds.—In this class we find a larger proportionate number<strong>of</strong> peculiar forms. Hypergerus and Ale<strong>the</strong>, belonging to <strong>the</strong>Timaliidse, or babblers, are perhaps allied to Malayan groupsParinia, a peculiar form <strong>of</strong> tit, is found only in Prince's IslandIxonotus is an abundant and characteristic form <strong>of</strong> Pycnonotidse;Frascria, Hypodes, Cuphopterus, and Chaunonutus, are peculiargenera <strong>of</strong> shrikes ;Picathartes is one <strong>of</strong> <strong>the</strong> many strange forms<strong>of</strong> <strong>the</strong> crow family ; Cinnyricinclus is a peculiar genus <strong>of</strong> sunbirds;Pholidornis is supposed to belong to <strong>the</strong> Oriental Dicseidse,or flower-peckers ; Waldenia is a recently-described new form<strong>of</strong> swallow ; Ligumus, a finch, Spermospiga, a weaver bird, andOnychognathus a starling, are also peculiar West African genera.Coming to <strong>the</strong> Picarire we have Verreauxia, a peculiar woodpecker; three peculiar genera <strong>of</strong> barbets (Megala?midse) ;<strong>the</strong> typical plantain-eaters (Musophaga) ; Myioceyx, a peculiargenus <strong>of</strong> kingfishers ; while Berenicornis is a genus <strong>of</strong> crestedhornbills, only found elsewhere in Malaya. <strong>The</strong> grey parrots,<strong>of</strong> <strong>the</strong> genus Psittacus, are confined to this sub-region, as aretwo peculiar genera <strong>of</strong> partridges, and three <strong>of</strong> guineafowl.We have also here a species <strong>of</strong> Pitta, one <strong>of</strong> <strong>the</strong> Orientalfamily <strong>of</strong> ground-thrushes ; and <strong>the</strong> Oriental paroquets,Palceornis, are found here as well as in Abyssinia and <strong>the</strong>Mascarene Islands.We thus find, both in <strong>the</strong> Mammalia and birds <strong>of</strong> West Africa,a special Oriental oreven Malayan element not present in <strong>the</strong>o<strong>the</strong>r parts <strong>of</strong> tropical Africa, although appearing again inMadagascar.In <strong>the</strong> Mammalia it is represented by <strong>the</strong> anthropoidapes ; by Colobus allied to Semncpi<strong>the</strong>cus, and by Cercocelmsallied to Macacus ; and especially by a form <strong>of</strong> <strong>the</strong> Malayanfamily <strong>of</strong> chevrotains (Tragulidae).<strong>The</strong> Malayan genus <strong>of</strong> otters,Aonyx, is also said to occur in West and South Africa. Inbirds we have special Oriental and Malayan affinities inAle<strong>the</strong>,Pholidornis, Berenicornis, Pitta, and Palmornis ; while <strong>the</strong>Oriental genus Treron has a wide range in Africa. We shall


;264 ZOOLOGICAL GEOGRAPHY. [part hi.endeavour to ascertain <strong>the</strong> meaning <strong>of</strong> this special relation ata subsequent stage <strong>of</strong> our inquiries.Plate V.— River Scene in West Africa, <strong>with</strong> CharacteristicAnimals.—Our artist has here well represented <strong>the</strong> luxurianceand beauty <strong>of</strong> a tropical forest; and <strong>the</strong> whole scene issuch as might be witnessed on <strong>the</strong> banks <strong>of</strong> one <strong>of</strong> <strong>the</strong>rivers <strong>of</strong> equatorial West Africa. On <strong>the</strong> right we see ared river-hog (Potamochcerus penicillatus), one <strong>of</strong> <strong>the</strong> handsomest<strong>of</strong> <strong>the</strong> swine family, and highly characteristic <strong>of</strong> <strong>the</strong>West African sub-region. In a tree overhead is <strong>the</strong> potto{Perodicticus potto), one <strong>of</strong> <strong>the</strong> curious forms <strong>of</strong> lemur confinedto West Africa. On <strong>the</strong> left is <strong>the</strong> remarkable Potamogalevelox, firstdiscovered by Du Chaillu,—an Insectivorousanimal, <strong>with</strong> <strong>the</strong> form and habits <strong>of</strong> an otter. On <strong>the</strong> o<strong>the</strong>rside <strong>of</strong> <strong>the</strong> river are seen a pair <strong>of</strong> gorillas {Troglodytes gorilla),<strong>the</strong> largest <strong>of</strong> <strong>the</strong> anthropoid apes.<strong>The</strong> bird on <strong>the</strong> wing is<strong>the</strong> Whyclah finch (Vidua paradisea),remarkable for <strong>the</strong> enormous plumes <strong>with</strong> which <strong>the</strong> tail<strong>of</strong> <strong>the</strong>male bird is decorated during <strong>the</strong> breeding season. <strong>The</strong> crestedbird overhead is one <strong>of</strong> <strong>the</strong> beautiful green touracos (TuracusmacrorhyncJius), belonging to <strong>the</strong> Musophagida?, or plantain-eaters,a family wholly African, and most abundant in <strong>the</strong> westernsub-region.Reptiles.—In this class we find a large number <strong>of</strong> peculiarforms ;13 genera <strong>of</strong> snakes, 3 <strong>of</strong> lizards, and 2 <strong>of</strong> tortoises beingconfined to <strong>the</strong> sub-region. <strong>The</strong> snakes are Pariaspis, Elapops,and Prosymna (Calamariidas), Rhamnophis, Herpetcthiops, andGrayia (Colubrida?,), Neusterophis and Limnophis (Homalopsidse),Simocephalus and Holurophis (Lycodontidae) ; Pclophilus (Pythonidie); Plapsoidea (Elapidae) ;and A<strong>the</strong>ris (Viperidae). <strong>The</strong>lizards are Balophia (Lepidosternidae) ; Otosaurus (Scincidse)Psilodactylus (Geckotidas). <strong>The</strong> tortoises, Cinyxis (Testudinidaa)and Tetrathyra (Trionichidse).Amphibia.—Of Amphibia, <strong>the</strong>re are 2 peculiar genera <strong>of</strong> treefrogs,Hylambatis and Hcmimantis, belonging to <strong>the</strong> Polypedatidse.


chap. xi.J THE ETHIOPIAN REGION. 265Here, too, we find some interesting <strong>relations</strong> <strong>with</strong> <strong>the</strong> Orientalregion on <strong>the</strong> one side, and <strong>the</strong> Neotropical on <strong>the</strong> o<strong>the</strong>r. <strong>The</strong>snakes <strong>of</strong> <strong>the</strong> family Homalopsidse have a wide range, in America,Europe, and all over <strong>the</strong> Oriental region, but are confined toWest Africa in <strong>the</strong> Ethiopian region. Dryiophis (Dryiophidoe)and Dipsadoboa (Dipsadidse) on <strong>the</strong> o<strong>the</strong>r hand, are genera <strong>of</strong>tropical America which occur also in West Africa. <strong>The</strong> family<strong>of</strong> lizards, Acontiadae, are found in West and. South Africa,Ceylon, and <strong>the</strong> Moluccas. <strong>The</strong> family <strong>of</strong> toads, Engystomidse,in West and South Africa and <strong>the</strong> whole Oriental region ;while<strong>the</strong> Phryniscidse inhabit tropical Africa and Java.Insects.—We have here a large number <strong>of</strong> peculiar genera.<strong>The</strong>re are 10 <strong>of</strong> butterflies, Lachnoptera, Amphidema, and Catunabelonging to <strong>the</strong> Nymphalidse, while four o<strong>the</strong>rs are Lycsenidae.<strong>The</strong> genus Euxan<strong>the</strong> is common to West Africa and Madagascar.Of Coleoptera <strong>the</strong>re are 53 peculiar genera ; 20 are Carabidse,2 Lucanidse, 12 Cetoniidse, 3 Prionidae, 16 Cerambycidse, and34 Lamiidse. Besides <strong>the</strong>se <strong>the</strong>re are 4 or 5 genera confined toWest Africa and Madagascar.Land Shells.—West Africa is very rich in land shells, but itdoes not appear to possess any well-marked genera, althoughseveral <strong>of</strong> <strong>the</strong> smaller groups or sub-genera are confined to it.Helicidse <strong>of</strong> <strong>the</strong> genera Nanina, Bulimimis and Achatina areabundant and characteristic.Islands <strong>of</strong> <strong>the</strong> West African Sub-region.—<strong>The</strong> islands in <strong>the</strong>Gulf <strong>of</strong> Guinea are, Fernando Po, very near <strong>the</strong> main land, <strong>with</strong>Prince's Island and St. Thomas, considerably fur<strong>the</strong>r away to <strong>the</strong>south-west.Fernando Po was once thought to be a remarkableinstance <strong>of</strong> an island possessing a very peculiar fauna, althoughclose to <strong>the</strong> main land and not divided from it by a deep sea.This, however, was due to our having obtained considerablecollections from Fernando Po, while <strong>the</strong> opposite coast wasalmost unknown. One after ano<strong>the</strong>r <strong>the</strong> species supposed to bepeculiar have been found on <strong>the</strong> continent, tillit becomes probable,that, as in <strong>the</strong> case <strong>of</strong> o<strong>the</strong>r islands similarly situated, itcontains no peculiar species whatever. <strong>The</strong> presence <strong>of</strong> numerousmammalia, among which are baboons, lemurs, Hyrax, andVol. L—19


266 ZOOLOGICAL GEOGRAPHY. [part hi.Anomalurus, shows that this island has probably once been unitedto <strong>the</strong> continent.Prince's Island, situated about 100 miles from <strong>the</strong> coast, hasno mammals, but between 30 and 40 species <strong>of</strong> birds. Of- <strong>the</strong>se7 are peculiar species ,\iz., Zostcrojjs Jiccdulina, Cupliopterus dohmi(a peculiar genus <strong>of</strong> Sylviidse), Sympledes princeps, Crithagrarvfilata, Columba chlorophosa, Peristera principalis, and Strixthomensis.In <strong>the</strong> Island <strong>of</strong> St. Thomas, situated on <strong>the</strong> equator about 150miles from <strong>the</strong> coast, <strong>the</strong>re are 6 peculiar species out <strong>of</strong> 30 knownbirds, viz., Scops leucopsis, Zosterops lugubris, Turdus olivace<strong>of</strong>uscus,Oriolus crassirostris, Sympledes sandi-thomce and Aplopcliasimplex ; also Strix thomensis in common <strong>with</strong> Prince's Island.<strong>The</strong> remainder are all found on <strong>the</strong> adjacent coasts. It is remarkablethat in Prince'sIsland <strong>the</strong>re are no birds <strong>of</strong> prey, anythat appear being driven <strong>of</strong>f by <strong>the</strong> parrots (Psittams erithacus)that abound <strong>the</strong>re ; whereas in St. Thomas and Fernando Po<strong>the</strong>y are plentiful.III.South-African Sub-region.This is <strong>the</strong> most peculiar and interesting part <strong>of</strong> Africa, butowing to <strong>the</strong> absence <strong>of</strong> existing barriers its limits cannot bewell defined. <strong>The</strong> typical portion <strong>of</strong> it hardly contains morethan <strong>the</strong> narrow strip <strong>of</strong> territory limited by <strong>the</strong> mountain rangewhich forms <strong>the</strong> boundary <strong>of</strong> <strong>the</strong> Cape Colony and Natal, whilein a wider sense it may be extended to include Mozambique. Itmay perhaps be best characterised as bounded by <strong>the</strong> Kalaharidesert and <strong>the</strong> Limpopo river. It is in <strong>the</strong> more limited district<strong>of</strong> <strong>the</strong> extreme south, that <strong>the</strong> wonderful Cape flora alone exists.Here are more genera and species, and more peculiar types <strong>of</strong>plants congregated toge<strong>the</strong>r, than in any o<strong>the</strong>r part <strong>of</strong> <strong>the</strong> globe<strong>of</strong> equal extent. <strong>The</strong>re are indications <strong>of</strong> a somewhat similarrichness and specialization in <strong>the</strong> zoology <strong>of</strong> this country; but<strong>animals</strong> are so much less closely dependent on soil and climate,that much <strong>of</strong> <strong>the</strong> original peculiarity has been obliterated, bylong continued interchange <strong>of</strong> species <strong>with</strong> so vast an area as


——chap, xi.] THE ETHIOPIAN REGION. 267that <strong>of</strong> Africa south <strong>of</strong> <strong>the</strong> equator. <strong>The</strong> extreme peculiarityaud isolation <strong>of</strong> <strong>the</strong> flora must not, however, be lost sight <strong>of</strong>, ifwe would correctly interpret <strong>the</strong> phenomena afforded by <strong>the</strong> <strong>distribution</strong><strong>of</strong> animal life on <strong>the</strong> African continent.Mammalia.—A much larger number <strong>of</strong> peculiar forms <strong>of</strong> mammalsare found here than in any <strong>of</strong> <strong>the</strong> o<strong>the</strong>r sub-regions,although it is far less in extent than ei<strong>the</strong>r <strong>of</strong> <strong>the</strong> three divisions<strong>of</strong> <strong>the</strong> continent. Among Insectivora we have <strong>the</strong>Chrysochloridae, or golden moles, consisting <strong>of</strong> two genera confinedto South Africa ; while <strong>the</strong> Macroscelididae, or elephant shrews,are also characteristically South African, although ranging as far asMozambique and <strong>the</strong> Zambezi, <strong>with</strong> one outlying species in NorthAfrica. <strong>The</strong> Viverridse are represented by three peculiar genera,Ariela, Cynictis, and Suricata. <strong>The</strong> Carnivora present someremarkable forms : Proteles, forming a distinct family allied to<strong>the</strong> hyaenas and weasels ; and two curious forms <strong>of</strong> CanidasMegalotis (<strong>the</strong> long-eared fox) and Lycaon (<strong>the</strong> hyaena-dog), <strong>the</strong>latter found also in parts <strong>of</strong> East Africa. Hydrogale is apeculiar form <strong>of</strong> Mustelidae ;Pelea one <strong>of</strong> <strong>the</strong> antelopes ; Dendromys,Malacothrix, and Mystromys are peculiar genera <strong>of</strong> <strong>the</strong>mouse family (Muridae) ; Bathyerges one <strong>of</strong> <strong>the</strong> mole-rats (Spalacidae); Pedetes, <strong>the</strong> Cape-hare, a remarkable form <strong>of</strong> jerboa ; andPetromys, one <strong>of</strong> <strong>the</strong> spiny-rats (Echimyidae). <strong>The</strong> remarkableOryderopus, or earth-pig, has one species in South and one inNorth East Africa. We have thus eighteen genera <strong>of</strong> mammaliaalmost or quite peculiar to South Africa.Birds.—<strong>The</strong>se do not present so many peculiar forms, yetsome are very remarkable. Chcctops is an isolated genus <strong>of</strong>thrushes (Turdidre). Lioptilus, one <strong>of</strong> <strong>the</strong> fruit-thrushes (Pycnonotidae).Pogonocichla, one <strong>of</strong> <strong>the</strong> fly-catchers; Urolestes, ashrike ; Promcrops, a sun-bird ; Philetwrns and Chera, weaverbirds;and three peculiar genera <strong>of</strong> larks Spizocorys, Heterocorys,and Tephrocorys, complete <strong>the</strong> list <strong>of</strong> peculiar types <strong>of</strong> Passeres.A wood-pecker, Geocolaptes, is nearly allied to a South Americangenus. <strong>The</strong> Cape-dove, (Ena, is confined to South and East Africaand Madagascar ; and Thalassornis is a peculiar form <strong>of</strong> duck.Several genera are also confined to West and South Africa ;


;268 ZOOLOGICAL GEOGRAPHY. [part hi.as Phyllastrephus (Pycnonotidse), Smithomis (Muscicapidse),Corvinella (Laniidse) ;Barbatula and Xyldbucco (Megalsemidse)Ceuthmochares, also in Madagascar, (Cuculidae) ;Typanistria(ColumbidEe). O<strong>the</strong>r remarkable forms, though widely spreadover Africa, appear to have <strong>the</strong>ir metropolis here, as Colius andIndicator. O<strong>the</strong>rs seem to be confined to South Africa andAbyssinia, as <strong>the</strong> curious Bwphaga (Sturnidse) ; and Apaloderma(Trogonidse). Machcerhamphus (Falconidse) is found onlyin South-West Africa, Madagascar, and <strong>the</strong> Malay Peninsula.Reptiles.—<strong>The</strong>re are 4 peculiar genera <strong>of</strong> smikes,—Typhline,belonging to <strong>the</strong> blind burrowing snakes, Typhlopidse ;Lamprophis(Lycodontidse) ; Cyrtophis and Pcecilophis (Elapidae), afamily which is chiefly Oriental and Australian. Of Lizards<strong>the</strong>re are 10 peculiar genera ; Monotrophis (Lepidosternidse), but<strong>with</strong> an allied form in Angola ; Cordylus, Pseudocordylus, Platysaurus,Cordylosaurus, Pleurostichus, and Saurophis, allgenera <strong>of</strong> Zonuridse ;Chamsesauridse ;peculiarGhamcesaura, forming <strong>the</strong> peculiar familyColopus and Rhopitropus (Geckotidae).Amphibia— Of Amphibia <strong>the</strong>re are 4 peculiar genera:Schismaderma (Bufonidae) ;Brachymerus (Engystomidse) ;Phrynobatrackusand Stenorhynchus (Kanidee). <strong>The</strong>se last are alliedto Oriental genera, and <strong>the</strong> only o<strong>the</strong>r Engystomidse are Orientaland Neotropical.Fresh-water Fish—Of fresh-water fishes <strong>the</strong>re is1 genus—Abrostomus—belongingto <strong>the</strong> carp family, peculiar to South Africa.Insects.—South Africa is excessively rich in insects, and <strong>the</strong>number <strong>of</strong> peculiar types surpasses that <strong>of</strong> any o<strong>the</strong>r part <strong>of</strong> <strong>the</strong>We can only here summarize <strong>the</strong> results.Lcpidoptera.—Of butterflies <strong>the</strong>re are 7 peculiar genera ;2region.belonging to <strong>the</strong> Satyridse, 1 to Acrseidas, 3 to Lycsenidae,and 1 to Hesperidee. Zeritis (Lycaanidse) is also characteristic<strong>of</strong> this sub-region, although 1 species occurs in West Africa.Coleoptera.— <strong>The</strong>se are very remarkable. In <strong>the</strong> family <strong>of</strong>CicindelidaB, or tiger-beetles, we have <strong>the</strong> extraordinary Manticoraand Platychile, forming a sub-family, whose nearest allies are inNorth America ;as well as Ophryodera and Dromica, <strong>the</strong> latteran extensive genus, which ranges as far north as Mozambique


—chap, xi.] THE ETHIOPIAN REGION. 269and Lake Ngami. Ano<strong>the</strong>r genus <strong>of</strong> tHis family, Jansenia, iscommon to South Africa and South India.In <strong>the</strong> large family <strong>of</strong> Carabidae,or ground-beetles, <strong>the</strong>re are17 peculiar South African genera, <strong>the</strong> most important beingCrcpidogaster, ffytrichopus, Arsinoe, and Piezia.Three o<strong>the</strong>rsEunostus, Glyphodactyla, and Megalonychus—are common toSouth Africa and Madagascar only. <strong>The</strong>re is also a genus iacommon <strong>with</strong> Java, and one <strong>with</strong> Australia.Of Lucanidae, or stag-beetles, <strong>the</strong>re are 3 peculiar genera ; <strong>of</strong>Cetoniidse, or rose-chafers, 14 ; and <strong>of</strong> Buprestidee, 2.In <strong>the</strong> great family <strong>of</strong> Longicorns <strong>the</strong>re are no less than 67peculiar genera—an immense number when we consider that <strong>the</strong>generally open character <strong>of</strong> <strong>the</strong> country, is such as is not usuallywell suited to this group <strong>of</strong> insects. <strong>The</strong>y consist <strong>of</strong> 5 peculiargenera <strong>of</strong> Prionidse, 25 <strong>of</strong> Cerambycidae, and 37 <strong>of</strong> Lamiidae.Summary <strong>of</strong> South-African Zoology.—Summarizing <strong>the</strong>se results,we find that South Africa possesses 18 peculiar genera <strong>of</strong>Mammalia, 12 <strong>of</strong> Birds, 18 <strong>of</strong> Eeptiles, 1<strong>of</strong> Fishes, 7 <strong>of</strong> Butterflies,and 107 <strong>of</strong> <strong>the</strong> six typical families <strong>of</strong> Coleoptera.Besidesthis large amount <strong>of</strong> speciality it contains many o<strong>the</strong>r groups,which extend ei<strong>the</strong>r to West Africa, to Abyssinia, or to Madagascaronly, a number <strong>of</strong> which are no doubt to be referred asoriginating here.We also find many cases <strong>of</strong> direct affinity <strong>with</strong><strong>the</strong> Oriental region, and especially <strong>with</strong> <strong>the</strong> Malay districts, ando<strong>the</strong>rs <strong>with</strong> Australia ; and <strong>the</strong>re are also less marked indications<strong>of</strong> a relation toAmerica.Atlantic Islands <strong>of</strong> <strong>the</strong> Ethiopian Region. St. Hehna.—<strong>The</strong>position <strong>of</strong> St. Helena, about 1,000 miles west <strong>of</strong> Africa and 16°south <strong>of</strong> <strong>the</strong> equator, renders it difficult to place it in ei<strong>the</strong>r <strong>of</strong><strong>the</strong> sub-regions ; and its scanty fauna has a general ra<strong>the</strong>r thanany special resemblance to that <strong>of</strong> Africa. <strong>The</strong> entire destruction<strong>of</strong> its luxuriant native forests by <strong>the</strong> introduction <strong>of</strong> goatswhich killed all <strong>the</strong> young trees (a destruction which was nearlycompleted two centuries ago) must have led to <strong>the</strong> extermination<strong>of</strong> most <strong>of</strong> <strong>the</strong> indigenous birds and insects. At present <strong>the</strong>re isno land bird that is believed to be really indigenous, and but one


;270 ZOOLOGICAL GEOGRAPHY. [pakt in.wader, a small plover {JEgialitis sandce-helence) which is peculiarto <strong>the</strong> island, but closely allied to African species. Numerousimported birds, such as canaries, Java sparrows, some Africanfinches, guinea-fowls, and partridges, are now wild. <strong>The</strong>re areno native butterflies, but a few introduced species <strong>of</strong> almostworld-wide range. <strong>The</strong> only important remnant <strong>of</strong> <strong>the</strong> originalfauna consists <strong>of</strong> beetles and land shells. <strong>The</strong> beetles are <strong>the</strong>more numerous and have been critically examined and describedby Mr. T. V. Wollaston, whose researchesislands are so well known.in <strong>the</strong> o<strong>the</strong>r AtlanticColeoptera <strong>of</strong> St. Helena.— Omitting those beetles which getintroduced everywhere through man's agency, <strong>the</strong>re are 59 species<strong>of</strong> Coleoptera known from St. Helena ;and even <strong>of</strong> <strong>the</strong>se <strong>the</strong>reare a few widely distributed speciesthat may have been introducedby man. It will be well, <strong>the</strong>refore, to confine ourselvesalmost wholly to <strong>the</strong> species peculiar to <strong>the</strong> island, and, <strong>the</strong>refore,almost certainly forming part <strong>of</strong> <strong>the</strong> endemic or original fauna.Of <strong>the</strong>se we find that 10 belong to genera which have a verywide range, and thus afford no indication <strong>of</strong> <strong>geographical</strong> affinity2 belong to genera which are characteristic <strong>of</strong> <strong>the</strong> Palamrcticfauna {Bembidium, Longitarsus) ;3 to African genera (Adoretus,Sciobius, Aspidomorpha) ; and two species <strong>of</strong> Calosoma are mostallied to African species. <strong>The</strong>re are also 4 African species,which may be indigenous in St. Helena. <strong>The</strong> peculiar genera,7 in number, are, however, <strong>the</strong> most interesting. We have firstHaplothorax, a large beetle allied to Carabus and Calosoma, though<strong>of</strong> a peculiar type. This may be held to indicate a remotePaloearctic affinity.Melissius, one <strong>of</strong> <strong>the</strong> Dynastidse, is allied toSouth African forms. .Microxylobius, one <strong>of</strong> <strong>the</strong> Cossonides (asub-family <strong>of</strong> Curculionidas) is <strong>the</strong> most important genus, comprisingas it does 13 species. It is, according to Mr. Wollaston,an altoge<strong>the</strong>r peculiar type, most allied to Pentarthrurn,, a genusfound in St. Helena, Ascension, and <strong>the</strong> south <strong>of</strong> England, anditself very isolated. Nesiotes, ano<strong>the</strong>r genus <strong>of</strong> Curculionidse,belongs to a small group, <strong>the</strong> allied genera forming which inhabitEurope, Madeira, and Australia. A third peculiar and isolatedgenus is Trachyphlwosoma. <strong>The</strong> Anthribidie are represented by


;chap, xi.] THE ETHIOPIAN REGION. 2712 genera, Notioxerms and Homceodera, which are altoge<strong>the</strong>rpeculiar and isolated, and contain 9 species.Thus no less than27 species, or more than half <strong>of</strong> <strong>the</strong> undoubtedly indigenousbeetles, belong to 5 peculiar and very remarkable genera <strong>of</strong>Ehyncophora.It appears from this enumeration, that<strong>the</strong> peculiar species asa whole, exhibit most affinity to <strong>the</strong> Ethiopian fauna ; next to<strong>the</strong> South European fauna ;and lastly to that <strong>of</strong> <strong>the</strong> islands <strong>of</strong><strong>the</strong> North Atlantic ;while <strong>the</strong>re is such a large amount <strong>of</strong> peculiarityin <strong>the</strong> most characteristic forms, that no special <strong>geographical</strong>affinity can be pointed out.Land Shells.—<strong>The</strong>se consist <strong>of</strong> about a dozen living species, andabout as many extinct found in <strong>the</strong> surface soil, and probablyexterminated by <strong>the</strong> destruction <strong>of</strong> <strong>the</strong> forests. <strong>The</strong> genera areSuccinea, Zonites, Helix, Bidimus, Pupa, and Achatina. <strong>The</strong>Bidimi (all now extinct but one) comprise one large, andseveral small species, <strong>of</strong> a peculiar type, most resembling formsnow inhabiting South America and <strong>the</strong> islands <strong>of</strong> <strong>the</strong> Pacific.Zonites is chiefly South European, but <strong>the</strong> o<strong>the</strong>r genera are <strong>of</strong>wide range, and none are peculiar to <strong>the</strong> island.<strong>The</strong> marine shells aremostly Mediterranean, or West Indianspecies, <strong>with</strong> some found in <strong>the</strong> Indian Ocean ;being peculiar to <strong>the</strong> island.Tristan d'Acunha.—This small island issituated nearly midwaybetween <strong>the</strong> Cape <strong>of</strong> Good Hope and <strong>the</strong>only 4 or 5 speciesmouth <strong>of</strong> <strong>the</strong> LaPlata, but it is ra<strong>the</strong>r nearer Africa than America, and a littlenearer still to St. Helena. An island so truly oceanic and <strong>of</strong> whoseproductions so little is known, cannot be placed in any region,and is only noticed here because it comes naturally after St. Helena.It is known to possess three peculiar land birds. One is a thrush(Ncsocichla eremita) whose exact affinities are not determined<strong>the</strong> o<strong>the</strong>r a small water-hen (Gallinula nesiotis) allied to ournative species, but <strong>with</strong> shorter and s<strong>of</strong>ter wings, which <strong>the</strong> birddoes not use for flight. A finch <strong>of</strong> <strong>the</strong> genus Crithagra showsAfrican affinities ;while ano<strong>the</strong>r recently described as Nesospizaaeunlim (Journ. fur Orn. 1873, p. 154) forms a new genus said toresemble more nearly some American forms.


272 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>The</strong> only known land-shells are 2 peculiar species <strong>of</strong> Balea, agenus only found elsewhere in Europe and Brazil.IV. Madagascar and <strong>the</strong> Mascarene Islands, or <strong>the</strong> MalagasySub-region.This insular sub-region is one <strong>of</strong> <strong>the</strong> most remarkable zoologicaldistricts on <strong>the</strong> globe, bearing a similar relation to Africaas <strong>the</strong> Antilles to tropical America, or New Zealand to Australia,but possessing a much richer fauna than ei<strong>the</strong>r <strong>of</strong> <strong>the</strong>se, and insome respects a more remarkable one even than New Zealand.It comprises, besides Madagascar, <strong>the</strong> islands <strong>of</strong> Mauritius,Bourbon, and Bodriguez, <strong>the</strong> Seychelles and Comoro islands.Madagascar itself is an island <strong>of</strong> <strong>the</strong> first class, being a thousandmiles long and about 250 miles in average width. It liesparallel to <strong>the</strong> coast <strong>of</strong> Africa, near <strong>the</strong> sou<strong>the</strong>rn tropic, and isseparated by 230 miles <strong>of</strong> sea from <strong>the</strong> nearest part <strong>of</strong> <strong>the</strong> continent,although a bank <strong>of</strong> soundings projecting from its westerncoast reduces this distance to about 160 miles. Madagascar isa mountainous island, and <strong>the</strong> greater part <strong>of</strong> <strong>the</strong> interior consists<strong>of</strong> open elevated plateaus ;but between <strong>the</strong>se and <strong>the</strong> coast <strong>the</strong>reintervene broad belts <strong>of</strong> luxuriant tropical forests. It is thisforest-district which has yielded most <strong>of</strong> those remarkable types<strong>of</strong> animal life which we shall have to enumerate ; and it isprobable that many more remain to be discovered. As all <strong>the</strong>main features <strong>of</strong> this sub-region are developed in Madagascar,we shall first endeavour to give a complete outline <strong>of</strong> <strong>the</strong> fauna<strong>of</strong> that country, and afterwards show how far <strong>the</strong> surroundingislands partake <strong>of</strong> its peculiarities.Mammalia.—<strong>The</strong> fauna <strong>of</strong> Madagascar is tolerably rich ingenera and species <strong>of</strong> mammalia, although <strong>the</strong>se belong to a verylimited number <strong>of</strong> families and orders. It is especially characterizedby its abundance <strong>of</strong> Lemuridaa and Insectivora ;it alsopossesses a few peculiar Carnivora <strong>of</strong> small size ; but most <strong>of</strong><strong>the</strong> o<strong>the</strong>r groups in which Africa is especially rich—apes andmonkeys, lions, leopards and hyaenas, zebras, giraffes, antelopes,elephants and rhinoceroses, and even porcupines and squirrels,are wholly wanting. No less than 40 distinct families <strong>of</strong> land


—CHAP. XI.] THE ETHIOPIAN REGION. 273mammals are represented on <strong>the</strong> continent <strong>of</strong> Africa, only 11 <strong>of</strong>which occur in Madagascar, which also possesses 3 familiespeculiar to itself. <strong>The</strong> following is a list <strong>of</strong> all <strong>the</strong> genera <strong>of</strong>Mammalia as yet known to inhabit <strong>the</strong> island :PRIMATES.


274 ZOOLOGICAL GEOGRAPHY. [part hisented by a single group in Africa, <strong>with</strong> two peculiar forms on<strong>the</strong> West coast.<strong>The</strong>y also re-appear under peculiar and isolatedforms in Sou<strong>the</strong>rn India and Malaya, and are evidently but <strong>the</strong>remains <strong>of</strong> a once wide-spread group, since in Eocene times <strong>the</strong>yinhabited North America and Europe, and very probably <strong>the</strong>whole nor<strong>the</strong>rn hemisphere.<strong>The</strong> Insectivora are ano<strong>the</strong>r group<strong>of</strong> high antiquity, widely scattered over <strong>the</strong> globe under anumber <strong>of</strong> peculiar forms ;but in no equally limited area representedby so many peculiar types as in Madagascar.South andWest Africa are also rich in this order.<strong>The</strong> Carnivora <strong>of</strong> Madagascar are mostly peculiar forms <strong>of</strong>Viverridas, or civets, a family now almost confined to <strong>the</strong>Ethiopian and Oriental regions, but which was abundant inEurope during <strong>the</strong> Miocene period.<strong>The</strong> Potamochcerus is a peculiar species only, which may beperhaps explained by <strong>the</strong> unusual swimming powers <strong>of</strong> swine,and <strong>the</strong> semi-aquatic habits <strong>of</strong> this genus, leading to an immi-<strong>of</strong> <strong>the</strong> o<strong>the</strong>r Mammalia.gration at a later period than in <strong>the</strong> case<strong>The</strong> same remark will apply to <strong>the</strong> small Hippopotamus, whichwas coeval <strong>with</strong> <strong>the</strong> great Struthious bird iEpiornis.Rodents are only represented by three peculiar forms <strong>of</strong>Muridse, but it isprobable that o<strong>the</strong>rs remain to be discovered.Birds.—Madagascar is exceedingly rich in birds, and especiallyin remarkable forms <strong>of</strong> Passeres. No less than 88 generaand 111 species <strong>of</strong> land-birds have been discovered, and everyyear some additions are being made to <strong>the</strong> list. <strong>The</strong> Africanfamilies <strong>of</strong> Passeres are almost all represented, only two beingabsent—Parid*and Fringillidse, both very poorly represented inAfrica itself. Among <strong>the</strong> Picarhe, however, <strong>the</strong> case is verydifferent, no less than 7 families being absent, viz.—Picidse,or woodpeckers ; Indieatoridre, or honey-guides ;Megaleemidae,or barbets ;Musophagidse, or plantain-eaters ; Coliidse, or colies;Bucerotidse, or hornbills ; and Irrisorida?, or mockers. Three <strong>of</strong><strong>the</strong>se arepeculiar to Africa, and all are well represented <strong>the</strong>re,so that <strong>the</strong>ir absence from Madagascar isa very remarkable fact.<strong>The</strong> number <strong>of</strong> peculiar genera in Madagascar constitutes one <strong>of</strong><strong>the</strong> main features <strong>of</strong> its ornithology, and many <strong>of</strong> <strong>the</strong>se are so


;CHAP. XI.] THE ETHIOPIAN REGION. 275isolated that it is very difficult to classify <strong>the</strong>m, and <strong>the</strong>yremain to this day a puzzle to ornithologists. In order toexhibit clearly <strong>the</strong> striking characteristics <strong>of</strong> <strong>the</strong> bird-fauna <strong>of</strong>this island, we shall first give a list <strong>of</strong> all <strong>the</strong> peculiar generaano<strong>the</strong>r, <strong>of</strong> <strong>the</strong> genera <strong>of</strong> which <strong>the</strong> species only are peculiarand, lastly, a list <strong>of</strong> <strong>the</strong> species which Madagascar possessesin common <strong>with</strong> <strong>the</strong> Africancontinent.Genera op Birds peculiar to Madagascar, or found elsewhereonly in <strong>the</strong> mascarene islands.Sylviid^e.1. Bernieria2. Ellisia3. Mystacornis ...4. Eroessa5. Gervasia


276 ZOOLOGICAL GEOGRAPHY. [PART III.Ethiopian or Oriental Genera which are represented inMadagascar by peculiar species.Turdid^e.Species.ALCEDINIDiE.Species.1. Bessonornis ...21. Corythorais ..22. Ispidina11SyLVIID.E.2. Acrocephalus ...3. Copsychus (Or.)4. Pratincola111Upupid^e.23. Upupa (?) ..PYCNONOTID.E.5. Hijpsipetes (Or.)Caprimulgid^:.24. Caprimulgus6. Andropadus ...Campephagid^:.7. Campephaga ...CYPSELIDiE.25. Cypselus26. Chaetura21DlCRURIDiE.Psittacid^e.8. Dicrurus27. Poliopsitta ..Muscicapid^:.9. TchitreaLa.niid^e.CoLUMBID^E.28. Treron29. Columba30. Turtur10. LaniariusPteroclid^e.Nectariniid^e.31. Pterocles11. Nectarinia ...PLOCEIID.E.12. Foudia13. Hypargos14. Spermestes ...211Tetraonid^e.32. Francolinus .,Phasianid^e.33. NumidaAlaudid^e.15. MirafraTURNICIDiE.34. TurnixMotacillid^e.16. MotacillaCuculid.e.17. Ceuthrnochares18. Centropus ...19. CuculusC0RACIID.E.20. Eurystomus ...111FALCONID.E.35. Polyboroides36. Circus37. Astur...38. Accipiter39. Buteo...40. Haliaeetus .41. Pernis42. Baza ...43. Cerchneis113111111


—CHAP. XI.] THE ETHIOPIAN KEGION. 277Strigid-e.44. A<strong>the</strong>ne45. ScopsSpecies.11PLATALEID.E.49. Ibis ...Species.Rallid-j:.46. Rallus47. Porzana31Podicipid^;.50. PodicepsScOLOPACID.dE.48. GallinagoTotal peculiar species <strong>of</strong> Eth. \ ,1 or Or. genera ... ... /fiSpecies <strong>of</strong> Birds common to Madagascar and Africa or Asia.1. Cisticola cursitans.2. Corvus scapulatus.3. Crithagra canicollis.4. Merops superciliosus.5. Collocalia fuciphaga.6. OSna capensis.7. Aplopelia tympanistria.8. Falco minor.9. Falco concolor.10. Milvus segyptius.11. Milvus migrans.12. Strix flammea.<strong>The</strong>se three tables show us an amount <strong>of</strong> speciality hardly tobe found in <strong>the</strong> birds <strong>of</strong> any o<strong>the</strong>r part <strong>of</strong> <strong>the</strong> globe. Out <strong>of</strong>111 land-birds in Madagascar, only 12 are identical <strong>with</strong> speciesinhabiting <strong>the</strong> adjacent continents, and most <strong>of</strong> <strong>the</strong>se belong topowerful-winged, or wide-ranging forms, which probably now<strong>of</strong>ten pass from one country to <strong>the</strong> o<strong>the</strong>r. <strong>The</strong> peculiar species—49 land-birds and 7 waders, or aquatics—are mostly wellmarkedforms <strong>of</strong> African genera. <strong>The</strong>re are, however, severalgenera (marked by italics) which have Oriental or Palsearctioaffinities, but not African, viz. Copsychus, Hypsipetes, Hypherpes,Alcrtrcenas, and Margaroperdix. <strong>The</strong>se indicate a closer approximationto <strong>the</strong> Malay countries than now exists.<strong>The</strong> table <strong>of</strong> 33 peculiar genera is <strong>of</strong> great interest. Most <strong>of</strong><strong>the</strong>se are well-marked forms, belonging to families which arefully developed in Africa; though it is singular that notone <strong>of</strong> <strong>the</strong> exclusively African families is represented inany way in Madagascar. O<strong>the</strong>rs, however, are <strong>of</strong> remote oraltoge<strong>the</strong>r doubtful affinities. Sittidce is Oriental and Palaearctic,but not Ethiopian. Oxylabes and Mystacornis are <strong>of</strong> doubtfulaffinities. Artainia and Cyanolanius still more so, and it isquite undecided what family <strong>the</strong>y belong to. Calicalicus isalmost equally obscure. Neodrcpanis, one <strong>of</strong> <strong>the</strong> most recentdiscoveries, seems to connect <strong>the</strong> Nectariniidse <strong>with</strong> <strong>the</strong> Pacific


278 ZOOLOGICAL GEOGRAPHY. [part hi.Depanidida?. Euryceros is a complete puzzle, having beenplaced <strong>with</strong> <strong>the</strong> hornbills, <strong>the</strong> starlings, or as a distinctfamily.Falculia is an exceedingly aberrant form <strong>of</strong> starling, long thoughtto be allied to Irrisor. Philepitta, forming a distinct family,(Paictidse), is most remarkable and isolated, perhaps <strong>with</strong> remoteSouth American affinities. Leptosoma is ano<strong>the</strong>r extraordinaryform, connecting <strong>the</strong> cuckoos <strong>with</strong> <strong>the</strong> rollers. Atelornis,Brachypteracias, and Geobiastes, are terrestrial rollers, <strong>with</strong> <strong>the</strong>form and colouring <strong>of</strong> Pitta. So many perfectly isolated andremarkable groups are certainly nowhere else to be found ; and<strong>the</strong>y fitly associate <strong>with</strong> <strong>the</strong> wonderful aye -aye (Chiromys),<strong>the</strong> insectivorous Centetidee, and carnivorous Cryptoproctaamong <strong>the</strong> Mammalia. <strong>The</strong>y speak to us plainly <strong>of</strong> enormousantiquity, <strong>of</strong> long-continued isolation ;and not less plainly <strong>of</strong>a lost continent or continental island, in which so many, andvarious, and peculiarly organized creatures, could have beengradually developed in a connected fauna, <strong>of</strong> which we havehere but <strong>the</strong> fragmentary remains.Plate VI— Illustrating <strong>the</strong> characteristic features <strong>of</strong> <strong>the</strong>Zoology <strong>of</strong> Madagascar.—<strong>The</strong> lemurs, which form <strong>the</strong> mostprominent feature in <strong>the</strong> zoology <strong>of</strong> Madagascar, being comparativelywell-known from <strong>the</strong> numerous specimens in ourzoological gardens ;and good figures <strong>of</strong> <strong>the</strong> Insectivorous generanot being available, we have represented <strong>the</strong> nocturnal andextraordinary aye-aye (Chiromys madagascariensis) to illustrateits peculiar and probably very ancient mammalian fauna ;while<strong>the</strong> river-hogs in <strong>the</strong> distance (Potamochcerus edwardsii) allied toAfrican species, indicate a later immigration from <strong>the</strong> mainlandthan in <strong>the</strong> case <strong>of</strong> most <strong>of</strong> <strong>the</strong> o<strong>the</strong>r Mammalia. <strong>The</strong>peculiar birds being far less generally known, we have figuredthree <strong>of</strong> <strong>the</strong>m. <strong>The</strong> largest is <strong>the</strong> Euryceros prevosti, here classed<strong>with</strong> <strong>the</strong> starlings, although itsremarkable bill and o<strong>the</strong>r peculiaritiesrender it probable that itshould form a distinct family.Its colours are velvety black and rich brown <strong>with</strong> <strong>the</strong> billpearly grey.<strong>of</strong> a<strong>The</strong> bird beneath (Vanga curvirostris) is one <strong>of</strong> <strong>the</strong>peculiar Madagascar shrikes whose plumage, variegated <strong>with</strong>green-black and pure white is very conspicuous ; while that in


PLATE VI.SCENE IN MADAGASCAR, WITH CHARACTERISTIC ANIMALS.


—chap, xi.] THE ETHIOPIAN REGION. 279<strong>the</strong> right hand corner is <strong>the</strong> Lcptosoma discolor, a bird whichappears to be intermediate between such very distinct familiesas <strong>the</strong> cuckoos and <strong>the</strong> rollers, and is <strong>the</strong>refore considered t<strong>of</strong>orm a family by itself. It is a coppery-green above and nearlywhite beneath, <strong>with</strong> a black bill and red feet. <strong>The</strong> fan-shapedplant on <strong>the</strong> left is <strong>the</strong> traveller's tree (Urania speciosa), one <strong>of</strong><strong>the</strong> peculiar forms <strong>of</strong> vegetation in this marvellous island.Reptiles.—<strong>The</strong>se present some very curious features,comparativelyfew <strong>of</strong> <strong>the</strong> African groups being represented, while <strong>the</strong>reare a considerable number <strong>of</strong> Eastern and even <strong>of</strong> Americanforms. Beginning <strong>with</strong> <strong>the</strong> snakes, we find, in <strong>the</strong> enormousfamily <strong>of</strong> Colubridse, none <strong>of</strong> <strong>the</strong> African types ; but instead <strong>of</strong><strong>the</strong>m three genera Herpetodryas, PMlodryas, and Heterodon—only found elsewhere in South and North America. <strong>The</strong>Psammophidse, which are both African and Indian, are representedby a peculiar genus, Mimophis. <strong>The</strong> Dendrophidse arerepresented by Ahcetvlla, a genus which is both African andAmerican. <strong>The</strong> Dryiophidse, which inhabit all <strong>the</strong> tropics butare most developed in <strong>the</strong> Oriental region, are represented bya peculiar genus, Langaha. <strong>The</strong> tropical Pythonidse are representedby ano<strong>the</strong>r peculiar genus, Sanzinia. <strong>The</strong> Lycodontidaeand Viperidse, so well developed in Africa, are entirely absent.<strong>The</strong> lizards are no less remarkable. <strong>The</strong> Zonuridse, abundantlydeveloped in Africa, are represented by one peculiargenus, Cicigna. <strong>The</strong> wide-spread Scincidae by ano<strong>the</strong>r peculiar<strong>The</strong> African Sepsidse, are represented by threegenus, Pygomeles.genera, two <strong>of</strong> which are African, and one, Amphiglossus, peculiar.<strong>The</strong> Acontiadse are represented by a species <strong>of</strong> <strong>the</strong> African genusAcontias. Of Scincidae <strong>the</strong>re is <strong>the</strong> wide-spread Euprepes. <strong>The</strong>Sepidse are represented by <strong>the</strong> African genera Seps and Scelotes.<strong>The</strong> Geckotidse are not represented by any purely Africangenera, but by Phyllodactylus, which is American and Australian;Hemidactyhis, which is spread over all <strong>the</strong> tropics ; bytwo peculiar genera ;and by Uroplatis, GecJcolepis, and Phelsuma,confined to Madagascar, Bourbon, and <strong>the</strong> Andaman Islands.<strong>The</strong> Agamidae, which are mostly Oriental and are represented in


—;280 ZOOLOGICAL GEOGRAPHY. [part hi.Africa by <strong>the</strong> single genus Agama, have here three peculiargenera, Trachcloptychus, Chalarodon, and Hoplurus. Lastly, <strong>the</strong>American Iguanidse are said to be represented by a species <strong>of</strong> <strong>the</strong>South American genus Opiums. <strong>The</strong> classification <strong>of</strong> Reptilesis in such an unsettled state that some <strong>of</strong> <strong>the</strong>se determinations<strong>of</strong> affinities are probably erroneous ; but it is not likely thatany corrections which may be required will materially affect <strong>the</strong>general bearing <strong>of</strong> <strong>the</strong> evidence, as indicating a remarkableamount <strong>of</strong> Oriental and American <strong>relations</strong>hip.<strong>The</strong> o<strong>the</strong>r groups are <strong>of</strong> less interest. Tortoises are representedby two African or wide-spread genera <strong>of</strong> Testudinidee,Teshido and Chersina, and by one peculiar genus, Pyxis ; and<strong>the</strong>re are also two African genera <strong>of</strong> Chelydidae.<strong>The</strong> Amphibia are not very well known. <strong>The</strong>y appear to beconfined to species <strong>of</strong> <strong>the</strong> wide-spread Ethiopian and Orientalgenera Hylarana, Polypedates, and Rappia (Polypedatidse)and Pyxicephahcs (Ranidse).Frcsh-ivater Fishes.—<strong>The</strong>se appear to be at present almostunknown. When carefully collected <strong>the</strong>y will no doubt furnishsome important facts.TJie Mascarene Islands.<strong>The</strong> various islands which surround Madagascar—Bourbon,Mauritius, Rodriguez, <strong>the</strong> Seychelles, and <strong>the</strong> Comoro Islands—all partake in a considerable degree <strong>of</strong> its peculiar fauna,while having some special features <strong>of</strong> <strong>the</strong>ir own.Indigenous Mammalia (except bats) are probably absent fromall <strong>the</strong>se islands (except <strong>the</strong> Comoros), although Lemur and Centetesare given as natives <strong>of</strong> Bourbon and Mauritius. <strong>The</strong>y have,however, perhaps been introduced from Madagascar. Lemurmayottensis, a peculiar species, is found in <strong>the</strong> Comoro Islands,where a Madagascar species <strong>of</strong> Viverra also occurs.Bourbon and Mauritius may be taken toge<strong>the</strong>r, as <strong>the</strong>y muchresemble each o<strong>the</strong>r. <strong>The</strong>y each possess species <strong>of</strong> a peculiargenus <strong>of</strong> Campephagidse, or caterpillar shrikes, Oxynotus; while<strong>the</strong> remarkable Fregilupus, belonging to <strong>the</strong> starling family,inhabits Bourbon, if it is not now extinct. <strong>The</strong>y also have


chap, xi.] THE ETHIOPIAN REGION." 281peculiar species <strong>of</strong> Pratincola, Hypsipetcs, Phedina, Tchitrea,Zosteraps, Foudia, Collocalia, and Coracopsis ;while Mauritius hasa very peculiar form <strong>of</strong> dove <strong>of</strong> <strong>the</strong> sub-genus Trocaza ;an Alectrainas,extinct <strong>with</strong>in <strong>the</strong> last thirty years ;Oriental genus <strong>of</strong> parroquets, Palceornis.and a species <strong>of</strong> <strong>the</strong><strong>The</strong> small and remoteisland <strong>of</strong> Rodriguez has ano<strong>the</strong>r Palceornis, as well asa peculiarFoudia, and a Drymceca <strong>of</strong> apparently Indian affinity.Coming to <strong>the</strong> Seychelle Islands, far to <strong>the</strong> north, we find <strong>the</strong>only mammal an Indian species <strong>of</strong> bat (Pteropus edivardsii).<strong>the</strong> twelve land-birds all but one are peculiar species, but all belongto genera found also in Madagascar, except one—a peculiar species<strong>of</strong> Palmomis. This is an Oriental genus, but found also in severalMascarene Islands and on <strong>the</strong> African continent. A species <strong>of</strong>black parrot {Coracopsistype {Foudia seychellarum)<strong>with</strong> Madagascar.<strong>The</strong>re are also two peculiar pigeons—a shortwingedTurtur and an Alcctrwnas.Most <strong>of</strong>Ofbarklayi) and a weaver bird <strong>of</strong> peculiarshow, however, a decided connection<strong>the</strong> birds <strong>of</strong> <strong>the</strong> Comoro Islands are Madagascar species,only two being African.Five are peculiar, belonging to <strong>the</strong>genera Nectarinia, Zosterops, Dicrurus, Foudia, and Alectrosnas.Reptiles are scarce. <strong>The</strong>re appear to be no snakes in Mauritiusand Bourbon, though some African species are said to befound in <strong>the</strong> Seychelle Islands. Lizards are fairly represented.Mauritiushas CryptoMepharus, an Australian genus <strong>of</strong> Gymnopthalmidse; Hemidactylus (a wide- spread genus) ; Percpus(Oriental and Australian)—both belonging to <strong>the</strong> Geckotidae.Bourbon has Heteropus, a Moluccan and Australian genus <strong>of</strong>Scincidae; Phelsuma (Geckotidae), and Chameleo, both found alsoin Madagascar ; as well as Pyxis, one <strong>of</strong> <strong>the</strong> tortoises. <strong>The</strong>Seychelles have <strong>The</strong>conyx, a peculiar genus <strong>of</strong> Geckotidae, andChameleo. Gigantic land-tortoises, which formerly inhabitedmost <strong>of</strong> <strong>the</strong> Mascarene Islands, now only survive inAldabra, asmall island north <strong>of</strong> <strong>the</strong> Seychelles. <strong>The</strong>se will be noticedagain fur<strong>the</strong>r on. Amphibia seem only to be recorded from <strong>the</strong>Seychelles, where two genera <strong>of</strong> tree-frogs <strong>of</strong> <strong>the</strong> family Polypedatidaeare found ; one {Megalixalus) peculiar, <strong>the</strong> o<strong>the</strong>r{Bappia) found also in Madagascar and Africa.Yol. I.—20


282 ZOOLOGICAL GEOGRAPHY. [part in.<strong>The</strong> few insect groups peculiar to <strong>the</strong>se islands will be notedwhen we deal <strong>with</strong> <strong>the</strong> entomology <strong>of</strong> Madagascar.Extinct fauna <strong>of</strong> <strong>the</strong> Mascarene Islands and Madagascar.—Beforequitting <strong>the</strong> vertebrate groups, we must notice <strong>the</strong> remarkablebirds which have become extinct in <strong>the</strong>se islands little morethan a century ago. <strong>The</strong> most celebrated is <strong>the</strong> dodo <strong>of</strong> <strong>the</strong>Mauritius (Didus ineptus), but an allied genus, Pezophaps, inhabitedRodriguez, and <strong>of</strong> both <strong>of</strong> <strong>the</strong>se almost perfect skeletonshave been recovered. O<strong>the</strong>r species probably existed in Bourbon.Remains <strong>of</strong> two genera <strong>of</strong> flightless rails have also been found,Aphanapteryx and Erythromachus ; and even a heron (Ardeamegacephala) which was short -winged and seldom flew ; whilein Madagascar <strong>the</strong>re lived a gigantic Struthious bird, <strong>the</strong>JEpyornis. Some fur<strong>the</strong>r details as to <strong>the</strong>se extinct forms willbe found under <strong>the</strong> respective families, Dididee, Rallidse, andiEpyornithidae, in <strong>the</strong> fourth part <strong>of</strong> this work ; and <strong>the</strong>ir bearingon <strong>the</strong> past history <strong>of</strong> <strong>the</strong> region will be adverted to in <strong>the</strong>latter part <strong>of</strong> this chapter. Dr. Gun<strong>the</strong>r has recently distinguishedfive species <strong>of</strong> fossil tortoises from Mauritius andRodriguez,—all <strong>of</strong> <strong>the</strong>m quite different from <strong>the</strong> living species<strong>of</strong> Aldabra.Insects.—<strong>The</strong> butterflies <strong>of</strong> Madagascar are not soremarkableas some o<strong>the</strong>r orders <strong>of</strong> insects. <strong>The</strong>re seems to be only onepeculiar genus, Heteropsis (Satyridse). <strong>The</strong> o<strong>the</strong>r genera areAfrican, leptoneura being confined to Madagascar and SouthAfrica. <strong>The</strong>re are some fine Papilios <strong>of</strong> uncommon forms. <strong>The</strong>most interesting lepidopterous insect, however, is <strong>the</strong> fine diurnalmoth {Urania), as all <strong>the</strong> o<strong>the</strong>r species <strong>of</strong> <strong>the</strong> genus inhabittropical America and <strong>the</strong> West Indian Islands.<strong>The</strong> Coleoptera have been better collected, and exhibit somevery remarkable affinities. <strong>The</strong>re is but one peculiar genus <strong>of</strong>Cicindelidse, Pogonostoma, which is allied to <strong>the</strong> South Americangenus, Ctenostoma. Ano<strong>the</strong>r genus, Peridexia, is common toMadagascar and South America. None <strong>of</strong> <strong>the</strong> important Africangenera are represented, except Eurymorpha ; while Meglaommais common to Madagascar and <strong>the</strong> Oriental region.In <strong>the</strong>Carabidre we have somewhat similar phenomena on a


chap, xi.] THE ETHIOPIAN REGION. 283wider scale. Such large and important African genera asPolyhirma and Anthia, are absent ; but <strong>the</strong>re are four generain common <strong>with</strong> South Africa, and two <strong>with</strong> West Africa ;whilethree o<strong>the</strong>rs are as much Oriental as African. One genus,Disirigus, is wholly Oriental ; and ano<strong>the</strong>r, Homalosoma, Australian.Colpodes, well developed in Bourbon and Mauritius, isOriental and South American.Of <strong>the</strong> peculiar genera, Sphccrostylishas South American affinities ; Microchila, Oriental ; <strong>the</strong>o<strong>the</strong>rs being related to widely distributed genera.<strong>The</strong> Lucanidse are few in number, and all haye African affinities.Madagascar is very rich in Cetoniidce, and possesses 20peculiar genera. Bothrorhina, and three o<strong>the</strong>r genera belongingto <strong>the</strong> Ichnostoma group, have wholly African <strong>relations</strong>. Doryscelisand Chromoptila are no less clearly allied to Orientalgenera. A series <strong>of</strong> eight peculiar genera belong to <strong>the</strong> Schizorhinidse,a family <strong>the</strong> bulk <strong>of</strong> which are Australian, while <strong>the</strong>re areonly a few African forms. <strong>The</strong> remaining genera appear to haveAfrican affinities, but few <strong>of</strong> <strong>the</strong> peculiarly African genera arerepresented.Glyciphana is characteristic <strong>of</strong> <strong>the</strong> Oriental region.<strong>The</strong> Buprestidse <strong>of</strong> Madagascar consist mainly <strong>of</strong> one largeand peculiar genus, Polybothris, alliedto <strong>the</strong> almost cosmopolitePsiloptera. Most <strong>of</strong> <strong>the</strong> o<strong>the</strong>r genera are both Ethiopian andOriental ; but Polycesta is mainly South American, and <strong>the</strong>remarkable and isolated genus Sponsor is confined to <strong>the</strong>Mauritius <strong>with</strong> a species in Celebes and New Guinea.<strong>The</strong> Longicorns are numerous and interesting, <strong>the</strong>re being noless than 24 peculiar genera. Two <strong>of</strong> <strong>the</strong> genera <strong>of</strong> Prionidseare very isolated, while a third, Closterus, belongs to a groupwhich is Malayan and American.Of <strong>the</strong> Cerambycidee, Philematium ranges to Africa and <strong>the</strong>West Indies ; Leptocera is only found eastward in Ceylon and<strong>the</strong> New Hebrides ;while Euporus is African. Of <strong>the</strong> peculiargenera, 2 are <strong>of</strong> African type ; 3 belong to <strong>the</strong> Leptura group,which are mostly Palaearctic and Oriental, <strong>with</strong> a few inSouth Africa ;while Philocalocera is allied to a South Americangenus.Among <strong>the</strong> Lamiidae <strong>the</strong>re are several wide-ranging and 7


:284 ZOOLOGICAL GEOGRAPHY. [part hi.African genera ; but Coptops is Oriental, and <strong>the</strong> OrientalPraonetha occurs in <strong>the</strong> Comoro Islands. Among <strong>the</strong> peculiargenera several have African affinities, but Tropidema belongs toa group which is Oriental and Australian ;Oopsis is found alsoin <strong>the</strong> Pacific Islands ;allied to Malayan and American genera.My<strong>the</strong>rgates, Sulemus, and Coedomcea, areGeneral Remarks on <strong>the</strong> Insect-fauna <strong>of</strong> Madagascar.—Taking<strong>the</strong> insects as a whole, we find <strong>the</strong> remarkable result that <strong>the</strong>iraffinities are largely Oriental, Australian, and South Americanwhile <strong>the</strong> African element is represented chiefly by specialSouth African or West African forms, ra<strong>the</strong>r than by such asare widely spread over <strong>the</strong> Ethiopian region. 1 In somefamilies—as Cetoniidse and Lamiidre—<strong>the</strong> African elementappears to preponderate ; in o<strong>the</strong>rs—as Cicindelidce—<strong>the</strong> SouthAmerican affinity seems strongest ; in Carabidse, perhaps <strong>the</strong>Oriental ; while in Buprestida3 and Cerambycidae <strong>the</strong> Africanand foreign elements seem nearly balanced. We must not imputetoo much importance to <strong>the</strong>se foreign alliances amonginsects, because we find examples <strong>of</strong> <strong>the</strong>m inevery country on<strong>the</strong> globe. <strong>The</strong> reason <strong>the</strong>y are so much more pronounced inMadagascar may be,that during long periods <strong>of</strong> time this islandhas served as a refuge for groups that have been dying out on<strong>the</strong> great continents ; and that, owing to <strong>the</strong> numerous deficiencies<strong>of</strong> a somewhat similar kind in <strong>the</strong> series<strong>of</strong> vertebratain Australia and South America, <strong>the</strong> same groups have <strong>of</strong>tenbeen able to maintain <strong>the</strong>mselves in all <strong>the</strong>se countries as wellas in Madagascar.It must be remembered too, that <strong>the</strong>se peculiaritiesin <strong>the</strong> Malagasy and Mascarene insect-fauna are but exaggerations<strong>of</strong> a like phenomenon on <strong>the</strong> mainland. Africa alsohas numerous affinities <strong>with</strong> South America, <strong>with</strong> <strong>the</strong> Malaycountries, and <strong>with</strong> Australia ;but <strong>the</strong>y do not bear anything likeso large a proportion to <strong>the</strong> whole fauna, and do not, <strong>the</strong>refore,attract so much attention. <strong>The</strong> special conditions <strong>of</strong> existenceand <strong>the</strong> long-continued isolation <strong>of</strong> Madagascar, will account formuch <strong>of</strong> this difference ;and it will evidently not be necessary1<strong>The</strong>re are also some special resemblances between <strong>the</strong> plants <strong>of</strong> Madagascarand South Africa, according to Dr. Kirk.


;chap, xi.] THE ETHIOPIAN REGION. 285to introduce, as some writers are disposed to do, a special landconnection or near approach between Madagascar and all <strong>the</strong>secountries, independently <strong>of</strong> Africa ; except perhaps in <strong>the</strong> case<strong>of</strong> <strong>the</strong> Malay Islands, as will be discussed fur<strong>the</strong>r on.Land-shells.-^ Madagascar and <strong>the</strong> adjacent islands are allrich in land-shells. <strong>The</strong> genera <strong>of</strong> Helicidas are Vitrina, Helix,Achatina, Columna (peculiar to Madagascar and West Africa),Buliminus, Cionella (chiefly Oriental and South American, butnot African), Pupa, Streptaxis, and Suecinea. Among <strong>the</strong> Operculatawe have Truncatella (widely scattered, but not African)Cyclotus (South American, Oriental, and South African) ;Cycloplwrus(mostly Oriental, <strong>with</strong> a few South African) ;Leptopoma(Oriental) ; Megalomastoma (Malayan and South American);Lithidion (peculiar to Madagascar, Socotra, and South-WestArabia) ; Otopoma (<strong>with</strong> <strong>the</strong> same range, but extending to WestIndia and New Ireland) ;Cyclostomus (widely spread but notAfrican) ;and Omphalotropis (wholly Oriental and Australian).We thus find <strong>the</strong> same general features reproduced in <strong>the</strong> landshellsas in <strong>the</strong> insects, and <strong>the</strong> same remarks will to a greatextent apply to both. <strong>The</strong> classification <strong>of</strong> <strong>the</strong> former is, however,by no means so satisfactory, and we have no extensiveand accurate general catalogues <strong>of</strong> shells, like those <strong>of</strong> Lepidopteraand Coleoptera, which have furnished us <strong>with</strong> suchvaluable materials for <strong>the</strong> comparison <strong>of</strong> <strong>the</strong> several faunas.On <strong>the</strong> probable Past History <strong>of</strong> <strong>the</strong>Ethiopian Region.Perhaps none <strong>of</strong> <strong>the</strong> great zoological regions <strong>of</strong> <strong>the</strong> earthpresent us <strong>with</strong> problems <strong>of</strong> greater difficulty or higherinterest than <strong>the</strong> Ethiopian. We find in it <strong>the</strong> evidence <strong>of</strong>several distinct and successive faunas, now intermingled ; and itis very difficult, <strong>with</strong> our present imperfect knowledge, to forman adequate conception <strong>of</strong> how and when <strong>the</strong> several changesoccurred. <strong>The</strong>re are, however, a few points which seem sufficientlyclear, and <strong>the</strong>se afford us a secure foundation in ourendeavour to comprehend <strong>the</strong> rest.Let us <strong>the</strong>n consider what are <strong>the</strong> main facts we have toaccount for.— 1. In Continental Africa, more especially in <strong>the</strong> south


286 ZOOLOGICAL GEOGRAPHY. [part. hi.and west, we find, along <strong>with</strong> much that is peculiar, a number <strong>of</strong>generashowing a decided Oriental, and o<strong>the</strong>rs <strong>with</strong> an equallystrong South American affinity ; this latter more particularly showingitself among reptiles and insects. 2. All over Africa, but moreespecially in <strong>the</strong> east, we have abundance <strong>of</strong> large ungulates andfelines—antelopes, giraffes, buffaloes, elephants, and rhinoceroses,<strong>with</strong> lions, leopards, and hysenas, all <strong>of</strong> types now or recentlyfound in India and "Western Asia. 3. But we also have to note<strong>the</strong> absence <strong>of</strong> a number <strong>of</strong> groups which abound in <strong>the</strong> abovenamedcountries, such as deer, bears, moles, and true pigs ; whilecamels and goats—characteristic <strong>of</strong> <strong>the</strong> desert regions just to<strong>the</strong> north <strong>of</strong> <strong>the</strong> Ethiopian—are equally wanting. 4. <strong>The</strong>re isa wonderful unity <strong>of</strong> type and want <strong>of</strong> speciality in <strong>the</strong> vastarea <strong>of</strong>our first sub-region extending from Senegal across to <strong>the</strong>east coast, and southward to <strong>the</strong> Zambezi; while West Africa andSouth Africa each abound in peculiar types. 5. We have <strong>the</strong>extraordinary fauna <strong>of</strong> Madagascar to account for, <strong>with</strong> itsevidentmain derivation from Africa, yet wanting all <strong>the</strong> largerand higher African forms ; its resemblances to Malaya and toSouth America ; and its wonderful assemblage <strong>of</strong> altoge<strong>the</strong>rpeculiar types.Here we find a secure starting-point, for we are sure thatMadagascar must have been separated from Africa before <strong>the</strong>assemblage <strong>of</strong> large <strong>animals</strong> enumerated above, had enteredit. Now, it is a suggestive fact, that all <strong>the</strong>se belong to typeswhich abounded in Europe and India about <strong>the</strong> Miocene period.It is also known, from <strong>the</strong> prevalence <strong>of</strong> Tertiary deposits over<strong>the</strong> Sahara and much <strong>of</strong> Arabia, Persia, and Nor<strong>the</strong>rn India,that during early Tertiary times a continuous sea from <strong>the</strong> Bay<strong>of</strong> Bengal to <strong>the</strong> British Isles completely cut <strong>of</strong>f all land communicationbetween Central and Sou<strong>the</strong>rn Africa on <strong>the</strong> oneside, and <strong>the</strong> great continent <strong>of</strong> <strong>the</strong> Eastern hemisphere on <strong>the</strong>o<strong>the</strong>r.When Africa was thus isolated, its fauna probably had acharacter somewhat analogous to that <strong>of</strong> South America at <strong>the</strong>same period.Most <strong>of</strong> <strong>the</strong> higher types <strong>of</strong> mammalian life wereabsent, while lemurs, Edentates, and Insectivora took <strong>the</strong>ir place.At this period Madagascar was no doubt united <strong>with</strong> Africa,


chap, xi.] THE ETHIOPIAN REGION. 287and helped to form a great sou<strong>the</strong>rn continent which must at onetime have extended eastward as far as Sou<strong>the</strong>rn India andCeylon ; and over <strong>the</strong> whole <strong>of</strong> this <strong>the</strong> lemurine type no doubtprevailed.During some portion <strong>of</strong> this period, South Temperate Africamust have had a much greater extension, perhaps indicated by<strong>the</strong> numerous shoals and rocks to <strong>the</strong> south and east <strong>of</strong> <strong>the</strong>Cape <strong>of</strong> Good Hope, and by <strong>the</strong> Crozets and Kerguelen Islandsfur<strong>the</strong>r to <strong>the</strong> south-east.This would have afforded means forthat intercommunion <strong>with</strong> Western Australia which is so clearlymarked in <strong>the</strong> flora, and to some extent also in <strong>the</strong> insects <strong>of</strong> <strong>the</strong>two countries ; and some such extension is absolutely requiredfor <strong>the</strong> development <strong>of</strong> that wonderfully rich and peculiartemperate flora and fauna, which, now crowded into a narrowterritory, is one <strong>of</strong> <strong>the</strong> greatest marvels <strong>of</strong> <strong>the</strong> organic world.During this early period, when <strong>the</strong> great sou<strong>the</strong>rn continents—South America, Africa, and Australia—were equally free from<strong>the</strong> incursions <strong>of</strong> <strong>the</strong> destructive felines <strong>of</strong> <strong>the</strong> north, <strong>the</strong>Struthious or ostrich type <strong>of</strong> birds was probably developed intoits existing forms. It is not at all necessary to suppose that<strong>the</strong>se three continents were at any time united, in order toaccount for <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong>se great terrestrial birds ; asthis may have arisen by at least two o<strong>the</strong>r easily conceivablemodes. <strong>The</strong> ancestral Struthious type may, like <strong>the</strong>Marsupial, have once spread over <strong>the</strong> larger portion <strong>of</strong> <strong>the</strong> globe ;but as higher forms,especially <strong>of</strong> Carnivora, became developed,it would be exterminated everywhere but in those regionswdiere it was free from <strong>the</strong>ir attacks.In each <strong>of</strong> <strong>the</strong>se it woulddevelope into special forms adapted to surrounding conditions; and<strong>the</strong> large size, great strength, and excessive speed <strong>of</strong> <strong>the</strong> ostrich,may have been a comparatively late development caused by itsexposure to <strong>the</strong> attacks <strong>of</strong> enemies which rendered such modificationnecessary. This seems <strong>the</strong> most probable explanation<strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> Struthious birds, and itis rendered almostcertain by <strong>the</strong> discovery <strong>of</strong> remains <strong>of</strong> this order in Europe inEocene deposits, and by <strong>the</strong> occurrence <strong>of</strong> an ostrich among <strong>the</strong>fossils <strong>of</strong> <strong>the</strong> Siwalik hills ;but it is just possible, also, that <strong>the</strong>


288 ZOOLOGICAL GEOGRAPHY. [part m.ancestral type may have been a bird capable <strong>of</strong> flight, and thatit spread from one <strong>of</strong>* <strong>the</strong> three sou<strong>the</strong>rn continents to <strong>the</strong> o<strong>the</strong>rsat <strong>the</strong> period <strong>of</strong> <strong>the</strong>ir near approach, and more or less completelylost <strong>the</strong> power <strong>of</strong> flight owing to <strong>the</strong> long continued absence<strong>of</strong> enemies.During <strong>the</strong> period we have been considering, <strong>the</strong> ancestors <strong>of</strong>existing apes and monkeys flourished (as we have seen inChapter VI.) along <strong>the</strong> whole sou<strong>the</strong>rn shores <strong>of</strong> <strong>the</strong> old Palaearcticcontinent ; and it seems likely that <strong>the</strong>y first enteredAfrica by means <strong>of</strong> a land connection indicated by <strong>the</strong> extensiveand l<strong>of</strong>ty plateaus <strong>of</strong> <strong>the</strong> Sahara, situated to <strong>the</strong> south-east <strong>of</strong> Tunisand reaching to a little north-west <strong>of</strong> Lake Tchad ;and at <strong>the</strong> sametime <strong>the</strong> elephant and rhinoceros type may have entered. Thiswill account for <strong>the</strong> curious similarity between <strong>the</strong> higher faimas<strong>of</strong> West Africa and <strong>the</strong> Indo-Malay sub-region, forowing to <strong>the</strong>present <strong>distribution</strong> <strong>of</strong> land and sea and <strong>the</strong> narrowing <strong>of</strong> <strong>the</strong>tropical zone since Miocene times, <strong>the</strong>se are now <strong>the</strong> onlylowland, equatorial, forest-clad countries, which were inconnection<strong>with</strong> <strong>the</strong> sou<strong>the</strong>rn shores <strong>of</strong> <strong>the</strong> old Palsearcticcontinent at<strong>the</strong> time <strong>of</strong> its greatest luxuriance and development. Thiswestern connection did not probably last long, <strong>the</strong> junction thatled to <strong>the</strong> greatest incursion <strong>of</strong> new forms, and <strong>the</strong> completechange in <strong>the</strong> character <strong>of</strong> <strong>the</strong> Africanbeen effected by way <strong>of</strong>fauna, having apparentlySyria and <strong>the</strong> shores <strong>of</strong> <strong>the</strong> Red Sea ata somewhat later date. By this route <strong>the</strong> old South-Palsearcticfauna, indicated by <strong>the</strong> fossils <strong>of</strong> Pikermi and <strong>the</strong> Siwalik Hills,poured into Africa; and finding <strong>the</strong>re a new and favourablecountry, almost wholly unoccupied by large Mammalia, increasedto an enormous extent, developed into new forms, and finallyoverran <strong>the</strong> whole continent.Before this occurred, however, a great change had taken placein <strong>the</strong> geography <strong>of</strong> Africa.It had gradually diminished on <strong>the</strong>south and east ; Madagascar had been left isolated ; while anumber <strong>of</strong> small islands, banks, and coral reefs in <strong>the</strong> IndianOcean alone remained to indicate <strong>the</strong> position <strong>of</strong> a once extensiveequatorial land. <strong>The</strong> Mascarene Islands appear to represent<strong>the</strong> portion which separated earliest, before any carnivora had


chap, xi.] THE ETHIOPIAN REGION. 289reached <strong>the</strong> country ; and it was in consequence <strong>of</strong> this totalexemption from danger, that severalgroups <strong>of</strong> birds altoge<strong>the</strong>rincapable <strong>of</strong> flight became developed here, culminating in <strong>the</strong>huge and unwieldy Dodo, and <strong>the</strong> more active Aphanapteryx.To <strong>the</strong> same cause may be attributed <strong>the</strong> development, in <strong>the</strong>seislands, <strong>of</strong> gigantic land-tortoises, far surpassing any o<strong>the</strong>rs nowliving on <strong>the</strong> globe. <strong>The</strong>y appear to have formerly inhabitedMauritius, Bourbon, and Bodriguez, and perhaps all <strong>the</strong>o<strong>the</strong>r Mascarene islands, but having been recklessly destroyed,now only survive in <strong>the</strong> small uninhabited Aldabra islandsnorth <strong>of</strong> <strong>the</strong> Seychelle group. <strong>The</strong> largest living specimen(5^ feet long) is now in our Zoological Gardens. <strong>The</strong> onlyo<strong>the</strong>r place where equally large tortoises (<strong>of</strong> an allied species)are found, is <strong>the</strong> Galapagos islands, where <strong>the</strong>y were equally freefrom enemies till civilized man came upon <strong>the</strong> scene ;who, partlyby using <strong>the</strong>m for food, partly by <strong>the</strong> introduction <strong>of</strong> pigs, whichdestroy <strong>the</strong> eggs, has greatly diminished <strong>the</strong>ir numbers andsize, and will probably soon wholly exterminate <strong>the</strong>m. It is acurious fact, ascertained by Dr. Giin<strong>the</strong>r, that <strong>the</strong> tortoises <strong>of</strong><strong>the</strong> Galapagos are morenearly related to <strong>the</strong> extinct tortoises <strong>of</strong>Mauritius than is <strong>the</strong> living tortoise <strong>of</strong> Aldabra. This wouldimply that several distinct groups or sub-genera <strong>of</strong>had a wide range over <strong>the</strong> globe, and that some <strong>of</strong>Testudo haveeach havesurvived in very distant localities. This is rendered quite conceivableby <strong>the</strong> known antiquity <strong>of</strong> <strong>the</strong> genus Testudo, whichdates back to at least <strong>the</strong> Eocene formation (in North America)<strong>with</strong> very little change <strong>of</strong> form. <strong>The</strong>se sluggish reptiles, solong-lived and so tenacious <strong>of</strong> life, may have remained unchanged,while every higher animal type around <strong>the</strong>m hasbecome extinct and been replaced by very different forms ; asin <strong>the</strong> case <strong>of</strong> <strong>the</strong> living Emys tectum, which is <strong>the</strong> sole survivor<strong>of</strong> <strong>the</strong> strange Siwalik fauna <strong>of</strong> <strong>the</strong> Miocene epoch. <strong>The</strong> ascertainedhistory <strong>of</strong> <strong>the</strong> genus and <strong>the</strong> group, thus affords a satisfactoryexplanation <strong>of</strong> <strong>the</strong> close affinity <strong>of</strong> <strong>the</strong> gigantic tortoises<strong>of</strong> Mauritius and <strong>the</strong> Galapagos.<strong>The</strong> great island <strong>of</strong> Madagascar seems to have remained longerunited <strong>with</strong> Africa, till some <strong>of</strong> <strong>the</strong> smaller and more active


290 ZOOLOGICAL GEOGRAPHY. [part in.carnivora had reached it; and we consequently find <strong>the</strong>re, nowholly terrestrial form <strong>of</strong> bird but <strong>the</strong> gigantic and powerfulJEpijomis, well able to defend itself against such enemies. Asalready intimated, we refer <strong>the</strong> South American element inMadagascar, not to any special connection <strong>of</strong> <strong>the</strong> two countriesindependently <strong>of</strong> Africa, but to <strong>the</strong> preservation <strong>the</strong>re <strong>of</strong> anumber <strong>of</strong> forms, some derived from America through Africa,o<strong>the</strong>rs <strong>of</strong> once almost cosmopolitan range, but which, owing to <strong>the</strong>severer competition, have become extinct on <strong>the</strong> African continent,while <strong>the</strong>y have continued to exist under modifiedformsin <strong>the</strong> two o<strong>the</strong>r countries.<strong>The</strong> depths <strong>of</strong> all <strong>the</strong> great oceans are now known to be sopr<strong>of</strong>ound, that we cannot conceive <strong>the</strong> elevation <strong>of</strong> <strong>the</strong>ir bedsabove <strong>the</strong> surface<strong>with</strong>out some corresponding depression elsewhere.And if, as is probable, <strong>the</strong>se opposite motions <strong>of</strong> <strong>the</strong>earth's crust usually take place in parallel bands, and are tosome extent dependent on each o<strong>the</strong>r, an elevation <strong>of</strong> <strong>the</strong> seabed could hardly fail to lead to <strong>the</strong>submergence <strong>of</strong> large tracts<strong>of</strong> existing continents ; and this is <strong>the</strong> more likely to occur onaccount <strong>of</strong> <strong>the</strong> great disproportion that we have seen existsbetween <strong>the</strong> mean height <strong>of</strong> <strong>the</strong> land and <strong>the</strong> mean depth <strong>of</strong> <strong>the</strong>ocean. Keeping this principle in view, we may, <strong>with</strong> someprobability, suggest <strong>the</strong> successive stages by which <strong>the</strong> Ethiopianregion assumed its present form, and acquired <strong>the</strong> strikingpeculiarities that characterise its several sub-regions. During<strong>the</strong> early period, when <strong>the</strong> rich and varied temperate flora <strong>of</strong> <strong>the</strong>Cape, and its hardly less peculiar forms <strong>of</strong> insects and <strong>of</strong> low typemammalia, were in process <strong>of</strong> development in an extensivesouth temperate land, we may be pretty sure that<strong>the</strong> east and much <strong>of</strong> <strong>the</strong> north <strong>of</strong> Africa was deep sea.<strong>the</strong> whole <strong>of</strong>At alater period, when this continent sank towards <strong>the</strong> south andeast, <strong>the</strong> elevation may have occurred which connected Madagascar<strong>with</strong> Ceylon ; and only at a still later epoch, when <strong>the</strong>Indian Ocean had again been formed, did central, eastern, andnor<strong>the</strong>rn Africa gradually rise above <strong>the</strong> ocean, and effect aconnection <strong>with</strong> <strong>the</strong> great nor<strong>the</strong>rn continent by way <strong>of</strong> Abyssiniaand Arabia. And if this last change took place <strong>with</strong>


chap, xi.] THE ETHIOPIAN REGION. 291tolerable rapidity, or if<strong>the</strong> elevatory force acted from <strong>the</strong> northtowards <strong>the</strong> south, <strong>the</strong>re would be a new and unoccupiedterritory to be taken possession <strong>of</strong> by immigrants from <strong>the</strong>north, toge<strong>the</strong>r <strong>with</strong> a few from <strong>the</strong> south and west. <strong>The</strong> morehighly -organized types from <strong>the</strong> great nor<strong>the</strong>rn continent, however,would inevitably prevail ; and we should thus haveexplained <strong>the</strong> curious uniformity in <strong>the</strong> fauna <strong>of</strong> so large anarea, toge<strong>the</strong>r <strong>with</strong> <strong>the</strong> absence from it <strong>of</strong> those peculiarEthiopian types which so abundantly characterise <strong>the</strong> o<strong>the</strong>rthree sub-regions.We may now perhaps see <strong>the</strong> reason <strong>of</strong> <strong>the</strong> singular absence fromtropical Africa <strong>of</strong> deer and bears ; for <strong>the</strong>se are both groupswhich live in fertile orwell-wooded countries, whereas <strong>the</strong> line<strong>of</strong> immigration from Europe to Africa was probably always, asnow, to a great extent a dry and desert tract, suited to antelopesand large felines, but almost impassable to deer and bears.find, too, that whereas remains <strong>of</strong> antelopes and giraffesWeaboundin <strong>the</strong> Miocene deposits <strong>of</strong> Greece, <strong>the</strong>re were no deer (whichare perhaps a somewhat later development) ; nei<strong>the</strong>r were <strong>the</strong>reany bears, but numerous forms <strong>of</strong> Eelidse, Viverridge, Mustelidse,and ancestral forms <strong>of</strong> Hycona, exactly suited to be <strong>the</strong>progenitors <strong>of</strong> <strong>the</strong> most prevalent types <strong>of</strong> modern AfricanZoology.<strong>The</strong>re appears to have been one o<strong>the</strong>r change in <strong>the</strong> geography<strong>of</strong> Africa and <strong>the</strong> Atlantic Ocean that requires notice.<strong>The</strong> ra<strong>the</strong>r numerous cases <strong>of</strong> close similarity in <strong>the</strong> insectforms <strong>of</strong> tropical Africa and America, seem to indicate somebetter means <strong>of</strong> transmission, at a not very remote epoch, thannow exists. <strong>The</strong> vast depth <strong>of</strong> <strong>the</strong> Atlantic, and <strong>the</strong> absence <strong>of</strong>any corresponding likeness in <strong>the</strong> vertebrate fauna, entirelynegative <strong>the</strong> idea <strong>of</strong> any union between <strong>the</strong> two countries ;but a moderate extension <strong>of</strong> <strong>the</strong>irshores towards each o<strong>the</strong>r isnot improbable, and this, <strong>with</strong> large islands in <strong>the</strong> place <strong>of</strong> <strong>the</strong>Cape Verd group, St. Paul's Eocks, and Fernando Noronha, toafford resting places in <strong>the</strong> Atlantic, wrould probably suffice toexplain <strong>the</strong> amount <strong>of</strong> similarity that actually exists.Our knowledge <strong>of</strong> <strong>the</strong> geology and palaeontology <strong>of</strong> Africa


292 ZOOLOGICAL GEOGRAPHY. [part hi.being so scanty, it would be imprudent to attempt any moredetailed explanation <strong>of</strong> <strong>the</strong> peculiarities <strong>of</strong> its existing fauna.<strong>The</strong> sketch now given is, it is believed, founded on a sufficientbasis <strong>of</strong> facts to render it not only a possible but a probableaccount <strong>of</strong> what took place ;and it is something gained to beable to show, that a large portion <strong>of</strong> <strong>the</strong> peculiarities andanomalies <strong>of</strong> so remarkable a fauna as that <strong>of</strong> <strong>the</strong> Ethiopianregion, can be accounted for by a series <strong>of</strong> changes <strong>of</strong> physicalgeography during <strong>the</strong> tertiary epoch, which can hardly be consideredextreme, or in any way unlikely to have occurred.


—;chap, xi.] THE ETHIOPIAN REGION. 293TABLES OF DISTRIBUTION.In drawing up <strong>the</strong>se tables showing <strong>the</strong> <strong>distribution</strong> <strong>of</strong> variousclasses <strong>of</strong> <strong>animals</strong> in <strong>the</strong> Ethiopian Region, <strong>the</strong> following sources<strong>of</strong> information have been chiefly relied on, in addition to <strong>the</strong>general treatises, monographs, and catalogues, used for <strong>the</strong> FourthPart <strong>of</strong> this work:Mammalia.—Blanford's Abyssinia ;Peters's Mozambique;Heuglin and Schweinfurth for North East Africa ; GrandidierSchlegel, &c, for Madagascar; <strong>the</strong> local lists given by Mr.Andrew Murray ; numerous papers by Fraser, Gray, Kirk,Mivart, Peters, Sclater, and Speke; and a MS. list <strong>of</strong> Bovida?from Sir Victor Brooke.Birds.—Finsch and Hartlaub for East Africa; Heuglin forNorth-East Africa ; Blanford for Abyssinia ; Layard for SouthAfrica ; Hartlaub for West Africa ; Dohrn for Princes IslandAndersson for Damaraland;and papers by Gurney, Hartlaub,Kirk, Newton, Peters, Sharpe, Sclater, Schlegel, and Pollen ;and a MS. list <strong>of</strong> Madagascar Birds from Mr. Sharpe.


.294 ZOOLOGICAL GEOGRAPHY. [PART I'TABLE I.FAMILIES OF ANIMALS INHABITING THE ETHIOPIAN REGION.Explanation.Names in italics show families peculiar to <strong>the</strong> region.Names inclosed thus ( ) barely enter <strong>the</strong> region, and are not considered properlyto belong to it.Numbers are not consecutive, but correspond to those in Part IV.Sub-regions.Order and Family.Hi:Kange beyond <strong>the</strong> Region.MAMMALIA.Primates.1. Simiidae2. Semnopi<strong>the</strong>cidse3. Cynopi<strong>the</strong>cidae6. Lemuridae8. Chiromyidce ...Cheiroptera.9. Pteropidae11. Rhinolophidae12. Vespertilionidae13. Noctilionidae . .Insectivora.15. Macroscelididae17. Erinaceidae ...18. Centetidce19. Potairwgalidce20. Chrysochloridce22. SoricidaeCarnivora.23. Felidae24. Cryptoproclidce25. Viverridae26. Protelidce27. Hyamidae28. Canida? ...29. Mustelidae33. Otariidae...Cetacea.36 to 41.SlRENIA.42. ManatidaeUngulata.43. Equidae ...,OrientalOrientalOriental,OrientalPalaearcticOriental, Australian<strong>The</strong> Eastern HemisphereCosmopoliteAll Tropical regionsSouth PalaearcticPalaearctic, OrientalGreater AntillesAll regions but Australian and NeotropicalAll regions but AustralianOriental, S. PalaearcticS. Palaearctic, IndiaAlmost cosmopoliteAll regions but AustralianAll temperate regionsOceanicNeotropical, Oriental, AustralianPalaearctic


CHAP. XI.] THE ETHIOPIAN EEGION. 295


.296 ZOOLOGICAL GEOGRAPHY. [PART IIISub-regions.Order and Family.38. Motacillidse ...47. Pittidse48. PaictidceH« (1CO —< *5 «3


CHAP. XI.] THE ETHIOPIAN REGION. 297


,298 ZOOLOGICAL GEOGRAPHY. [PART 111.Sub-regions.Order and Family.S-g


CHAP. XI.] THE ETHIOPIAN REGION. 299


300 ZOOLOGICAL GEOGRAPHY. [PART III.TABLELIST OF GENERA OF TERRESTRIAL MAMMALIA AND BIRDSINHABITING TEE ETHIOPIAN REGION.II.Explanation.Names in italics show genera peculiar to <strong>the</strong> region.Names inclosed thus (...)show genera which just enter <strong>the</strong> region, but are not consideredproperly to belong to it.Genera which undoubtedly belong to <strong>the</strong> region are numbered consecutively.MAMMALIA.Order, Family, andGenus.


CHAP. XI.] THE ETHIOPIAN REGION. 301Order, Family, andGenus.


1302 ZOOLOGICAL GEOGRAPHY. [part III.jOrder, Family, andGenus.*•hr


CHAP. XI.j THE ETHIOPIAN REGION. 303Order, Family, andGenus.


304 ZOOLOGICAL GEOGRAPHY. [PART 111.Order, Family, andGenus.


CHAP. XI.] THE ETHIOPIAN KEGION. 305Order, Family, andGenus.


306 ZOOLOGICAL GEOGRAPHY. [part III.Order, Family, andGenus.© a>Range <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.Timaliid^:.33. Chatarrhsea34. Crateropus35. Hypergerus36. Cichlctdusa37. Alc<strong>the</strong>38. Oxylabcs1171342AbyssiniaAll AfricaW. AfricaW. and E. AfricaW. AfricaMadagascarOriental, PalestineN. Africa, PersiaClNCLID.®. [?]39. MesitesMadagascarT ROGLODYTID.E . ['.


CHAP. XI.] THE ETHIOPIAN KEGION. 307


308 ZOOLOGICAL GEOGRAPHY. [part III.


CHAP. XI.] THE ETHIOPIAN REGION. 309Order, Family, andGenus.Range <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.154. Falculia155. Fregilupus ...MadagascarBourbonAlaudid>e.156. Alauda157. Spizocorys158. Galerida159. Calendula(Melanocorypha160. Certhilauda ...161. Alaemon162. Heterocorys ...163. Mirafra164. Ammomanes ..165. Mcgalophonus166. Tephrocorys ...167. Pyrrhulauda ..31421331104526Abyssinia and S. W. AfricaSouth AfricaNorth <strong>of</strong> tropical AfricaAbyssinia, S. AfricaAbyssinia)South AfricaSouth AfricaSouth AfricaSouth Africa, MadagascarAfrican desertsTropical and S. AfricaS. AfricaTropical and S. AfricaPalaearctic,Palaearctic,IndianIndianPalaearctic genusS. EuropeS. PalaearcticOriental, AustralianS. Palaearctic, IndianOriental, Canary IslandsMotacillim:.168. Budytes169. Anthus170. Macronyx104<strong>The</strong> whole regionTropical and S. AfricaTropical and S. AfricaPalaearctic, Oriental,AustralianAll regions, exc. AustraliaPITTIDjE.171. PittaW. AfricaOriental, AustralianPAICTID.E.172. Philepitta ...MadagascarPICARIJE.PICID.E.173. Verreauxia ...174. Dendropicus ...175. Campe<strong>the</strong>ra ...176. Geocolaptes ...114141W. AfricaTropical and S. AfricaTropical and S. AfricaSouth AfricaYxtngid^:.177. YunxN. E. Africa, S. AfricaPalaearcticiNDICATORICffi.178. IndicatorTropical and S. AfricaOrientalMEGAL.EMID.E.179. Pogonorhynchus180. Buccanodon ...181. Stadolwma ...182. Barbatula183. Xylobucco ...184. Gymnobitcco ...185. Trachyphonus14119336Tropical and S. AfricaWest AfricaWest AfricaWest and South AfricaWest and South AfricaWest AfricaTropical and South AfricaMusophagid,*:.186. Musophaga ...West Africa


.310 ZOOLOGICAL GEOGRAPHY. [PART 111.Order, Family, andGenus.Range <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.187. Turacus188. Schizorhis ...106Tropical and S. AfricaTropical and S. AfricaCoLIIDjE.189. ColiusTropical and S.AfricaCuCULIDiE.190. Ceuthmochares191. Coua192. Cochlothraustes193. Centropus194. Cuculus195. Chrysococcyx196. Coccystes29181076Africa and MadagascarMadagascarMadagascarAfrica and MadagascarAfrica and MadagascarTropical and S. AfricaTropical and S. AfricaOriental, AustralianPalsearc, Orien., Austral.Oriental, AustralianS. Palsearctic, OrientalLEPTOSOMIDiE.197. Leptosomus ..MadagascarCORACIIDJE.198. Coracias199. Eurystomus ...200. Atelomis201. Brachypteracias202. GeobiastesAfrica and MadagascarAfrica and MadagascarMadagascarMadagascarMadagascarS. Palsearctic, OrientalOriental, AustralianMeropid^;.203. Merops204. Melittophagus115Africa and MadagascarTropical and S. AfricaS.Palaear., Orien., Austral.TROGONID.E.205. Apaloderma ...Tropical and S.AfricaALCEDINID.E.206. Alcedo207. Corythornis ...208. Ceryle209. Myioceyx210. Ispidina. . ...211. Halcyon2 W. Africa, Abyssinia, Natal3 Africa and Madagascar1 W. Africa, Abyssinia, Natal2 West Africa4 Africa and Madagascarin Africa,Prince's Is., St. ThomePalsear., Orien., Austral.American, PalsearcticS.Palsear., Orien. Austral.Bucerotiixe.212. Berenicornis . .213. Tockus214. Bycanistes215. Bucorvus112West AfricaTropical and S.Tropical and S.Tropical and S.AfricaAfricaAfricaMalayaUpupid^!.216. UpupaAfrica and MadagascarS. Palsearctic, OrientalIrrisorid^e.217. lrrisor12Africa and Madagascar


CHAP. XI.] THE ETHIOPIAN REGION. 311Order, Family, andGenus.


312 ZOOLOGICAL GEOGEAPHY. [PART III.Order, Family, andGenus.


CHAP. XI.] THE ETHIOPIAN REGION. 313Order, Family, andGenus.6 9 Range <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.PANDIONIDiE.287. Pandion...Strigid-e.All AfricaCosmopolite288. A<strong>the</strong>ne ...289. Bubo ...290. Scotopelia291. Scops ...292. Syrnium293. Asio294. Strix ...Africa and Madagascar, Rodriquez(extinct)Africa and MadagascarWest and S. Africa to ZambesiW. and S. Africa, Madagascar,Comoro IslandsAfricaN. E. and S. AfricaAfrica and MadagascarPalsearctic,AustralianCosmopoliteAlmost CosmopolitePalcearctic,AmericanCosmopoliteCosmopoliteOriental,Oriental,Peculiar or very Characteristic Genera <strong>of</strong> Wading or Swimming Birds.GRALL^E.


CHAPTEE XII.THE ORIENTAL REGION.This region is<strong>of</strong> comparatively small extent, but it has a verydiversified surface, and is proportionately very rich. <strong>The</strong> desertson <strong>the</strong> north-west <strong>of</strong> India are <strong>the</strong> debatable land thatseparates it from <strong>the</strong> Palsearctic and Ethiopian regions. <strong>The</strong>great triangular plateau which forms<strong>the</strong> peninsula <strong>of</strong> India is<strong>the</strong> poorest portion <strong>of</strong> <strong>the</strong> region, owing in part to its arid climateand in part to its isolated position ;for <strong>the</strong>re can be little doubtthat in <strong>the</strong> later Tertiary period it was an island, separated by anarm <strong>of</strong> <strong>the</strong> sea (now forming <strong>the</strong> valleys <strong>of</strong> <strong>the</strong> Ganges andIndus) from <strong>the</strong> luxuriantHimalayan and Burmese countries.Its sou<strong>the</strong>rn extremity, <strong>with</strong> Ceylon, has a moister climate andmore luxuriant vegetation, and exhibits indications <strong>of</strong> a formerextension southwards, <strong>with</strong> a richer and more peculiar fauna,partly Malayan and partly Mascarene in its character. <strong>The</strong>whole sou<strong>the</strong>rn slopes <strong>of</strong> <strong>the</strong> Himalayas, <strong>with</strong> Burmah, Siam andWestern China, as well as <strong>the</strong> Malay peninsula and <strong>the</strong> Indo-Malay islands, are almost everywhere covered <strong>with</strong> tropicalforests <strong>of</strong> <strong>the</strong> most luxuriant character, which abound in variedand peculiar forms <strong>of</strong> vegetable and animal life.<strong>The</strong> flora andfauna <strong>of</strong> this extensive district are essentially <strong>of</strong> one typethroughout; yet it may be usefully divided into <strong>the</strong> Indo-Chinese and <strong>the</strong> Malayan sub-regions, as each possesses anumber <strong>of</strong> peculiar or characteristic <strong>animals</strong>. <strong>The</strong> former subregion,besides having many tropical and sub-tropical types<strong>of</strong> its own, also possesses a large number <strong>of</strong> peculiarly modifiedtemperate forms on <strong>the</strong> mountain ranges <strong>of</strong> its nor<strong>the</strong>rn


10-«8JOMiS


;chap, xii.] THE ORIENTAL REGION. 315boundary, which are wholly wanting in <strong>the</strong> Malayan subregion.<strong>The</strong> Philippine islands are best classed <strong>with</strong> <strong>the</strong> Indo-Malay group, although <strong>the</strong>y are strikingly deficient in manyMalayan types, and exhibit an approach to <strong>the</strong> Celebesian division<strong>of</strong> <strong>the</strong> Austro-Malay sub-region.Zoological Characteristics <strong>of</strong> <strong>the</strong> Oriental Region.—<strong>The</strong> OrientalEegion possesses examples <strong>of</strong> 35 families <strong>of</strong> Mammalia, 71 <strong>of</strong>Birds, 35 <strong>of</strong> Eep tiles, 9 <strong>of</strong> Amphibia, and 13 <strong>of</strong> Fresh-waterFishes. Of <strong>the</strong>se 163 families, 12 are peculiar to <strong>the</strong> region;namely, Tarsiidae, Galeopi<strong>the</strong>cidse, and Tupaiidse among Mammalia,while iEluridse, though confined to <strong>the</strong> higher Himalayas,may perhaps <strong>with</strong> more justice be claimed by <strong>the</strong> Palaearctic region;Liotrichidse, Phyllornithidae, and Eurylaemidae among birdsXenopeltidse (extending, however, to Celebes), Uropeltidae, andAcrochordidse among reptiles ;Luciocephalidse, Ophiocephalidseand Mastacembelidae among fresh-water fishes.A number <strong>of</strong>ando<strong>the</strong>r families are abundant, and characteristic <strong>of</strong> <strong>the</strong> region ;it possesses many peculiar and characteristic genera, which mustbe referred to somewhat more in detail.Mammalia.—<strong>The</strong> Oriental region is richin quadrumana, andis especially remarkable for its orang-utans and long-armed apes(Simia, Hylohates, and Siamanga) ; its abundance <strong>of</strong> monkeys<strong>of</strong> <strong>the</strong> genera Presbytes and Macacus ; its extraordinary long-nosedmonkey (Presbytes nasalis) ;its Lemuridse {Nycticebus and Loris);and its curious genus Tarsius, forming a distinct family <strong>of</strong>lemurs. All <strong>the</strong>se quadrumanous genera are confined to it,except Tarsius which extends as far as Celebes. It possessesmore than 30 genera <strong>of</strong> bats, which are enumerated in<strong>the</strong> lists given at <strong>the</strong> end <strong>of</strong> this chapter. In Insectivora it isvery rich, and possesses several remarkable forms, such as <strong>the</strong>flying lemur (Galcopi<strong>the</strong>cus) ; <strong>the</strong> squirrel-like TupaiidsB consisting<strong>of</strong> three genera ; and <strong>the</strong> curious Gymnura allied to <strong>the</strong> hedgehogs.In Carnivora, it is especially rich in many forms <strong>of</strong> civets(Viverridse), possessing 10 peculiar genera, among which Prionodonand Cynogale are remarkable; numerous Mustelidae, <strong>of</strong>which Gymnopus, Mydaus, Aonyx and Helictis are <strong>the</strong> most conspicuous;JElurus, a curious animal, cat-like in appearance but


316 ZOOLOGICAL GEOGRAPHY. [part hi.more allied to <strong>the</strong> bears, forming a distinctfamily <strong>of</strong> Carnivora,and confined to <strong>the</strong> high forest-districts <strong>of</strong> <strong>the</strong> Eastern Himalayasand East Thibet ;Melursus and Helardos, peculiar forms <strong>of</strong>bears ;,Platanista, a dolphin peculiar to <strong>the</strong> Ganges and Indus.Among Euminants it has <strong>the</strong> beautiful chevrotain, forming<strong>the</strong> genus Tragulus in <strong>the</strong> family Tragulidse ; <strong>with</strong> one peculiargenus and three peculiar sub-genera <strong>of</strong> true deer. <strong>The</strong> Antilopinreand Caprine are few, confined to limited districts and notcharacteristic <strong>of</strong> <strong>the</strong> region ;but <strong>the</strong>re are everywhere wild cattle<strong>of</strong> <strong>the</strong> genera Bihos and Bubalus, which, <strong>with</strong> species <strong>of</strong> Rhinocerosand Blephas, form a prominent feature in <strong>the</strong> fauna. <strong>The</strong> Eodentsare less developed than in <strong>the</strong> Ethiopian region, but several forms<strong>of</strong> squirrels everywhere abound, toge<strong>the</strong>r <strong>with</strong> some species <strong>of</strong>porcupine ; and <strong>the</strong> Edentata are represented by <strong>the</strong> scaly manis.Birds.—<strong>The</strong> families and genera <strong>of</strong> birds which give a characterto Oriental lands, are so numerous and varied, that wecan here only notice <strong>the</strong> more prominent and more remarkable.<strong>The</strong> Timaliida?, represented by <strong>the</strong> babblers (Garrulax, Pomatorhinus,Timalia, &c), are almost everywhere to be met <strong>with</strong>, andno less than 21 genera are peculiar to <strong>the</strong> region ; <strong>the</strong> elegantfork-tailed Enicurus and rich blue Myiophonus, though comparativelyscarce, are characteristic <strong>of</strong> <strong>the</strong> Malayan and Indo-Chinese faunas ; <strong>the</strong> elegant little " hill-tits " (Liotrichidse)abound in <strong>the</strong> same part <strong>of</strong> <strong>the</strong> region ;<strong>the</strong> green bulbuls {Phyllornis)are found everywhere ;as are various forms <strong>of</strong> Pycnonotidse,<strong>the</strong> black and crimson " minivets " (Pericrocohis), and <strong>the</strong>glossy " king-crows " (Dicrurus) ;Urocissa, Platylophus and Dendrocittaare some <strong>of</strong> <strong>the</strong> interesting and characteristic forms<strong>of</strong> <strong>the</strong> crow family ; sun-birds (Netariniida?) <strong>of</strong> at least threegenera are found throughout <strong>the</strong> region, as are <strong>the</strong> beautiful littleflower-peckers (Dicreidse), and some peculiar forms <strong>of</strong> weaverbirds(Ploceus and Mania). Of <strong>the</strong> starling family, <strong>the</strong> mostconspicuous are <strong>the</strong> glossy mynahs (Eulabes). <strong>The</strong> swallowshrikes(Artamus) are very peculiar, as are <strong>the</strong> exquisitelycoloured pittas (Pittidse), and <strong>the</strong> gaudy broad-bills (Eurylsemidse).Leaving <strong>the</strong> true Passeres, we find woodpeckers,barbets, and cuckoos everywhere, <strong>of</strong>ten <strong>of</strong> peculiar and re-


chap, xii.] THE ORIENTAL REGION. 317markable forms; among <strong>the</strong> bee-eaters we have <strong>the</strong> exquisiteNyctiornis <strong>with</strong> its pendent neck-plumes <strong>of</strong> blue or scarlet;brilliant kingfishers and strangely formed hornbills aboundeverywhere ; while brown-backed trogons <strong>with</strong> red and orangebreasts, though far less frequent, are equally a feature <strong>of</strong> <strong>the</strong>Ornithology. Next we have <strong>the</strong> frog-mou<strong>the</strong>d goatsuckers {Battrachostomus),and <strong>the</strong> whiskered swifts {Devdrochelidon), bothwide-spread, remarkable, and characteristic groups <strong>of</strong> <strong>the</strong> Orientalregion. Coming to <strong>the</strong> parrot tribe, we have only <strong>the</strong> long-tailedPalceornis and <strong>the</strong> exquisite little Zoriculus, as characteristicgenera. We now come to <strong>the</strong> pigeons, among which <strong>the</strong> fruiteatinggenera Treron and Carpophaga are <strong>the</strong> most conspicuous.<strong>The</strong> gallinaceous birds <strong>of</strong>fer us some grand forms, such as<strong>the</strong> peacocks (Pavo) ; <strong>the</strong> argus pheasants (Argusianus) ; <strong>the</strong> firebackedpheasants {Euplocanuis) ;and <strong>the</strong> jungle-fowl (Gallus), allstrikingly characteristic ;and <strong>with</strong> <strong>the</strong>se we may close our sketch,since <strong>the</strong> birds <strong>of</strong> prey and <strong>the</strong> two Orders comprising <strong>the</strong>waders and swimmers <strong>of</strong>fer nothing sufficiently remarkable tobe worthy <strong>of</strong> enumeration here.Reptiles.—Only <strong>the</strong> more abundant and characteristic groupswill here be noticed. In <strong>the</strong> serpent tribe, <strong>the</strong> Oligodontidse,a small family <strong>of</strong> ground-snakes; <strong>the</strong> Homalopsidae, or freshwatersnakes ; <strong>the</strong> Dendrophidse, or tree-snakes ; <strong>the</strong> Dryiophidse,or whip-snakes ; <strong>the</strong> Dipsadidse, or nocturnal tree-snakes ; <strong>the</strong>Lycodontidse or fanged ground-snakes ; <strong>the</strong> Pythonidre, or rocksnakes;<strong>the</strong> Elapidye, or venomous colubrine snakes (including<strong>the</strong> " cobras ") ; and <strong>the</strong> Crotalidse, or pit-vipers, are all abundantand characteristic, ranging over nearly <strong>the</strong> whole region, and presentinga great variety <strong>of</strong> genera and species.Among lizards, <strong>the</strong>Varanidaj or water-lizards ; <strong>the</strong> Scincidse or " scinks ; " <strong>the</strong> Geckotidre,or geckoes ; and <strong>the</strong> Agamidae, or eastern iguanas ; are <strong>the</strong>most universal and characteristic groups.Among crocodiles <strong>the</strong>genus Crocodilus is widely spread, Oavialis being characteristic<strong>of</strong> <strong>the</strong> Ganges. Among Chelonia, or shielded reptiles, forms <strong>of</strong>fresh-water Testudinidse and Trionychidse (s<strong>of</strong>t tortoises) aretolerably abundant.Amphibia.—<strong>The</strong> only abundant and characteristic groups <strong>of</strong>


318 ZOOLOGICAL GEOGRAPHY. [part in.this class are toads <strong>of</strong> <strong>the</strong> family Engystornidse ;tree-frogs <strong>of</strong><strong>the</strong> family Polypedatidse ; and several genera <strong>of</strong> true frogs,Ranida?.Fresh-water Fishes.—<strong>The</strong> more remarkable and characteristicfishes inhabiting <strong>the</strong>fresh waters <strong>of</strong> <strong>the</strong> Oriental region belongto <strong>the</strong> following families: Nandidee, Labyrinthici, Ophiocephalidse,SiluridEe, and Cyprinid* ;<strong>the</strong> last being specially abundant.<strong>The</strong> sketch here very briefly given, must be supplemented byan examination <strong>of</strong> <strong>the</strong> tables <strong>of</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong> genera <strong>of</strong> all<strong>the</strong> Mammalia and Birds inhabiting <strong>the</strong> region. We will nowbriefly summarize <strong>the</strong> results.Summary <strong>of</strong> <strong>the</strong> Oriental Vertebrata.—-<strong>The</strong> Oriental regionpossesses examples <strong>of</strong> 163 families <strong>of</strong> Vertebrata <strong>of</strong> which 12are peculiar, a proportion <strong>of</strong> a little more than one-fourteenth<strong>of</strong> <strong>the</strong> whole.Out <strong>of</strong> 118 genera <strong>of</strong> Mammalia 54 seem to be peculiar to<strong>the</strong> region, equal to a proportion <strong>of</strong> ^Vor a little less than half.Of Land-Birds <strong>the</strong>re are 342 genera <strong>of</strong> which 165 are peculiar,bringing <strong>the</strong> proportion very close to a half.In <strong>the</strong> Ethiopian region <strong>the</strong> proportion <strong>of</strong> peculiar formsboth <strong>of</strong> Mammalia and Birds is greater ; a fact which is notsurprising when we consider <strong>the</strong> long continued isolation <strong>of</strong> <strong>the</strong>latter region—an isolation which is even now very complete,owing to <strong>the</strong> vast extent <strong>of</strong> deserts intervening between it and<strong>the</strong> Palsearctic region ; while <strong>the</strong> Oriental and Palsearctic were,during much <strong>of</strong> <strong>the</strong> Tertiary epoch, hardly separable.Insects.Lcpidoptcra.—We can only glance hastily at <strong>the</strong> more prominentfeatures <strong>of</strong> <strong>the</strong> wonderfully rich and varied butterflyfauna<strong>of</strong> <strong>the</strong> Oriental region. In <strong>the</strong> first family Danaidse, <strong>the</strong>genera Danais and Euplcea are everywhere abundant, and <strong>the</strong>latter especially forms a conspicuous feature in <strong>the</strong> entomologicalaspect <strong>of</strong> <strong>the</strong> country; <strong>the</strong> large "spectre-butterflies"(Rcstia) are equally characteristic <strong>of</strong> <strong>the</strong> Malayan sub-region.Satyridse, though abundant are not very remarkable, Debts,Melanitis, Mycalesis, and Ypthima being <strong>the</strong> most characteristic


chap, xii.] THE ORIENTAL REGION. 319genera. Morphidae are well represented by <strong>the</strong> genera Amathusia,Zeuxidia, Discophora, and Thaumantis, some <strong>of</strong> <strong>the</strong>species <strong>of</strong> which almost equal <strong>the</strong> grand South AmericanMorphos. <strong>The</strong> Nymphalidse furnish us <strong>with</strong> a host <strong>of</strong> characteristicgenera, among <strong>the</strong> most remarkable <strong>of</strong> which are,Terinos, Adolias, Cethosia, Cyrestis, Limenitis, and Nymphalis,all abounding in beautiful species. Among <strong>the</strong> LycsenidsB area number <strong>of</strong> fine groups, among which we may mention Ilerda,Myrina, Deudoryx, Aphneus, lolaics, and AmUypodia, ascharacteristicexamples. <strong>The</strong> Pieridse furnish many fine forms, suchas Thyca, Iphias, <strong>The</strong>stias, Eronia, Prioneris, and Dercas, <strong>the</strong> lasttwo being peculiar. <strong>The</strong> Papilionidae are unsurpassed in <strong>the</strong>world, presenting such grand genera as Teinopalpus and Bhutanitis;<strong>the</strong> yellow-marked Omithoptercc ; <strong>the</strong> superb "Brookiana;"<strong>the</strong> elegant Leptocercus; and Papilios <strong>of</strong> <strong>the</strong> "Coon," "Philoxenus,"" Menmon," " Protenor," and especially <strong>the</strong> ' green-andgold-dusted' " Paris " groups.<strong>The</strong> Moths call for no special observations, except to notice<strong>the</strong> existence in Nor<strong>the</strong>rn India <strong>of</strong> a number <strong>of</strong> forms whichresemble in a striking manner some <strong>of</strong> <strong>the</strong> most remarkable<strong>of</strong> <strong>the</strong> above mentioned groups <strong>of</strong> <strong>the</strong> genus Papilio, especially<strong>the</strong> "Protenor" group, which <strong>the</strong>re is reason to believe isprotected by a peculiar smell or taste like <strong>the</strong> Eeliconias andDanaidse.Coleoptera.—<strong>The</strong> most characteristic Oriental form <strong>of</strong> <strong>the</strong>Cicindelidse or tiger beetles, is undoubtedly <strong>the</strong> elegant genusCollyris, which is found over <strong>the</strong> whole region and is almostconfined to it. Less abundant, but equally characteristic, is <strong>the</strong>wingless ant-like Tricondyla. Two small genera Apteroessa andDromicidia are confined to <strong>the</strong> Indian Peninsula, while <strong>The</strong>ratesonly occurs in <strong>the</strong> Malayan sub-region.<strong>The</strong> Carabidas, or ground carnivorous beetles, are so numerousthat we can only notice a few <strong>of</strong> <strong>the</strong> more remarkable andcharacteristic forms. <strong>The</strong> wonderful Mormolyce <strong>of</strong> <strong>the</strong> Indo-Malay sub-region, stands pre-eminent for singularity in <strong>the</strong>entire family. Thyreopterus, Orthogonius, Gatascopus, and Pericallusare very characteristic forms, as well as Planetes and


320 ZOOLOGICAL GEOGKAPHY. [part hi.Distrigus, <strong>the</strong> latter having a single species in Madagascar.<strong>The</strong>re are 80 genera <strong>of</strong> this family peculiar to <strong>the</strong> region, 10<strong>of</strong> which have only been found in Ceylon.Among <strong>the</strong> Lucanidae, or stag-beetles, Lucanus, Odontolabris,and Cladognathus are <strong>the</strong> most characteristic forms. Sixteengenera inhabit <strong>the</strong> region, <strong>of</strong> which 7 are altoge<strong>the</strong>r peculiar,while three o<strong>the</strong>rs only extend eastward to <strong>the</strong> Austro-Malayansub-region.<strong>The</strong> beautiful CetoniidaB, or rose-chafers, are well representedby Rhomborhina, Heterorhina, Clinteria, Macronota, Agestrata,Chalco<strong>the</strong>a and maDy fine species <strong>of</strong> Cetonia. <strong>The</strong>re are 17peculiar genera, <strong>of</strong> which Mycteristes, Phcedimus, Plectrone, andRhagopteryx, are Malayan ; while Narycius, Clerota, Bombodes,and Chiloldba are Indian.In Buprestidse—those elongate metallic-coloured beetles whoseelytra are used as ornaments in many parts <strong>of</strong> <strong>the</strong> world—thisregion stands pre-eminent, in its gigantic Catoxantha, its fineChrysochroa, its Indian Sternocera, its Malayan Chalcophoraand Belionota, as well as many o<strong>the</strong>r beautiful forms. Itpossesses 41 genera, <strong>of</strong> which 14 are peculiar to it, <strong>the</strong> restbeing generally <strong>of</strong> wide range or common to <strong>the</strong> Ethiopian andAustralian regions.In <strong>the</strong> extensive and elegant group <strong>of</strong> Longicorns, <strong>the</strong> Orientalregion is only inferior to <strong>the</strong> Neotropical. It possesses 360genera, 25 <strong>of</strong> which are Prionidse, 117 Cerambicidse, and 218Lamiidse;—about 70 per cent, <strong>of</strong> <strong>the</strong> whole being peculiar.<strong>The</strong> most characteristic genera are RhapMdopodus and JEgosomaamong Prionidse ; Neocerambyx, Euryarthrum, Pachyteria, Acrocyrta,Tetraommatus, Chloridolum, and Polyzonus among Cerambycidse;and Ccelosterna, Rhytidophora, Batocera, Agelasta, andAsta<strong>the</strong>s among Lamiidse.Of remarkable forms in o<strong>the</strong>r families,we may mention <strong>the</strong>gigantic horned Chalcosoma among Scaraba?ida3 ; <strong>the</strong> metallicCampsosternus amoDg Elateridae ;<strong>the</strong> handsome but anomalousTrictenotoma forming a distinct family; <strong>the</strong> gorgeous Pachyrhynchi<strong>of</strong> <strong>the</strong> Philippine Islands among Curculionidse ;Diurus


chap, xii.] THE ORIENTAL REGION. 321among Brenthidre ; <strong>with</strong> an immense number and variety <strong>of</strong>Anthotribidse, Heteromera, Malacoderma, and Phytophaga.<strong>The</strong> Oriental Sub-regions.<strong>The</strong> four sub-regions into which we have divided <strong>the</strong> Orientalregion, are very unequal in extent, and perhaps more so inproductiveness, but <strong>the</strong>y each have well-marked special features,and serve well to exhibit <strong>the</strong> main zoological characteristics <strong>of</strong><strong>the</strong> region. As <strong>the</strong>y are all tolerably well defined and <strong>the</strong>irfaunas comparatively well-known, <strong>the</strong>ir characteristics will begiven <strong>with</strong> ra<strong>the</strong>r more than usual detail.7". Hindostan, or Indian Sub-region.This includes <strong>the</strong> whole peninsula <strong>of</strong> India from <strong>the</strong> foot <strong>of</strong> <strong>the</strong>Himalayas on <strong>the</strong> north to somewhere near Seringapatam on <strong>the</strong>south, <strong>the</strong> boundary <strong>of</strong> <strong>the</strong> Ceylonese sub-region being unsettled.<strong>The</strong> deltas <strong>of</strong> <strong>the</strong> Ganges and Brahmaputra mark its easternlimits, and it probably reaches to about Cashmere in <strong>the</strong> northwest,and perhaps to <strong>the</strong> valley <strong>of</strong> <strong>the</strong> Indus fur<strong>the</strong>r south ; but<strong>the</strong> great desert tract to <strong>the</strong> east <strong>of</strong> <strong>the</strong> Indus forms a transitionto <strong>the</strong> south Palaearctic sub-region. Perhaps on <strong>the</strong> whole <strong>the</strong>Indus may be taken as a convenient boundary. Many Indiannaturalists, especially Mr. Blyth and Mr. Blanford, are impressed<strong>with</strong> <strong>the</strong> <strong>relations</strong> <strong>of</strong> <strong>the</strong> greater part <strong>of</strong> this sub-region to <strong>the</strong>Ethiopian region, and have proposed to divide it into severalzoological districts dependent on differences <strong>of</strong> climate and vegetation,and characterized by possessing faunas more or less alliedei<strong>the</strong>r to <strong>the</strong> Himalayan or <strong>the</strong> Ethiopian type. But <strong>the</strong>se subdivisionsappear far too complex to be useful to <strong>the</strong> general student,and even were <strong>the</strong>y proved to be natural, would be beyond<strong>the</strong> scope <strong>of</strong> this work. I agree, however, <strong>with</strong> Mr. Elwes inthinking that <strong>the</strong>y really belong to local ra<strong>the</strong>r than to <strong>geographical</strong><strong>distribution</strong>, and confound " station " <strong>with</strong> " habitat."Wherever <strong>the</strong>re is a marked diversity <strong>of</strong> surface and vegetation<strong>the</strong> productions <strong>of</strong> a country will correspondingly differ; <strong>the</strong>groups peculiar to forests,for example, will be absent from open


.322 ZOOLOGICAL GEOGRAPHY. [part hi.plains or arid deserts. It happens that <strong>the</strong> three great OldWorld regions are separated from each o<strong>the</strong>r by a debatable landwhich is chiefly <strong>of</strong> a desert character ; hence we must expect t<strong>of</strong>ind a resemblance between <strong>the</strong> inhabitants <strong>of</strong> such districts ineach region. We also find a great resemblance between <strong>the</strong> aquaticbirds <strong>of</strong> <strong>the</strong> three regions ; and as we generally give little weightto <strong>the</strong>se in our estimate <strong>of</strong> <strong>the</strong> degree <strong>of</strong> affinity <strong>of</strong> <strong>the</strong> faunas <strong>of</strong>different countries, so we should not count <strong>the</strong> desert fauna as <strong>of</strong>equal weight <strong>with</strong> <strong>the</strong> more restricted and peculiar types whichare found in <strong>the</strong> fertile tracts,—in <strong>the</strong> mountains and valleys, andespecially in <strong>the</strong> primeval forests. <strong>The</strong> supposed preponderance<strong>of</strong> exclusively Ethiopian groups <strong>of</strong> Mammalia and Birds in thissub-region, deserves however a close examination, in order toascertain how far <strong>the</strong> facts really warrant such an opinion.Mammalia.—<strong>The</strong> following list<strong>of</strong> <strong>the</strong> more important genera<strong>of</strong> Mammalia which range over <strong>the</strong> larger part <strong>of</strong> this sub-regionwill enable naturalists to form an independent judgment as to<strong>the</strong> preponderance <strong>of</strong> Ethiopian, or <strong>of</strong> Oriental and Palsearctictypes, in this, <strong>the</strong> most important <strong>of</strong> all <strong>the</strong> classes <strong>of</strong> <strong>animals</strong>for <strong>geographical</strong> <strong>distribution</strong>.Range <strong>of</strong> <strong>the</strong> Genera <strong>of</strong> Mammalia which Inhabit <strong>the</strong> Sub-regionOF HlNDOSTAN.1. Presbytes ... Oriental only.2. Macacus ... Oriental only.3. Erinaceus ... Palsearctic genus.4. Sorex Widely distributed.5. Felis ... ... Almost Cosmopolitan.6. Cynselurus ... Ethiopian and S. Palsearctic.7. Viverra . . Ethiopian and Oriental to China and Malaya.8. Viverricula ... Oriental only.9. Paradoxurus ... Oriental only.10. Herpestes ... Ethiopian, S. Palsearctic, and Oriental to Malaya.11. Calogale ... Ethiopian, Oriental to Cambodja.12. Tseniogale ... Oriental.13. Hysena ... Palsearctic and Ethiopian (a Palsearctic species.)14. Canis ... ... Palsearctic and Oriental to Malaya.15. Cuon Oriental to Malaya.16. Vulpes ... Very wide range.17. Lutra ... ... Oriental and Palsearctic.18. Mellivora ... Ethiopian.19. Melursus ... Oriental only ; family not Ethiopian.20. Sus ... ... Palsearctic and Oriental, not Ethiopian.21. Tragulus ... Oriental.


chap, xii.] THE ORIENTAL REGION. 32322. Cervus


;;324 ZOOLOGICAL GEOGRAPHY. [part hi.open to observation; and that <strong>the</strong>se are <strong>of</strong>ten <strong>of</strong> wide-spreadtypes, and some few almost exclusively African. <strong>The</strong> facts headduced do not, however, seem to have satisfied <strong>the</strong> objectorsand as <strong>the</strong> subject is an important one, I will here give lists<strong>of</strong> all <strong>the</strong> genera <strong>of</strong> Passeres, Picariae, Psittaci, Columbee, andGalling, which inhabit <strong>the</strong> sub-region, leaving out those whichonly just enter <strong>with</strong>in its boundaries from adjacent sub-regions.<strong>The</strong>se are arranged under four heads:— 1. Oriental genera; whichare ei<strong>the</strong>r wholly confined to, or strikingly prevalent in, <strong>the</strong>Oriental region beyond <strong>the</strong> limits <strong>of</strong> <strong>the</strong> Indian peninsula. 2.Genera <strong>of</strong> Wide Ptange ; which are fully as much entitled to beconsidered Oriental or Paleearctic as Ethiopian, and cannot beheld to prove any Ethiopian affinity. 3. Palsearctic genera;which are altoge<strong>the</strong>r or almost absent from <strong>the</strong> Ethiopian region.4. Ethiopian genera ; which are confined to, or very prevalentin, <strong>the</strong> Ethiopian region, whence <strong>the</strong>y extend into <strong>the</strong> Indianpeninsula but not over <strong>the</strong> whole Oriental region. <strong>The</strong> last are<strong>the</strong> only ones which, can be fairly balanced against those <strong>of</strong> <strong>the</strong>first list, in order to determine <strong>the</strong> character <strong>of</strong> <strong>the</strong> fauna.1. Oriental Genera in Central India.Geocichla, Orthotomtcs, Prinia, Megalums, Abrornis, Larvivora,Copsychus, Kittacincla, PomatorJiinus, Malacocercus, Chatarrhcea,Layardia, Garrulax, Trochalopteron, Pellorneum, Dumetia,Pyctoris, Alcippe, Myiophonus, Sitta, Dendropliila, Phyllornis,lora, Hypsipetcs, Pericrocotus, Graucalus, Volvocivora, Chibia,Chaptia, Irena, Erythrosterna, Hewipus, Hemichelidon, Niltava,Cyomis, Eumyias, Hypothymis, Myicdestcs, Tephrodornis, Dendrocitta,Arachnechthra, Nectarophila, Arachno<strong>the</strong>ra, Dicocum, Piprisoma,Mania, Eulabes, Pastor, Acrido<strong>the</strong>res, Sturnia, Stumopastor,Artamus, Nemoricola, Pitta, Yungipicus, Clirysocolaptes, Hemicircus,Oecinus, Mtdleripicus, Bracliypternus, Tiga, Micropternus,Megala?ma, Xantliolccma, Rhopodytes, Taccocoua, Sumiculus,Hierococcyx, Eudynamnis, Nyctiomis, Harpactes, Pelargopsis,Ceyx, Hydrocissa, Meniceros, Batrachostomus, Dendrochelidon,Collocalia, Palceornis, Treron, Carpophaga, Chalcophaps, Ortygornis,Perdix, Pavo, Gallus, Gallopcrdix — 87 genera; and


chap, xii.] THE ORIENTAL REGION. 325one peculiar genus, Salpornis, whose affinities are Palrearctic orOriental.2. Genera <strong>of</strong> Wide Eange occurring in Central India.Tardus, Monticola, Drymceca, Cisticola, Acrocephalus, Phylloscopus,Pratincola, Parus, Pycnonotus, Criniger, Oriolus, Dicrurus,Tchitrea, Zanitis, Corvus, Zosterops, Hirundo, Cotyle, Passer,Ploceus, Estrilda, Alauda, Galandrella, Mirafra, Ammomanes,Motacilla, Anthus, Picus,Yunx, Centropus, Cuculus, Chrysoccocyx,Cqccystes, Coracias, Eurystomus, Merop>s, Alcedo, Ceryle, Halcyon,Upupa, Caprimtdgus, Cypselus, Chcetura, Columba,Turtur, Pterocles,Coturnix, Turnix;—48 genera.3. Pal^earctic Genera occurring in Central India.Hypolais, Sylvia, Ourruca, Cyanecula, Calliope, Chelidon, Euspiza,Emhcriza, Galerita, Calobates, Corydalla;— 11 genera.4. Ethiopian Genera occurring in Central India.Tliamnobia, Pyrrhulauda, Pterocles, Francolinus ;— 4 genera.A consideration <strong>of</strong> <strong>the</strong> above lists shows us, that <strong>the</strong> Hindostansub-region is by no means so poor in forms <strong>of</strong> bird-life as isgenerally supposed (and as I had myself anticipated, it wouldprove to be), possessing, as it does, 151 genera <strong>of</strong> land-birds,<strong>with</strong>out counting <strong>the</strong> Accipitres. It must also set at rest <strong>the</strong>question <strong>of</strong> <strong>the</strong> zoological affinities <strong>of</strong> <strong>the</strong> district, since a preponderance<strong>of</strong> 88 genera, against 4, cannot be held to be insufficient,and cannot be materially altered by any corrections indetails that may be proposed or substantiated. Even <strong>of</strong> <strong>the</strong>sefour, only <strong>the</strong> first two are exclusively Ethiopian, Pterocles andFrancolinus both being Palsearctic also. It is a question, indeed,whe<strong>the</strong>r anywhere in <strong>the</strong> world an outlying sub-region canbe found, exhibiting less zoological affinity for <strong>the</strong> adjacentregions ;and we have here a striking illustration <strong>of</strong> <strong>the</strong> necessity<strong>of</strong> deciding all such cases, not by examples, which may be sochosen as to support any view, but by carefully weighing andcontrasting <strong>the</strong> whole <strong>of</strong> <strong>the</strong> facts on which <strong>the</strong> solution <strong>of</strong> <strong>the</strong>


;326 ZOOLOGICAL GEOGRAPHY. [part hi.problem admittedly depends. It will, perhaps, be said that agreat many <strong>of</strong> <strong>the</strong> 88 genera above given are very scarce and verybut this is certainly not <strong>the</strong> case <strong>with</strong> <strong>the</strong> majority <strong>of</strong> <strong>the</strong>mlocal ;and even where it is so, that does not in any degree affect <strong>the</strong>irvalue as indicating zoo-<strong>geographical</strong> affinities. It is <strong>the</strong> presence<strong>of</strong> a type in a region, not its abundance orscarcity, that is<strong>the</strong> important fact ; and when we have to do, as we have here,<strong>with</strong> many groups whose habits and mode <strong>of</strong> life necessarilyseclude <strong>the</strong>m from observation, <strong>the</strong>ireven be a fact.supposed scarcity may notReptiles and Amphibia.—Pieptiles entirely agree <strong>with</strong> Mammaliaand Birds in <strong>the</strong> main features <strong>of</strong> <strong>the</strong>ir <strong>distribution</strong>.Out <strong>of</strong> 17 families <strong>of</strong> snakes inhabiting Hindostan, 16 rangeover <strong>the</strong> greater part <strong>of</strong> <strong>the</strong> entire region, and only two canbe supposed to show any Ethiopian affinity. <strong>The</strong>se are <strong>the</strong>Psammophidre and Erycidee, both desert-haunting groups, andalmost as much South Palsearctic as African. <strong>The</strong> genus Tropidococcyxis peculiar to <strong>the</strong> sub-region, and Aspidura, Passeritaand Cynopliis to <strong>the</strong> peninsula and Ceylon ; while a large number<strong>of</strong> <strong>the</strong> most characteristic genera, as Dipsas, Simotes, Bungarus,Naja, Trimeresurus, Lycodon and Python, are characteristicallyOriental.Of <strong>the</strong> six families <strong>of</strong> lizards all have a wide range <strong>The</strong>genera Eumeces, Pentadactylus, Gecko, Eublepharis, and Draco, arecharacteristically or wholly Oriental ; Ophiops and Uromastixare Palrearctic ; while Cliammlcon is <strong>the</strong> solitary case <strong>of</strong> decidedEthiopian affinity.Of <strong>the</strong> Amphibia not a single family exhibits special Ethiopianaffinities.II.Sub-region <strong>of</strong> Ceylon and South-India.<strong>The</strong> Island <strong>of</strong> Ceylon is characterised by such striking peculiaritiesin its animal productions, as to render necessary itsseparation from <strong>the</strong> peninsula <strong>of</strong> India as a sub-region ;but itis found that most <strong>of</strong> <strong>the</strong>se special features extend to <strong>the</strong> Neilgherriesand <strong>the</strong> whole sou<strong>the</strong>rn mountainous portion <strong>of</strong> India,and that <strong>the</strong> two must be united inany zoo-<strong>geographical</strong> pro-


—chap, xii.] THE ORIENTAL REGION. 327vince. <strong>The</strong> main features <strong>of</strong> this division are,—<strong>the</strong> appearance<strong>of</strong> numerous <strong>animals</strong> allied to forms only found again in <strong>the</strong>Himalayas or in <strong>the</strong> Malayan sub-region, <strong>the</strong> possession <strong>of</strong>several peculiar generic types, and an unusual number <strong>of</strong>peculiar species.Mammalia.—Among Mammalia <strong>the</strong> most remarkable formis Loris, a genus <strong>of</strong> Lemurs altoge<strong>the</strong>r peculiar to <strong>the</strong> subregion;several peculiar monkeys <strong>of</strong> <strong>the</strong> genus Presbytes ; <strong>the</strong>Malayan genus Tupaia ; and Platacanthomys, a peculiar genus<strong>of</strong> Muridse.Birds.—Among birds it has Ochromela, a peculiar genus <strong>of</strong>flycatchers ;Phomicopliaes (Cuculidae) and Drymocataphus (Timaliidae),both Malayan forms ; a species <strong>of</strong> Myiophonus whosenearest ally is in Java; Trochalopteron, Brachypteryx, Bucerosand Loriculus, which are only found elsewhere in <strong>the</strong> Himalayasand Malayana. It also possesses about 80 peculiar species <strong>of</strong>birds, including a large jungle fowl, one owl and two hornbills.Reptiles.—It is however by its Eeptiles, even more than by itshigher vertebrates, that this sub-region is clearly characterised.Among snakes it possesses an entire family, Uropeltidae, consisting<strong>of</strong> 5 genera and 18 species altoge<strong>the</strong>r confined to it,Rhinophisand Uropeltis in Ceylon, Silybura, Plecturus and Melanophidiumin Sou<strong>the</strong>rn India. Four o<strong>the</strong>r genera <strong>of</strong> snakes, Haplocercus,Cercaspis, Peltopelor, and Hypnale are also peculiar; Chersydrusis only found elsewhere in Malaya ; while Aspidura, Passerita,and Cyjiophis, only extend to Hindostan ; and species <strong>of</strong> Eryx,PJchis, and Psammophis show an affinity <strong>with</strong> Ethiopian andPalasarctic forms. Among lizards several genera <strong>of</strong> Agamidceare peculiar, such as Otocryptis, Lyricoephalus, Ceratophora, Cophotis,Salea, Sitana and Charasia. In <strong>the</strong> family Acontiadae,Nessia is peculiar to Ceylon, while a species <strong>of</strong> <strong>the</strong> Africangenus Acontias shows an affinity for <strong>the</strong> Ethiopian region.Amphibia.—<strong>The</strong> genera <strong>of</strong> Amphibians that occur here aregenerally <strong>of</strong> wide range, but Nannophrys, Haphbatrachus,and Cacopus are confined to <strong>the</strong> sub-region ; while Megalophrysis Malayan, and <strong>the</strong> species found in Ceylon also inhabitJava.


328 ZOOLOGICAL GEOGRAPHY. [part hi.Insects.—<strong>The</strong> insects <strong>of</strong> Ceylon also furnish some curiousexamples <strong>of</strong> its distinctness from Hindostan, and its affinity <strong>with</strong>Malaya. Among its butterflies we find Papilio jophon, closelyallied to P. antiphus <strong>of</strong> Malaya. <strong>The</strong> remarkable genus Hestia, socharacteristic <strong>of</strong> <strong>the</strong> Malay archipelago, only occurs elsewhere on<strong>the</strong> mountains <strong>of</strong> Ceylon ; while its Cynthia and Parthcnos areAmongclosely allied to, if not identical <strong>with</strong>, Malayan species.Coleoptera we have yet more strikiDg examples. <strong>The</strong> highly characteristicMalayan genus Tricondyla is represented in Ceylon byno less than 10 species ;and among Longicorns we find <strong>the</strong> generaTetraommatus, Thranius, Cacia, Praonetha, Ropica, and Serixia,allexclusively Malayan or only just entering <strong>the</strong> Indo-Chinesepeninsula, yet all represented in Ceylon, while not a singlespecies occurs in any part <strong>of</strong> India or <strong>the</strong> Himalayas.<strong>The</strong> Past History <strong>of</strong> Ceylon and South-India as indicated by itsFauna.—In our account <strong>of</strong> <strong>the</strong> Ethiopian region we have alreadyhad occasion to refer to an ancient connection between this subregionand Madagascar, in order to explain <strong>the</strong> <strong>distribution</strong> <strong>of</strong><strong>the</strong> Lemurine type, and some o<strong>the</strong>r curious affinities between <strong>the</strong>two countries. This view is supported by <strong>the</strong> geology <strong>of</strong> India,which shows us Ceylon and South India consisting mainly <strong>of</strong>granitic and old metamorphic rocks, while <strong>the</strong> greater part <strong>of</strong> <strong>the</strong>peninsula, forming our first sub-region, is <strong>of</strong> tertiary formation,<strong>with</strong> a few isolated patches <strong>of</strong> secondary rocks. It is evident<strong>the</strong>refore, that during much <strong>of</strong> <strong>the</strong> tertiary period, Ceylon andSouth India were bounded on <strong>the</strong> north by a considerable extent<strong>of</strong> sea, and probably formed part <strong>of</strong> an extensive sou<strong>the</strong>rn continentor great island. <strong>The</strong> very numerous and remarkable cases<strong>of</strong> affinity <strong>with</strong> Malaya, require however some closer approximationto <strong>the</strong>se islands, which probably occurred at a later period.When, stilllater, <strong>the</strong> great plains and table-lands <strong>of</strong> Hindostanwere formed, and a permanent land communication effected <strong>with</strong><strong>the</strong> rich and highly developed Himalo-Chinese fauna, a rapid immigration<strong>of</strong> new types took place, and many <strong>of</strong> <strong>the</strong> less specialisedforms <strong>of</strong> mammalia and birds (particularly those <strong>of</strong> ancientAmong reptiles and insects <strong>the</strong>Ethiopian type) became extinct.competition was less severe, or <strong>the</strong> older forms were too well


chap, xii.] THE ORIENTAL REGION. 329adapted to local conditions to be expelled ; so that it is among<strong>the</strong>se groups alone that we find any considerable number, <strong>of</strong> whatare probably <strong>the</strong> remains <strong>of</strong> <strong>the</strong> ancient fauna <strong>of</strong> a now submergedsou<strong>the</strong>rn continent.777. Himalayan or Indo-Chinese Suh-region.This, which is probably <strong>the</strong> richest <strong>of</strong> all<strong>the</strong> sub-regions, andperhaps one <strong>of</strong> <strong>the</strong> richest <strong>of</strong> all tracts <strong>of</strong> equal extent on <strong>the</strong>face <strong>of</strong> <strong>the</strong> globe, is essentially a forest-covered, mountainouscountry, mostly <strong>with</strong>in <strong>the</strong> tropics, but on its nor<strong>the</strong>rn marginextending some degrees beyond it, and rising in a continuousmountain range till it meets and intercalates <strong>with</strong> <strong>the</strong> Manchuriansub-division <strong>of</strong> <strong>the</strong> Palsearetic region. <strong>The</strong> peculiarmammalia, birds and insects <strong>of</strong> this sub-region begin to appearat <strong>the</strong> very foot <strong>of</strong> <strong>the</strong> Himalayas, but Dr. Gun<strong>the</strong>r has shownthat many <strong>of</strong> <strong>the</strong> reptiles characteristic <strong>of</strong> <strong>the</strong> plains <strong>of</strong> Indiaare found to a height <strong>of</strong> from 2,000 to 4,000 feet.In Sikhim, which may be taken as a typical example <strong>of</strong> <strong>the</strong>Himalayan portion <strong>of</strong> <strong>the</strong> sub-region, it seems to extend to analtitude <strong>of</strong> little less than 10,000 feet, that being <strong>the</strong> limit <strong>of</strong> <strong>the</strong>characteristic Timaliidse or babbling thrushes ;while <strong>the</strong> equallycharacteristic Pycnonotidse, or bulbuls, and Treronidse, or thickbilledfruit-pigeons, do not, according to Mr. Blanford, reachquite so high. We may perhaps take 9,000 feet as a goodapproximation over a large part <strong>of</strong> <strong>the</strong> Himalayan range; butit is evidently not possible to define <strong>the</strong> line <strong>with</strong> any greatprecision. Westward, <strong>the</strong> sub-region extends in diminishingbreadth, till it terminates in or near Cashmere, where <strong>the</strong> fauna<strong>of</strong> <strong>the</strong> plains <strong>of</strong> India almost meets that <strong>of</strong> <strong>the</strong> Palaearcticregion, at a moderate elevation. Eastward, it reaches into EastThibet and North-westChina, where Pere David has found alarge number <strong>of</strong> <strong>the</strong> peculiar types <strong>of</strong> <strong>the</strong> Eastern Himalayas.fauna, in general features identical, extends over Burmah andSiam to South China; mingling <strong>with</strong> <strong>the</strong> Palaearctic fauna in<strong>the</strong> mountains south <strong>of</strong> <strong>the</strong> Yang-tse-kiang river, and <strong>with</strong>that <strong>of</strong> Indo-Malaya in Tenasserim, and to a lesser extent inSou<strong>the</strong>rn Siam and Cochin China.Yol. I.—23A


—;330 ZOOLOGICAL GEOGRAPHY. [part in.Zoological Characteristics <strong>of</strong> <strong>the</strong> Himalayan Sub-region.—Taking this sub-region as a whole, we find it to be characterisedby 3 genera <strong>of</strong> mammalia (<strong>with</strong>out counting bats), and44 genera <strong>of</strong> land-birds, which are altoge<strong>the</strong>r peculiar to itand by 13 genera <strong>of</strong> mammalia and 36 <strong>of</strong> birds, which itpossesses in common <strong>with</strong> <strong>the</strong> Malayan sub-region; andbesides <strong>the</strong>se it has almost all <strong>the</strong> genera before enumeratedas " Oriental," and several o<strong>the</strong>rs <strong>of</strong> wide range, more especiallya number <strong>of</strong> Palrearetic genera which appear in <strong>the</strong> higherHimalayas. <strong>The</strong> names <strong>of</strong> <strong>the</strong> more characteristic genera areas follows :Peculiar Himalo-Chinese Genera.Mammalia.— Urva, Arctonyx, JElurus.Birds. — Suya, Sorites, Chaimarrhornis, Tarsiger, Oreicola,Acanthoptila, Grammatoptila, Trochaloptcron,Actinodura, Sibia,Suthora, Paradoxomis, Chlenasicus, Tesia, Rimator, sEgithaliscus,Cephalopyrus, Liothrix, Siva, Minla, Proparus, Cutia, Yuhina,Ixtdus, Myzornis, Erpornis, Hemixus, Chibia, Niltava, Anthipes,Chelidorhynx, Urocissa, Pachyglossa, Hctcrura, Hosmatospiza,Ampeliceps, Saroglossa, Psarisomus, Serilophus, Vivia, Hyopicus,Gicinulus, Aceros,Ceriomis.Genera common to <strong>the</strong> Himalo-Chinese and MalayanSub-regions.Mammalia. — Hylobates, Nycticebus, Viverricula, Prionodon,Arctitis, Pagama, Arctogale, Cuon, Gymnopus, Aonyx, Helictis,Rhinoceros, Nemorhedus, Rhizomys.Birds.— Oreocincla, Notodela, Janthocincla, Timalia, Stachyris,Mixornis, Trichastoma, JEnicurus, Pncepyga, Melanochlora, Allotrius,Microscelis, Iole, Analcipus, Cochoa, Rhringa, Xanthopygia,Hylocharis, Cissa, Temnurus, Cryjjsirhina, Chalcosletha, Anthreptes,Chalcoparia, Cymbirhynchus, Hydrornis, Sasia, Venilia,Indicator, Carcincutes, Lyncornis, Macropygia, Argusianus Polyplectron,Ruplocamus, Phodilus.Plate VII. Scene in Nepal, <strong>with</strong> Characteristic HimalayanAnimals.— Our illustration contains figures <strong>of</strong> two mammals


Tm:aa #liFf;**!.'' liF* -H


—;chap, xii.] THE ORIENTAL REGION. 331and two birds, characteristic <strong>of</strong> <strong>the</strong> higher woody region <strong>of</strong> <strong>the</strong>Himalayas. <strong>The</strong> lower figure on <strong>the</strong> left is <strong>the</strong> Helictis nepalensis,confined to <strong>the</strong> Eastern Himalayas, and belonging to a genus <strong>of</strong><strong>the</strong> weasel family which is exclusively Oriental. It is marked<strong>with</strong> white on a grey-brown ground. Above it is <strong>the</strong> remarkablePanda (sElurus fulgent) , a beautiful animal <strong>with</strong> a glossy fur<strong>of</strong> a reddish colour, darker feet, and a white somewhat cat-likeface. It is distantly allied to <strong>the</strong> bears, and more nearly to <strong>the</strong>American racoons, yet <strong>with</strong> sufficient differences to constitutea distinct family. <strong>The</strong> large bird on <strong>the</strong> tree, is <strong>the</strong> hornedTragopan (Ceriomis satyra), one <strong>of</strong> <strong>the</strong> fine Himalayan pheasants,magnificently spotted <strong>with</strong> red and white, and ornamented <strong>with</strong>fleshy erectile wattles and horns, <strong>of</strong> vivid blue and red colours.<strong>The</strong> bird in <strong>the</strong> foreground is <strong>the</strong> Ibidorhynchus stru<strong>the</strong>rsii, arare and curious wader, allied to <strong>the</strong> curlews and sandpipers buthaving <strong>the</strong> bill and feet red. It frequents <strong>the</strong> river-beds in <strong>the</strong>higher Himalayas, but has also been found in Thibet.itReptiles.—Very few genera <strong>of</strong> reptiles are peculiar to thissub-region, all <strong>the</strong> more important ranging into <strong>the</strong> Malayislands. Of snakes <strong>the</strong> following are <strong>the</strong> more characteristicgenera :Typhline, Cylindrophis, Xenopeltis,Calamaria, Xenelaphis,Hypsirhina, Fordonia, several small genera <strong>of</strong> Homalopsidse{Herpeton and Hipistes being characteristic <strong>of</strong> Burmah andSiam) Psammodynastes, Oonyosoma, Chrysopelea, Tragops, Dipsas,Pareas, Python, Bungarus, Naja, Callophis, and Trimeresurus.Naja reaches 8,000 feet elevation in <strong>the</strong> Himalayas, Tropidonotus9,000 feet, Ablabes 10,000 feet, and Simotes 15,000 feet.Of lizards, Pseudopus has one species in <strong>the</strong> Khasya hillswhile <strong>the</strong> o<strong>the</strong>r inhabits South-east Europe ; and <strong>the</strong>re are twosmall genera <strong>of</strong> Agamidaa peculiar to <strong>the</strong> Himalayas, whileDraco and Calotes have a wide range and Acanthosaurd, Dilophyrus,Physignathiis, and Liolepis are found chiefly in <strong>the</strong>Indo-Chinese peninsula.<strong>The</strong>re are several genera <strong>of</strong> Scincidaeand <strong>the</strong> extensive genus <strong>of</strong> wall-lizards, Gecko, ranges over <strong>the</strong>whole region.Of Amphibia, <strong>the</strong> peculiar forms are not numerous.Ichthyophis


—;;332 ZOOLOGICAL GEOGRAPHY. [part hi.a genus <strong>of</strong> Ceciliadse, is peculiar to <strong>the</strong> Khasya Hills ; Tylotritron(Salamandridae) to Yunan in Western China, and perhapsbelongs to <strong>the</strong> Paleearctie region.Of <strong>the</strong> tail-less Batrachians, Glyphoglossus is found in PeguXenoplvys in <strong>the</strong> Eastern Himalayas; while Gallula, Ixalus,Rhacophorus, Hylurana, Oxyglossus, and Phrynoglossus, are commonto <strong>the</strong> Himalo-Chinese and Malayan sub-regions.Of <strong>the</strong> lizards, Colotes, Barycephalus, and Hinulia,—and <strong>of</strong><strong>the</strong> Batrachia, Bufo,—are found at above 11,000 feet elevation in<strong>the</strong> Himalayas.Insects.— So little has been done in working out <strong>the</strong> insectfaunas <strong>of</strong> <strong>the</strong> separatesub-regions, that <strong>the</strong>y cannot be treatedin detail, and <strong>the</strong> reader is referred to <strong>the</strong> chapter on <strong>the</strong> <strong>distribution</strong><strong>of</strong> insects in <strong>the</strong> part <strong>of</strong> this work devoted to GeographicalZoology. A few particulars may, however, be given as to<strong>the</strong> butterflies, which have been more systematically collected intropical countries than any o<strong>the</strong>r order <strong>of</strong> insects. <strong>The</strong> Himalayanbutterflies, especially in <strong>the</strong> eastern portions <strong>of</strong> <strong>the</strong> rangein Assam and <strong>the</strong> Khasya Hills—are remarkably fineand veryabundant; yet all <strong>the</strong> larger groups extend into <strong>the</strong> Malayansub-region, many to Ceylon, and a considerable proportion evento Africa and Austro-Malaya. <strong>The</strong>re are a large number <strong>of</strong>peculiar types, but most <strong>of</strong> <strong>the</strong>m consist <strong>of</strong> few or singlespecies.Such are Neope, Orenoma, and Rhaphicera, gener a <strong>of</strong> SatyridseEnispe (Morphidse) ; Hestina, Penthcma, and Abrota (Nymphalidse); Dodona (Erycinidse) ; Ilerda (Lycsenidse) ; Calinaga, Teinopalpus,and Bhutanitis (Papilionidse). Its more prominent featuresare, however, derived from what may be termed Malayan,or even Old. World types, such as Euplcca, among Danaidse;Amatliusia, Chrome, and Thaumantis, among Morphidse ; Euripus,Diadema, Athyma, Limenitis, and Adolias, among Nymphalidse,Zemeros and Taxila among Erycinidse; Amblypodia, Miletus,Ilerda, and Myrina, among Lycasnidte ;Thyca, Prioneris, Dercas,Iphias, and <strong>The</strong>stias among Pieridse ; and Papilios <strong>of</strong> <strong>the</strong>" Amphrisius," " Coon," " Philoxenus," " Protenor" " Paris," and" Sarpedon " groups. In <strong>the</strong> Himalayas <strong>the</strong>re is an unusualabundance <strong>of</strong> large and gorgeous species <strong>of</strong> <strong>the</strong> genus Papilio,


chap, xii.] THE ORIENTAL REGION. 333and <strong>of</strong> large and showy Nymphalidae, Morphidae, and Danaidae,which render it, in favoured localities, only second to SouthAmerica for a display <strong>of</strong> this form <strong>of</strong> beauty and variety ininsect life.Among <strong>the</strong> o<strong>the</strong>r orders <strong>of</strong> insects in which <strong>the</strong> Himalayasare remarkably rich, we may mention large and brilliant Cetoniidae,cliiefly <strong>of</strong> <strong>the</strong> genus Rhomborhima ;a magnificent Lamellicorn,Uuchirus macleayii, allied to <strong>the</strong> gigantic long-armedbeetle (E. longimanus) <strong>of</strong> Amboyna ; superb moths <strong>of</strong> <strong>the</strong>families Agaristidas and Sesiidae ; elegant and remarkable Fulgoridae,and strange forms <strong>of</strong> <strong>the</strong> gigantic Phasmidae ;most <strong>of</strong>which appear to be <strong>of</strong> larger size or <strong>of</strong> more brilliant coloursthan <strong>the</strong>ir Malayan allies.Islands <strong>of</strong> <strong>the</strong> Indo-Chinese Sub-region.—A few importantislands belong to this sub-region, <strong>the</strong> Andamans, Formosa, andHainan being <strong>the</strong>most interesting.Andamans.—<strong>The</strong> only mammalia are a few rats and mice, aParadoxurus, and a pig supposed to be a hybrid race,— all <strong>of</strong>which may have been introduced by man's agency. <strong>The</strong>birds <strong>of</strong> <strong>the</strong> Andaman Islands have been largely collected, noless than 155 species having been obtained ; and <strong>of</strong> <strong>the</strong>se 17,(all land-birds) are peculiar. <strong>The</strong> genera are all found on <strong>the</strong>continent, and are mostly characteristic <strong>of</strong> <strong>the</strong> Indo-Chinesefauna, to which most <strong>of</strong> <strong>the</strong> species belong. Eeptiles are alsotolerably abundant ; about 20 species are known, <strong>the</strong> majoritybeing found also on <strong>the</strong> continent, while a few are peculiar.<strong>The</strong>re are also a few Batrachia, and some fresh-water fishes, closelyresembling those <strong>of</strong> Burmah. <strong>The</strong> absence <strong>of</strong> such mammaliaas monkeys and squirrels, which abound on <strong>the</strong> mainland, andwhich are easily carried over straits or narrow seas by floatingtrees, is sufficient pro<strong>of</strong> that <strong>the</strong>se islands have not recentlyformed part <strong>of</strong> <strong>the</strong> continent. <strong>The</strong> birds are mostly such asmay have reached <strong>the</strong> islands while in <strong>the</strong>ir present <strong>geographical</strong>position ;and <strong>the</strong> occurrence <strong>of</strong> reptiles and fresh-water fishes,said to be identical in species <strong>with</strong> those <strong>of</strong> Burmah, must bedue to <strong>the</strong> facilities, which some <strong>of</strong> <strong>the</strong>se <strong>animals</strong> undoubtedly


'334 ZOOLOGICAL GEOGRAPHY. [part hi.possess, for passing over a considerable width <strong>of</strong> sea.We mustconclude, <strong>the</strong>refore, that <strong>the</strong>se islands do not owe <strong>the</strong>ir existingfauna to an actual union <strong>with</strong> <strong>the</strong> mainland ; but it is probablethat <strong>the</strong>y may have been formerly more extensive, andhave <strong>the</strong>n been less distant from <strong>the</strong> continent than at <strong>the</strong>present time.<strong>The</strong> Nicobar Islands, usually associated <strong>with</strong> <strong>the</strong> Andamans,are less known, but present somewhat similar phenomena. <strong>The</strong>yare, however, more Malayan in <strong>the</strong>ir fauna, and seem properlyto belong to <strong>the</strong> Indo-Malay sub-region.Formosa.—This island has been carefully examined by Mr.Swinhoe, who found 144 species <strong>of</strong> birds, <strong>of</strong> which 34 are peculiar.<strong>The</strong>re is one peculiar genus, but <strong>the</strong> rest are all Indo-Chinese,though some <strong>of</strong> <strong>the</strong> species are more allied to Malayan than toChinese or Himalayan forms. About 30 species <strong>of</strong> mammaliawere found in Formosa, <strong>of</strong> which 11 are peculiar species, <strong>the</strong> restbeing ei<strong>the</strong>r Chinese or Himalayan. <strong>The</strong> peculiar species belongto <strong>the</strong> genera Taipei, Helictis, Sciuropterus, Pteromys, Mus, Sus,Cervus, and Capricornis. A few lizards and snakes <strong>of</strong> continentalspecies have also been found.<strong>The</strong>se facts clearly indicate<strong>the</strong> former connection <strong>of</strong> Formosa <strong>with</strong> China and Malaya, aconnection which is rendered <strong>the</strong> more probable by <strong>the</strong> shallowsea which still connects all <strong>the</strong>se countries.Hainan.—<strong>The</strong> island <strong>of</strong> Hainan, on <strong>the</strong> south coast <strong>of</strong> China,is not so well known in proportion, though Mr. Swinhoe collected172 species <strong>of</strong> birds, <strong>of</strong> which 130 were land-birds. Of<strong>the</strong>se about 20 were peculiar species ;<strong>the</strong> remainder being ei<strong>the</strong>rChinese, Himalayan, or Indo-Malayan. Mr. Swinhoe also obtained24 species <strong>of</strong> mammalia, all being Chinese, Himalayan,or Indo-Malayan species except a hare, which is peculiar. Thisassemblage <strong>of</strong> <strong>animals</strong> would imply that Hainan, as might beanticipated from its position,has been more recently separatedfrom <strong>the</strong> continent than <strong>the</strong> more distant island <strong>of</strong> Formosa.IV. Indo-Malaya, or <strong>the</strong> Malayan Sub-region.This sub-region, which is almost wholly insular (includingonly <strong>the</strong> Malayan peninsula on <strong>the</strong> continent <strong>of</strong> Asia), is equal, if


chap, xil.] THE ORIENTAL REGION. 335not superior, in <strong>the</strong> variety and beauty <strong>of</strong> its productions, to thatwhich we have just been considering. Like Indo-China, it is a region<strong>of</strong> forests, but it is more exclusively tropical ; and it is <strong>the</strong>reforedeficient in many <strong>of</strong> those curious forms <strong>of</strong> <strong>the</strong> temperatezone <strong>of</strong> <strong>the</strong> Himalayas, which seem to have been developed fromPalsearctic ra<strong>the</strong>r than from Oriental types.Here alone, in <strong>the</strong>Oriental region, are found <strong>the</strong> most typical equatorial forms <strong>of</strong>lifeorganisms adapted to a climate characterised by uniform butnot excessive heat, abundant moisture, and no marked departurefrom <strong>the</strong> average meteorological state, throughout <strong>the</strong> year. <strong>The</strong>sefavourable conditions <strong>of</strong> life only occur in three widely separateddistricts <strong>of</strong> <strong>the</strong> globe—<strong>the</strong> Malay archipelago, Western Africa,and equatorial South America. Hence perhaps it is, that <strong>the</strong>tapir and <strong>the</strong> trogons <strong>of</strong> Malacca should so closely resemble those<strong>of</strong> South America ;and that <strong>the</strong> great anthropoid apes and crestedhornbills <strong>of</strong> Western Africa, should find <strong>the</strong>ir nearest allies inBorneo and Sumatra.Although <strong>the</strong> islands which go to form this sub -regionare <strong>of</strong>ten separated from each o<strong>the</strong>r by a considerable expanse<strong>of</strong> sea, yet <strong>the</strong>ir productions in general <strong>of</strong>fer no greaterdifferences than those <strong>of</strong> portions <strong>of</strong> <strong>the</strong> Indo-Chinese subregionseparated by an equal extent <strong>of</strong> dry land. <strong>The</strong> explanationis easy, however, when we find that <strong>the</strong> sea whichseparates <strong>the</strong>m is a very shallow one, so shallow that an elevation<strong>of</strong> only 300 feet would unite Sumatra, Java, and Borneo intoone great South-eastern prolongation <strong>of</strong> <strong>the</strong> Asiatic continent-As we know that our own country has been elevated and depressedto a greater amount than this, at least twice in recentgeological times, we can have no difficulty inadmitting similarchanges <strong>of</strong> level in <strong>the</strong> Malay archipelago, where <strong>the</strong> subterraneanforces which bring about such changes are still atwork, as manifested by <strong>the</strong> great chain <strong>of</strong> active volcanoes inSumatra and Java.Pro<strong>of</strong>s <strong>of</strong> somewhat earlier changes <strong>of</strong> levelare to be seen in <strong>the</strong> Tertiary coal formations <strong>of</strong> Borneo, whichdemonstrate a succession <strong>of</strong> elevations and subsidences, <strong>with</strong> asmuch certainty as if we had historical record <strong>of</strong> <strong>the</strong>m.It is not necessary to suppose, nor is it probable, that all <strong>the</strong>se


336 ZOOLOGICAL GEOGRAPHY. l^rt hi.great islands were recently united to <strong>the</strong> continent, and that<strong>the</strong>ir separation took place by one general subsidence <strong>of</strong> <strong>the</strong>whole. It is more consonant <strong>with</strong> what we know <strong>of</strong> such matters,that <strong>the</strong> elevations and depressions were partial, varyingin <strong>the</strong>ir points <strong>of</strong> action and <strong>of</strong>ten recurring ; sometimes extendingone part <strong>of</strong> an island, sometimes ano<strong>the</strong>r ; now joiningan island to <strong>the</strong> main land, now bringing two islands intocloserproximity. <strong>The</strong>re is reason to believe that sometimes an interveningisland has sunk or receded and allowed o<strong>the</strong>rs which itbefore separated to effect a partial union independently <strong>of</strong> it. Ifwe recognise <strong>the</strong> probability that such varied and <strong>of</strong>ten-renewedchanges <strong>of</strong> level have occurred, we shall be better able to understandhow certain anomalies <strong>of</strong> <strong>distribution</strong> in <strong>the</strong>se islands mayhave been brought about. We will now endeavour to sketch <strong>the</strong>general features <strong>of</strong> <strong>the</strong> zoology <strong>of</strong> this interesting district, and<strong>the</strong>n proceed to discuss some <strong>of</strong> <strong>the</strong> <strong>relations</strong> <strong>of</strong> <strong>the</strong> islands toeach o<strong>the</strong>r.Mammalia.—We have seen that <strong>the</strong> Indo-Chinese sub-regionpossesses 13 species <strong>of</strong> mammalia in common <strong>with</strong> <strong>the</strong> Indo-Malay sub-region, and 4 o<strong>the</strong>rs peculiar to itself, besides oneEthiopian and several Oriental and Palsearctic forms <strong>of</strong> widerange. Of this latter class <strong>the</strong> Malay islands have comparativelyfew, but <strong>the</strong>y possess no less than 14 peculiar genera, viz.Simia, Siamanga, Tarsius, Galcopi<strong>the</strong>cus, Hylomys, Ptilocerns,Gymnura, Cynogale, Hemigalea, Arctogale, Barangia, Mydans,Hdarctos, and Tapirus. <strong>The</strong> islands also possess tigers, deer,wild pigs, wild cattle, elephants, <strong>the</strong> scaly ant-eater, and most<strong>of</strong> <strong>the</strong> usual Oriental genera ; so that <strong>the</strong>y are on <strong>the</strong> wholefully as rich as, if not richer than, any part <strong>of</strong> Asia ;a fact veryunusual in island faunas, and very suggestive <strong>of</strong> <strong>the</strong>ir reallycontinental nature.Plate VIII. Scene in Borneo <strong>with</strong> Characteristic MalayanQuadrupeds.—<strong>The</strong> Malayan fauna is so rich and peculiar thatwe devote two plates to illustrate it. We have here a group <strong>of</strong>mammalia, such as might be seen toge<strong>the</strong>r in <strong>the</strong> vast forests <strong>of</strong>Borneo. In <strong>the</strong> foreground we have <strong>the</strong> beautiful deer-likeChevrotain [Tragulus javanicus). <strong>The</strong>se are delicate little


PLATE VIII.A FOREST IN BORNEO, WITH CHARACTERISTIC MAMMALIA.


;chap, xii.] THE ORIENTAL REGION. 337<strong>animals</strong> whose body is not larger than a rabbit's, <strong>the</strong>nce <strong>of</strong>tencalled " mouse-deer." <strong>The</strong>y were formerly classed <strong>with</strong> <strong>the</strong>" musk-deer," owing to <strong>the</strong>ir similar tusk-like upper caninesbut <strong>the</strong>ir anatomy shows <strong>the</strong>m to form quite a distinct family,having more resemblance to <strong>the</strong> camels. On <strong>the</strong> branch aboveis <strong>the</strong> curious fea<strong>the</strong>r-tailed Tree-Shrew (Ptilocerus lowii), a smallinsectivorous animal altoge<strong>the</strong>r peculiar to Borneo.Above thisis <strong>the</strong> strange little Tarsier {Tarsius spectrum), one <strong>of</strong> <strong>the</strong> lemursconfined to <strong>the</strong> Malay islands, but so distinct from all o<strong>the</strong>rs asto constitute a separate family. <strong>The</strong> o<strong>the</strong>r small <strong>animals</strong> are<strong>the</strong>Flying Lemurs (Galceopi<strong>the</strong>cus volans) formerly classed <strong>with</strong><strong>the</strong> lemurs, but now considered to belong to <strong>the</strong> Insectivora.<strong>The</strong>y have a very large expansion <strong>of</strong> <strong>the</strong> skin connecting <strong>the</strong>fore and hind limbs and tail, and are able to take long flightsfrom one tree to ano<strong>the</strong>r, and even to rise over obstacles in <strong>the</strong>ircourse by <strong>the</strong> elevatory power <strong>of</strong> <strong>the</strong> tail-membrane. <strong>The</strong>yfeed chiefly on leaves, and have a very s<strong>of</strong>t and beautifullymarbled fur.In <strong>the</strong> distance is <strong>the</strong> Malayan tapir (Tapirus indicus), arepresentative <strong>of</strong> a group <strong>of</strong> <strong>animals</strong> now confined to <strong>the</strong> largerMalay islands and tropical America, but which once rangedover <strong>the</strong> greater part <strong>of</strong> temperate Europe.Birds.—Owing to several <strong>of</strong> <strong>the</strong> families consisting <strong>of</strong> veryobscure and closeby alliedspecies, which have never been criticallyexamined and compared by a competent ornithologist,<strong>the</strong> number <strong>of</strong> birds inhabiting this sub-region is uncertain.From <strong>the</strong> best available materials <strong>the</strong>re appear to be somewhatless than 650 species <strong>of</strong> land-birds actually known, or excluding<strong>the</strong> Philippine Islands somewhat less than 600. <strong>The</strong>larger part <strong>of</strong> <strong>the</strong>se are peculiar species, but mostly allied tothose <strong>of</strong> Indo-China ; 36 <strong>of</strong> <strong>the</strong> genera, as already stated, beingcommon to <strong>the</strong>se two sub-regions. <strong>The</strong>re are, however, no lessthan 46 genera which are peculiarly or wholly Indo Malayanand, in many cases, have no close affinity <strong>with</strong> o<strong>the</strong>r Orientalgroups. <strong>The</strong>se peculiar genera are as follows :— Timalia, Malacopteron,Macronus, Napoilwra,Turdinus, and Trichixos—genera


—;;338 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>of</strong> Timaliidse ;Eupetes, a most remarkable form, perhaps alliedto Enicurus, and Cinclus ;Rhabclomis (Certhiidae) found only in<strong>the</strong> Philippines; Psaltria, a diminutive bird <strong>of</strong> doubtful affinities,provisionally classed among <strong>the</strong> tits(Paridae); form's (Pycnonotidse);Lalagc (Campephagidse) extending eastward to <strong>the</strong> PacificIslands ;Pycnosphrys, Philcntoma (Muscicapidae) ;Lanicllus,a beautiful bird doubtfully classed <strong>with</strong> <strong>the</strong>Platylophus and Pityriasis, <strong>the</strong>shrikes (Laniidse)latter a most anomalous formperhaps a distinct family, at present classed <strong>with</strong> <strong>the</strong> jays, inCorvidse ; Prionochihis, a curious form classed <strong>with</strong> Dicseidse ;Eryihrura (Ploceidse), extending eastwards to <strong>the</strong> Fiji Islands;Oymnops, Calornis, (Sturnidse); Eurylcemus, Corydon, and Calyptomena(Eurylcemidse) ;Eucichla, <strong>the</strong> longest tailed and mostelegantly marked <strong>of</strong> <strong>the</strong> Pittidee ;Rdnwardtipicus and Miglyptes(Picidse) ;Psilopogon and Calorhamphus, (Megalaemida?) ; Rhinococcyx,Dasylophus, Lepidogrammus, Cdrpococcyx, Zanclostomus,Poliococcyx, Phinortha, (Cuculidse) ;Berenicornis, Caldo, Cranorhinas,Pcnclopidcs, Rhinopldx, (Bucerotidre) ; Psittinus, (Psittacidae); Ptilopus, Phapitreron, (Columbidae) ; Rollulus, (Treronidre); Machccrhamphus, (Falconida?). Many <strong>of</strong> <strong>the</strong>se genera areabundant and wide-spread, while some <strong>of</strong> <strong>the</strong> most characteristicHimalayan genera, such as Larvivora, Garrulax, Hypsipetes,Pomatorhinus, and Dcndrocitta, are here represented by onlya few species.Among <strong>the</strong> groups that are characteristic <strong>of</strong> <strong>the</strong> Malayansub-region, <strong>the</strong> Timaliidse and Pycnonotida? stand pre-eminent; <strong>the</strong>former represented chiefly by <strong>the</strong> genera Timalia, Malacopteron,Macronus, and Trichastoma, <strong>the</strong> latter by Criniger, Microscelis,and many forms <strong>of</strong> Pycnonotus. <strong>The</strong> Muscicapidse, Dicruridse,Campephagidffi, Ploceidse, and Nectariniidse are also welldeveloped ; as well as <strong>the</strong> Pittidse, and <strong>the</strong> Euryleemidae, <strong>the</strong>limited number <strong>of</strong> species <strong>of</strong> <strong>the</strong> latter being compensated bya tolerable abundance <strong>of</strong> individuals. Among <strong>the</strong> Picarioe aremany conspicuous groups ; as, woodpeckers (Picidae) ; barbets(Megalaemidse) ; trogons (Trogonidse) ; kingfishers (Alcedinida3)and hornbills (Bucerotidpe) ; five families which are perhaps <strong>the</strong>most conspicuous in <strong>the</strong> whole fauna.Lastly come <strong>the</strong> pigeons


chap, xii.] THE OEIENTAL REGION. 339(Colunibida?), and <strong>the</strong> pheasants (Phasianidae), which are fairlyrepresented by such fine genera as Treron, Ptilopus, Euplocamus,and Argusianus. A few forms whose affinities are Australianra<strong>the</strong>r than Oriental, help to give a character to <strong>the</strong> ornithology,though none <strong>of</strong> <strong>the</strong>m are numerous. <strong>The</strong> swallow-shrikes(Artamus); <strong>the</strong> wag-tail fly-catchers (Rhipidura); <strong>the</strong> green fruitdoves(Ptilopus) ;and <strong>the</strong> mound-makers (Mcgapodius), are <strong>the</strong>chief <strong>of</strong> <strong>the</strong>se.<strong>The</strong>re are a few curious examples <strong>of</strong> remote <strong>geographical</strong>alliances that may be noted. First, we have a direct Africanconnection in Machairhamphus, a genus <strong>of</strong> hawks, and Berenicornis,a genus <strong>of</strong> hornbills ; <strong>the</strong> only close allies being, in <strong>the</strong> formercase in South, and in <strong>the</strong> latter in West Africa. <strong>The</strong>n we havea curious Neotropical affinity, indicated by Garpococcyx, a largeBornean ground-cuckoo, whose nearest ally is <strong>the</strong> genus Neomorphus<strong>of</strong> South America; and by <strong>the</strong> lovely green-colouredCalyptomena which seems unmistakably allied to <strong>the</strong> orano-ecolouredRupicola, or " Cock <strong>of</strong> <strong>the</strong> rock," in general structureand in <strong>the</strong> remarkable form <strong>of</strong> crest, a resemblance which hasbeen noticed by many writers.In <strong>the</strong> preceding enumeration <strong>of</strong> Malayan genera severalare included which extend into<strong>the</strong> Austro-Malay Islands, ourobject, at present, being to show <strong>the</strong> differences and <strong>relations</strong> <strong>of</strong><strong>the</strong> two chief Oriental sub-regions.Plate IX, A Malayan Forest <strong>with</strong> some <strong>of</strong> its peculiar Birds.—Our second illustration <strong>of</strong> <strong>the</strong> Malayan fauna is devoted to itsbird-life ;and for this purpose we place our scene in <strong>the</strong> Malaypeninsula, where birds are perhaps more abundant and moreinteresting, than in any o<strong>the</strong>r part <strong>of</strong> <strong>the</strong> sub-region. Conspicuousin <strong>the</strong> foreground is <strong>the</strong> huge Ehinoceros Hornbill(Buceros rhinoceros), one <strong>of</strong> <strong>the</strong> most characteristic birds <strong>of</strong> <strong>the</strong>Malayan forests, <strong>the</strong> flapping <strong>of</strong> whose wings, as it violentlybeats <strong>the</strong> air to support its heavy body, may be heard a mile<strong>of</strong>f. On <strong>the</strong> ground behind, is <strong>the</strong> Argus pheasant (Argusianusgiganteus) whose beautifully ocellated wings have been<strong>the</strong> subject <strong>of</strong> a most interesting description in Mr. Darwin'sDescent <strong>of</strong> Man. <strong>The</strong> wing-fea<strong>the</strong>rs are here so enormously


340 ZOOLOGICAL GEOGRAPHY. [pabt hi.developed for display (as shown in our figure) that <strong>the</strong>ybecome almost, if not quite, useless for <strong>the</strong>ir original purpose <strong>of</strong>flight ;yet <strong>the</strong> colours are so sober, harmonizing completely<strong>with</strong> <strong>the</strong> surrounding vegetation, and <strong>the</strong> bird is so wary, thatin <strong>the</strong> forests where it abounds an old hunter assured me he hadnever been able to see a specimen till it was caught in hissnares. It is interesting to note, that during <strong>the</strong> display <strong>of</strong> <strong>the</strong>plumage <strong>the</strong> bird's head is concealed by <strong>the</strong> wings from aspectator in front, and, contrary to what usually obtains amongpheasants, <strong>the</strong> head is entirely unadorned, having nei<strong>the</strong>r crestnor a particle <strong>of</strong> vivid colour,—a remarkable confirmation <strong>of</strong>Mr. Darwin's views, that gayly coloured plumes are developedin <strong>the</strong> male bird for <strong>the</strong> purpose <strong>of</strong> attractive display in<strong>the</strong> breeding season. <strong>The</strong> long-tailed bird on <strong>the</strong> right isone <strong>of</strong> <strong>the</strong> Drongo-shrikes (Bhringa remifer), whose long baretail-fea<strong>the</strong>rs, <strong>with</strong> an oar-like web at <strong>the</strong> end, and blueblaekglossy plumage, render it a very attractive object as itflies after its insect prey. On <strong>the</strong> left is ano<strong>the</strong>r singular bird<strong>the</strong> great Broad-bill (Corydon sumatranus), <strong>with</strong> dull and sombreplumage, but <strong>with</strong> a beak more like that <strong>of</strong> a boat-bill than <strong>of</strong> afruit-eating passerine bird. Over all, <strong>the</strong> white-handed Gibbon(Hylobates lar) swings and gambols among <strong>the</strong> topmost branches<strong>of</strong> <strong>the</strong> forest.Reptiles and Amphibia.—<strong>The</strong>se are notsufficiently known tobe <strong>of</strong> much use for our present purpose. Most <strong>of</strong> <strong>the</strong> generabelong to <strong>the</strong> continental parts <strong>of</strong> <strong>the</strong> Oriental region, or have awide range. Of snakes Bhabdosoma, Typhlocalamus, Tetragonosoma,Acrochordus, and Atropos, are <strong>the</strong> most peculiar, and <strong>the</strong>reare several peculiar genera <strong>of</strong> Homalopsidse. Of Oriental genera,Gylindrophis, Xenopeltes, Calamaria, Hypsirhina, Psammodynastcs,Gonyosoma, Tragops, Dipsas, Pareas, Python, Bungarus,Naja, and Callophis are abundant; as well as Simotcs, Ablabes.Tropidonotus, and Dendrophis, which are widely distributed.Among lizards Hydrosaurus and Gecko are common; <strong>the</strong>re aremany isolated groups <strong>of</strong> Scincidse; while Draco, Calotes, andmany forms <strong>of</strong> Agamidse, some <strong>of</strong> which are peculiar, abound.Among <strong>the</strong> Amphibia, toads and frogs <strong>of</strong> <strong>the</strong> genera Micrhyla,


PLATE IX.A MALAYAN FOREST, WITH SOME OF ITS PECULIAR BIRDS.


chap, xii.] THE ORIENTAL REGION. 341Kalophrynus, Ansonia, and Pscudobufo, are peculiar :while <strong>the</strong>Oriental Mcgalophrys, Ixalus, Ehac&phorus, and Hylorana areabundant and characteristic.Fishes.—<strong>The</strong> fresh-water fishes <strong>of</strong> <strong>the</strong> Malay archipelagohave been so well collected and examined by <strong>the</strong> Dutchnaturalists, that <strong>the</strong>y <strong>of</strong>fer valuable indications <strong>of</strong> zoo-<strong>geographical</strong>affinity ; and <strong>the</strong>y particularly well exhibit <strong>the</strong>sharply defined limits <strong>of</strong> <strong>the</strong> region, a large number <strong>of</strong> Orientaland even Ethiopian genera extending eastward as far as Javaand Borneo, but very rarely indeed sending a single speciesfur<strong>the</strong>r east, to Celebes or <strong>the</strong> Moluccas. Thirteen families <strong>of</strong>fresh-water fishes are found in <strong>the</strong> Indo-Malay sub-region. Of<strong>the</strong>se <strong>the</strong> Scienidse and Symbranchidae have mostly a widerange in <strong>the</strong> tropics. Ophiocephalidse are exclusively Oriental,reaching Borneo and <strong>the</strong> Philippine islands. <strong>The</strong> Mastacembelidseare also Oriental, but one species is found as far as Ceram.Of <strong>the</strong> ISTandidse, 3 genera range over <strong>the</strong> whole region. <strong>The</strong>Labyrinthici extend from Africa through <strong>the</strong> Oriental region toAmboyna. <strong>The</strong> single species constituting <strong>the</strong> family Luciocephalidaeisconfined to Borneo and <strong>the</strong> small islands <strong>of</strong> Bilitonand Banca. Of <strong>the</strong> extensive family Siluridse 17 genera areOriental and Malayan, and 11 are Malayan exclusively; andnot one <strong>of</strong> <strong>the</strong>se appears to pass beyond <strong>the</strong> limits <strong>of</strong> <strong>the</strong> subregion.<strong>The</strong> Cyprinidse <strong>of</strong>fer an equally striking example, 23genera ranging eastward to Java and Borneo and not onebeyond; 14 <strong>of</strong> <strong>the</strong>se being exclusively Malayan. It must beremembered that this is not from any want <strong>of</strong> knowledge <strong>of</strong> <strong>the</strong>countries far<strong>the</strong>r east, as extensive collections have also beenmade in Celebes, <strong>the</strong> Moluccas, and Timor ; so that <strong>the</strong> facts <strong>of</strong><strong>distribution</strong> <strong>of</strong> fresh-water fishes come, most unexpectedly, t<strong>of</strong>ortify that division <strong>of</strong> <strong>the</strong> archipelago into two primaryregions, which was founded on a consideration <strong>of</strong> mammaliaand birds only.Insects.—Few countries in <strong>the</strong> world can present a richer andmore varied series <strong>of</strong> insects than <strong>the</strong> Indo-Malay islands, andwe can only here notice a few <strong>of</strong> <strong>the</strong>ir more striking peculiaritiesand more salient features.


;342 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>The</strong> butterflies <strong>of</strong> this sub-region, according to <strong>the</strong> best estimatethat can be formed, amount to about 650 described species,a number that will yet, no doubt, be very considerably increased.<strong>The</strong> genera which appear to be peculiar to it are Erites (Satyridse); Zeuxidia (Morphida?) ; Amnosia, Xanthotcenia, andTancccia (Nymphalidre). <strong>The</strong> groups which are most characteristic<strong>of</strong> <strong>the</strong> region, ei<strong>the</strong>r from <strong>the</strong>ir abundance in individualsor species, or from <strong>the</strong>ir size and beauty, are—<strong>the</strong> rich darkcolouredEuplcea ; <strong>the</strong> large semi-transparent Hcstia; <strong>the</strong> plaincolouredMycalesis, which replace our meadow-brown butterflies{Hipparchia) ;<strong>the</strong> curious Elymnias, which <strong>of</strong>ten closely resembleEupla?as ; <strong>the</strong> large and handsome Thamantis and Zeuxidia,which take <strong>the</strong> place <strong>of</strong> <strong>the</strong> giant Morphos <strong>of</strong> South America<strong>the</strong> Cethosia, <strong>of</strong> <strong>the</strong> brightest red, and marked <strong>with</strong> a curiouszigzag pattern ; <strong>the</strong> velvety and blue-glossed Terinos ; <strong>the</strong> paleand delicately-streaked Cyrcstis; <strong>the</strong> thick-bodied and boldlycoloured Adolias ; <strong>the</strong> small wine-coloured Taxila ; <strong>the</strong> fine blueAmblypodia ; <strong>the</strong> beautiful Thyca, elegantly marked underneath<strong>with</strong> red and yellow, which representour common whitebutterflies and are almost equally abundant ; <strong>the</strong> pale blueEronia, and <strong>the</strong> large red-tipped Iphias. <strong>The</strong> genus Papiliois represented by a variety <strong>of</strong> fine groups ; <strong>the</strong> large Ornithoptera,<strong>with</strong> satiny yellow under -wings ; <strong>the</strong> superb green-marked" brookeana ; " <strong>the</strong> " paradoxa" group, <strong>of</strong>ten closely resembling<strong>the</strong> Euplfleas that abound in <strong>the</strong> same district ; <strong>the</strong> " paris "group richly dusted <strong>with</strong> golden-green specks ; <strong>the</strong> " helenus "group <strong>with</strong> wide-spreading black and white wings ; <strong>the</strong> blackand crimson " polydorus " group ; <strong>the</strong> " memnon " group, <strong>of</strong> <strong>the</strong>largest size and richly-varied colours ;and <strong>the</strong> " eurypilus "group, elegantly banded or spotted <strong>with</strong> blue or green :all <strong>the</strong>seare so abundant that some <strong>of</strong> <strong>the</strong>m are met <strong>with</strong> in every walk,and are a constant delight to <strong>the</strong> naturalist who has <strong>the</strong> privilege<strong>of</strong> observing <strong>the</strong>m in <strong>the</strong>ir native haunts.<strong>The</strong> Coleoptera are far lessprominent and require to be carefullysought after ; but <strong>the</strong>y <strong>the</strong>n well repay <strong>the</strong> collector. Asaffording some measure <strong>of</strong> <strong>the</strong> productiveness <strong>of</strong> <strong>the</strong> tropics ininsect life it will not be out <strong>of</strong> place togive a few notes <strong>of</strong> <strong>the</strong>


chap, xii.] THE ORIENTAL REGION. 343number <strong>of</strong> species collected by myself in some <strong>of</strong> <strong>the</strong> bestlocalities. At Singapore 300 species <strong>of</strong> Coleoptera were collectedin 15 days, and in a month <strong>the</strong> number had increased to520 ; <strong>of</strong> which 100 were Longicorns and 140 Ehyncophora. AtSarawak in Borneo I obtained 400 species in 15 days, and600 in a month. In two months this number had increased toabout 850, and in three months to 1,000 species.This was <strong>the</strong>most prolific spot I ever collected in, especially for Longicornswhich formed about one-fifth <strong>of</strong> all <strong>the</strong> species <strong>of</strong> beetles. In<strong>the</strong> Aru Islands in one month, I obtained only 235 species <strong>of</strong>Coleoptera, and about 600 species <strong>of</strong> insects <strong>of</strong> all orders ; andthis may be taken as a fair average, in localities where nospecially favourable conditions existed. On <strong>the</strong> average 40 to60 species <strong>of</strong> Coleoptera would be a good day's collecting ; 70exceptionally good ; while <strong>the</strong> largest number ever obtained inone day was 95, and <strong>the</strong> majority <strong>of</strong> <strong>the</strong>se would be veryminute insects. It must be remembered, however, that manyvery common species were passed over, yet had every species met<strong>with</strong> been collected, not much more than 100 species would everhave been obtained in one day'scollecting <strong>of</strong> four or five hours.<strong>The</strong>se details may afford an interesting standard <strong>of</strong> comparisonfor collectors in o<strong>the</strong>r parts <strong>of</strong> <strong>the</strong> world.Of Cicindelidse <strong>the</strong> most peculiarly Malayan form is<strong>The</strong>rates,found always on leaves in <strong>the</strong> forests in <strong>the</strong> same localities as<strong>the</strong> more widely spread Collyris.Indo-Malayan.Five genera <strong>of</strong> this family are<strong>The</strong> Carabidse, though sufficiently plentiful, are mostly <strong>of</strong> smallsize, and not conspicuous in any way.But <strong>the</strong>re is one strikingexceptionin <strong>the</strong> purely Malayan genus Mormolyce, <strong>the</strong> largestand most remarkable <strong>of</strong> <strong>the</strong> whole family. It is nocturnal,resting during <strong>the</strong> days on <strong>the</strong> under side <strong>of</strong> large boleti in <strong>the</strong>virgin forest. Pericallus and Catascopus are among <strong>the</strong> fewgenera which are at all brillantly coloured.Buprestidse are abundant, and very gay ; <strong>the</strong> genus Belionotabeing perhaps one <strong>of</strong> <strong>the</strong> most conspicuousand characteristic.<strong>The</strong> giant Catoxantha is, however, <strong>the</strong> most peculiar, thoughcomparatively scarce. Chrysochroa and Chalcophora are also


344 ZOOLOGICAL GEOGRAPHY. [part hi.abundant and characteristic. Out <strong>of</strong> <strong>the</strong> 41 Oriental genera 21are Malayan, and 10 <strong>of</strong> <strong>the</strong>se are not found in <strong>the</strong> o<strong>the</strong>r subregions.In Lucanidae <strong>the</strong> Malay islands are rich, 14 out <strong>of</strong> <strong>the</strong> 16Oriental genera occurring <strong>the</strong>re, and 3 being peculiar. <strong>The</strong>reare many fine species <strong>of</strong> Odontolabris, which may be considered<strong>the</strong> characteristic genus <strong>of</strong> <strong>the</strong> sub-region.<strong>The</strong> Cetoniidae are well represented by 16 genera and about 120species. <strong>The</strong> genera Mycteristes, Phccdimus, Plectrone, Euremina,Rhagopteryx and Ccntrognathus are peculiar, while Agestrata,Ghalcothca, and Macronota are abundant and characteristic.<strong>The</strong> Longicorns, as in all continental forest regions near <strong>the</strong>equator, are very abundant and in endlessly varied forms. Noless than 55 genera containing about 200 species are peculiar tothis sub-region, <strong>the</strong> Cerarnbycidse being much <strong>the</strong> most numerous.Euryarthrum, Coelosterna, Ayelasta,and Asta<strong>the</strong>s may be consideredas most characteristic ; but to name <strong>the</strong> curious and interestingforms would be to give a list <strong>of</strong> half <strong>the</strong> genera.For<strong>the</strong> <strong>relations</strong> <strong>of</strong> <strong>the</strong> Longicorns <strong>of</strong> <strong>the</strong> Indo-Malay, and those <strong>of</strong><strong>the</strong> Austro-Malay region, <strong>the</strong> reader is referred to <strong>the</strong> chaptef on<strong>the</strong> <strong>distribution</strong> <strong>of</strong> insects in <strong>the</strong> succeeding part <strong>of</strong> this work.Terrestrial Mollusca.—<strong>The</strong> Philippine islands are celebratedas being one <strong>of</strong> <strong>the</strong> richest parts <strong>of</strong> <strong>the</strong> world for land shells,about 400 species being known. <strong>The</strong> o<strong>the</strong>r islands <strong>of</strong> <strong>the</strong> subregionare far less rich, not more than about 100 species havingyet been described from <strong>the</strong> whole <strong>of</strong> <strong>the</strong>m. Helix and Bulimusboth abound in species in <strong>the</strong> Philippines, whereas <strong>the</strong>latter genus is very scarce in Borneo and Java.Ten genera <strong>of</strong>Helicidse inhabit <strong>the</strong> sub-region ; Pfeifferia is found in <strong>the</strong>Philippines and Moluccas, while <strong>the</strong> large genus Cochlostyla isalmost peculiar to <strong>the</strong> Philippines.Of <strong>the</strong> Operculata <strong>the</strong>re arerepresentatives <strong>of</strong> 20 genera, <strong>of</strong> which Dermatoma and Pupinellaare peculiar, while Registoma and Callia extend to <strong>the</strong>Australian region. Cycloplwrus, Leptopoma, and Pupina areperhaps <strong>the</strong> most characteristic genera.


—chap, xii.] THE ORIENTAL REGION.. 345<strong>The</strong> Zoological Relations <strong>of</strong> <strong>the</strong> Several Islands <strong>of</strong> <strong>the</strong> Inch-MalaySub -region.Although we have grouped <strong>the</strong> Philippine islands <strong>with</strong> <strong>the</strong>Indo-Malay sub-region, to which, as we shall see, <strong>the</strong>y undoubtedlybelong, yet most <strong>of</strong> <strong>the</strong> zoological characteristics wehave just sketched out, apply more especially to <strong>the</strong> o<strong>the</strong>r groups<strong>of</strong> islands and <strong>the</strong> Malay peninsula. <strong>The</strong> Philippine islandsstand, to Malaya proper, in <strong>the</strong> same relation that Madagascardoes to Africa or <strong>the</strong> Antilles to South America ; that is, <strong>the</strong>yare remarkable for <strong>the</strong> absence <strong>of</strong> whole families and generawhich everywhere characterise <strong>the</strong> remainder <strong>of</strong> <strong>the</strong> district.<strong>The</strong>y are, in fact, truly insular, while <strong>the</strong> o<strong>the</strong>r islands are reallycontinental in all <strong>the</strong> essential features <strong>of</strong> <strong>the</strong>irnatural history.Before, <strong>the</strong>refore, we can conveniently compare <strong>the</strong> separateislands <strong>of</strong> Malaya 1<strong>with</strong> each o<strong>the</strong>r, we must first deal <strong>with</strong> <strong>the</strong>Philippine group, showing in what its speciality consists, andwhy it must be considered apart from <strong>the</strong> sub-region to whichit belongs.Mammals <strong>of</strong> <strong>the</strong> Philippine Islands.—<strong>The</strong> only mammalia recordedas inhabiting <strong>the</strong> Philippine Islands are <strong>the</strong> following :Quadrumana. 1. Macacus cynomolgus.2. Cynopi<strong>the</strong>cus niger. Dr. Semper doubts this beinga Philippine species.Lemuroidea. 3. Tarsius spectrum.Insectivora. 4. Galeopi<strong>the</strong>cus philippinensis.5. Tupaia (species). On Dr. Semper's authority.Carnivora. 6. Viverra tangalunga.7. Paradoxurus philippensis.Ungulata. 8. Sus (species). On Dr. Semper's authority.9. Cervus mariannus.10. Cervus philippensis.11. Cervus alfredi.12. Bos (species). "Wild cattle ;perhaps introduced.Rodentia. 13. Phlseonrys cummingii.14. Scuirus philippinensis.Also 24 species, belonging to 17 genera, <strong>of</strong> bats.1As so many typical Malay groups are absent only from <strong>the</strong> Philippines, Ihave adopted <strong>the</strong> term " Malaya," to show <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong>se, using<strong>the</strong> term " Indo-Malaya" when <strong>the</strong> range <strong>of</strong> <strong>the</strong> group includes <strong>the</strong>Philippines.This must be remembered when consulting <strong>the</strong> tables <strong>of</strong> <strong>distribution</strong>at <strong>the</strong> end <strong>of</strong> this chapter.Vol. I.—24.


346 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>The</strong> foregoing list, although small, contains an assemblage <strong>of</strong>species which are wholly Oriental in character, and several <strong>of</strong>which {Tardus, Galeopi<strong>the</strong>cus, Tupaia) are characteristic andhighly peculiar Malayan forms. At <strong>the</strong> same time <strong>the</strong>se islandsare completely separated from <strong>the</strong> rest <strong>of</strong> Malaya by <strong>the</strong> totalabsence <strong>of</strong> Semnopi<strong>the</strong>cus, Hylobates, Fclis,Hclarctos, Rhinoceros,Manis, and o<strong>the</strong>r groups constantly found in <strong>the</strong> great Indo-Malay islands and peninsula <strong>of</strong> Malacca. We find apparentlytwo sets <strong>of</strong> <strong>animals</strong> : a more ancient series, represented by <strong>the</strong>deer, Galeopi<strong>the</strong>cus, and squirrel, in which <strong>the</strong> species are distinctfrom any o<strong>the</strong>rs ; and a more recent series, represented byMacacus cynomolgus, and Viverra tangalunga, identical <strong>with</strong>common Malayan <strong>animals</strong>. <strong>The</strong> former indicate <strong>the</strong> earliestperiod when <strong>the</strong>se volcanic islands were connected <strong>with</strong> somepart <strong>of</strong> <strong>the</strong> Malayan sub-region, and <strong>the</strong>y show that this wasnot geologically remote, since no peculiar generic types havebeen preserved or differentiated. <strong>The</strong> latter may indicate ei<strong>the</strong>r<strong>the</strong> termination <strong>of</strong> <strong>the</strong> period <strong>of</strong> union, or merely <strong>the</strong> effects <strong>of</strong>introduction by man. <strong>The</strong> reason why a larger number <strong>of</strong>mammalian forms were not introduced and established, wasprobably because <strong>the</strong> union was effectedonly <strong>with</strong> some smallislands, and from <strong>the</strong>se communicated to o<strong>the</strong>r parts <strong>of</strong> <strong>the</strong>archipelago ; or it may well be that later subsidences extinguishedsome <strong>of</strong> <strong>the</strong> forms that had established <strong>the</strong>mselves.Birds <strong>of</strong> <strong>the</strong> Philippine Islands.— <strong>The</strong>se have been carefullyinvestigated by Viscount Walden, in a paper read before <strong>the</strong>Zoological Society <strong>of</strong> London in 1873, and we are thus furnished<strong>with</strong> ample information on <strong>the</strong> <strong>relations</strong> <strong>of</strong> this importantportion <strong>of</strong> <strong>the</strong> fauna.<strong>The</strong> total number <strong>of</strong> birds known to inhabit <strong>the</strong> Philippinesis 219, <strong>of</strong> which 106 are peculiar. If, however, following ourusual plan, we take only <strong>the</strong> land-birds, we find <strong>the</strong> numbersto be 159 species, <strong>of</strong> which 100 are peculiar ;an unusually largeproportion for a group <strong>of</strong> islands so comparatively near tovarious parts <strong>of</strong> <strong>the</strong> Oriental and Australian regions. <strong>The</strong>families <strong>of</strong> birds which are more especially characteristic <strong>of</strong> <strong>the</strong>Indo-Malay sub-region are about 28 in number, and examples


—chap, xii.] THE ORIENTAL REGION. 347<strong>of</strong> all <strong>the</strong>se are found in <strong>the</strong> Philippines except four, viz., Cinclidse,Phyllornithidse, Eurylsemidae, and Podargidse. <strong>The</strong> onlyMalayan are, Cacatuidse and Megapodiidae. Yet although <strong>the</strong>birds are unmistakably Malayan, as a whole, <strong>the</strong>re are, as in<strong>the</strong> mammalia (though in a less degree), marked deficiencies <strong>of</strong>most characteristic Malayan forms. Lord Walden gives a list<strong>of</strong> no less than 69 genera thus absent ; but it will be sufficien<strong>the</strong>re to mention such wide-spread and specially Indo-Malaygroupsas,Malacopteron,Eurylcemus, Nydiornis, Arachnotliera, Geocichla,Timalia, Pomatorhinus, Phyllornis, lora, CrinigerEnicurus, Chaptia, Tchitrea, Dendrocitta, Eulahes, Palceornis,Miglyptes, Tiga, and Euplocamus. <strong>The</strong>se deficiencies plainlyshow <strong>the</strong> isolated character <strong>of</strong> <strong>the</strong>that it has never formed a part <strong>of</strong> thatPhilippine group, and implyIndo-Malayan extension<strong>of</strong> <strong>the</strong> continent which almost certainly existed when <strong>the</strong>yPhilippine families which are, o<strong>the</strong>rwise, exclusively Austro-peculiarMalayan fauna was developed ; or that, if it has been sounited, it has been subsequently submerged and broken up tosuch an extent, as to cause <strong>the</strong> extinction <strong>of</strong> many <strong>of</strong> <strong>the</strong> absenttypes.It appears from Lord "Walden's careful analysis, that 31 <strong>of</strong> <strong>the</strong>Philippine species occur in <strong>the</strong> Papuan sub-region, and 47 inCelebes ;69 occur also in India, and 75 in Java. This last factiscurious, since Java is <strong>the</strong> most remote <strong>of</strong> <strong>the</strong> Malayan islands,but it is found to arise almost wholly from <strong>the</strong> birds <strong>of</strong> thatisland being better known, since only one species, Xantholcemarosea, is confined to <strong>the</strong> Philippine Islands and Java.<strong>The</strong> wading and swimming birds are mostly <strong>of</strong> wide-spreadforms, only 6 out <strong>of</strong> <strong>the</strong> 60 species being peculiar to <strong>the</strong> Philippinearchipelago. Confining ourselves to <strong>the</strong> land-birds, and combiningseveral <strong>of</strong> <strong>the</strong> minutely subdivided genera <strong>of</strong> Lord Walden'spaper so as to agree <strong>with</strong> <strong>the</strong> arrangement adopted in thiswork, we find that <strong>the</strong>re are 112 genera <strong>of</strong> land-birds representedin <strong>the</strong> islands. Of <strong>the</strong>se, 50 are eithsr cosmopolitan, <strong>of</strong>wide range, or common to <strong>the</strong> Oriental and Australian regions,and may be put aside as affording few indications <strong>of</strong> <strong>geographical</strong>affinity. Of <strong>the</strong> remaining 62 no less than 40 are exclusively


——348 ZOOLOGICAL GEOGRAPHY. [part hi.or mainly Oriental, and most <strong>of</strong> <strong>the</strong>m are genera which rangewidely over <strong>the</strong> region, only two (Philentoma and Rolhdus) beingexclusively Malayan, and two o<strong>the</strong>rs {Megalurus and Malacocircus)more especially Indian or continental. Five o<strong>the</strong>r genera, thoughhaving a wide range, are typically Palrearctic, and have reached<strong>the</strong> islands through North China. <strong>The</strong>y are, Monticola, Acrocephalus,Phylloscopus, Calliope, and Passer ; <strong>the</strong> two first havingextended <strong>the</strong>ir range southward into <strong>the</strong> Moluccas. <strong>The</strong> peculiarlyAustralian genera are only 12, <strong>the</strong> majority being characteristicPapuan and Moluccan forms ; such as Gampephaga,Alcyone, Cacatua, Tanygnathus, Ptilopus, Janthcenas, Phlogamas,and Megapodius. One is peculiar to Celebes (Prioniturus) ; oneto <strong>the</strong> Papuan group (Cyclopsitta) ; and one is chiefly Australian(Gerygone). <strong>The</strong> beautiful little parroquets forming <strong>the</strong> genusLoriculus, are characteristic <strong>of</strong> <strong>the</strong> Philippines, which possess 5species, a larger number than occurs in any o<strong>the</strong>r group <strong>of</strong>islands, though <strong>the</strong>y range from India to New Guinea. <strong>The</strong>re remainsix peculiar genera Bhabdornis, an isolated form <strong>of</strong> creepers(Certhiida?) : Gymnops, a remarkable bareheaded bird belongingto <strong>the</strong> starlings (Stumidae); Dasylophus, and Lepidogrammus,remarkable genera <strong>of</strong> cuckoos (Cuculidse) ; Penelopides, a peculiarhornbill, and Phapitreron, a genus <strong>of</strong>pigeons. Besides <strong>the</strong>se<strong>the</strong>re are four o<strong>the</strong>r types (here classed as sub-genera, but consideredto be distinct by Lord Walden) which are peculiar to<strong>the</strong> Philippines. <strong>The</strong>se are Pseudoptynx, an owl <strong>of</strong> <strong>the</strong> genusA<strong>the</strong>ne; Pseudolcdage, a sub-genus oiLalage; Zeocephus, a subgenusoiTchiirea;and Ptilocolpa, included under Garpophaga.When we look at <strong>the</strong> position <strong>of</strong> <strong>the</strong> Philippine group, connectedby <strong>the</strong> Bashee islands <strong>with</strong> Formosa, by Palawan and <strong>the</strong>Sooloo archipelago <strong>with</strong> Borneo, and by <strong>the</strong> Tulour and o<strong>the</strong>rislets <strong>with</strong> <strong>the</strong> Moluccas and Celebes, we have little difficulty inaccounting for <strong>the</strong> peculiarities <strong>of</strong> its bird fauna. <strong>The</strong> absence<strong>of</strong> a large number <strong>of</strong> Malayan groups would indicate that <strong>the</strong>actual connection <strong>with</strong> Borneo, which seems necessary for <strong>the</strong>introduction <strong>of</strong> <strong>the</strong> Malay types <strong>of</strong> mammalia, was not <strong>of</strong> longduration ; while <strong>the</strong> large proportion <strong>of</strong> wide-spread continentalgenera <strong>of</strong> birds would seem to imply that greater facilities had


chap, xii.1 THE ORIENTAL REGION. 349once existed for immigrationfrom Sou<strong>the</strong>rn China, perhaps bya land connection through Formosa, at which time <strong>the</strong> ancestors<strong>of</strong> <strong>the</strong> peculiar forms <strong>of</strong> deer entered <strong>the</strong> country. It may indeedbe objected that our knowledge <strong>of</strong> <strong>the</strong>se islands is fartoo imperfect to arrive at any satisfactory conclusions as to <strong>the</strong>irformer history ; but although many more species no doubtremain to be discovered, experience shows that <strong>the</strong> broad characters<strong>of</strong> a fauna are always determined by a series <strong>of</strong> collectionsmade by different persons, at various localities, and at differenttimes, even when more imperfect than those <strong>of</strong> <strong>the</strong>Philippinebirds really are. <strong>The</strong> isolated position, and <strong>the</strong> volcanic structure<strong>of</strong> <strong>the</strong> group, would lead us to expect <strong>the</strong>m to be somewhatless productive than <strong>the</strong> Moluccas, close to <strong>the</strong> rich and variedPapuan district,—or than Celebes, <strong>with</strong> its numerous indications<strong>of</strong> an extensive area and great antiquity; and taking into account<strong>the</strong> excessive poverty <strong>of</strong> its mammalian fauna, which is certainto be pretty well known, I am inclined to believe that no futurediscoveries will materially alter <strong>the</strong> character <strong>of</strong> Philippineornithology, as determined from <strong>the</strong> materials already at ourcommand.Java.—Following <strong>the</strong> same plan as we have adopted infirstdiscussing <strong>the</strong> Philippine islands, and separating <strong>the</strong>m from <strong>the</strong>body <strong>of</strong> <strong>the</strong> sub-region on account <strong>of</strong> special peculiarities, wemust next take Java, as possessing marked individuality, and asbeing to some extent more isolated in its productions than <strong>the</strong>remaining great islands.Java is well supplied <strong>with</strong> indigenousmammalia, possessingas nearly as can be ascertained 55 genera and 90 species.None<strong>of</strong> <strong>the</strong>se genera are peculiar, and only about 5 <strong>of</strong> <strong>the</strong> species,—3 quadrumana, a deer and a wild pig. So far <strong>the</strong>n <strong>the</strong>re isnothing remarkable in itsfauna, but on comparing it <strong>with</strong> that <strong>of</strong><strong>the</strong> o<strong>the</strong>r great islands, viz., Borneo and Sumatra, and <strong>the</strong> Malaypeninsula, we find an unmistakable deficiency <strong>of</strong> characteristicforms, <strong>the</strong> same in kind asthat we have just commented on in<strong>the</strong> case <strong>of</strong> <strong>the</strong> Philippines, though much less in degree. First,taking genera which are found in all three <strong>of</strong> <strong>the</strong> above-named


;350 ZOOLOGICAL GEOGRAPHY. [part hi.localities and which must <strong>the</strong>refore be held to be typical Malayangroups, <strong>the</strong> following are absent from Java : Viverra,Gyrmiopus, Lutra, Helarctos, Tapirus, Elephas, and Gymnura;while <strong>of</strong> those known to occur in two, and which, owing to ourimperfect knowledge, may very probably one day be discovered in<strong>the</strong> third, <strong>the</strong> following are equally wanting : Simia, Siamanga,Hemigalea, Paguma, Rhinoscinrus, and Rhizomys. It may besaid this is only negative evidence, but in <strong>the</strong> case <strong>of</strong> Java it ismuch more, because this island is not only <strong>the</strong> best known <strong>of</strong>any in <strong>the</strong> archipelago, but <strong>the</strong>re is perhaps no portion <strong>of</strong>British India <strong>of</strong> equal extent so well known. It is one <strong>of</strong> <strong>the</strong>oldest <strong>of</strong> <strong>the</strong> Dutch possessions and <strong>the</strong> seat <strong>of</strong> <strong>the</strong>ir colonialgovernment ;good roads traverse it in every direction, and experiencednaturalists have been resident in various parts <strong>of</strong>itfor years toge<strong>the</strong>r, and have visited every mountain and everyforest, aided by bands <strong>of</strong> diligent native collectors. Weshould be almost as likely to find new species <strong>of</strong> mammaliain Central Europe as in Java ; and <strong>the</strong>refore <strong>the</strong> absence <strong>of</strong>such <strong>animals</strong> as <strong>the</strong> Malay bear, <strong>the</strong> elephant, tapir, gymnura,and even less conspicuous forms, must be accepted as apositive fact.In <strong>the</strong> o<strong>the</strong>r islands <strong>the</strong>re are still vast tracts <strong>of</strong> forest in <strong>the</strong>hands <strong>of</strong> natives and utterly unexplored, and any similar absencein <strong>the</strong>ir case will prove little;yet on making <strong>the</strong> same comparisonin <strong>the</strong> case <strong>of</strong> Borneo, <strong>the</strong> most peculiar and <strong>the</strong> leastknown <strong>of</strong> <strong>the</strong> o<strong>the</strong>r portions <strong>of</strong> <strong>the</strong> sub-region, we find only2 genera absent which 'are found in <strong>the</strong> three o<strong>the</strong>r divisions,and only 3 which are found in two o<strong>the</strong>rs. A fact to be notedalso is, that <strong>the</strong> only genus found in Java but not in o<strong>the</strong>r parts<strong>of</strong> <strong>the</strong> sub-region (Helirtis) occurs again in North India; andthat some Javan species, as Rhinoceros javanicus, and Lepus kurgosaoccur again in <strong>the</strong> Indo-Chinese sub-region, but not in <strong>the</strong>Malayan.Among <strong>the</strong> birds we meet <strong>with</strong> facts <strong>of</strong> a similar importand though <strong>the</strong> absence <strong>of</strong> certain types from Java isnot quiteso certain as among <strong>the</strong> mammalia, this is more than balancedby <strong>the</strong> increased number <strong>of</strong> such deficiencies, so that if a few


—;chap, xii.] THE ORIENTAL REGION. 351shouid be proved to be erroneous, <strong>the</strong> main result will remainunaltered.Java possesses about 270 species <strong>of</strong> land birds, <strong>of</strong> which about40 are peculiar to it. <strong>The</strong>re are, however, very few peculiargenera, Laniellus, a beautiful spotted shrike, being <strong>the</strong> mostdistinct, while Cochoa and Psaltria are perhaps not different from<strong>the</strong>ir Indian allies. <strong>The</strong> island has however a marked individualityin two ways—in <strong>the</strong> absence <strong>of</strong> characteristic Malayantypes, and in <strong>the</strong> presence <strong>of</strong> a number <strong>of</strong> forms not yet foundin any <strong>of</strong> <strong>the</strong> o<strong>the</strong>r Malay islands, but having <strong>the</strong>ir nearest alliesin various parts <strong>of</strong> <strong>the</strong> Indo-Chinese sub-region.<strong>The</strong> following16 genera are all found in Malacca, Sumatra, and Borneo,but are absent from Java: Setomis, Temnurus, Dendrocitta,Corydon, Calyptomena, Venilia, Beinwardtipicus, Coloramplms,Bhinortha, Nydiomis, Cranorhinus, Psittinus, Polyplectron, Argusianus,Euplocamus, and Rolhdus. <strong>The</strong> following 9 are knownfrom two <strong>of</strong> <strong>the</strong> above localities, and will very probably befound in <strong>the</strong> third, but are absent from, and not likely tooccur in, Java : Trichixos, Eupetes, Melanochlora, Chaptia, Pityriasis,Lyncornis, Carpococcyx, Poliococcyx, and Ehinoplax. Wehave thus 25 typically Malayan genera which are not knownto occur in Java.<strong>The</strong> following genera, on <strong>the</strong> o<strong>the</strong>r hand, do not occur inany<strong>of</strong> <strong>the</strong> Malayan sub-divisions except Java, and <strong>the</strong>y all occuragain, or under closely allied forms, in <strong>the</strong> Indo-Chinese subregion: Brachypteryx (allied species in Himalayas) ; Zoo<strong>the</strong>raallied species in Aracan) ; Notodela (allied species in Pegu)Pnoepyga (allied species in Himalayas) ;Allotrius (allied speciesin <strong>the</strong> Himalayas) ; Cochoa (allied species in <strong>the</strong> Himalayas)Crypsirhina (allied species in Burmah) ; Estrilda (allied speciesin India) ;Psaltria (allied genusJZgithaliscus—in Himalayas)Pavo muticus and Harpactes oreskios (same species in Siamand Burmah) ; Cecropis striolata (same species in Java andFormosa, and allied species in India).Here we have 12 instances <strong>of</strong> very remarkable <strong>distribution</strong>,and considering that <strong>the</strong>re are nearly as many birds known fromSumatra and Borneo as from Java,and considerably more from


——352 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>the</strong> Malay peninsula, it is not likely that many <strong>of</strong> <strong>the</strong>se wellmarked forms will be discovered in <strong>the</strong>se countries.<strong>The</strong>re are also a considerable number <strong>of</strong> species <strong>of</strong> birdscommon to Malacca, Sumatra, and Borneo, but represented inJava by distinct though closely allied species. Such are,Venilia malaccensis (represented in Java by) V. miniata.Drymocataphus nigrocapitatus „ „ D. capistratusMalacopteron coronatum „M. rujifrons.Irena cyanea „Ploceus baya „Loriculus galgulus „Ptilopus jambu „I. turcosa.P. hypoxantha.L. pusillus.P. porphyreus.Now if we look at our map <strong>of</strong> <strong>the</strong> region, and consider <strong>the</strong>position <strong>of</strong> Java <strong>with</strong> regard to Borneo, Sumatra, and <strong>the</strong> Indo-Chinese peninsula, <strong>the</strong> facts just pointed out appear mostanomalous and perplexing. First, we have Java and Sumatraforming one continuous line <strong>of</strong> volcanoes, separated by a verynarrow strait, and <strong>with</strong> all <strong>the</strong> appearance <strong>of</strong> having formed onecontinuous land ;yet <strong>the</strong>ir productions differ considerably, andthose <strong>of</strong> Sumatra show <strong>the</strong> closest resemblance to those <strong>of</strong>Borneo, an island ten times fur<strong>the</strong>r <strong>of</strong>f than Java and differingwidely in <strong>the</strong> absence <strong>of</strong> volcanoes or any continuous range <strong>of</strong>l<strong>of</strong>ty mountains. <strong>The</strong>n again, not only does Java differ from<strong>the</strong>se two, but it agrees <strong>with</strong> a country beyond <strong>the</strong>m botha country from which <strong>the</strong>y seem to have a much better chanceto have been supplied by immigration than Java has, and tohave (almost necessarily)participated, even more largely, in <strong>the</strong>benefits <strong>of</strong> any means <strong>of</strong> transmission capable <strong>of</strong> reaching <strong>the</strong>latter island. Yet more; whatever changes have occurred tobring about <strong>the</strong> anomalous state <strong>of</strong> things that exists must havebeen, zoologically and geologically, recent ; for <strong>the</strong> strange crossaffinitiesbetween Java and <strong>the</strong> Indo-Chinese continent (inwhich Sumatra and Borneo have not participated), as well asthat between Malacca, Sumatra, and Borneo (in which Java hasnot participated) areexhibited, in many cases by community <strong>of</strong>species, in o<strong>the</strong>rs by <strong>the</strong> presence <strong>of</strong> very closely allied forms<strong>of</strong> <strong>the</strong> same genera, <strong>of</strong> mammalia and birds.Now we know that


chap, xii.] THE ORIENTAL REGION. 353<strong>the</strong>se higher <strong>animals</strong> become replaced by allied species muchmore rapidly than <strong>the</strong> mollusca; and it is also pretty certainthat <strong>the</strong> modification by which this replacement is effectedtakes place more rapidly when <strong>the</strong> two sets <strong>of</strong> individuals areisolated from each o<strong>the</strong>r, and especially when <strong>the</strong>y are restrictedto islands, where <strong>the</strong>y are necessarily subject to distinct andpretty constant conditions, both physical and organic. Itbecomes <strong>the</strong>refore almost a certainty, that Siam and Java on<strong>the</strong> one hand, and Sumatra, Borneo, and Malacca on <strong>the</strong> o<strong>the</strong>rmust have been brought into some close connexion, not earlierthan <strong>the</strong> newer Pliocene period ; but while <strong>the</strong> one set <strong>of</strong>countries were having <strong>the</strong>ir meeting, <strong>the</strong> o<strong>the</strong>r must have beenby some means got out <strong>of</strong> <strong>the</strong> way. Before attempting toindicate <strong>the</strong> mode by which this might have been effected inaccordance <strong>with</strong> what we know <strong>of</strong> <strong>the</strong> physical geography,geology, and vegetation <strong>of</strong> <strong>the</strong> several islands, it will be as wellto complete our sketch <strong>of</strong> <strong>the</strong>ir zoological <strong>relations</strong> to eacho<strong>the</strong>r, so as ascertain <strong>with</strong> some precision, what are <strong>the</strong> facts<strong>of</strong> <strong>distribution</strong> which we have to explain.Malacca, Sumatra, and Borneo.—After having set apart <strong>the</strong>Philippine Islands and Java, we have remaining two greatislands and a peninsula, which, though separated by considerablearms <strong>of</strong> <strong>the</strong> sea, possess a fauna <strong>of</strong> wonderful uniformityhaving all <strong>the</strong> typical Malayan features in <strong>the</strong>ir fulldevelopment. , <strong>The</strong>ir unity is indeed so complete, that we canfind hardly any groups <strong>of</strong> sufficient importance by which todifferentiate <strong>the</strong>m from each o<strong>the</strong>r ; and we feel no confidencethat future discoveries may not takeaway what speciality <strong>the</strong>ypossess. One after ano<strong>the</strong>r, species or genera once peculiar toBorneo or Sumatra have been found elsewhere; and this hasgone to such an extent in birds, that hardly a peculiar genusand very few peculiar species are left in ei<strong>the</strong>r island. Borneohowever is undoubtedly <strong>the</strong> most peculiar. It possesses threegenera <strong>of</strong> Mammalia not found elsewhere : Cynogale, a curiouscarnivore allied to <strong>the</strong> otters ; <strong>with</strong> Dendrogale and Ptilocerus,small insectivora allied to Tupaia. It has Simla, <strong>the</strong> Orang-


—354 ZOOLOGICAL GEOGRAPHY. [part iit.utan, and Paguma, one <strong>of</strong> <strong>the</strong> Viverridge, in common <strong>with</strong>Sumatra ;as well as Bhinosciurus, a peculiar form <strong>of</strong> squirrel, andHemigalea, one <strong>of</strong> <strong>the</strong> Viverridge, in common <strong>with</strong> Malacca.Sumatra has only one genus not found in any o<strong>the</strong>r Malayandistrictin North India.Nemorhedus, a form <strong>of</strong> antelope which occurs againIt also has Siamanga in common <strong>with</strong> Malacca,Mydaus <strong>with</strong> Java, and Rhizomys <strong>with</strong> India.<strong>The</strong> Malay Peninsulaseems to have no peculiar forms <strong>of</strong> Mammalia, thoughit is rich in all <strong>the</strong> characteristic Malay types.<strong>The</strong> hats <strong>of</strong> <strong>the</strong> various islands have been very unequallycollected, 36 species being recorded from Java, 23 from Sumatra,but only 16 each from Borneo and Malacca.Leaving <strong>the</strong>se out<strong>of</strong> consideration, and taking into account <strong>the</strong> terrestrial mammalsonly, we find that Java is <strong>the</strong> poorest in species, whileBorneo, Sumatra, and Malacca are tolerably equal ;<strong>the</strong> numbersbeing 55, 62, 66, and 65 respectively. Of <strong>the</strong>se we find that<strong>the</strong> species confined to each island or district are (in <strong>the</strong> sameorder) 6, 16, 5, and 6. It thus appears that Borneo is, in itsmammalia, <strong>the</strong> most isolated and peculiar ;next comes Sumatra,and <strong>the</strong>n Malacca and Java, as shown by <strong>the</strong> following table.PeculiarSpecies.16566This result differs from that which we have arrived atmore detailed consideration <strong>of</strong> <strong>the</strong> fauna <strong>of</strong> Java; and itby <strong>the</strong>servesto show that <strong>the</strong> estimate <strong>of</strong> a country by <strong>the</strong> number <strong>of</strong> itspeculiar genera and species alone, may not always represent itstrue zoological importance or its most marked features. Java,as we have seen, is differentiated from <strong>the</strong> o<strong>the</strong>r three districtsand byby <strong>the</strong> absence <strong>of</strong> numerous types common to <strong>the</strong>m all,its independent continental <strong>relations</strong>. Borneo is also well distinguishedby its peculiar genera and specific types, yet it is at<strong>the</strong> same time more closely related to Sumatra and Malaccathan is Java. <strong>The</strong> two islands have evidently had a verydifferent history, which a detailed knowledge <strong>of</strong> <strong>the</strong>ir geology


;chap, xii.] THE ORIENTAL REGION. 355would alone enable us to trace. Should we ever arrive at afair knowledge <strong>of</strong> <strong>the</strong> physical changes that have resulted in<strong>the</strong> present condition, we shall almost certainly find that many<strong>of</strong> <strong>the</strong> differences and anomalies <strong>of</strong> <strong>the</strong>ir existing fauna andflora will be accounted for.In Birds we hardly find anything to differentiate Borneo andSumatra in any clear manner.Pityriasis and Carpococcyx, oncethought peculiar to <strong>the</strong> former, are now found also in <strong>the</strong> latterand we have not a single genus left to characterize Borneo exceptScluvaneria a peculiar fly-catcher, and Indicator, an African andIndian group not known to occur elsewhere in <strong>the</strong> Malaysub-region.Sumatra as yet alone possesses Psil&pogon, a remarkableform <strong>of</strong> barbet, but we may well expect that itwill be soonfound in <strong>the</strong> interior <strong>of</strong> Borneo or Malacca ; it also has Berenicornis,an African form <strong>of</strong> hornbill. <strong>The</strong> Malay Peninsulaappears to have no genus peculiar to it, but it possessessome Chinese and Indian forms which do not pass into <strong>the</strong>islands.As to <strong>the</strong> species, our knowledge <strong>of</strong> <strong>the</strong>m is at presentvery imperfect. <strong>The</strong> Malay Peninsula is perhaps <strong>the</strong> bestknown, but it is probable that both Sumatra and Borneo arequite as rich in species. With <strong>the</strong> exception <strong>of</strong> <strong>the</strong> generanoted above, and two or three o<strong>the</strong>rs as yet found in two islandsonly, <strong>the</strong> three districts we arenow considering may be said tohave an almost identical bird-fauna, consisting largely <strong>of</strong> <strong>the</strong>same species and almost wholly <strong>of</strong> <strong>the</strong>se toge<strong>the</strong>r <strong>with</strong>closelyallied species <strong>of</strong> <strong>the</strong> same genera. <strong>The</strong>re are no well-markedgroups which especially characterise one <strong>of</strong> <strong>the</strong>se islands ra<strong>the</strong>rthan <strong>the</strong> o<strong>the</strong>r, so that even <strong>the</strong> amount <strong>of</strong> speciality whichBorneo undoubtedly exhibits as regards mammalia, is onlyfaintly shown by its birds. <strong>The</strong> Pittidae may perhaps be namedas <strong>the</strong> most characteristic Bornean group, that island possessingmost beautiful birds <strong>of</strong> an unusually beautiful family.six species, three <strong>of</strong> which are peculiar to it and are among <strong>the</strong>Yet Sumatrapossesses two peculiar, and hardly less remarkable species.In o<strong>the</strong>r classes <strong>of</strong> vertebrates, in insects, and in land-shells,our knowledge is far too imperfect touseful comparison between <strong>the</strong> faunas.allow <strong>of</strong> our making any


356 ZOOLOGICAL GEOGRAPHY. [part Hi,Banca.—We must, however note <strong>the</strong> fact <strong>of</strong> peculiar speciesoccurring in Banca, a small island close to Sumatra, and thus<strong>of</strong>fering ano<strong>the</strong>r problem in <strong>distribution</strong>. A squirrel (Sciurusbangkanus) is allied to three species found in Malacca, Sumatra,and Borneo respectively, but quite as distinct from <strong>the</strong>m all<strong>the</strong>y are from each o<strong>the</strong>r. More curious are <strong>the</strong> two species <strong>of</strong>Pitta peculiar to Banca ; one, Pitta megarhynchus, is allied to<strong>the</strong> P. brachyurus, which inhabitsas<strong>the</strong> whole sub-region and extendsto Siam and China, but differs from it in its very large billand differently coloured head ; <strong>the</strong> o<strong>the</strong>r, P. bangkanus, is alliedto P. cucullatus, which extends from Nepal to Malacca, and toP. sordidus, which inhabits both Borneo and Sumatra as well as<strong>the</strong> Philippines.We have here, on a small scale, asomewhat similar problemto that <strong>of</strong> Java, and as this is comparatively easy <strong>of</strong>solution wewill consider it first. Although, on <strong>the</strong> map, Banca is so veryclose to Sumatra, <strong>the</strong> observer on <strong>the</strong> spot at once sees that <strong>the</strong>proximity has been recently brought about. <strong>The</strong> whole sou<strong>the</strong>astcoast <strong>of</strong> Sumatra isa great alluvial plain, hardly yet raisedabove <strong>the</strong> sea level, and half flooded in <strong>the</strong> wet season. It isplainly a recent formation, caused by<strong>the</strong> washing down into ashallow sea <strong>of</strong> <strong>the</strong> debris from <strong>the</strong> grand range <strong>of</strong> volcanicmountains 150 miles distant. Banca, on <strong>the</strong> o<strong>the</strong>r hand is,though low, a rugged and hilly island, formedalmost wholly <strong>of</strong>ancient rocks <strong>of</strong> apparently volcanic origin, and closely resemblingparts <strong>of</strong> <strong>the</strong> Malay Peninsula and <strong>the</strong> intervening chain <strong>of</strong>small islands. <strong>The</strong>re is every appearance that Banca onceformed <strong>the</strong> extremity <strong>of</strong> <strong>the</strong> Peninsula, at which time it wouldprobably have been separated from Sumatra by 50 or 100 miles<strong>of</strong> sea.Its productions should, <strong>the</strong>refore, most resemble those <strong>of</strong>Singapore and Malacca, and <strong>the</strong> few peculiar speciesit possesseswill be due to <strong>the</strong>ir isolation in a small tract <strong>of</strong> country, surroundedby a limited number <strong>of</strong> animal and vegetable forms, andsubject to <strong>the</strong> influence <strong>of</strong> a peculiar soil and climate. <strong>The</strong>parent species existing in such large tracts as Borneo or Sumatra,subjected to more varied conditions <strong>of</strong> soil, climate,vegetation, food, and enemies, would preserve, almost or quite


chap, xii.] THE ORIENTAL REGION. 357unchanged, <strong>the</strong> characteristics which had been developed undernearly identical conditions when <strong>the</strong> great island formed part<strong>of</strong> <strong>the</strong> continent. Geology teaches us that similar changes in<strong>the</strong> forms <strong>of</strong> <strong>the</strong> higher vertebrates have taken place during <strong>the</strong>Post-Tertiary epoch ; and <strong>the</strong>re are o<strong>the</strong>r reasons for believingthat, under such conditions <strong>of</strong> isolation as in Banca, <strong>the</strong>change may have required but a very moderate period, evenreckoned in years. We will now return to <strong>the</strong> more difficultproblem presented by <strong>the</strong> peculiar continental <strong>relations</strong> <strong>of</strong> Java,as already detailed.Probable Recent Geographical Changes in <strong>the</strong> Indo-MalayIslands.—Although Borneo is by far <strong>the</strong> largest <strong>of</strong> <strong>the</strong> Indo-Malay islands, yet its physical conformation is such that, werea depression to occur <strong>of</strong> one or two thousand feet, it would bereduced to a smaller continuous area than ei<strong>the</strong>r Sumatra orJava. Except in its nor<strong>the</strong>rn portion it possesses no l<strong>of</strong>tymountains, while alluvial valleys <strong>of</strong> great extent penetrate faiinto its interior. A very moderate depression, <strong>of</strong> perhaps 500feet, would convert it into an island shaped something like Celebes;and its mountains are <strong>of</strong> so small an average elevation, andconsist so much <strong>of</strong> isolated hills and detached ranges, that adepression <strong>of</strong> 2,000 feet would almost certainly break it up intoa group <strong>of</strong> small islands, <strong>with</strong> a somewhat larger one to <strong>the</strong>north.Sumatra (and to a less extent Java) consists <strong>of</strong> an almostcontinuous range <strong>of</strong> l<strong>of</strong>ty mountains, connected by plateaus from3,000 to 4,000 feet high ; so that although a depression <strong>of</strong> 2,000feet would greatly diminish <strong>the</strong>ir size, it would probably leave<strong>the</strong> former a single island, while <strong>the</strong> latter would be separatedinto two principal islands <strong>of</strong> still considerable extent. <strong>The</strong> enormousamount <strong>of</strong> volcanic actionin <strong>the</strong>se two islands, and <strong>the</strong>great number <strong>of</strong> conical mountains which must have been slowlyraised, chiefly by ejected matter, to <strong>the</strong> height <strong>of</strong> 10,000 and12,000 feet, and whose shape indicates that <strong>the</strong>y have been formedabove water, renders it almost certain that for long periods<strong>the</strong>y have not undergone submersion to any considerable extent.In Borneo, however, we have no such evidences. No volcano,


358 ZOOLOGICAL GEOGRAPHY. [part in.active or extinct, is known in its entire area ; while extensivebeds <strong>of</strong> coal <strong>of</strong> tertiary age, in every part <strong>of</strong> it, prove that it hasbeen subject to repeated submersions, at no distant date geologically.An indication, if not a pro<strong>of</strong>, <strong>of</strong> still more recent submersionis to be found in <strong>the</strong> great alluvial valleys which on<strong>the</strong> south and south-west extend fully 200 miles inland, while<strong>the</strong>y are to a less degree a characteristic feature all round <strong>the</strong>island. <strong>The</strong>se swampy plains have been formed by <strong>the</strong> combinedaction <strong>of</strong> rivers and tides ; and <strong>the</strong>y point clearly to an immediatelypreceding state <strong>of</strong> things, when that which is even nowbarely raised above <strong>the</strong> ocean, was more or less sunk below it.<strong>The</strong>se various indications enable us toclaim, as an admissibleand even probable supposition, that at some epoch during <strong>the</strong>Pliocene period! <strong>of</strong> geology, Borneo, as we now know it, did notexist ;but was represented by a mountainous island at its presentnor<strong>the</strong>rn extremity, <strong>with</strong> perhaps a few smaller islets to <strong>the</strong>south. We thus have a clear opening from Java to <strong>the</strong> SiamesePeninsula ;and as <strong>the</strong> whole <strong>of</strong> that sea is less than 100 fathomsdeepj <strong>the</strong>re is no difficulty in supposing an elevation <strong>of</strong> landconnecting <strong>the</strong> two toge<strong>the</strong>r, quite independent <strong>of</strong> Borneo on <strong>the</strong>one hand and Sumatra on <strong>the</strong> o<strong>the</strong>r.This union did not probablylast long ; but it was sufficient to allow <strong>of</strong> <strong>the</strong> introductioninto Java <strong>of</strong> <strong>the</strong> Rhinoceros javanicus, and that group <strong>of</strong> Indo-Chinese and Himalayan species <strong>of</strong> mammalia and birds whichit alone possesses. When this ridge had disappeared by subsidence,<strong>the</strong> next elevation occurred a little more to <strong>the</strong> east,and produced <strong>the</strong> union <strong>of</strong> many islets which, aided by subaerialdenudation, formed <strong>the</strong> present island <strong>of</strong> Borneo. It isprobable that this elevation was sufficiently extensive to uniteBorneo for a time <strong>with</strong> <strong>the</strong> Malay Peninsula and Sumatra, thushelping to produce that close resemblance <strong>of</strong> genera and even <strong>of</strong>species, which <strong>the</strong>se countries exhibit, and obliterating muchot <strong>the</strong>ir former speciality, <strong>of</strong> which, however, we have stillsome traces in <strong>the</strong> long-nosed monkey and Ptilocerus <strong>of</strong>Borneo, and <strong>the</strong> considerable number <strong>of</strong> genera both <strong>of</strong> mammaliaand birds confined to two only out <strong>of</strong> <strong>the</strong>three divisions<strong>of</strong> typical Malaya. <strong>The</strong> subsidence which again divided <strong>the</strong>se


;chap, xii.] THE ORIENTAL REGION. 359countries by arms <strong>of</strong> <strong>the</strong> sea ra<strong>the</strong>r wider than at present,might have left Banca isolated, as already referred to, <strong>with</strong> itsproportion <strong>of</strong> <strong>the</strong> common fauna to be, in a few instancessubsequently modified.Thus we are enabled to understand how <strong>the</strong> special <strong>relations</strong><strong>of</strong> <strong>the</strong> species <strong>of</strong> <strong>the</strong>se islands to each o<strong>the</strong>r may have beenbrought about To account for <strong>the</strong>ir more deep-seated andgeneral zoological features, we must go far<strong>the</strong>r back.Probable Origin <strong>of</strong> <strong>the</strong> Malayan Fauna.—<strong>The</strong> typical Malayanfauna is essentially an equatorial one, and must have beenelaborated in an extensive equatorial area. This ancient landalmost certainly extended northward over <strong>the</strong> shallow sea as faras<strong>the</strong> island <strong>of</strong> Palawan, <strong>the</strong> Paracels shoals and even Hainan.To <strong>the</strong> east, it may at one time have included <strong>the</strong> Philippinesand Celebes, but not <strong>the</strong> Moluccas. To <strong>the</strong> south it was limitedby <strong>the</strong> deep sea beyond Java. It included all Sumatra and <strong>the</strong>Nicobar islands, and <strong>the</strong>re is every reason to believe that itstretched out also to <strong>the</strong> west so as to include <strong>the</strong> central peak<strong>of</strong> Ceylon, <strong>the</strong> Maldive isles, and <strong>the</strong> Cocos islands west <strong>of</strong>Sumatra. We should <strong>the</strong>n have an area as extensive as SouthAmerica to 15° south latitude, and well calculated to developthat luxuriant fauna and flora which has since spread to <strong>the</strong>Himalayas. <strong>The</strong> submergence <strong>of</strong> <strong>the</strong> western half <strong>of</strong> this area(leaving only a fragment in Ceylon) would greatly diminish <strong>the</strong>number <strong>of</strong> <strong>animals</strong> and perhaps extinguish some peculiar typesbut <strong>the</strong> remaining portion would still form a compact and extensivedistrict, twice as large as <strong>the</strong> peninsula <strong>of</strong> India, over <strong>the</strong>whole <strong>of</strong> which a uniform Malayan fauna would prevail.<strong>The</strong>first important change would be <strong>the</strong> separation <strong>of</strong> Celebes ;and':his was probably effected by a great subsidence, forming <strong>the</strong> deepstrait that now divides that island from Borneo. During <strong>the</strong>process Celebesitself was no doubt greatly submerged, leavingonly a few islands in which were preserved that remnant <strong>of</strong> <strong>the</strong>ancient Malayan fauna that now constitutes one <strong>of</strong> its moststriking and anomalous features. <strong>The</strong> Philippine area wouldnext be separated, and perhaps be almost wholly submerged ;or


;360 ZOOLOGICAL GEOGRAPHY. [past hi.broken up into many small volcanic islets in which a limitednumber <strong>of</strong> Malayan types alone survived. Such a condition <strong>of</strong>things will account for <strong>the</strong> very small variety <strong>of</strong> mammalia compared<strong>with</strong> <strong>the</strong> tolerably numerous genera <strong>of</strong> birds, that nowcharacterise its fauna ; while both here and in Celebes we findsome <strong>of</strong> <strong>the</strong> old Malayan types preserved, which, in <strong>the</strong> extendedarea <strong>of</strong> <strong>the</strong> Sunda Isleshave been replaced by more dominantforms.<strong>The</strong> next important change would be <strong>the</strong> separation <strong>of</strong> Javaand here also no doubt a considerable submergence occurred,rendering <strong>the</strong> island an unsuitable habitation for <strong>the</strong> variousMalay types whose absence forms one <strong>of</strong> its conspicuous features.It has since remained permanently separated from <strong>the</strong> o<strong>the</strong>rislands, and has no doubt developed some peculiar species, whileit may have preserved some ancient forms which in <strong>the</strong> largerarea have become changed. From <strong>the</strong> fact that a number <strong>of</strong> itsspecies are confined ei<strong>the</strong>r to <strong>the</strong> western or <strong>the</strong> eastern half <strong>of</strong><strong>the</strong> island, it is probable that it long continued as two islands,which have become united at a comparatively recent period.It has also been subjected to <strong>the</strong> immigration <strong>of</strong> Indo-Chineseforms, as already referred to in <strong>the</strong> earlier part <strong>of</strong> this sketch.We have thus shown how <strong>the</strong> main zoological features <strong>of</strong> <strong>the</strong>several sub-divisions <strong>of</strong> <strong>the</strong> Malayan sub-region may beaccounted for, by means <strong>of</strong> a series <strong>of</strong> suppositions as to pastchanges which, though for <strong>the</strong> most part purely hypo<strong>the</strong>tical,are always in accordance <strong>with</strong> what we know both <strong>of</strong> <strong>the</strong>physical geography and <strong>the</strong> zoology <strong>of</strong> <strong>the</strong> districts in questionand those which surround <strong>the</strong>m.It may also be remarked, thatwe know, <strong>with</strong> a degree <strong>of</strong> certainty which may be called absolute,that alternate elevation and subsidence is <strong>the</strong> normal state <strong>of</strong>things all over <strong>the</strong> globe ; that it was <strong>the</strong> rule in <strong>the</strong> earliestgeological epochs, and that it has continued down to <strong>the</strong>historical era. We know too, that <strong>the</strong> amount <strong>of</strong> elevation andsubsidence that can be proved to have occurred again and againin <strong>the</strong> same area, is <strong>of</strong>ten much greater than is required for <strong>the</strong>changes here speculated on,—while <strong>the</strong> time, required for suchchanges is certainly less than that necessitated by <strong>the</strong> changes


chap, xii.] THE ORIENTAL REGION. 361<strong>of</strong> specific and generic forms which have coincided <strong>with</strong>, and beento a large extent dependent on <strong>the</strong>m. We have, <strong>the</strong>refore, truecauses at work, and our only suppositions have been as to howthose causes could have brought about <strong>the</strong> results which wesee ; and however complex and unlikely some <strong>of</strong> <strong>the</strong> supposedchanges may seem to <strong>the</strong> reader, <strong>the</strong> geologist who has made a<strong>study</strong> <strong>of</strong> such changes, as recorded in <strong>the</strong> crust <strong>of</strong> <strong>the</strong> earth,will not only admit <strong>the</strong>m to be probable, but will be inclinedto believe that <strong>the</strong>y have really been far more complex andmore unexpected than any supposition we can make about<strong>the</strong>m.<strong>The</strong>re is one o<strong>the</strong>r external relation <strong>of</strong> <strong>the</strong> Malayan faunaabout which it may be necessary to say a few words. I havesupposed <strong>the</strong> greatestwestward extension <strong>of</strong> <strong>the</strong> Malayan areato be indicated by <strong>the</strong> Maldive islands, but some naturalistswould extend it to include Madagascar in order to account for<strong>the</strong> range <strong>of</strong> <strong>the</strong> Lemuridse.Such an extension would, however,render it difficult to explain <strong>the</strong> very small amount <strong>of</strong> correspondence<strong>with</strong> a pervading diversity, between <strong>the</strong> Malayan andMalagasy faunas. It seems more reasonable to suppose anapproximation <strong>of</strong> <strong>the</strong> two areas, <strong>with</strong>out actual union havingever occurred. This approximation would have allowed <strong>the</strong>interchange <strong>of</strong> certain genera <strong>of</strong> birds, which are common to<strong>the</strong> Oriental Eegion and <strong>the</strong> Mascarene islands, but it wouldhave been too recent to account for <strong>the</strong> diffusion<strong>of</strong> <strong>the</strong> lemurswhich belong to distinct genera and even distinct families.Thisprobably dates back to a much earlier period, when <strong>the</strong> lemurinetype had a wide range over <strong>the</strong> nor<strong>the</strong>rn hemisphere. Subjectedto <strong>the</strong> competition <strong>of</strong> higher forms, <strong>the</strong>se imperfectlydeveloped groups have mostly died out, except a few isolatedexamples, chiefly found in islands, and a few groups in Africa.In our discussion <strong>of</strong> <strong>the</strong> origin <strong>of</strong> <strong>the</strong> Ethiopian fauna, wehave supposed that a close connection once existed betweenMadagascar and Ceylon.This was during a very early tertiaryepoch ; and if, long after it had ceased and <strong>the</strong> fauna <strong>of</strong> Ceylonand South India had assumed somewhat more <strong>of</strong> <strong>the</strong>ir presentcharacter, we suppose <strong>the</strong> approximation or union <strong>of</strong> CeylonYol. I.—25


—302 ZOOLOGICAL GEOGRAPHY. [part hi.and Malaya to have taken place, we shall perhaps be able toaccount for most <strong>of</strong> <strong>the</strong> special affinities <strong>the</strong>y present, <strong>with</strong> <strong>the</strong>least amount <strong>of</strong> simultaneous elevation <strong>of</strong> <strong>the</strong> ocean bed ; whichit must always be remembered, requires a corresponding depressionelsewhere to balance it.Concluding Remarks on <strong>the</strong> Oriental Region.—We have alreadyso fully discussed <strong>the</strong> internal and external <strong>relations</strong> <strong>of</strong> <strong>the</strong>several sub-regions, that little more need be said. <strong>The</strong> rich andvaried fauna which inhabited Europe at <strong>the</strong> dawn <strong>of</strong> <strong>the</strong> tertiaryperiod,—as shown by <strong>the</strong> abundant remains <strong>of</strong> mammaliawherever suitable deposits <strong>of</strong> Eocene age have been discovered,proves, that an extensive Palsearctic continent <strong>the</strong>n existed;and <strong>the</strong> character <strong>of</strong> <strong>the</strong> floraisand fauna <strong>of</strong> <strong>the</strong> Eocene depositsso completely tropical, that we may be sure <strong>the</strong>re was <strong>the</strong>n nobarrier <strong>of</strong> climate between it and <strong>the</strong> Oriental region. At thatearly period <strong>the</strong> nor<strong>the</strong>rn plains <strong>of</strong> Asia were probably underwater, while <strong>the</strong> great Thibetan plateau and <strong>the</strong> Himalayan range,had not risen to more than a moderate height, and would havesupported a luxuriant sub*-tropical flora and fauna.<strong>The</strong> UpperMiocene deposits <strong>of</strong> nor<strong>the</strong>rn and central India, and Burmah,agree in <strong>the</strong>ir mammalian remains <strong>with</strong> those <strong>of</strong> central andsou<strong>the</strong>rn Europe, whileclosely allied forms <strong>of</strong> elephant, hysena,tapir, rhinoceros, and Chalico<strong>the</strong>rium have occurred in NorthChina; leading us to conclude that one great fauna <strong>the</strong>nextended over much <strong>of</strong> <strong>the</strong> Oriental and Palsearctic regions.Perim island at <strong>the</strong> mouth <strong>of</strong> <strong>the</strong> Eed Sea, where similarremains are found, probably shows <strong>the</strong> sou<strong>the</strong>rn boundary <strong>of</strong>this part <strong>of</strong> <strong>the</strong> old Palsearctic region in <strong>the</strong> Miocene period.Towards <strong>the</strong> equator <strong>the</strong>re would, <strong>of</strong> course, be some peculiargroups ; but we can hardly doubt, that, in that wonderful timewhen even <strong>the</strong> lands that stretched out fur<strong>the</strong>st towards <strong>the</strong>pole, supported a luxuriant forest vegetation, substantially onefauna ranged over <strong>the</strong> whole <strong>of</strong> <strong>the</strong> great eastern continent <strong>of</strong> <strong>the</strong>nor<strong>the</strong>rn hemisphere. During <strong>the</strong> Pliocene period, however, aprogressive change went on which resulted in <strong>the</strong> completedifferentiation <strong>of</strong> <strong>the</strong> Oriental and Palsearctic faunas. <strong>The</strong>


chap, xii.] THE ORIENTAL EEGION. 363causes <strong>of</strong> this change were <strong>of</strong> two kinds. <strong>The</strong>re was a great<strong>geographical</strong> and physical revolution effected by <strong>the</strong> elevation<strong>of</strong> <strong>the</strong> Himalayas and <strong>the</strong> Thibetan plateau, and, probably at<strong>the</strong> same time, <strong>the</strong> northward extension <strong>of</strong> <strong>the</strong> great Siberianplains. This alone would produce an enormous change <strong>of</strong>climate in all <strong>the</strong> extra-tropical part <strong>of</strong> Asia, and inevitablylead to a segregation <strong>of</strong> <strong>the</strong> old fauna into tropical and temperate,and a modification <strong>of</strong> <strong>the</strong> latter so as to enable it tosupport a climate far more severe than it had previously known.But it is almost certain that, concurrently <strong>with</strong> this, <strong>the</strong>re wasa change going on <strong>of</strong> acosmical nature, leading to an alteration<strong>of</strong> <strong>the</strong> climate <strong>of</strong> <strong>the</strong> nor<strong>the</strong>rn hemisphere from equable toextreme, and culminating in that period <strong>of</strong> excessive cold whichdrove <strong>the</strong> last remnants <strong>of</strong> <strong>the</strong> old sub-tropical fauna beyond<strong>the</strong> limits <strong>of</strong> <strong>the</strong> Palaearctic region. From that time, <strong>the</strong> Orientaland <strong>the</strong> Ethiopian regions alone contained <strong>the</strong> descendants <strong>of</strong> many<strong>of</strong> <strong>the</strong> most remarkable types which had previously flourishedover all Europe and Asia ; but <strong>the</strong> early history <strong>of</strong> <strong>the</strong>se tworegions, and <strong>the</strong> peculiar equatorial types developed in each,sufficiently separate <strong>the</strong>m, as we have already shown. <strong>The</strong>Malayan sub-region is tLat in which characteristic Orientaltypes are now best developed, and where <strong>the</strong> fundamental contrast<strong>of</strong> <strong>the</strong> Oriental, as compared <strong>with</strong> <strong>the</strong> Ethiopian andPaleearctic regions, is most distinctly visible.


364 ZOOLOGICAL GEOGEAPHY. [part in.TABLES OF DISTRIBUTION.In constructing <strong>the</strong>se tables, showing <strong>the</strong> <strong>distribution</strong> <strong>of</strong>various classes <strong>of</strong> <strong>animals</strong> in <strong>the</strong> Oriental region, <strong>the</strong> followingsources <strong>of</strong> information have been chiefly relied on, in additionto <strong>the</strong> general treatises, monographs, catalogues, &c, used for<strong>the</strong> compilation <strong>of</strong> <strong>the</strong> Fourth Part <strong>of</strong> this work.Mammalia.—Jerdon's Indian Mammalia ;Kelaart's Fauna <strong>of</strong>Ceylon; Horsfield and Moore's Catalogue <strong>of</strong> <strong>the</strong> East IndiaMuseum ; Swinhoe's Catalogue <strong>of</strong> Chinese Mammalia ; S.Miiller's Zoology <strong>of</strong> <strong>the</strong> Indian Archipelago ;Dr. J. E. Gray'slist <strong>of</strong> Mammalia <strong>of</strong> <strong>the</strong> Malay Archipelago (Voyage <strong>of</strong> Samarang); and papers by Anderson, Blyth, Cantor, Gray, Peters,Swinhoe, &c.Birds.—Jerdon's Birds <strong>of</strong> India ; Horsfield and Moore's Catalogue;Holdsworth's list <strong>of</strong> Ceylon Birds ; Schlegel's Catalogue<strong>of</strong> <strong>the</strong> Leyden Museum ; Swinhoe on <strong>the</strong> Birds <strong>of</strong> China, Formosa,and Hainan ; Salvadori on <strong>the</strong> Birds <strong>of</strong> Borneo ; LordWalden on <strong>the</strong> Birds <strong>of</strong> <strong>the</strong> Philippine Islands ; and papersby Blyth, Blanford, Elwes, Elliot, Stoliczka, Sclater, Sharpe,Swinhoe, Verreaux,and Lord Walden.Reptiles.—Giin<strong>the</strong>r's Eeptiles <strong>of</strong> British India; papers bysame author,and by Dr. Stoliczka.


chap, xii.] THE ORIENTAL EEGION. 365TABLE I.FAMILIES OF ANIMALS INHABITING THE ORIENTAL REGION.Explanation.Names in italics show families peculiar to <strong>the</strong> region.Numbers correspond <strong>with</strong> those in Part I V".Names enclosed thus ( ) barely enter <strong>the</strong> region, and are not considered really tobelong to it.Sub-regions.Order and Family.o .•a c.5-3•c.sE? Range beyond <strong>the</strong> Region.MAMMALIA.Primates.1. Simiidae2. Semnopi<strong>the</strong>cida?3. Cynopi<strong>the</strong>cidae6. Lemuridae7. TarsiidceW. AfricaTropical AfricaAll Africa, S. PalaearcticEthiopianCelebesChiroptera.9. Pteropidae11. Rhinolophidae12. Vespertilionidse13. Noctilionidae ...Ethiopian, Australian<strong>The</strong> Eastern HemisphereCosmopoliteTropical regionsInsectivora.14. Galeopi<strong>the</strong>cidce16. Tupaiidce17. Erinaceidse21. Talpidae22. SoricidaePalaearctic, S. AfricaPalaearctic, NearcticPalaearctic,Ethiopian, N. AmericaCarnivora.23. Felidae ...25. Viverridae27. HyaenidaeAll regions but AustralianEthiopian, S. PalaearcticEthiopian, S. Palaearctic28. Canidpe ...29. Mustelidse31. iEluridae ...32. Ursidae ...All regions but Australian [?]All regions but AustralianPalaearcticPalaearctic, Nearctic, ChiliCetacea.OceanicSlRENIA.42. ManatidaeEthiopian, N. PacificUngulata.43. (Equidae)...Palaearctic, Ethiopian


.i366 ZOOLOGICAL GEOGEAPHY. [PART III.Sub-regions.Order and Family.aoo =3•o.S'OeSO >-.S3Range beyond <strong>the</strong> Region.44. Tapiridae...45. Rinocerotidae47. Suidae ...49. Tragulidse50. Cervidae ...52. Bovidae53. ElephantidaeNeotropicalEthiopianPalaearctic, Ethiopian, NeotropicalW. AfricaAll regions but Ethiopian and AustralianAll regions but Australian and NeotropicalEthiopianRodentia.55. Muridae ...56. Spalacidae61. Sciuridse ...67. Hystricidse70. LeporidaeCosmopolite, excl. OceaniaPalaearctic, EthiopianAll regions but AustralianS. Palaearctic, EthiopianAll regions but AustralianEdentata.72. ManididseEthiopianBIRDS.Passeres.1. Turdidse2. Sylviidae3. Timaliidse4. Panuridae5. Cinclidae6. Troglodytidse .8. Certhiidae9. Sittidae ...10. Paridse.Almost CosmopoliteAlmost CosmopoliteEthiopian, AustralianPalaearcticNot Ethiopian or AustralianAmerican and PalaearcticPalaearctic, Nearctic, AustralianPalaearctic, Nearctic, Australian, Madagascar<strong>The</strong> Eastern Hemisphere and North America11. Liotrichidos ..12. Fhyllomithidce13. Pycnonotidse ..14. Oriolidae15. Campephagidae16. Dicruridae17. Muscicapida? ...18. Pachycephalidae19. Laniidae20. Corvidae23. Nectariniidae ...24. Dicaeidae30. Hirundinidae ...33. Fringillidae ...34. Ploceidae35. Sturnidse36. Artaniidae37. Alaudidae38. Motacillidae ...Ethiopian, Moluccas<strong>The</strong> Eastern HemisphereEthiopian, AustralianEthiopian, Australian<strong>The</strong> Eastern HemisphereAustralian<strong>The</strong> Eastern Hemisphere and North AmericaCosmopoliteEthiopian, AustralianEthiopian, Australian]CosmopoliteAll regions but AustralianEthiopian, Australian<strong>The</strong> Eastern HemisphereAustralianAll regions but NeotropicalCosmopolite43. Eurylannidce ...47. PittidaeEthiopian, Australian


CHAP. XII.] THE ORIENTAL REGION. 367Sub-regions.Order and Family.o .ri ad =3 1 *aw 75Range beyond <strong>the</strong> Region.PlCARiai.51. Picidae52. Yungidas53. Indicatoridae54. Megalaemidae58. Cuculidae62. Coraciidae63. Meropidae66. Trogonidae67. Alcedinidae68. Bucerotidae69. Upupidae71. Podargidae73. Caprimulgidae74. Cypselidae ...Psittaci.76. (Cacatuidffi) ...78. PalaeornithidaeColumba;.84. Columbidse ...Galling.86. Pteroclidae ...87. Tetraonidae ...88. Phasianidae89. Turnicidaj90. MegapodiidaeAll regions but AustralianPalaearcticEthiopianEthiopian, NeotropicalCosmopoliteEthiopian, AustralianEthiopian, AustralianNeotropical, EthiopianCosmopoliteEthiopian, Austro-MalayanEthiopian, S. PalaearcticAustralianCosmopoliteCosmopoliteAustralianEthiopian, Austro-MalayanCosmopoliteEthiopian, PalaearcticEastern Hemisphere and North AmericaEthiopian, Palaearctic, North AmericaEthiopian, Australian, S. PalaearcticAustralianAccipitres.94. Vulturidae ..96. Falconidae ...97. Pandionidae ...98. StrigidaeAll regions but AustralianCosmopoliteCosmopoliteCosmopoliteGrall.5:.99. Rallidae100. Scolopacidae103. Parridae...104. Glareolidae105. Charadriidae106. Otididae...107. Gruidae ..113. Ardeidaj114. Plataleidae115. Ciconiidae117. PhaenicopteridaeCosmopoliteCosmopoliteTropical regionsEastern HemisphereCosmopoliteEastern HemisphereAll regions but NeotropicaCosmopoliteAlmost CosmopoliteAlmost CosmopoliteEthiopian, Neotropical, S. Palaearctic


368 ZOOLOGICAL GEOGEAPHY. [PART III.


jCHAP. XII.THE ORIENTAL REGION. 369


370 ZOOLOGICAL GEOGRAPHY. [PART III.


,CHAP. XII.] THE ORIENTAL REGION. 371TABLEGENERA OF TERRESTRIAL MAMMALIA AND BIRDS INHABITINGTHE ORIENTAL REGION.II.Explanation.Names in italics show genera peculiar to <strong>the</strong> region.Names inclosed thus (...)show genera which just enter <strong>the</strong> region, but are not consideredproperly to belong to it.Genera truly belonging to <strong>the</strong> region are numbered consecutively.MAMMALIA.Order, Family, audGenus.3 °Range <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong>Region.PRIMATES.SlMIID^E.1. Simia2. Hylobates3. SiamangaBorneo and SumatraSylhet to Java and S. GhinaMalacca and SumatraSemnopi<strong>the</strong>cid.®.4. PresbytesCYNOPITHECID.E.5. Macacus6. Cynopi<strong>the</strong>cus .28221Simla to Aracan and E. Thibet,Ceylon, and Java<strong>The</strong> whole regionPhilippinesMoupin, Palsearctic [?]S. PalsearcticCelebes(Sub-Order)LEMUROIDEA.Lemtjkid^!.7. Nycticebus8. Loris ... .E. Bengal to Java, and S. ChinaCeylon and S. IndiaTAKSIID.E.9. TarsiusCHIROPTERA.Pteropim:.Sumatra, Borneo and PhilippinesN. Celebes10. Pteropus11. Xantharpyia .12. Cynopterus13. Megcerops ... .14. Macroglossus .15. Harpyia ... .<strong>The</strong> whole region<strong>The</strong> whole region<strong>The</strong> whole regionSumatraJava, Borneo, PhilippinesPhilippinesTropics <strong>of</strong> E. Hemisp.Austro-Malaya, Ethiop.S. PalsarcticTropical AfricaAustro-MalayaAustro-MalayaRhinolophid-s;.16. Aquias ... .2 Nepal to Java


372 ZOOLOGICAL GEOGRAPHY. [PART III.Order, Family, andGenus..'3o &"A ^Range <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.17. Phyllotis18. Rhinoloplius ...19. Hipposideros ...20. Phyllorhina ...21. Asellia22. Petalia23. Coslops24. Rhinovoma25. Megadernia26. Nycteris110Philippines<strong>The</strong> whole region<strong>The</strong> whole regionIndo-Malay subregionJava, SumatraJavaIndia (Bengal)All India<strong>The</strong> whole regionJavaWanner parts <strong>of</strong> E. Hem.Austro-MalayaAustro-Malaya, TropicalAfricaAmboyna, EgyptEgypt, PalestineTernate, N. EthiopianEthiopianVESPERTILIONIDiE.27. Scotophilus28. "Vespertilio29. Keriovula30. TrilatUus31. Noctulina32. Miniopteris33. Murina34. Nycticejus35. Harpiocephalus36. Taphozous37. Myotis38. Plecotus ..-. ...39. Barbastellus ..40. Nyctophilus ..10128233<strong>The</strong> whole region<strong>The</strong> whole region<strong>The</strong> whole regionIndo-MalayaNepal to PhilippinesJava, Philippines, and ChinaHimalayas to JavaAll IndiaJava and Philippines<strong>The</strong> whole regionHimalayasOarjeelingHimalayasMussoorieAustral., Nearc, Neotrop.CosmopoliteS. Africa, N. China??S. Africa, S. Palsearctic,Australian?Trop. Africa, Temp. Amer.Ethiop. , Austro-Malayan,NeotropicalTimor, S. PaleearcticEuropeAustralianNoctilionid^!.41. Chiromcles42. Nyctinomus ..Indo-Malaya, Siam<strong>The</strong> whole regionMadagascar, AmericaINSECTIVORA.GaLEOPITHECIDjE.43. Galeopi<strong>the</strong>cus ..TUPAIID.E.14. Tupaia45. Hylomys46. PtilocerusIndo-Malay and Philippines,excl. JavaS. and E. <strong>of</strong> India to BorneoTenasserim to Java and BorneoBorneoErinaceid^e.47. Erinaceus48. GymnuraHindostan and FormosaMalacca, Sumatra, BorneoPalsearctic, S. AfricaTALPID.E.49. TalpaHimalayas to Assam, & FormosaPalaearcticSoricid^:.50. Sorex20<strong>The</strong> whole regionAll regions but Austral,and S. America


CHAP. XII.] THE ORIENTAL REGION. 37^1Order, Family, andGenus.


374 ZOOLOGICAL GEOGRAPHY. [PAET III.Order, Family, andGenus.


CHAP. XII.] THE ORIENTAL REGION. 375Order, Family, andGenus.


376 ZOOLOGICAL GEOGRAPHY. [part hi.


CHAP. XII.] THE ORIENTAL REGION. 377Order, Family, andGenus.


:378 ZOOLOGICAL GEOGKAPHY. [PART III,Order, Family, andGenus.81. Siva82. Minla83. Proparus84. Allotrius85. Cutia86. Yuhina87. Ixulus88. Myzornis34Himalayas -.—3,000—7,000 ftNepal to E. Thibet ; moderateheightsN. W. Himalayas to E. Thibethi §hN. W. Himalayas to TenassenmE. Thibet and JavaNepal and SikhimHimalayas to E. Thibet, highDarjeeling to TenasserimNepal and SikhimPerhaps also PalsearcticPerhaps Palse arcticPhyllornithid^;.89. Phyllomis ...90. Iora91. Erpomis1052<strong>The</strong> whole region ; excludingChina and Philippines<strong>The</strong> whole reg. , excl. PhilippinesNepal and HainanPycnonotid^i.92. Microscelis93. Pycnonotus ..94. Hemixxis95. Hypsipetes ...96. Criniger97. Setomis98. MeOriolid^;.99. Oriolus100. AnalcipusCaMPEPHAGIDjE.101. Pcricrocotus ...102. Graucalus103. Campephaga ...104. Volvocivora ...105. Lalage106. Cochoa54021511341232271723Burmah, China, Malaya<strong>The</strong> whole regionHimalayas and Hainan<strong>The</strong> whole regionIndia, Ceylon, Malaya, HainanMalacca, Sumatra, BorneoAracan and Malaya<strong>The</strong> whole regionHimalayas, Malaya, Formosa,HainanJapanEthiopianMadagascarAfrica, MoluccasPahearc. Ethiopian, Celebes,Flores<strong>The</strong> whole regionLombock ;migrant<strong>the</strong> Amoor,India, Ceylon, Malaya, Philippines,AustralianHainan and FormosaPhilippine IslandsCelebes to N. Guinea<strong>The</strong> whole reg., excl. PhilippinesMalaya and PhilippinesCelebes to Pacific Is.Himalayas and JavaDicrurid^;.107. Dicrurus108. Bhringa109. Chibia110.. Chaptia111. IrenaMuscicapice.112. Muscicapula ..113. Erythrostema172133<strong>The</strong> whole regionHimalayas to Burmah and JavaIndia to ChinaIndia to Borneo and FormosaS. India and Ceylon, Assam toMalaya and PhilippinesCashmere to "W. China, S. India<strong>The</strong> whole region, excludingPhilippinesEthiop. and AustralianPekin in summerPalsearctic and Mada-gascar


—?nAP. xii.] THE ORIENTAL REGION. 379Order, Family, andGenus.114. Xanthpygia ...115. Hemipus116. Pycnophrys ...117. Heniiehelidon118. Niltava119. Cyornis120. Cyanoptila ...121. Eumyias122. Siphia123. Anthipes124. Schwaneria ...125. Hypothymis ...126. Rhipidura ..127. Chelidorkynx128. Cryptolopha ...21133141Malacca to ChinaIndia and CeylonJavaN. India to Ceylon, and China ;? PhilippinesHimalayas to W. China<strong>The</strong> whole regionHainan to Japan<strong>The</strong> whole reg. , excl. PhilippinesN. W. India, Ceylon, Formosa,E. ThibetNepalBorneo<strong>The</strong> whole regionAll India and Ceylon, Malaya,PhilippinesN. India<strong>The</strong> whole regionN. China and JapanEastern AsiaCelebes and TimorJapan and N. ChinaCelebesAustralianCelebes129. Tchitrea130. Philentoma ...PACHYCEPHALID.E.131. Hylocharis ...<strong>The</strong> whole regionMalaya and PhilippinesAracan to Malaya & PhilippinesN. China, and Japan,Flores, EthiopianCelebes, TimorLANIID.E.132. Lanius133. Lanielhcs . .134. Tephrodorms...CORVID-E.135. Pityriasis136. Platylophus ..137. Garrulus138. Cissa139. Urocissa140. Temnurus ...141. Dendrocitta ...142. Crypsirhina ...143. Nucifraga144. Pica145. Corvus(Fregilus1615<strong>The</strong> whole regionJavaIndia, Ceylon, andHanianMalaya ;Borneo, SumatraMalayaHimalayas, S. China, FormosaHimalayas and Aracan to JavaN. "VV. Himalayas, Ceylon, Burmah,China, FormosaMalaya and Cochin ChinaAll India to S. China, Formosa,and SumatraJava and BurmahHimalayas and E. Thibet ;8,000—10,000 feetChina and Himalayas <strong>of</strong> Bcetan<strong>The</strong> whole regionHimalayas, high)Nearc, Palsearc, Ethiop.PalsearcticN. China and JapanPalaearctic genusPalaearctic and NearcticCosmop., excl. S. Am.Palaearctic genusNECTARIXIID-ffil.146. ^Ethopaga147. Chalcostetha...148. Arachno<strong>the</strong>ra149. Arachnec<strong>the</strong>ra13112Himalayas to "W. China & Java,Central IndiaMalaya and Siam<strong>The</strong> whole reg., excl. Philippines<strong>The</strong> whole region, excl.ChinaCelebesCelebes to New GuineaCelebes, Lombock, NewGuineaCelebes to New Iceland


380 ZOOLOGICAL GEOGRAPHY. [PART III.Order, Family, andGenus.


.CHAP. XII.] THE ORIENTAL REGION. 381Order,Family, andGenus.c ^Range <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.ARTAMIDjE.180. Artamus<strong>The</strong> whole regionAustralianAlaudid^e.(Otocorys181. Alauda182. Galerita183. Calandrella ...(Melanocorypha184. Mirafra185. Ammomanes...186. Pyrrhulauda...N. India, in winter)India and ChinaCentral IndiaIndia and BurinahN. W. India)India, Ceylon, and JavaCentral IndiaIndia and CeylonPalaearctic and NearcticPalaearctic and EthiopianPalaearcticPalaearctic and EthiopianPalaearcticEthiopianPalaearctic and EthiopianEthiopianMOTACILLID^!.187. Motacilla188. Budytes189. Calobates190. Nemoricola ...191. Authus192. Corydalla193. Heterura62113India and Ceylon to China andPhilippinesChina and Philippines<strong>The</strong> whole regionIndia, Ceylon, and MalayaIndia and China<strong>The</strong> whole regionHimalayasPalaearctic and EthiopianPalaearctic & Ethiopian,MoluccasPalaearcticCosmopolitePalaearctic, AustralianEURYLEMID-E.194. Eurylocmm ...195. Serilophus196. Psarisomus ..197. Corydon198. Cymbirhynchus199. Calyptomcna . .MalayaHimalayasHimalayasMalacca, Sumatra, BorneoAracan, Siam, and MalayaMalacca, Sumatra, BorneoPlTTIDiE200. Pitta ...201. Eucichla202. Hydrornis1133<strong>The</strong> whole regionMalayaHimalayas and MalayaAustralian,EthiopianPICAKI.E.PlCIDiE.203. Vivia204. Sasia205. Picus206. Hyopicus207. Yungipicus ...208. Reinwardtipicus209. Vcnilia210. Chrysocolaptes211. Hcmicercus ...212. Gecinus21411212512N. W. Himalayas to E. Thibet,3,000-5.000 ft.Nepal to Malaya and Borneo<strong>The</strong> whole region, excl. PhilippinesHimalayas<strong>The</strong> whole regionPenang to Sumatra and BorneoNepal to Sumatra and BorneoIndia, Ceylon, Malaya, PhilippinesMalabar, Pegu to MalayaAll India and Ceylon to Peguand MalayaPalaearctic, AmericanN. ChinaN. China, Japan, CelebesPalaearctic


.<strong>The</strong>382 ZOOLOGICAL GEOGRAPHY. [PART III.Order, Family, andGenus.© QJ! olRange <strong>with</strong>in <strong>the</strong> Region.Rang? beyond <strong>the</strong> Region.213. Mulleripicus . .214.215.216.217.218.BrachyplernusTigaGecinulusMiglyptesMicropternus...Yttngid^.219. YunxMalabar, Aracan to Malaya and CelebesPhilippinesIndia, Ceylon, and ChinaIndia to MalayaS. Himalayas to BurmahMalayaIndia and Ceylon, to Borneo andS. ChinaCentral and S.ChinaPalpearctic, SAfricaINDICATORID.E.220. IndicatorHimalayas and BorneoEthiopianMEGAL.EMID.aS.221. Megalcema ...222. Xantholccma ...223. Psilopogon224. Coloramphus...27412<strong>The</strong> whole region, exel. PhilippinesAll India and Ceylon to Peguand MalayaSumatraMalacca, Sumatra and BorneoCuCTJLIDiE.225. Phcenicopliaes226. Rhinococcyx ...227. Dasylophus ...228. Lepidogrammus219. Carpococcyx ...230. Zanclostomus ..231. Rlwpodytes ...232. Taccocoua233. Poliococcyx ...234. Rhinortha235. Centropus236. Cuculus237. Cacomantis ...238. Chrysococcyx239. Surniculus ...240. Hierococcyx ...241. Coccystes242. Eudynamis ...1111117411141095262 ICeylonJavaPhilippine IslandsPhilippine IslandsBorneo, SumatraMalayaNepal to Ceylon, Hainan andMalayaAll India, Ceylon, MalaccaMalacca, Sumatra, BorneoMalacca, Sumatra, Borneo<strong>The</strong> whole region<strong>The</strong> whole region<strong>The</strong> whole region<strong>The</strong> whole regionIndia, Ceylon and Malaya<strong>The</strong> whole region<strong>The</strong> whole region, excl. Philippineswhole regionEthiopian, AustralianPalsearc, Ethiop., Aust.AustralianEthiopian, AustralianCelebes, N. China andAmoorlandEthiopianAustralianCoraciim:.243. Coracias244. Eurystomus ...MEROPID.E.245. Nyctiomis2 India, Ceylon and Burmah<strong>The</strong> whole regionS. India to Himalayas, Burmah,Sumatra, and BorneoEthiopian, S. PalaearcticCelebesEthiopian, Australian


JCHAP. XII.] THE ORIENTAL REGION. 383Order, Family,Genus.and°.2O eoRange <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.GO246. MeropsTrogonuxe.<strong>The</strong> whole regionS. Palsearctic, Ethiopian,Australian247. Harpactes10<strong>The</strong> whole region, excl.ChinaALCEDINID.E.248. Halcyon249. Pelargopsis ...250. Carcineutes ...251. Ceyx252. Alcedo253. Alcyone254. CeryleBUCEROTID.E.255. Buceros256. Hydrocissa ...257. Berenicornis ...258. Calao259. Aceros260. Cranorrhinus261. Penclopides ...262. Rhinoplax263. Meniceros10726<strong>The</strong> whole region<strong>The</strong> whole region, excl. ChinaBurmah, Siam, and MalayaIndia and Ceylon, Malaya andPhilippines<strong>The</strong> whole regionPhilippinesIndia to S. ChinaS. Palcearctic, Ethopian,AustralianCelebes and TimorMoluccas & New Guin.Palaearctic, Ethiopian,Austro-MalayanAustralian genusEthiopian, S. Palsearctic,AmericanNepal to Malaya, S. India,PhilippinesIndia, Ceylon and MalayaSumatraTenasserim, MalayaW. AfricaAustro-MalayaS. E. HimalayasMalacca to Borneo and PhilippinesCelebesPhilippinesSumatra, BorneoIndia and Ceylon to TenasserimUpUPIDjE.264. UpupaIndia,Ceylon and BurmahEthiopian, S. PalsearcticPoDARGIDiE.265. BatraehostomusIndia,Ceylon and MalayaMoluccasCaprimulgid^e.266. Caprimulgus ...267. Lyncornis134<strong>The</strong> whole regionBurmah, Malaya, & Philippines<strong>The</strong> Eastern HemisphereCelebesCypselid^e.268. Cypselus269. Dendrochelidon270. Collocalia ...271. Chsetura<strong>The</strong> region, excl. Philippines <strong>The</strong> 01dWorld& S.Am er.Ceylon, India, Malaya, Philipp. Austro-Malaya<strong>The</strong> whole regionMadagascar, Moluccas,Ceylon, India, Malaya, HainanPacific IslandsAmerica, AfricaPSITTACI.CACATTTID2E.(CacatuaPhilippines)Australian genusPAL.EORNITHID.E.272. Palaeornis14N. "W. India to Ceylon, Siam &MalayaEthiopian


N.384 ZOOLOGICAL GEOGRAPHY. [part nr.Order,Family, andGenus.273. Prioniturus274. Cyclopsitta275. Psittimos276. Tanygnathus277. LoriculusCOLUMB.E.COLUMBID.E.278. Treron .... .279. Ptilopus... .280. Carpophaga .281. Columba282. Janthsenas283. Macropygia284. Turtur ...285. Chalcophaps286. Phapitreron287. Caloenas...288. Phlegoenas289. GeopeliaGALLING.Pteroclim:.290. Pterocles21310736822121Philippine IslandsPhilippine IslandsMalaya, excl. JavaPhilippine IslandsCeylon, India, Malaya,pinesPhilip-<strong>The</strong> whole regionMalaya and PhilippinesIndia and Ceylon to Hainan andPhilippinesCeylon and India to TenasserimPhilippine, Andaman & NicobarIslandsNepal, Java, Hainan, Philippines<strong>The</strong> whole regionIndia, Ceylon, Malaya, Hainan,Philippines, FormosaPhilippine IslandsNicobar and Philippine IslandsPhilippine and Sooloo IslandsPhilippine Islands, JavaCentral and S.IndiaCelebesPapuan IslandsAustro-MalayaCelebes and Moluccas,FloresEthiopian, MoluccasAustralianAustralianPalsearc, Ethiop., Amer.Japan, Moluccas to SauioanIslandsAustro-Malaya, AustraliaOldWorld, Austro-Malay.Austro-Malaya, AustraliaAustro-MalayaAustro-Mal. & PolynesiaAustro-Malaya &Austral.S. Palsearctic. Ethiopian.TetraonipjE.291. Francolinus292. Ortygornis293. Perdix ...294. Coturnix295. Rollulus...(Caccabis3312921Ceylon and India to S. ChinaCeylon to Himalayas, Sumatra &BorneoIndia, Malaya, Philippines, China<strong>The</strong> whole regionMalacca, Siam, Borneo, Philipp.W. Himalayas)S. Palsearctic, EthiopianPalaearctie<strong>The</strong> Eastern HemispherePalsearctic genusPhasianid*.296. Pavo297. Argiisianus ...298. Polypledron ...(Lophophorus(Tetraophasis299. Ceriornis(Pucrasia300. Phasianus301. Emplocamus ..302. Gallus303. Galloperdix ..Ceylon to Himalayas, S.W. Chinaand Java4 Siam, Malacca, Borneo5 Upper Assam to S. W. China &Sumatra3 Cashmere and E. Thibet)1 E. Thibet)'5 W. Himalayas to W. China3 N. W. Himalayas to N. Chinaand Mongolia)3 W. Himalayas, S. China, Formosa13 N. W. Himalayas to ChinaSumatra and Borneo4 <strong>The</strong> region, excl. China3 Central India to CeylonPalsearctic genusPalsearctic genusS. E. PalsearcticPala?arctic genusS. PalsearcticCelebes and Timor


CHAP. XII.] THE ORIENTAL REGION. 385Order,Family,Genus.andC O)Range <strong>with</strong>in thtTURNICID.E.304. TurnixMegapodiid^!.305. Megapodius ..ACCIPITRES.VULTURID^I.306. Vultur307. Gyps308. Pscud-ogyps ...309. Neophron ...Falconid^;.310. Circus311. Astur312. Accipiter313. Buteo314. Aquila315. Nisaetus316. Lophotriorchis317. Neopus318. Spizaetus319. Circaetus320. Spilornis321. Butastur322. Haliseetus ...323. Haliastur324. Milvus325. Elanus326. Machserhamphus327. Pernis328. Baza329. Hicrax330. Poliohierax ...331. Falco332. CerchneisPandionid^e.333. Pandion334. PolioaetusStrigid^.335. A<strong>the</strong>ne ...336. Ninox337. Bubo ...338. Ketupa ...339. Scops


386 ZOOLOGICAL GEOGRAPHY. [PART III.Order, Family, andGenus.


. tmiftxttozsoo . i1A V ST RAlVi N K EC 1 () NScale lm . b.-1.000 miles \ut«-/M&rqui-sas orWraJmOM Is.^ArchiprfagoEXPLANATIONTeir*stmd ContoaMifrom Sea tarl (.. 100(> fat. Z500 . . -.*«•.UIMi'»tX>l>ioooo saooo .HTuMiummuo 100 soStnnf.rdi rinyin^inil fiVnM L.wAv,Sp «Y..rk !'*, *B.«fk. . 1 Bi


CHAPTER XIII.THE AUSTRALIAN EEGION.<strong>The</strong> Australian is <strong>the</strong> great insular region <strong>of</strong> <strong>the</strong> earth. As awhole it is one <strong>of</strong> <strong>the</strong> best marked, and has even been consideredto be equal in zoological value to all <strong>the</strong> rest <strong>of</strong> <strong>the</strong> globe ; butits separate portions are very heterogeneous, and <strong>the</strong>ir limitssometimes ill-defined. Its central and most important massesconsist <strong>of</strong> Australia and New Guinea, in which <strong>the</strong> main features<strong>of</strong> <strong>the</strong> region are fully developed. To <strong>the</strong> north-west it extends toCelebes, in which a large proportion <strong>of</strong> <strong>the</strong> Australian charactershave disappeared, while Oriental types are mingled <strong>with</strong> <strong>the</strong>mto such an extent that it is ra<strong>the</strong>r difficult to determine where tolocate it. To <strong>the</strong> south-east it includes New Zealand, which isin some respects so peculiar, that it has even been proposed toconstitute it a distinct region. On <strong>the</strong> east it embraces <strong>the</strong>whole <strong>of</strong> Oceania to <strong>the</strong> Marquesas and Sandwich Islands, whosevery scanty and <strong>of</strong>ten peculiar fauna, must be affiliated to <strong>the</strong>general Australian type.Australia is <strong>the</strong> largest tract <strong>of</strong> land in <strong>the</strong> region, beingseveral times more extensive than all <strong>the</strong> o<strong>the</strong>r islands combined,and it is here that <strong>the</strong> greatest variety <strong>of</strong> peculiar types havebeen developed. This island-continent, being situated in <strong>the</strong>track <strong>of</strong> <strong>the</strong> sou<strong>the</strong>rn desert zone, and having no central mountainsto condense <strong>the</strong> vapours from <strong>the</strong> surrounding ocean, has alarge portion <strong>of</strong> its interior so parched up and barren as to bealmost destitute <strong>of</strong> animal life. <strong>The</strong> most extensive tract <strong>of</strong>fertile and well-watered country is on <strong>the</strong> east and south east,


388 ZOOLOGICAL GEOGRAPHY. [part hi.where a fine range <strong>of</strong> mountains reaches, in <strong>the</strong> Colony <strong>of</strong> Victoria,<strong>the</strong> limits <strong>of</strong> perpetual snow. <strong>The</strong> west coast also possessesmountains <strong>of</strong> moderate height, but <strong>the</strong> climate is very dry andhot. <strong>The</strong> nor<strong>the</strong>rn portion is entirely tropical, yet it nowherepresents <strong>the</strong> luxuriance <strong>of</strong> vegetation characteristic <strong>of</strong> <strong>the</strong> greatisland <strong>of</strong> New Guinea immediately to <strong>the</strong> north <strong>of</strong> it. Taken asa whole, Australia is characterized by an arid climate and a deficiency<strong>of</strong> water; conditions which have probably long prevailed,and under which its very peculiar fauna and flora have been developed.This fact will account for some <strong>of</strong> <strong>the</strong> marked differencesbetween it and <strong>the</strong> adjacent sub-regions <strong>of</strong> New Guineaand <strong>the</strong> Moluccas, where <strong>the</strong> climate is moist, and <strong>the</strong> vegetationluxuriant ;<strong>of</strong>,and <strong>the</strong>se divergent features must never be lost sightin comparing <strong>the</strong> different portions <strong>of</strong> <strong>the</strong> Australian region.In Tasmania alone, which is however, essentially a detachedportion <strong>of</strong> Australia, a more uniform and moister climate prevails;but it is too small a tract <strong>of</strong> land, and has been toorecently severed from its parent mass to have developed aspecial fauna.<strong>The</strong> Austro-Malay sub-region (<strong>of</strong> which New Guinea is <strong>the</strong>central and typical mass) is strikingly contrasted <strong>with</strong> Australia,being subjected to purely equatorial conditions,—a high, butuniform temperature, excessive moisture, and a luxuriant forestvegetation, exactly similar in general features to that whichclo<strong>the</strong>s <strong>the</strong> Indo-Malay Islands, and <strong>the</strong> o<strong>the</strong>r portions <strong>of</strong> <strong>the</strong>great equatorial forest zone.Such a climate and vegetation, being<strong>the</strong> necessary result <strong>of</strong> its <strong>geographical</strong> position, must haveexisted from remote geological epochs <strong>with</strong> but little change, andmust <strong>the</strong>refore have pr<strong>of</strong>oundly affected all <strong>the</strong> forms <strong>of</strong> lifewhich have been developed under <strong>the</strong>ir influence. Around NewGuinea as a centre are grouped a number <strong>of</strong> important islands,more or less closely agreeing <strong>with</strong> it in physical features, climate,vegetation, and forms <strong>of</strong> life. In most immediate connection weplace <strong>the</strong> Aru Islands, Mysol and Waigiou, <strong>with</strong> Jobie and <strong>the</strong>o<strong>the</strong>r Islands in Geelvinck Bay, all <strong>of</strong> which are connected <strong>with</strong>it by shallow seas ; <strong>the</strong>y possess one <strong>of</strong> its most characteristicgroups, <strong>the</strong> Birds <strong>of</strong> Paradise, and have no doubt only recently (in


;chap, xiii.] THE AUSTRALIAN REGION. 389a geological sense) been separated from it.In <strong>the</strong> next rank come<strong>the</strong> large islands <strong>of</strong> <strong>the</strong> Moluccas on <strong>the</strong> west, and <strong>the</strong> rangeterminating in <strong>the</strong> Solomon Islands on <strong>the</strong> east, both <strong>of</strong> whichgroups possess a clearly Papuan fauna, although deficient inmany <strong>of</strong> <strong>the</strong> most remarkable Papuan types.All <strong>the</strong>se islands agree closely <strong>with</strong> New Guinea itself inbeing very mountainous, and covered <strong>with</strong> a luxuriant forestvegetation ; but to <strong>the</strong> south-west we find a set <strong>of</strong> islands extendingfrom Timor to Lombock, which agreemore nearly <strong>with</strong>Australia, both in climate and vegetation; being arid andabounding in eucalypti, acacias, and thickets <strong>of</strong> thorny shrubs.<strong>The</strong>se, like <strong>the</strong> Moluccas, are surrounded by deep sea, and it isdoubtful whe<strong>the</strong>r <strong>the</strong>y have ei<strong>the</strong>r <strong>of</strong> <strong>the</strong>m been actuallyconnected<strong>with</strong> New Guinea or Australia in recent geological timesbut <strong>the</strong> general features <strong>of</strong> <strong>the</strong>ir zoology oblige us to unite all<strong>the</strong>se islands <strong>with</strong> New Guinea as forming <strong>the</strong> Austro-Malaysub-division <strong>of</strong> <strong>the</strong> Australian region. Still fur<strong>the</strong>r west however,we have <strong>the</strong> large island <strong>of</strong> Celebes, whose position is verydifficult to determine. It is mountainous, but has also extensiveplains and low lands. Its climate is somewhat arid in <strong>the</strong> south,where <strong>the</strong> woods are <strong>of</strong>ten scattered and thorny, while in <strong>the</strong>north it is moister, and <strong>the</strong> forests are luxuriant.It is surroundedby deep seas, but also by coralline and volcanic islets, indicatingformer elevations and subsidences.Its fauna presents <strong>the</strong> mostpuzzling <strong>relations</strong>, showing affinities to Java, to <strong>the</strong> Philippines,to <strong>the</strong> Moluccas, to New Guinea, to continental India, and even toAfrica;so that it is almost impossible to decide whe<strong>the</strong>r to placeit in <strong>the</strong> Oriental or <strong>the</strong> Australian region. On <strong>the</strong> whole <strong>the</strong>preponderance <strong>of</strong> its <strong>relations</strong> appears to be <strong>with</strong> <strong>the</strong> latter,though itis undoubtedly very anomalous, and may, <strong>with</strong> almostas much propriety, be classed <strong>with</strong> <strong>the</strong> former. This will bebetterunderstood when we come to discuss its zoological peculiarities.<strong>The</strong> next sub-region consists <strong>of</strong> <strong>the</strong> extensive series <strong>of</strong> islandsscatteredover <strong>the</strong> Pacific, <strong>the</strong> principal groups being <strong>the</strong> SandwichIslands, <strong>the</strong> Marquesas and Society Islands, <strong>the</strong> Navigators',Friendly, and Fiji Islands. New Caledonia and <strong>the</strong> New


—;390 ZOOLOGICAL GEOGRAPHY. [part hi.Hebrides have ra<strong>the</strong>r an uncertain position, and it is difficulttodecide whe<strong>the</strong>r to class <strong>the</strong>m <strong>with</strong> <strong>the</strong> Austro-Malay Islands,<strong>the</strong> Pacific Islands,"or Australia.<strong>The</strong>. islands <strong>of</strong> <strong>the</strong> west Pacific,north <strong>of</strong> <strong>the</strong> equator, also probably come into this region,although <strong>the</strong> Ladrone Islands may belong to<strong>the</strong> Philippinesbut as <strong>the</strong> fauna <strong>of</strong> all <strong>the</strong>se small islets is very scanty, andvery little known, <strong>the</strong>y are not at present <strong>of</strong> much importance.<strong>The</strong>re remains <strong>the</strong> islands <strong>of</strong> New Zealand, <strong>with</strong> <strong>the</strong> surroundingsmall islands, as far as <strong>the</strong> Auckland, Chatham, and NorfolkIslands. <strong>The</strong>se are situated in <strong>the</strong> south temperateforest-zone. <strong>The</strong>y are mountainous, and have a moist, equable,and temperate climate. <strong>The</strong>y are true oceanic islands, and <strong>the</strong>totalabsence <strong>of</strong> mammalia intimates that <strong>the</strong>y have not beenconnected <strong>with</strong> Australia or any o<strong>the</strong>r continent in recent geologicaltimes. <strong>The</strong> general character <strong>of</strong> <strong>the</strong>ir zoology, no lessthan <strong>the</strong>ir botany, affiliates <strong>the</strong>m however, to Australia as portions<strong>of</strong> <strong>the</strong> same zoological region.General Zoological Characteristics <strong>of</strong> <strong>the</strong> Australian Region.—For <strong>the</strong> purpose <strong>of</strong> giving an idea <strong>of</strong> <strong>the</strong> very peculiar andstriking features which characterise <strong>the</strong> Australian region, itwill be as well at first to confine ourselves to <strong>the</strong> great centralland masses <strong>of</strong> Australia and New Guinea, where those featuresare manifested in <strong>the</strong>ir greatest force and purity, leaving <strong>the</strong>various peculiarities and anomalies <strong>of</strong> <strong>the</strong> outlying islands to bedealt <strong>with</strong> subsequently.Mammalia.—<strong>The</strong> Australian region is broadly distinguishedfroni all <strong>the</strong> rest <strong>of</strong> <strong>the</strong> globe by <strong>the</strong> entire absence <strong>of</strong> all <strong>the</strong>orders <strong>of</strong> non-aquatic mammalia that abound in <strong>the</strong> OldWorld, except two—<strong>the</strong> winged bats (Chiroptera), and <strong>the</strong> equallycosmopolite roderijts (Ptodentia). Of <strong>the</strong>se latter however, onlyone family is represented—<strong>the</strong> Muridse — (comprising <strong>the</strong> ratsand mice), and <strong>the</strong> Australian representatives <strong>of</strong> <strong>the</strong>se are all <strong>of</strong>small or moderate size—a suggestive fact in appreciating <strong>the</strong> truecharacter <strong>of</strong> <strong>the</strong> Australian fauna.In place <strong>of</strong> <strong>the</strong> Quadrumana,Carnivora, and Ungulates, which abound in endless varietyin all <strong>the</strong> o<strong>the</strong>r regions under equally favourable conditions,Australia possesses two new orders (or perhaps sub-classes)


;chap, xiii.] THE AUSTKALIAN REGION. 391Marsupialia and Monotremata, found nowhere else on <strong>the</strong> globeexcept a single family <strong>of</strong> <strong>the</strong> former in America. <strong>The</strong> Marsupialsare wonderfully developed in Australia, where <strong>the</strong>y existin <strong>the</strong> most diversified forms, adapted to different modes <strong>of</strong> life.Some are carnivorous, some herbivorous ; some arboreal, o<strong>the</strong>rsterrestrial. <strong>The</strong>re are insect-eaters, root-gnawers, fruit-eaters,honey-featers, leaf or grass-feeders. Some resemble wolves,o<strong>the</strong>rs nVarmots, weasels, squirrels, flying squirrels, dormice orjerboas. <strong>The</strong>y are classed in six distinct families, comprisingabout thirty genera, and subserve most <strong>of</strong> <strong>the</strong> purposes ineconomy <strong>of</strong> nature, fulfilled<strong>the</strong>in o<strong>the</strong>r parts <strong>of</strong> <strong>the</strong> world by verydifferent groups ;yet <strong>the</strong>y all possess common peculiarities <strong>of</strong>structure and habits which show that <strong>the</strong>y are members <strong>of</strong>one stock, and have no real affinity <strong>with</strong> <strong>the</strong> Old-World formswhich <strong>the</strong>y <strong>of</strong>ten outwardly resemble.<strong>The</strong> o<strong>the</strong>r order, Monotremata, is only represented by two rareand very remarkable forms, Ornitlwrhynchus and Echidna, probably<strong>the</strong> descendants <strong>of</strong> some <strong>of</strong> those earlier developments <strong>of</strong>mammalian life which in every o<strong>the</strong>r part <strong>of</strong> <strong>the</strong> globe have longbeen extinct.<strong>The</strong> bats <strong>of</strong> Australia all belong to Old-World genera andpossess no features <strong>of</strong> special interest, a result <strong>of</strong> <strong>the</strong> wanderinghabits <strong>of</strong> <strong>the</strong>se aerial mammals.<strong>The</strong> Eodents are more interesting.<strong>The</strong>y are all more or less modified forms <strong>of</strong> mice or rats.Some belong to <strong>the</strong> widely distributed genus Mus, o<strong>the</strong>rs to fourallied genera, which may be all modifications <strong>of</strong> some commonOld-World form. <strong>The</strong>y spread all over Australia, and alliedspecies occur in Celebes and <strong>the</strong> Papuan Islands; so that althoughnot yet known from <strong>the</strong> Moluccas, <strong>the</strong>reetettfeMhat some <strong>of</strong> <strong>the</strong>m exist <strong>the</strong>re.can be littleBirds.—<strong>The</strong> typical Australian region, as above defined, isalmost as well characterized by its birds, as by its mammaliabut in this case <strong>the</strong> deficiencies are less conspicuous, while <strong>the</strong>peculiar and characteristic families are numerous and important.<strong>The</strong> most marked deficiency as regards wide-spread families, is<strong>the</strong> total absence <strong>of</strong> Fringillidse (true finches), Picidse (woodpeckers),Vulturidse (vultures), and Phasianidee (pheasants)


392 ZOOLOGICAL GEOGRAPHY. [part in.and among prevalent Oriental groups, Pycnonotidse (bulbuls),Phyllornithidse (green bulbuls), and Megalsemidse (barbets) arefamilies whose absence is significant. Nine families are peculiarto <strong>the</strong> region, or only just pass its limits in <strong>the</strong> case <strong>of</strong> singlespecies. <strong>The</strong>se are Paridiseidse (paradise-birds), Meliphagidse(honey-suckers),Menuridse (lyre-birds), Atrichidae (scrub-birds),Cacatuidas (cockatoos), Platycercidse (broad-tailed and grassparoquets),Trichoglossidse (brush-tongued paroquets, Megapodiidae(mound-makers), and Casuariidae (cassowaries).<strong>The</strong>re arealso eight very characteristic families, <strong>of</strong> which four,—Pachycephalias(thick-headed shrikes), Campephagidse (caterpillarshrikes), Dicseidae (flower-peckers), and Artamidae (swallowshrikes)—arefeebly represented elsewhere, while <strong>the</strong> o<strong>the</strong>r four—Ploceidae (weaver-finches), Alcsedinidse (kingfishers), Poclargidae(frog-mouths), and Columbidse(pigeons)—although widely distributed,are here unusually abundant and varied, and (exceptin <strong>the</strong> case <strong>of</strong> <strong>the</strong> Ploceidse) better represented in <strong>the</strong> Australianthan in any o<strong>the</strong>r region.Of all <strong>the</strong>se <strong>the</strong> Meliphagidse (honeysuckers)are <strong>the</strong> most peculiarly and characteristically Australian.This family abounds in genera and species ;it extends into everypart <strong>of</strong> <strong>the</strong> region from Celebes and Lombock on <strong>the</strong> west, to<strong>the</strong> Sandwich Islands, Marquesas, and New Zealand on <strong>the</strong> east,while not a single species overpasses its limits, <strong>with</strong> <strong>the</strong> exception<strong>of</strong> one {Ptilotis lirribata) which abounds in all <strong>the</strong> islands <strong>of</strong><strong>the</strong> Timorese group, and has crossed <strong>the</strong> narrow strait fromLombock to Baly ; but this can hardly be considered to impugn<strong>the</strong> o<strong>the</strong>rwise striking fact <strong>of</strong> wide diffusion combined <strong>with</strong>strict limitation, which characterizes it.This family is <strong>the</strong> moreimportant, because, like <strong>the</strong> Trichoglossida3 or brush-tonguedparoquets, it seems to have been developed in co-ordination <strong>with</strong>that wealth <strong>of</strong> nectariferousflowering shrubs and trees which isone <strong>of</strong> <strong>the</strong> marked features <strong>of</strong> Australian vegetation.It probablyoriginated in <strong>the</strong> extensive land-area <strong>of</strong> Australia itself, and<strong>the</strong>nce spread into all <strong>the</strong> tributary islands, where it has becomevariously modified, yet always in such close adaptation to <strong>the</strong>o<strong>the</strong>r great features <strong>of</strong> <strong>the</strong> Australian fauna, that it seems unableto maintain itself when subject to <strong>the</strong> competition <strong>of</strong> <strong>the</strong> more


chap, xin.] THE AUSTRALIAN REGION. 393varied forms <strong>of</strong> life in <strong>the</strong> Oriental region ;to which, possessinggreat powers <strong>of</strong> flight, some species must occasionally have emigrated.Its presence or absence serves <strong>the</strong>refore to define andlimit <strong>the</strong> Australian region <strong>with</strong> a precision hardly to beequalled in <strong>the</strong>birds.case <strong>of</strong> any o<strong>the</strong>r region or any o<strong>the</strong>r family <strong>of</strong><strong>The</strong> Trichoglossidse, as already intimated, are ano<strong>the</strong>r <strong>of</strong> <strong>the</strong>sepeculiarly organized Australian families,— parrots <strong>with</strong> an extensilebrush-tipped tongue, adapted to extract <strong>the</strong> nectar andpollen from flowers. <strong>The</strong>se are also rigidly confined to thisregion, but <strong>the</strong>y do not range so completely over <strong>the</strong> whole <strong>of</strong> it,being absent from New Zealand (where however <strong>the</strong>y are representedby a closely allied form Nestor),and from <strong>the</strong> SandwichIslands. <strong>The</strong> Paradiseidse (birds <strong>of</strong> paradise and allies) areano<strong>the</strong>r remarkable family, confined to <strong>the</strong> Papuan group <strong>of</strong>Islands, and <strong>the</strong> tropical parts <strong>of</strong> Australia. <strong>The</strong> Megapodiidse(or mound-builders) are ano<strong>the</strong>r most remarkable and anomalousgroup <strong>of</strong> birds, no doubt specially adapted to Australian conditions<strong>of</strong> existence. <strong>The</strong>ir peculiarity consists in <strong>the</strong>ir layingenormous eggs (at considerable intervals <strong>of</strong> time) and burying<strong>the</strong>m ei<strong>the</strong>r in <strong>the</strong> loose hot sand <strong>of</strong> <strong>the</strong> beach above high-watermark, or in enormous mounds <strong>of</strong> leaves, sticks, earth, and refuse<strong>of</strong> all kinds, ga<strong>the</strong>red toge<strong>the</strong>r by <strong>the</strong> birds, whose feet andclaws are enlarged and streng<strong>the</strong>ned for <strong>the</strong> work. <strong>The</strong> warmth<strong>of</strong> this slightly fermenting mass hatches <strong>the</strong> eggs ; when <strong>the</strong>young birds work <strong>the</strong>ir way out, and <strong>the</strong>nceforth take care <strong>of</strong><strong>the</strong>mselves, as <strong>the</strong>y are able to run quickly, and even to fly shortdistances, as soon as <strong>the</strong>y are hatched.This may perhaps be anadaptation to <strong>the</strong> peculiar condition <strong>of</strong> so large a portion <strong>of</strong>Australia, in respect to prolonged droughts and scanty watersupply,entailing a periodical scarcity <strong>of</strong> all kinds <strong>of</strong> food.In sucha country <strong>the</strong> confinement <strong>of</strong> <strong>the</strong> parents to one spot during <strong>the</strong>long period <strong>of</strong> incubation would <strong>of</strong>ten lead to starvation, and <strong>the</strong>consequent death <strong>of</strong> <strong>the</strong> <strong>of</strong>fspring.But <strong>the</strong> same birds <strong>with</strong> freepower to roam about, might readily maintain <strong>the</strong>mselves.Thispeculiar constitution and habit, which enabled <strong>the</strong> Megapodii tomaintain an existence under <strong>the</strong> unfavourable conditions <strong>of</strong> <strong>the</strong>irVol. I.—27


394 ZOOLOGICAL GEOGRAPHY. [part hi.original habitat gives <strong>the</strong>m a great advantage in <strong>the</strong> luxuriantislands <strong>of</strong> <strong>the</strong> Moluccas, to which <strong>the</strong>y have spread. <strong>The</strong>re<strong>the</strong>y abound to a remarkable extent, and <strong>the</strong>ir eggs furnish aluxurious repast to <strong>the</strong> natives. <strong>The</strong>y have also reached many<strong>of</strong> <strong>the</strong> smallest islets, and have spread beyond <strong>the</strong> limits<strong>of</strong> <strong>the</strong>region to <strong>the</strong> Philippines, and North-Western Borneo, as wellas to <strong>the</strong> remote Nicobar Islands.<strong>The</strong> Platycercidse, or broad-tailed paroquets, are ano<strong>the</strong>rwide-spread Australian group, <strong>of</strong> weak structure but gorgeouslycoloured, ranging from <strong>the</strong> Moluccas to New Zealand and <strong>the</strong>Society Islands, and very characteristic <strong>of</strong> <strong>the</strong> region, to which<strong>the</strong>y are strictly confined. <strong>The</strong> Cockatoos have not quite sowide a range, being confined to <strong>the</strong> Austro-Malayan and Australiansub-regions, while one species extends into <strong>the</strong> PhilippineIslands. <strong>The</strong> o<strong>the</strong>r two peculiar families are more restricted in<strong>the</strong>ir range, and will be noticed under <strong>the</strong> sub-regions to which<strong>the</strong>y respectively belong.Of <strong>the</strong> characteristic families, <strong>the</strong> Pachycephalidae, or thickheadedshrikes, are especially Australian, ranging over all <strong>the</strong>region, except New Zealand ; while only a single species hasspread into <strong>the</strong> Oriental, and one <strong>of</strong> doubtful affinity to <strong>the</strong>Ethiopian region. <strong>The</strong> Artamidae, or swallow-shrikes, are alsoalmost wholly confined to <strong>the</strong> region, one species only extendingto India. <strong>The</strong>y range to <strong>the</strong> Fiji Islands on <strong>the</strong> east, but onlyto Tasmania on <strong>the</strong> south. <strong>The</strong>se two families must be consideredas really peculiar to Australia. <strong>The</strong> Podargidae, or frogmouths—large,thick-billed goat- suckers—are strange birds veiycharacteristic <strong>of</strong> <strong>the</strong> Australian region, although <strong>the</strong>y haverepresentatives in <strong>the</strong> Oriental and Neotropical regions. Campephagidae(caterpillar-shrikes) also abound, but <strong>the</strong>y are fairlylepresented both in India and Africa.<strong>The</strong> Ploceidae, or weaverbirds,are <strong>the</strong> finches <strong>of</strong> Australia, and present a variety <strong>of</strong>interesting and beautiful forms.We now come to <strong>the</strong> kingfishers, a cosmopolitan family <strong>of</strong>birds, yet so largely developed in <strong>the</strong> Australian region as todeserve special notice. Two-thirds <strong>of</strong> all <strong>the</strong> genera are foundhere, and no less than 10 out <strong>of</strong> <strong>the</strong> 19 genera in <strong>the</strong> family are


chap, xiii.] THE AUSTRALIAN REGION. 395peculiar to <strong>the</strong> Australian region.Ano<strong>the</strong>r <strong>of</strong> <strong>the</strong> universally distributedfamilies which have <strong>the</strong>ir metropolis here, is that <strong>of</strong> <strong>the</strong>Columbidse or pigeons.Three-fourths <strong>of</strong> <strong>the</strong> genera have representativesin <strong>the</strong> Australian region, while two-fifths <strong>of</strong> <strong>the</strong> whole areconfined to it ;o<strong>the</strong>r two regions combined.and it possesses as many species <strong>of</strong> pigeons as anyIt also possesses <strong>the</strong> most remarkableforms, as exemplified in <strong>the</strong> great crowned pigeons(Goura)and <strong>the</strong> hook-billed Didunculus, while <strong>the</strong> green fruit-pigeons(Ptilopus) are sometimes adorned <strong>with</strong> colours vying <strong>with</strong> those<strong>of</strong> <strong>the</strong> gayest parrots or chatterers.This enormous development<strong>of</strong> a family <strong>of</strong> birds so defenceless as <strong>the</strong> pigeons, whose rudenests expose <strong>the</strong>ir eggs and helpless young to continual danger,may perhaps be correlated, as I have suggested elsewhere (Ibis,1865, p. 366), <strong>with</strong> <strong>the</strong> entire absence <strong>of</strong> monkeys, cats, lemurs,weasels, civets and o<strong>the</strong>r arboreal mammals, which prey on eggsand young birds. <strong>The</strong> very prevalent green colour <strong>of</strong> <strong>the</strong> upperpart <strong>of</strong> <strong>the</strong>ir plumage, may be due to<strong>the</strong> need <strong>of</strong> concealmentfrom <strong>the</strong>ir only enemies,—birds <strong>of</strong> prey ; and this is renderedmore probable by <strong>the</strong> fact that it is among <strong>the</strong> pigeons <strong>of</strong> <strong>the</strong>small islands <strong>of</strong> <strong>the</strong> Pacific (where hawks and <strong>the</strong>ir allies are exceedinglyscarce) that we alone meet <strong>with</strong> specieswhose entireplumage is a rich and conspicuous yellow. Where <strong>the</strong> need <strong>of</strong>concealment is least, <strong>the</strong> brilliancy <strong>of</strong> colour has attained itsmaximum. We may <strong>the</strong>refore look upon <strong>the</strong> genus Ptilopus,<strong>with</strong> its fifty species whose typical coloration is green, <strong>with</strong>patches <strong>of</strong> bright blue, red, or yellow on <strong>the</strong> head and breast,as a special development suited to <strong>the</strong> tropical portion <strong>of</strong> <strong>the</strong>Australian region, to which it is almost wholly confined.It will be seen from <strong>the</strong> sketch just given, that <strong>the</strong> ornithologicalfeatures <strong>of</strong> <strong>the</strong> Australian region are almost as remarkableas those presented by its Mammalian fauna ; and from <strong>the</strong>fuller development attained by <strong>the</strong> aerial class <strong>of</strong> birds, muchmore varied and interesting. None <strong>of</strong> <strong>the</strong> o<strong>the</strong>r regions <strong>of</strong> <strong>the</strong>earth can <strong>of</strong>fer us so many families <strong>with</strong> special points <strong>of</strong>interest in structure, or habits, or general <strong>relations</strong>. <strong>The</strong>paradise-birds, <strong>the</strong> honeysuckers, <strong>the</strong> brush-tongued paroquets,<strong>the</strong> mound-builders, and <strong>the</strong> cassowaries—all strictly peculiar


396 ZOOLOGICAL GEOGRAPHY. [part hi.to <strong>the</strong> region—<strong>with</strong> such remarkable developments as we haveindicated in <strong>the</strong> kingfishers and pigeons, place <strong>the</strong> Australianregion in <strong>the</strong> first rank for <strong>the</strong> variety, singularity, and interest<strong>of</strong> its birds, and only second to South America as regardsnumbers and beauty.Reptiles.—In Eeptiles <strong>the</strong>peculiarity <strong>of</strong> <strong>the</strong> main Australianregion is less marked, although <strong>the</strong> fauna is sufficiently distinct.<strong>The</strong>re is no family <strong>of</strong> snakes confined to <strong>the</strong> region, but manyAboutpeculiar genera <strong>of</strong> <strong>the</strong> families Pythonidse and Elapidse.two-thirds <strong>of</strong> <strong>the</strong> Australian snakes belong to <strong>the</strong> latter family,and are poisonous ;so that although <strong>the</strong> Crotalidse and Viperidseare absent, <strong>the</strong>re are perhaps a larger proportion <strong>of</strong> poisonousto harmless snakes than in any o<strong>the</strong>r part <strong>of</strong> <strong>the</strong> world.Accordingto Mr. Gerard Krefft <strong>the</strong> proportion varies considerably in <strong>the</strong>different colonies.In Victoria, New South Wales, and Queensland<strong>the</strong> proportion is about two to one ; in West Australiathree to one ; and in South Australia six to one. In Tasmania<strong>the</strong>re are only 3 species and all are poisonous. <strong>The</strong> number<strong>of</strong> species, as in o<strong>the</strong>r parts <strong>of</strong> <strong>the</strong> world, seems to increase*<strong>with</strong> temperature. <strong>The</strong> 3 in Tasmania have increased to 12in Victoria, 15 in South Australia and <strong>the</strong> same in WestAustralia ; 31 in New South Wales, and 42 in sub-tropicalQueensland.<strong>The</strong> lizards <strong>of</strong> Australia have lately been catalogued by Dr.Gtin<strong>the</strong>r in <strong>the</strong> concluding part <strong>of</strong> <strong>the</strong> " Voyage <strong>of</strong> <strong>the</strong> Erebusand Terror," issued in 1875. <strong>The</strong>y belong to 8 families, 3 <strong>of</strong>which are peculiar ; 57 genera <strong>of</strong> which 36 are peculiar ; andabout 140 species, all but 2 or 3 <strong>of</strong> which are peculiar. <strong>The</strong>scinks and geckoes form <strong>the</strong> great bulk <strong>of</strong> <strong>the</strong> Australianlizards, <strong>with</strong> a few Agamidae,<strong>The</strong> three peculiar families are <strong>the</strong>Gymnopthalmidfe, and Varanidse.Pygopodidse, Aprasiidse andLialidae; comprising only 4 genera and 7 species. <strong>The</strong> aboveall belong to Australia proper. Those <strong>of</strong> <strong>the</strong> o<strong>the</strong>r sub-regionsare few in number and will be noticed under <strong>the</strong>ir, respectivelocalities.<strong>The</strong>y will perhaps bring up <strong>the</strong> number <strong>of</strong> genera to70. West and South Australia seem to <strong>of</strong>fer much peculiarityin <strong>the</strong>ir lizards; <strong>the</strong>se districts possessing 12 peculiar genera,


——chap, xiii.] THE AUSTRALIAN REGION. 397while a much smaller number are confined to <strong>the</strong> East andSouth-East, or to <strong>the</strong> North.Among <strong>the</strong> fresh-water turtles <strong>of</strong> <strong>the</strong> family Chelydida?<strong>the</strong>re are three peculiar generaChelodina, Chelemys, and Elseya,all from Australia.Amphibia.—No tailed amphibians are known from <strong>the</strong> wholeregion, but no less than eleven <strong>of</strong> <strong>the</strong> families <strong>of</strong> tail-lessBatrachians(toads and frogs) are known to inhabit some part oro<strong>the</strong>r <strong>of</strong> it. A peculiar family (Xenorhinidse), consisting <strong>of</strong>a single species, is found in New Guinea; <strong>the</strong> true toads(Bufonidae) are only represented by a single species<strong>of</strong> a peculiargenus in Australia, and by a Bufo in Celebes. Nine <strong>of</strong> <strong>the</strong>families are represented in Australia itself, and <strong>the</strong> followinggenera are peculiar to it :Pseudophryne (Phryniscidse),Pachybatrachus,and Chelydobatrachus (Engystomydee) ; Helioporus(Alytidse) ; Pelodyras and Chirodyras (Pelodryadse) ; Notaden(Bufonidse).Fresh-water Fish.—<strong>The</strong>re is only one peculiar family <strong>of</strong> freshwaterfishes in this region—<strong>the</strong> Gadopsidse—represented by asingle genus and species. <strong>The</strong> o<strong>the</strong>r species <strong>of</strong> Australia belongto <strong>the</strong> families Trachinidse, A<strong>the</strong>rinidse, Mugillidae, Siluridse,Homalopterse, Haplochitonidse, Galaxidae, Osteoglossidse, Symbranchidee,and Sirenoidei ; most <strong>of</strong> <strong>the</strong> genera being peculiar.<strong>The</strong> large and widely-distributed families, Cyprinodontidae andCyprinidse, are absent. <strong>The</strong> most remarkable fish is <strong>the</strong> recentlydiscovered Ceratodus, allied to <strong>the</strong> Lepidosiren <strong>of</strong> TropicalAmerica, and Protopterus <strong>of</strong> Tropical Africa, <strong>the</strong> three speciesconstituting <strong>the</strong> Sub-class Dipnoi, remains <strong>of</strong> which have beenfound fossil in <strong>the</strong> Triassic formation.Summary <strong>of</strong> Australian Vertebrata.—In order to completeour general sketch <strong>of</strong> Australian zoology, and to afford materialsfor comparison <strong>with</strong> o<strong>the</strong>r regions, we will here summarize <strong>the</strong><strong>distribution</strong> <strong>of</strong> Vertebrata in <strong>the</strong> entire Australian region, asgiven in detail in <strong>the</strong> tables at <strong>the</strong> end <strong>of</strong> this chapter. Whenan undoubted Oriental family or genus extends to Celebes onlywe do not count it as belonging to <strong>the</strong> Australian region, thatisland being so very anomalous and intermediate in character.


398 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>The</strong> Australian region, <strong>the</strong>n, possesses examples <strong>of</strong>18 families<strong>of</strong> Mammalia, 8 <strong>of</strong> which are peculiar ; 71 <strong>of</strong> Birds, 16 beingpeculiar; 31 <strong>of</strong> Eeptiles, 4 being peculiar ;11 <strong>of</strong> Amphibia, <strong>with</strong>1 peculiar; and 11 <strong>of</strong> Fresh-water fish, <strong>with</strong> 1 peculiar. In all,142 families <strong>of</strong> Vertebrates, 30 <strong>of</strong> which are almost or quiteconfined to it, or between one-fourth and one-fifth <strong>of</strong> <strong>the</strong> wholenumber.<strong>The</strong> genera <strong>of</strong> Mammalia, occurring <strong>with</strong>in <strong>the</strong> limits <strong>of</strong> thisregion are 70, <strong>of</strong> which 45 are almost, or quite, confined to it.Of Land-Birds <strong>the</strong>re are 296 genera, 196 <strong>of</strong> which are equallylimited. <strong>The</strong> proportion is in both cases very nearly fiveeighths.This shows a considerable deficiency both in families <strong>of</strong> Vertebratesand genera <strong>of</strong> Mammalia, as compared <strong>with</strong> <strong>the</strong> Orientaland Ethiopian regions ; while in genera <strong>of</strong> Birds it is a littlesuperior to <strong>the</strong> latter in total numbers, and considerably so in<strong>the</strong> proportion <strong>of</strong> peculiar types.Supposed Land Connection between Australia and South America.We may now consider how far <strong>the</strong> different classes and orders<strong>of</strong> vertebrates afford indications that during past ages <strong>the</strong>rehas been some closer connection between Australia and SouthAmerica than that which now exists.Among Mammalia we have <strong>the</strong> remarkable fact <strong>of</strong> a group<strong>of</strong> marsupials inhabiting South America, and extending eveninto <strong>the</strong> temperate regions <strong>of</strong> North America, while <strong>the</strong>y arefound in no o<strong>the</strong>r part <strong>of</strong> <strong>the</strong> globe beyond <strong>the</strong> limits <strong>of</strong> <strong>the</strong>Australian region ; and this has <strong>of</strong>ten been held to be evidence<strong>of</strong> a former connection between <strong>the</strong> two countries. A preliminaryobjection to this view is, that <strong>the</strong> opossums seem to bera<strong>the</strong>r a tropical group, only one species reaching as far as 42°south latitude on <strong>the</strong> west coast <strong>of</strong> South America ; but whateverevidence we have which seems to require a former union<strong>of</strong> <strong>the</strong>se countries shows that it took place, if at all, towards <strong>the</strong>ircold sou<strong>the</strong>rn limits, <strong>the</strong> tropical faunas on <strong>the</strong> whole showingno similarity.This is not a very strong objection, since climatesmay have changed in <strong>the</strong> south to as great an extent as we


;chap, xni.] THE AUSTRALIAN REGION. 39$know <strong>the</strong>y have in <strong>the</strong> north. Perhaps a more important considerationis, that Didelphys is a family type unknown in Australia; and this implies that <strong>the</strong> point <strong>of</strong> common origin is veryremote in geological time. But <strong>the</strong> most conclusive fact is thatin <strong>the</strong> Eocene and Miocene periods this very family, Didelphyidse,existed in Europe, while it only appeared in Americain <strong>the</strong> Post-pliocene or perhaps <strong>the</strong> Pliocene period ; so that itis really an Old-World group, which, though long since extinctin its birthplace, has survived in America, to which country itis a comparatively recent emigrant. Primeval forms <strong>of</strong> marsupialswe know abounded in Europe during much <strong>of</strong> <strong>the</strong> Secondaryepoch, and no doubt supplied Australia <strong>with</strong> <strong>the</strong> ancestors <strong>of</strong><strong>the</strong> present fauna. It is clear, <strong>the</strong>refore, that in this ease <strong>the</strong>reis not a particle <strong>of</strong> evidence for any former union betweenAustralia and South America ; while it is almost demonstratedthat both derived <strong>the</strong>ir marsupials from a common source in <strong>the</strong>nor<strong>the</strong>rn hemisphere.Birds <strong>of</strong>fer us more numerous but less clearly defined cases <strong>of</strong>this kind. Among Passeres, <strong>the</strong> wonderful lyre bird (Menura)is believed by some ornithologists to be decidedly allied to <strong>the</strong>South American Pteroptochidae, while o<strong>the</strong>rs maintain thatit is altoge<strong>the</strong>r peculiar, and has no such affinity. <strong>The</strong> AustralianPachycephalia? have also been supposed to find <strong>the</strong>irnearest allies in <strong>the</strong> American Vireonida?, but this is, perhaps,equally problematical. That <strong>the</strong> mound-makers (Megapodiidse)<strong>of</strong> <strong>the</strong> Australian region are more nearly allied to <strong>the</strong> SouthAmerican curassows ^Cracidse) than to any o<strong>the</strong>r family, is perhapsbetter established ; but if proved, it is probably due, as in<strong>the</strong> case <strong>of</strong> <strong>the</strong> marsupials, to <strong>the</strong> survival <strong>of</strong> an ancient andonce wide-spread type, and thus lends no support to <strong>the</strong> <strong>the</strong>ory<strong>of</strong> a land connection between <strong>the</strong> two regions. A recent author,Pr<strong>of</strong>essor Garrod, classes Phaps and o<strong>the</strong>r Australian genera <strong>of</strong>pigeons along <strong>with</strong> Zenaida and allied South American formsbut here again <strong>the</strong> affinity, if it exists, is so remote that <strong>the</strong> explanationalready given will suffice to account for it. <strong>The</strong>reremain only <strong>the</strong> penguins <strong>of</strong> <strong>the</strong> genus Eudyptes ; and <strong>the</strong>sehave almost certainly passed from one region to <strong>the</strong> o<strong>the</strong>r, but


;400 ZOOLOGICAL GEOGEAPHY. [part hi.no actual land connection is required for birds which can crossconsiderable arms <strong>of</strong> <strong>the</strong> sea.Eeptiles again seem to <strong>of</strong>fer no more support to <strong>the</strong> view thando mammalia or birds. Among snakes <strong>the</strong>re are no families incommon that have not a very wide <strong>distribution</strong>. Among lizards<strong>the</strong> Gymnopthalmidae are <strong>the</strong> only family that favour <strong>the</strong>notion, since <strong>the</strong>y are found in Australia and South America,but not in <strong>the</strong> Oriental region. Yet <strong>the</strong>y occur in both <strong>the</strong>PalaBarctic and Ethiopian regions, and <strong>the</strong>ir <strong>distribution</strong> is altoge<strong>the</strong>rtoo erratic to be <strong>of</strong> any value in a case <strong>of</strong> this kindand <strong>the</strong> same remarks apply to <strong>the</strong> tortoises <strong>of</strong> <strong>the</strong> familyChelydidse.<strong>The</strong> Amphibia, however, furnishus <strong>with</strong> some more decidedfacts. We have first <strong>the</strong> family <strong>of</strong> tree-frogs, Pelodryade, confinedto <strong>the</strong> two regions ,Litoria, a genus <strong>of</strong> <strong>the</strong> family Hylidsepeculiar to Australia, but <strong>with</strong> one species in Paraguay ;and in<strong>the</strong> family Discoglossidse, <strong>the</strong> Australian genus Chiroleptes hasits nearest ally in <strong>the</strong> Chilian genus Calyptocephalus.Fresh-water fishes give yet clearer evidence.Three groups areexclusively found in <strong>the</strong>se two regions ;Aphritis, a fresh-watergenus <strong>of</strong> Trachinidse, has one species in Tasmania and twoo<strong>the</strong>rs in Patagonia ;<strong>the</strong> Haplochitonidre inhabit only Terra delFuego, <strong>the</strong> Falkland Islands and South Australia; while <strong>the</strong>genus Galaxias (forming <strong>the</strong> family Galaxidse) is confined toSouth Temperate America, Australia, and New Zealand. Wehave also <strong>the</strong> genus Osteoglossum confined to <strong>the</strong> tropicalrivers <strong>of</strong> Eastern South America, <strong>the</strong> Indo-Malay Islands andAustralia.It is important here to notice that <strong>the</strong> heat-loving Eeptiliaafford hardly any indications <strong>of</strong> close affinity between <strong>the</strong> tworegions, while <strong>the</strong> cold-enduring amphibia and fresh-waterfish, <strong>of</strong>fer <strong>the</strong>m in abundance. Taking this fact in connection<strong>with</strong> <strong>the</strong> absence <strong>of</strong> all indications <strong>of</strong> close affinityamong <strong>the</strong> mammalia and terrestrial birds, <strong>the</strong> conclusion seemsinevitable that <strong>the</strong>re has been no land-connection between <strong>the</strong>two regions <strong>with</strong>in <strong>the</strong> period <strong>of</strong> existing species, genera, orfamilies. Yet some interchange <strong>of</strong> amphibia and fresh-water


chap, xiii.] THE AUSTRALIAN REGION. 401fishes, as <strong>of</strong> plants and insects, has undoubtedly occurred, butthis has been effected by o<strong>the</strong>r means. If we look at a globewe see at once how this interchange may have taken place.Immediately south <strong>of</strong> Cape Horn we have <strong>the</strong> South ShetlandIslands and Graham's land, which is not improbably continuous,or nearly so, <strong>with</strong> South Victoria land immediately to <strong>the</strong> south<strong>of</strong> New Zealand. <strong>The</strong> intervening space is partly occupiedby <strong>the</strong> Auckland, Campbell, and Macquaries' Islands, which,<strong>the</strong>re is reason to believe are <strong>the</strong> relics <strong>of</strong> a great sou<strong>the</strong>rn extension<strong>of</strong> New Zealand. At all events <strong>the</strong>y form points whichwould aid <strong>the</strong> transmission <strong>of</strong> many organisms ;<strong>of</strong> <strong>the</strong> Macquaries' group, Emerald Island, is<strong>the</strong> outlying islets <strong>of</strong> Victoria land.a considerable time in <strong>the</strong> air,and <strong>the</strong> far<strong>the</strong>stonly 600 miles from<strong>The</strong> ova <strong>of</strong> fish will surviveand <strong>the</strong> successful transmission <strong>of</strong>salmon ova to New Zealand packed in ice, shows how far <strong>the</strong>ymight travel on icebergs. Now <strong>the</strong>re is evidently some meansby which ova or young fishes are carried moderate distances, from<strong>the</strong> fact that remote alpine lakes and distinct river systems <strong>of</strong>tenhave <strong>the</strong> same species.<strong>of</strong> fresh water on <strong>the</strong>ir surfaces ;Glaciers and icebergs generally have poolsand whatever cause transmits fishto an isolated pond might occasionally stock <strong>the</strong>se pools, and bythis means introduce <strong>the</strong> fishes <strong>of</strong> one sou<strong>the</strong>rn island intoano<strong>the</strong>r. Batrachians, which are equally patient <strong>of</strong> cold, mightbe transported by similar means ; while, as Mr. Darwin has sowell shown, {Origin <strong>of</strong> Species, 6th Ed. p. 345) <strong>the</strong>re are variousknown modes by which plants might be transmitted, and weneed not <strong>the</strong>refore be surprised that botanists find a muchgreater similarity between <strong>the</strong> production <strong>of</strong> <strong>the</strong> several Sou<strong>the</strong>rnlands and islands, than do zoologists. It is important to noticethat, however this intercommunication was effected, ithas continueddown to <strong>the</strong> epoch <strong>of</strong> existing species ; for Dr. Giin<strong>the</strong>rfinds <strong>the</strong> same species <strong>of</strong> fresh-water fish (Galaxias attenuatus)inhabiting Tasmania, New Zealand, <strong>the</strong> Falkland Islands, andTemperate South America ; while ano<strong>the</strong>r species is common toNew Zealand and <strong>the</strong> Auckland Islands. We cannot believethat a land connection has existed between all <strong>the</strong>se remotelands <strong>with</strong>in <strong>the</strong> period <strong>of</strong> existence <strong>of</strong> this one species <strong>of</strong> fish,


402 ZOOLOGICAL GEOGRAPHY. [part hi.not only on account <strong>of</strong> what we know <strong>of</strong> <strong>the</strong> permanence <strong>of</strong>continents and deep oceans, hut because such a connection musthave led to much more numerous and important cases<strong>of</strong> similarity<strong>of</strong> natural productions than we actually find. And if<strong>with</strong>in <strong>the</strong> life <strong>of</strong> species such interchange may have takenplace across seas <strong>of</strong> greater or less extent, still more easy is itto understand, how, <strong>with</strong>in <strong>the</strong> life <strong>of</strong> genera and families, a number<strong>of</strong> such interchanges may have occurred ;yet always limitedto those groups whose conditions <strong>of</strong> life render transmissionpossible. Had an actual land connection existed <strong>with</strong>in <strong>the</strong>temperate zone, or during a period <strong>of</strong> warmth in <strong>the</strong> Antarcticregions, <strong>the</strong>re would have been no such strict limitations to <strong>the</strong>inter-migration <strong>of</strong> <strong>animals</strong>.that floating ice has had some share in <strong>the</strong>It may be held to support <strong>the</strong> viewtransmission <strong>of</strong> fishand amphibia, when -we find that in <strong>the</strong> case <strong>of</strong> <strong>the</strong> narrowtropical sea dividing Borneo from Celebes and <strong>the</strong> Moluccas, noproportionate amount <strong>of</strong> transmission has taken place, butnumerous species, genera, and whole families, terminate abruptlyat what we have o<strong>the</strong>r reasons for believing to be <strong>the</strong> fur<strong>the</strong>stlimits <strong>of</strong> an ancient continent. We can hardly suppose, however,that this mode <strong>of</strong> transmission would have sufficed forsuch groups as tree-frogs, which are inhabitants <strong>of</strong> <strong>the</strong> moretemperate or even warm portions <strong>of</strong> <strong>the</strong> two sou<strong>the</strong>rn lands.Some <strong>of</strong> <strong>the</strong>se cases may perhaps be explained by <strong>the</strong> supposition<strong>of</strong> a considerable extent <strong>of</strong> land in <strong>the</strong> South-Temperate andAntarctic regions now submerged, and by a warm or temperateclimate analogous to that which prevailed in <strong>the</strong> Arctic regionsduring some part <strong>of</strong> <strong>the</strong> Miocene epoch ; while o<strong>the</strong>rs may bedue to cases <strong>of</strong> survival in <strong>the</strong> two areas <strong>of</strong> once wide-spreadgroups, a view supported in <strong>the</strong> case <strong>of</strong> <strong>the</strong> Amphibia by <strong>the</strong>erratic manner in which many <strong>of</strong> <strong>the</strong> groups are spread over<strong>the</strong> globe.From an examination <strong>of</strong> <strong>the</strong> facts presented by <strong>the</strong> variousclasses <strong>of</strong> vertebrates, we are, <strong>the</strong>n, led to <strong>the</strong> conclusion,that <strong>the</strong>re is no evidence <strong>of</strong> a former land-connection be,tween <strong>the</strong> Australian and Neotropical regions ;but that <strong>the</strong>various scattered resemblances in <strong>the</strong>ir natural productions


chap, xiii.] THE AUSTKALIAN REGION. 403that undoubtedly occur, are probably due to three distinctcauses.First, we have <strong>the</strong> American Didelphyida?, among Mammals,and <strong>the</strong> Cracidse, among birds, allied respectively to <strong>the</strong> Marsupialsand <strong>the</strong> Megapodiidsp, <strong>of</strong> Australia. This is probably morea coincidence than an affinity,due to <strong>the</strong> preservation <strong>of</strong> ancientwide-spread types in two remote areas, each cut <strong>of</strong>f from <strong>the</strong>great nor<strong>the</strong>rn continental masses, inwhich higher forms wereevolved leading to <strong>the</strong> extinction <strong>of</strong> <strong>the</strong> lower types. In each<strong>of</strong> <strong>the</strong>se sou<strong>the</strong>rn isolated lands <strong>the</strong> original type would undergoa special development ; in <strong>the</strong> one case suited to an arborealexistence, in <strong>the</strong> o<strong>the</strong>r to a life among arid plains.<strong>The</strong> second case is that <strong>of</strong> <strong>the</strong> tree-frogs, and <strong>the</strong> genus Osteorjlossumamong fishes ; and is most likely due to <strong>the</strong> extensionand approximation <strong>of</strong> <strong>the</strong> two sou<strong>the</strong>rn continents, and <strong>the</strong> existence<strong>of</strong> some intermediate lands, during a warm period whenfacilities would be afforded for <strong>the</strong> transmission <strong>of</strong> a few organismsby <strong>the</strong> causes which have led to <strong>the</strong> exceptional diffusion<strong>of</strong> fresh-water productions in all parts <strong>of</strong> <strong>the</strong> world. As howeverOsteoglossum occurs also in <strong>the</strong> Sunda Islands, this may be a case<strong>of</strong> survival <strong>of</strong> a once wide-spread group.<strong>The</strong> third case is that <strong>of</strong> <strong>the</strong> same genera and even species <strong>of</strong>fish, and perhaps <strong>of</strong> frogs, in <strong>the</strong> two countries ; which maybe due to transmission from island to island by <strong>the</strong> aid <strong>of</strong> floatingice, <strong>with</strong> or <strong>with</strong>out <strong>the</strong> assistance <strong>of</strong> more interveninglands than now exist.Having arrived at <strong>the</strong>se conclusions from a consideration <strong>of</strong><strong>the</strong> vertebrata, we shall be in a position to examine how far<strong>the</strong> same causes will explain, or agree <strong>with</strong>, <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong>invertebrate groups, or elucidate any special difficulties we maymeet <strong>with</strong> in <strong>the</strong> <strong>relations</strong> <strong>of</strong> <strong>the</strong> sub-regions.Insects.<strong>The</strong> insects <strong>of</strong> <strong>the</strong> Australian region are as varied, and insome respects as peculiar as its higher forms <strong>of</strong> life. As wehave already indicated in our sketch <strong>of</strong> <strong>the</strong> Oriental region, avast number <strong>of</strong> forms inhabit <strong>the</strong> Austro-Malay sub-region


—404 ZOOLOGICAL GEOGRAPHY. [part hi.which are absent from Australia proper. Such <strong>of</strong> <strong>the</strong>se as arecommon to <strong>the</strong> Malay archipelago as a whole, have been alreadynoted;we shall here confine ourselves more especially to <strong>the</strong>groups peculiar to <strong>the</strong> region, which are almost all ei<strong>the</strong>rAustralian or Austro-Malayan, <strong>the</strong> Pacific Islands and NewZealand being very poor in insect life.Lepidoptera.—Australia itselfis poor in butterflies, except inits nor<strong>the</strong>rn and more tropical parts, where green Ornithopterceand several o<strong>the</strong>r Malayan forms occur. In South Australia<strong>the</strong>re are less than thirty-five species, whereas in Queensland <strong>the</strong>reare probably over a hundred. <strong>The</strong> peculiar Australian formsare few. In <strong>the</strong> family Satyridae, Xenica and Heteronympha,<strong>with</strong> Hypocista extending to New Guinea ; among <strong>the</strong> Lycaenidae,Ogyris and Utica are confined to Australia proper, andRypochrysops to <strong>the</strong> region ; and in Papilionidae, <strong>the</strong> remarkableEiirycus is confined to Australia, but is allied to Euryades,a genus found in Temperate South America (La Plata), and to<strong>the</strong> Pamassius <strong>of</strong> <strong>the</strong> North-Temperate zone.<strong>The</strong> Austro-Malay sub-region has more peculiar forms. Hamadryas,a genus <strong>of</strong> Danaidae, approximates to some South Americanforms ; Hyades and Hyantis are remarkable groups <strong>of</strong> Morphidae;Mynes and Prothoe are fine Nymphalidae, <strong>the</strong> former extending toQueensland ;Dicallaneura, a genus <strong>of</strong> Erycinidae, and Elodina,<strong>of</strong> Pieridae, are also peculiar forms. <strong>The</strong> fine ^Egeus group <strong>of</strong>Papilio, and Priamus group <strong>of</strong> Ornithoptera, also belong exclusivelyto this region.Xois is confined to <strong>the</strong> Fiji Islands, Bletogona to Celebes, andAcropthalmia to New Zealand, all genera <strong>of</strong> Satyridae. Seventeengenera in all are confined to <strong>the</strong> Australian region.Among <strong>the</strong> Sphingina, Pollanisus, a genus <strong>of</strong> Zygaenidae, isAustralian ;also four genera <strong>of</strong> Castniidae Synemon, Euschemon,Damias, and Cocytia, <strong>the</strong> latter being confined to <strong>the</strong> Papuanislands. <strong>The</strong> occurrence <strong>of</strong> this o<strong>the</strong>rwise purely South Americanfamily in <strong>the</strong> Australian region, as well as <strong>the</strong> affinity <strong>of</strong> Eurycusand Euryades noticed above, is interesting ; but as we have seenthat <strong>the</strong> genera and families <strong>of</strong> insects are more permanent thanthose <strong>of</strong> <strong>the</strong> higher <strong>animals</strong>, and as <strong>the</strong> groups in question are


;chap, xin.] THE AUSTRALIAN REGION. 405confined to <strong>the</strong> warmer parts <strong>of</strong> both countries, <strong>the</strong>y may be bestexplained as cases <strong>of</strong> survival <strong>of</strong> a once wide-spread type, andmay probably date back to <strong>the</strong> period when <strong>the</strong> ancestors <strong>of</strong> <strong>the</strong>Marsupials and Megapodii were cut <strong>of</strong>f from <strong>the</strong> rest <strong>of</strong> <strong>the</strong> world.Goleoptera.—<strong>The</strong> same remark applies here as in <strong>the</strong> Lepidoptera,respecting <strong>the</strong> affinity <strong>of</strong> <strong>the</strong> Austro-Malay fauna to that <strong>of</strong>Indo-Malay Islands; but Australia proper is much richer inbeetles than in butterflies, and exhibits much more speciality.Although <strong>the</strong> o<strong>the</strong>r two parts <strong>of</strong> <strong>the</strong> Australian region (Polynesiaand New Zealand) are very poor in beetles, it will, never<strong>the</strong>less,on <strong>the</strong> whole compare favourably w T ith any <strong>of</strong> <strong>the</strong> regions except<strong>the</strong> very richest.Cicindelidce are not very abundant.<strong>The</strong>rates and Tricondylaare <strong>the</strong> characteristic genera in Austro-Malaya, but are absentfrom Australia, where we have Tctracha as <strong>the</strong> most characteristicgenus, <strong>with</strong> one species <strong>of</strong> Megacephala and two <strong>of</strong> Distypsidera,a genus which is found also in New Zealand and some<strong>of</strong> <strong>the</strong> Pacific Islands. <strong>The</strong> occurrence <strong>of</strong> <strong>the</strong> South Americangenus, Tetracha, may perhaps be due to a direct transfer bymeans <strong>of</strong> intervening lands during <strong>the</strong> warm sou<strong>the</strong>rn periodbut considering <strong>the</strong> permanence <strong>of</strong> coleopterous forms (as shownby <strong>the</strong> Miocene species belonging almost wholly to existinggenera), it seems more probable that it is a case <strong>of</strong> <strong>the</strong> survival<strong>of</strong> a once wide -spread group.Carabidse are well represented, <strong>the</strong>re being no less than 94peculiar genera, <strong>of</strong> which 19 are confined to New Zealand.<strong>The</strong>Australian genera <strong>of</strong> most importance are Carenum (68 species),Promecoderus (27 species), Silphomorpha (32 species), Adelotopus(27 species), Scaraphites (25 species), Notonomus (18 species),Gnathoxys (12 species), Eutoma (9 species), JEnigma (15 species),Lacordairea (8 species), Pamborus (8 species), Catadromm (4 species),—<strong>the</strong>latter found in Australia and Celebes. Common toAustralia and New Zealand are Mecodema (14 species), Hornalosoma(32 species), Bicrochile (12 species), and Scopodes(5 species).<strong>The</strong> larger genera, confined to New Zealand only, are Metaglymma(8 species), and Demetrida (3 species). <strong>The</strong> curious genusPseudomorpha (10 species), is divided between California, Brazil,


—406 ZOOLOGICAL GEOGRAPHY. [part hi.and Australia ; and <strong>the</strong> Australian genera, Addotopus, Silphomorpha,and Sphallomorpha, form <strong>with</strong> ita distinct tribe <strong>of</strong> Coleoptera.<strong>The</strong>se being all confined to <strong>the</strong> warmer regions, and havingso scattered a <strong>distribution</strong>, are no doubt <strong>the</strong> relics <strong>of</strong> a widespreadgroup. <strong>The</strong> Australian genus, Promecoderus, has, however,closely allied genera (Cascelius and its allies), inChili andPatagonia; while two small genera confined to <strong>the</strong> AucklandIslands (Heterodactylus and Pristancydus) are allied to a groupfound only in Terra- del-Fuego and <strong>the</strong> Falkland Islands,(Migadops) ;and in <strong>the</strong>se cases we may well believe that a directtransmission has taken place by some <strong>of</strong> <strong>the</strong> various meansalready indicated.In Lucanidaa, Australia is only moderately rich, having 7peculiar genera. <strong>The</strong> most important are Ceratognathus and Bhyssonotus,confined to Australia ; Lissotes to Australia and NewZealand ; Lamprima to Australia and Papua. Mitopliyllus andDendroblax inhabit New Zealand only ;while Syndesus is foundin Australia, New Caledonia, and tropical South America.<strong>The</strong> beautiful Cetoniidse are poorly represented, <strong>the</strong>re beingonly 3 peculiar genera ;Schizorhina, mainly Australian, butextending to Papua and <strong>the</strong> Moluccas ; Anacamptorhina, confinedto New Guinea, and Sternoplus to Celebes.Lomaptera isvery characteristic <strong>of</strong> <strong>the</strong> Austro-Malay Islands. This almosttropical family shows no approximations between <strong>the</strong> Australianand Neotropical faunas.In Buprestidae, <strong>the</strong> Australian region is <strong>the</strong> richest, possessingno less than 47 genera, <strong>of</strong> which 20 are peculiar to it. Of <strong>the</strong>se, 15are peculiar to Australia itself, <strong>the</strong> most important being Stigw,odera(212 species), Ethon (13 species), and Nascio (3 species) ;Cisseis (17 species), and <strong>the</strong> magnificent Galodema (3 species),are common to Australia and Austro-Malaya ; while Sambus(10 species) and Anthaxomorpha (4 species), <strong>with</strong> some smallergroups, are peculiarly Austro-Malayan. In this family occurseveral points <strong>of</strong> contact <strong>with</strong> <strong>the</strong> Neotropical region. Stigmoderais said to have a species in Chili, while <strong>the</strong>re areundoubtedlyseveral allied genera in Chili and South Temperate America.<strong>The</strong> genus Curis has 5 Australian and 3 Chilian species, and


—chap, xiii.] THE AUSTRALIAN REGION. 407Acherusia has 2 species in Brazil, 1 in Australia. <strong>The</strong>se resemblancesmay probably have arisen from intercommunicationduring <strong>the</strong> warm sou<strong>the</strong>rn period, when floating timber wouldoccasionally transmit a few larvae <strong>of</strong> this family from island toisland across <strong>the</strong> antarctic seas. When <strong>the</strong> cold period returned,<strong>the</strong>y would spread northward, and become more or less modifiedunder <strong>the</strong> new physical conditions and organic competition, towhich <strong>the</strong>y were subjected.We now come to <strong>the</strong> very important group <strong>of</strong> Longicorns, inwhich <strong>the</strong> Australian region as a whole, is very rich, possessing360 genera, <strong>of</strong> which 263 are peculiar to it. Of <strong>the</strong>se about 50are confined to <strong>the</strong> Austro-Malay Islands, 12 to New Zealand,and <strong>the</strong> remainder to Australia proper <strong>with</strong> Tasmania. Of <strong>the</strong>genera confined to, or highly characteristic <strong>of</strong> Australia, <strong>the</strong>following are <strong>the</strong> most important :Cnemoplites,belonging to<strong>the</strong> Prionidae ; Phoracantha, to <strong>the</strong> Cerambycidse ; Zygocera,Hebecerus, Symphyletes, and Rhytidophora, to <strong>the</strong> Lamiida?.Confined to <strong>the</strong> Austro-Malay Islands are Tethionea (Cerambycidse): 1 mesisternus, Arrhenotus, Micracantha, and Sybra(Lamiidae) ;but <strong>the</strong>re are also such Malayan genera as BatoceraGnoma, Praonetha, and Sphenura, which are very abundant in<strong>the</strong> Austro-Malay sub-region. A species <strong>of</strong> each <strong>of</strong> <strong>the</strong> Australiangenera, Zygocera, Syllitus, and Pseudocephalus, is said tooccur in Chili, and one <strong>of</strong> <strong>the</strong> tropical American genus, Hammatochccrus,in tropical Australia ; an amount <strong>of</strong> resemblancewhich, as in <strong>the</strong> case <strong>of</strong> <strong>the</strong> Buprestidae, may be imputedto trans-oceanic migration during <strong>the</strong> Sou<strong>the</strong>rn warm period.This concludes our illustrations <strong>of</strong> <strong>the</strong> <strong>distribution</strong> <strong>of</strong> some <strong>of</strong><strong>the</strong> more important groups <strong>of</strong> Australian insects ; and it will beadmitted that we have not met <strong>with</strong> any such an amount <strong>of</strong>identity <strong>with</strong> <strong>the</strong> fauna <strong>of</strong> Temperate South America, as torequire us to modify <strong>the</strong> conclusions we arrived at from a consideration<strong>of</strong> <strong>the</strong> vertebrategroups.Land-Shells.—<strong>The</strong> <strong>distribution</strong> <strong>of</strong> many <strong>of</strong> <strong>the</strong> larger genera<strong>of</strong> land-shells is very erratic, while o<strong>the</strong>rs are exceedingly restricted,so that it requires an experienced conchologist toinvestigate <strong>the</strong> affinities <strong>of</strong> <strong>the</strong> several groups, and thus work


;408 ZOOLOGICAL GEOGRAPHY. [part hi.out <strong>the</strong> important facts <strong>of</strong> <strong>distribution</strong>. All that can be donehere is to note <strong>the</strong> characteristic and peculiar genera, and anyo<strong>the</strong>rs presenting features <strong>of</strong> special interest.In <strong>the</strong> great family <strong>of</strong> <strong>the</strong> snails (Helicidse), <strong>the</strong> only generastrictly confined to <strong>the</strong> region are, Partula, now containing above100 species, and ranging over <strong>the</strong> Pacific from <strong>the</strong> Solomon Isleson <strong>the</strong> west, to <strong>the</strong> Sandwich Islands and Tahiti on <strong>the</strong> eastand Achatinella, now containing nearly 300 species, and whollyconfined to <strong>the</strong> Sandwich Islands. Pfeiffcria is confined to <strong>the</strong>Philippine Islands and Moluccas ;Cochlostyla to <strong>the</strong> Indo-MalayIslands and Australia ;Bulimus occurs in most <strong>of</strong> <strong>the</strong> insulargroups, including New Zealand, but is absent from Australia.Among <strong>the</strong> Aciculidse, <strong>the</strong> widely-scattered Truncatella is <strong>the</strong>only genus represented.Among DiplommatinidaB, Diplommatinais <strong>the</strong> characteristic genus, ranging over <strong>the</strong> whole region,and found elsewhere as far as India, <strong>with</strong> one species inTrinidad. <strong>The</strong> extensive family Cyclostomidse, is not wellrepresented. Seven genera reach <strong>the</strong> Austro-Malay Islands,one <strong>of</strong> which, Eegistoma, is confined to <strong>the</strong> Philippines, Moluccas,New Caledonia, and <strong>the</strong> Marshall Islands. Omphalotropisis <strong>the</strong> most characteristic genus, ranging over <strong>the</strong> whole regionGallia is confined to <strong>the</strong> Philippines, Ceram, and Australia;Bealia to New Zealand and <strong>the</strong> Marquesas. <strong>The</strong> genus Helicinaalone represents <strong>the</strong> Helicinidse, and is found in <strong>the</strong> whole regionexcept New Zealand. <strong>The</strong> number <strong>of</strong> species known fromAustralia is perhaps about 300; while <strong>the</strong> Polynesian sub-region,according to Mr. Harper Pease, contains over 600 ;<strong>the</strong> Austro-Malay Islands will furnish probably 200; and New Zealandabout 100 ; making a total <strong>of</strong> about 1,200 species for <strong>the</strong> wholeregion.AUSTEALIAN SUR-EEGIONS.Few <strong>of</strong> <strong>the</strong> great zoological regions comprise four divisions sostrongly contrasted as <strong>the</strong>se, or which present so many interestingproblems. We have first <strong>the</strong> Austro-Malay Islands, an equatorialforest-region teeming <strong>with</strong> varied and beautiful forms <strong>of</strong> lifenext we have Australia itself, an island-continent <strong>with</strong> its satellite


chap, xiii.] THE AUSTRALIAN REGION. 409Tasmania, both tropical and temperate, but for <strong>the</strong> most partarid, yet abounding in peculiar forms in all <strong>the</strong> classes <strong>of</strong> <strong>animals</strong>;<strong>the</strong>n come <strong>the</strong> Polynesian Islands, ano<strong>the</strong>r luxuriant region <strong>of</strong>tropical vegetation, yet excessively poor in most <strong>of</strong> <strong>the</strong> highergroups <strong>of</strong> <strong>animals</strong> as well as in some <strong>of</strong> <strong>the</strong> lower ; and lastly,we have New Zealand, a pair <strong>of</strong> temperate forest-clad islandsfar in <strong>the</strong> sou<strong>the</strong>rn ocean, <strong>with</strong> a very limited yet strange andalmost wholly peculiar fauna. We have now to consider <strong>the</strong>general features and internal <strong>relations</strong><strong>of</strong> <strong>the</strong> faunas <strong>of</strong> each <strong>of</strong><strong>the</strong>se sub-regions, toge<strong>the</strong>r <strong>with</strong> any external <strong>relations</strong> whichhave not been discussed while treating <strong>the</strong> region asI. Austro-Malayan Sub-region.a whole.<strong>The</strong> central mass on which almost every part <strong>of</strong> this subregionis clearly dependent, is <strong>the</strong> great island <strong>of</strong> New Guinea,inhabited by <strong>the</strong> Papuan race <strong>of</strong> mankind ; and this, <strong>with</strong> <strong>the</strong>surrounding islands, which are separated from it by shallow seasand possess its most marked zoological features, are termed Papua,A little fur<strong>the</strong>r away lie <strong>the</strong> important groups <strong>of</strong> <strong>the</strong> Moluccason one side and <strong>the</strong> Eastern Papuan Islands on <strong>the</strong> o<strong>the</strong>r, whichpossess a fauna mainly derivative from New Guinea, yet wantingmany <strong>of</strong> its distinctive types ; and, in <strong>the</strong> case <strong>of</strong> <strong>the</strong> Moluccaspossessing many groups which are not Australian, but derivedfrom <strong>the</strong> adjacent Oriental region. To <strong>the</strong> south <strong>of</strong> <strong>the</strong>se wehave <strong>the</strong> Timor group, whose fauna is clearly derivative, fromAustralia, from Java, and from <strong>the</strong> Moluccas. Lastly comesCelebes, whose fauna is most complex and puzzling, and, so faras we can judge, not fundamentally derivative from any <strong>of</strong> <strong>the</strong>surrounding islands.Papua, or <strong>the</strong> Neiv &uinea Group.—New Guinea is verydeficient in Mammalia as compared <strong>with</strong> Australia, though thisapparent poverty may, in part, depend on our very scanty knowledge.As yet only four <strong>of</strong> <strong>the</strong> Australian families <strong>of</strong> Marsupialsare known to inhabit it, <strong>with</strong> nine genera, several <strong>of</strong> whichare peculiar. It also possesses a peculiar form <strong>of</strong> wild pig;but as yetno o<strong>the</strong>r non-marsupial terrestrial mammal has beendiscovered, except a rat, described by Dr. Gray as UromysVol. L—28


—410 ZOOLOGICAL GEOGRAPHY. [part hi.aruensis, but about <strong>the</strong> locality <strong>of</strong> which <strong>the</strong>re seems somedoubt. 1 Omittiug bats, <strong>of</strong> which our knowledge is very imperfect,<strong>the</strong> Papuan Mammals are as follows :Family.


..CHAP. XIII.] THE AUSTRALIAN REGION. 411This Papuan fauna is so interesting and remarkable, that itseems advisable to give lists <strong>of</strong> <strong>the</strong>se several classes <strong>of</strong> generictypes.I. Genera occurring in <strong>the</strong> Papuan Islands which are characteristic<strong>of</strong> <strong>the</strong> Australian region (89). Those marked <strong>with</strong> anasterisk are exclusively Papuan.Sylviidae...CerthiidaeSittidse ...Oriolidae . .CampephagidaeDicruridaeMudcicapidsePachycephalidae.Laniidae ...Corvidae ...ParadiseidaeMeliphagidseNectariniidaeArtamidaePittidae ...CuculidaeAlcedinidaePodargidaeCaprimulgidaeCacatuidaePlatycercidaePalseornithidaeTrichoglossidaeNestoridaeColumbidaeMegapodiidaeFalconidaeCasuariidaeMalurus, Gerygone, Petroica, Orthonyx.Climactcris.Sittella.Mimeta.Graucalus, Lalage.*Chcvtorhyuchus.*Peltops, Monarcha, *Leucophantes, Micrmca,Sisura, Myiagra, *Mach(erirhynchus, Bhipidura,*Todopsis.Pachycephala.*Rectes.Cracticus, *Gymnocorvtis.*Paradisea, *Manucodia, *Astrapia, *Parotia,*Lophorina, *Diphyllodes, *Xanthomelus,*Cicinnurus, *Paradigalla, *Epimachus,*l)repanomis, *Seleucides, Ptilorlds,JElurozdus,*Amblyornis.Myzomela, Entomophila. Glicyphila, Ptilotis,*Melidectes, *Melipotes, *Melirrhophetes,Anthochcera, Philemon, *Euthyrhynchus,Melithreptes.Chalcostetha, *Cosmetira.Artamus.*Mclampitta.*Caliechthrus.Alcyone, *Syma, Dacelo, *Tanysiptera,*Melidora.Podargus, ^Ego<strong>the</strong>les.Eurostopodus.Cacatua, *Microglossus, Licmetis, *Nasiterna.AprosmictusTanygnathus, Eclectus, Ge<strong>of</strong>froyus, *Cyclopsitta.Trichoglossus, *Charmosyna, Eos, Lorius.*Dasyptilus.Ptilopus, Carpophaga, Ianthoenas, Reinwardtcenas,*Trugon, *Henicophaps, Phlogoenas,*Otidiphaps, *Goura.Talegallus, Megapodius.* Henicopernis.Casuarius.<strong>The</strong> chief points <strong>of</strong> interest here are <strong>the</strong> richness and specialization<strong>of</strong> <strong>the</strong> parrots, pigeons, and kingfishers; <strong>the</strong> wonderful paradise-birds; <strong>the</strong> honeysuckers ; and some remarkable flycatchers.


——412 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>The</strong> most prominent deficiencies, as compared <strong>with</strong> Australia,are in Sylviidse, Timaliidse, Ploceidse, Platycercidse,and Falconidse.II. <strong>The</strong> genera which are characteristic <strong>of</strong> <strong>the</strong> whole MalayArchipelago are <strong>the</strong> following (10) :1. Erythrura ... (Ploceida?) 6. Loriculus ... (Psittacidee)2. Pitta (Pittidse) 7. Macropygia ... (Columbidse)3. Ceyx ... ... (Alcedinidse) 8. Chalcophaps ... „4. Calao ... (Bucerotidse) 9. Calcenas ... „5. Dendrochelidon (Cypselidee) 10. Baza ... ... (Falconidae)III. <strong>The</strong> curious set <strong>of</strong> genera apparently <strong>of</strong> Indo-Malayanorigin, but unknown in <strong>the</strong> Moluccas, are as follows :1. Eupetes ... (Cinclidse) 4. Arachno<strong>the</strong>ra (Nectariniidse)2. Alcippe ... (Timaliida?) 5. Prionochilus... (Dicseidse)3. Pomatorhinus „ 6. EulaLes ... (Sturnidse)<strong>The</strong> above six birds are very important as indicating pastchanges in <strong>the</strong> Austro-Malay Islands,and we must say a fewwords about each. (1) Eupetes is very remarkable, since <strong>the</strong>New Guinea birds resemble in all important characters thatwhich is confined to Malacca and Sumatra. <strong>The</strong>y are probably<strong>the</strong> survivors <strong>of</strong> a once wide-spread Malayan group.(2) Alcippe or Drymocataphus (for in which genus <strong>the</strong> birdsshould be placed is doubtful) seems ano<strong>the</strong>r clear case <strong>of</strong>a typical Indo-Malayan form occurring in New Guinea andJava, but in no intervening island.(3) Pomatorhinus is a mostcharacteristic Himalayan and Indo-Malayan genus, occurringagain in New Guinea and alsoin Australia, but in no intermediateisland. <strong>The</strong> New Guinea bird seems as nearly related toOriental as Australian species. (4) Arachno<strong>the</strong>ra is exactlyparallel to Alcippe, occurring nowhere east <strong>of</strong> Borneo except inNew Guinea. (5) Prionochilus, a small black bird, sometimesclassed as a distinct genus, but evidently allied to <strong>the</strong> Prionochili <strong>of</strong><strong>the</strong> Indo-Malay Islands.(6) Eulahes, <strong>the</strong> genus which contains<strong>the</strong> well known Mynahs <strong>of</strong> India, extends east <strong>of</strong> Java as far as<strong>The</strong> two NewPlores, but is not found in Celebes or <strong>the</strong> Moluccas.Guinea species are sometimes classed in different genera, but <strong>the</strong>yare undoubtedly allied to <strong>the</strong> Mynahs <strong>of</strong> India and Malaya.We find <strong>the</strong>n, that while <strong>the</strong> ornithology <strong>of</strong> New Guinea is


chap, xiii.] THE AUSTRALIAN REGION. 413preeminently Australian in character and possesses many peculiardevelopments <strong>of</strong> Australian types, it has also—as might be expectedfrom its <strong>geographical</strong> position, its climate, and its vegetation—receivedan infusion <strong>of</strong> Malayan forms. But while onegroup <strong>of</strong> <strong>the</strong>se isspread over <strong>the</strong> whole Archipelago, and occasionallybeyond it, <strong>the</strong>re is ano<strong>the</strong>r group which presents <strong>the</strong>unusual and interesting feature <strong>of</strong> discontinuous <strong>distribution</strong>,jumping over a thousand miles <strong>of</strong> island-studded sea from Javaand Borneo to New Guinea itself. It is a parallel case to that<strong>of</strong> Java in <strong>the</strong> Oriental region, which we have already discussed,but <strong>the</strong> suggested explanation in that case is more difficult toapply here. <strong>The</strong> recent soundings by <strong>the</strong> Challenger show us,that although <strong>the</strong> several islands <strong>of</strong> <strong>the</strong> Moluccas are surroundedby water from 1,200 to 2,800 fathoms deep, yet <strong>the</strong>se seas forminclosed basins <strong>with</strong> rims not more than from 400 to 900fathoms deep, suggesting <strong>the</strong> idea <strong>of</strong> great lakes or inland seaswhich have sunk down bodily <strong>with</strong> <strong>the</strong> surrounding land, or thatenormous local and restricted elevations and subsidences havehere occurred. We have also <strong>the</strong> numerous small islands and coralbanks south <strong>of</strong> Celebes and eastward towards Timor-Laut and <strong>the</strong>Aru Islands, indicating great subsidence ; and it is possible that<strong>the</strong>re was an extension <strong>of</strong> Papua to<strong>the</strong> west, approaching sufficientlynear to Java to receive occasional straggling birds <strong>of</strong> Indo-Malay type, altoge<strong>the</strong>r independent <strong>of</strong> <strong>the</strong> Moluccas to <strong>the</strong> north.Bright Colours and Ornamental Plumage <strong>of</strong> New Guinea Birds.—One <strong>of</strong> <strong>the</strong> most striking features <strong>of</strong> Papuan ornithology is <strong>the</strong>large proportion which <strong>the</strong> handsome and bright-coloured birdsbear to <strong>the</strong> more obscure species. That this is really <strong>the</strong> casehas been ascertained by going over my own collections, made atAru and New Guinea, and comparing <strong>the</strong>m <strong>with</strong> my collectionmade at Malacca—a district remarkable for <strong>the</strong> number <strong>of</strong> handsomebirds it produces. Using, as nearly as possible, <strong>the</strong> samestandard <strong>of</strong> beauty, about one-third <strong>of</strong> <strong>the</strong> Malacca birds may beclassed as handsome, 1 while in Papua <strong>the</strong> proportion comes outexactly one-half.This is due, in part to <strong>the</strong> great abundance <strong>of</strong>1 I also find about this proportion in my Amazonian collections, evencounting all <strong>the</strong> humming-birds, parrots, and toucans as handsome birds.


;414 ZOOLOGICAL GEOGRAPHY. [part hi.parrots, cockatoos, and lories,almost all <strong>of</strong> which are beautifuland <strong>of</strong> pigeons, more than half <strong>of</strong> which are very beautiful ; aswell as to <strong>the</strong> numerous kingfishers, most <strong>of</strong> which are excessivelybrilliant. <strong>The</strong>n we have <strong>the</strong> absence <strong>of</strong> thrushes, and <strong>the</strong> verysmall numbers <strong>of</strong> <strong>the</strong> warblers, shrikes, and Timaliidse, which aredull-coloured groups ;and, lastly, <strong>the</strong> presence <strong>of</strong> numerous gaypittas, flycatchers, and <strong>the</strong> unequalled family <strong>of</strong> paradise-birds.A large number <strong>of</strong> birds adorned <strong>with</strong> metallic plumage is also amarked feature <strong>of</strong> this fauna, more than a dozen genera being sodistinguished. Among <strong>the</strong> remarkable forms are Peltops, a flycatcher,long classed asone <strong>of</strong> <strong>the</strong> Indo-Malayan Eurylsemidae,which it resembles both in bill and coloration; Machcerirhynchus,curious little boat-billed flycatchers ;and Toclopsis, a group <strong>of</strong> terrestrialflycatchers <strong>with</strong> <strong>the</strong> brilliant colours <strong>of</strong> Pitta or Malurus.<strong>The</strong> paradise-birds present <strong>the</strong> most wonderful developments <strong>of</strong>plumage and <strong>the</strong> most gorgeous varieties <strong>of</strong> colour, to be foundamong passerine birds. <strong>The</strong> great whiskered-swift, <strong>the</strong> handsomestbird in <strong>the</strong> entire family, has its head-quarters here. Among kingfishers<strong>the</strong> elegant long-tailed Tanysvpterm are preeminent, whe<strong>the</strong>rfor singularity or beauty.Among parrots, New Guinea possesses<strong>the</strong> great black cockatoo, one <strong>of</strong> <strong>the</strong> largest and most singular birdsin <strong>the</strong> order ; Nasiterna, <strong>the</strong> smallest <strong>of</strong> known parrots ; andCharmosyna, perhaps <strong>the</strong> most elegant. Lastly, among <strong>the</strong>pigeons we have <strong>the</strong> fine crowned- pigeons, <strong>the</strong> largest and mostremarkable group <strong>of</strong> <strong>the</strong> order.Plate X. Illustrating <strong>the</strong> Ornithology <strong>of</strong> New G'ttinea.—<strong>The</strong>wonderful ornithological fauna we have just sketched, couldonly be properly represented in a series <strong>of</strong> elaborate colouredplates. We are obliged here to confine ourselves to representinga few <strong>of</strong> <strong>the</strong> more remarkable types <strong>of</strong> form, as samples <strong>of</strong> <strong>the</strong>great number that adorn this teeming bird-land. <strong>The</strong> largecentral figure is <strong>the</strong> fine twelve-wired paradise-bird {Seleucidesalbus), one <strong>of</strong> <strong>the</strong>most beautiful and remarkable <strong>of</strong> <strong>the</strong> family.Its general plumage appears, at first sight, to be velvety blackbut on closer examination, and by holding <strong>the</strong> bird in variouslights, it is found that every part <strong>of</strong> it glows <strong>with</strong> <strong>the</strong> most exquisitemetallic tints—rich bronze, intense violet, and, on <strong>the</strong>


-^PLATE X.z^f-^


—chap, xiii.] THE AUSTRALIAN REGION. 415edges <strong>of</strong> <strong>the</strong> breast-fea<strong>the</strong>rs, brilliant green.An immense tuft <strong>of</strong>dense plumes <strong>of</strong> a fine orange-buff colour, springs from each side<strong>of</strong> <strong>the</strong> body, and six <strong>of</strong> <strong>the</strong>se on each side terminate in a blackcurled rachis orshaft, which form a perfectly unique adornmentto this lovely bird. To appreciate this wonderful family (<strong>of</strong>which no good mounted collection exists) <strong>the</strong> reader shouldexamine <strong>the</strong> series <strong>of</strong> plates in Mr. Elliot's great work on <strong>the</strong>Paradiseidse, where every species is figured <strong>of</strong> <strong>the</strong> size <strong>of</strong> life,and<strong>with</strong> a perfection <strong>of</strong> colouring that leaves little to be desired.Below <strong>the</strong> Seleucides is one <strong>of</strong> <strong>the</strong> elegant racquet-tailedking-hunters (Tanysiptera galatea) whose plumage <strong>of</strong> vivid blueand white, and coral-red bill,combined <strong>with</strong> <strong>the</strong> long spatulatetail, renders this bird one <strong>of</strong> <strong>the</strong> most attractive <strong>of</strong> <strong>the</strong> interestingfamily <strong>of</strong> kingfishers. On a high branch is seated <strong>the</strong> littlePapuan parroquet (Charmosyna papiiensis), one <strong>of</strong> <strong>the</strong> Trichoglossidse,or brush-tongued parrots,—richly adorned in red andyellow plumage, and <strong>with</strong> an unusually long and slender tail.<strong>the</strong> ground is <strong>the</strong> well-known crowned pigeon (Goura coronata,)a genus which is wholly confined to New Guinea and a few <strong>of</strong><strong>the</strong> adjacent islands. One <strong>of</strong> <strong>the</strong> very few Papuan mammals, atree-kangaroo (Dendrolagus inustus), is seated on a high branch.It is interesting, as an arboreal modification <strong>of</strong> a family which inAustralia is purely terrestrial ; and as showing how very littlealteration <strong>of</strong> form or structure is needed to adapt an animal tosuch a different mode <strong>of</strong> life.OnReptiles and Amphibia.—Of <strong>the</strong>se classes comparatively littleis at present known, but <strong>the</strong>re is evidence that <strong>the</strong> same intermixture<strong>of</strong> Oriental and Australian forms that occurs in birds andinsects, is also found here. Dr. A. B. Meyer, <strong>the</strong> translator <strong>of</strong> thiswork into German, and well known for his valuable discoveries inNew Guinea, has kindly furnished me <strong>with</strong> a manuscript list <strong>of</strong>Papuan reptiles, from which most <strong>of</strong> <strong>the</strong> information I am ableto give is derived.Of Snakes, 24 genera are known, belonging to 11 families. Six<strong>of</strong> <strong>the</strong> genera are Oriental, Calamaria, Cerberus, Clirysopelea,Lycodon, Chersydrus, and Ophiophagus. Four are Australian,


———416 ZOOLOGICAL GEOGRAPHY. [part hi.— Morelia, Liasis, Diemenia, and Acanthophis ; while four o<strong>the</strong>rsare more especially Papuan, Dibamus (Typhlopidse), Brachyorros—asub-genus <strong>of</strong> <strong>the</strong> wide-spread Rhabdosoma (Calamariidae),found also in Timor ; Nardoa and Enygrus (Pythonidse), rangingfrom <strong>the</strong> Moluccas to <strong>the</strong> Fiji Islands. <strong>The</strong> rest are ei<strong>the</strong>r commonto <strong>the</strong> Oriental and Australian regions or <strong>of</strong> wide range.Of Lizards also, 24 genera are recorded, belonging to 5 families.Three only are peculiarly Oriental,Eumeces, Tiaris, and Nycteridium; but ano<strong>the</strong>r, Oonyocephalus, is Malayan, ranging fromJava and Borneo to <strong>the</strong> Pelew Islands. Three are Australian,Gyclodus, Heteropus, and Gehyra ; while six are especially Papuan,— Kcneuxia (extending to <strong>the</strong> Philippines), Mania, Garlia (toNorth Australia), Lipinia (to <strong>the</strong> Philippine Islands), and Tribolonotus,—all belonging to <strong>the</strong> Scincidae ;and Arua belongingto <strong>the</strong> Agamidee. We must add Cryptobhpliarus, which is confinedto <strong>the</strong> Australian region, except a species in Mauritius.<strong>The</strong> o<strong>the</strong>r genera have a wider <strong>distribution</strong>.<strong>The</strong> preponderant Oriental element in <strong>the</strong> snakes as compared<strong>with</strong> <strong>the</strong> lizards, issuggestive <strong>of</strong> <strong>the</strong> dispersal <strong>of</strong> <strong>the</strong> former beingdependent on floating trees, or even on native canoes, which foran unknown period have traversed <strong>the</strong>se seas, and in whichvarious species <strong>of</strong> snakes <strong>of</strong>ten secrete <strong>the</strong>mselves. Tins seems<strong>the</strong> more probable, as snakes are usually more restricted in <strong>the</strong>irrange than lizards, and exhibit less numerous examples <strong>of</strong> widespreadgenera and species. <strong>The</strong> o<strong>the</strong>r orders <strong>of</strong> reptiles presentno features <strong>of</strong> interest.Of Amphibia only 8 genera are known, belonging to 6 families.Eana, Hylarana, and Hyla are wide-spread genera, <strong>the</strong> formerbeing, however, absent from Australia. Hyperolius, Pclodryas,Litoria, and Aderophrys are Australian; while Platymantis isPolynesian, <strong>with</strong> a species in <strong>the</strong> Philippine Islands. Henceit appears that <strong>the</strong> amphibia, so far as yet known, exhibitno Oriental affinity; and this is a very suggestive fact. Wehave seen (p. 29) that salt water is almost a complete barrierto <strong>the</strong> dispersal <strong>of</strong> <strong>the</strong>se creatures ; so that <strong>the</strong> wholly Australiancharacter <strong>of</strong> <strong>the</strong> Papuan batrachia is what we mightexpect, if, as here advocated, no actualland connection between


chap, xiii.] THE AUSTRALIAN REGION. 417<strong>the</strong> Oriental and Australian regions, has probably occurred during<strong>the</strong> entire Tertiary and Post-tertiary periods.Insects.—<strong>The</strong> general character <strong>of</strong> <strong>the</strong> Papuan insects hasbeen sufficiently indicated in our sketch <strong>of</strong> <strong>the</strong> Entomology <strong>of</strong><strong>the</strong> region. We will here only add, that <strong>the</strong> metallic lustre soprevalent among <strong>the</strong> birds, is also apparent in such insects asSphingnotus mirabilis,a, most brilliant metallic Longicorn ; Lomapteraivallacei and Anacamptorhina fidgicla, Cetonii <strong>of</strong> intenselustre ; Calodema ivallacei among <strong>the</strong> Buprestidae ;and <strong>the</strong> elegantblue Eaplioli among <strong>the</strong> weevils. Even among mothswe have Cocytia durvillii, remarkable for its brilliant metalliccolours.<strong>The</strong> Moluccas.—<strong>The</strong> islands <strong>of</strong> Gilolo, Bouru, and Ceram, <strong>with</strong>several smaller islands adjacent, toge<strong>the</strong>r <strong>with</strong> Sanguir, andperhaps Tulour or Salibaboo to <strong>the</strong> north-west, and <strong>the</strong> islandsfrom Ke to Timor-Laut to <strong>the</strong> south-east, form <strong>the</strong> group <strong>of</strong> <strong>the</strong>Moluccas or Spice- Islands, remarkable for <strong>the</strong> luxuriance <strong>of</strong><strong>the</strong>ir vegetation and <strong>the</strong> extreme beauty <strong>of</strong> <strong>the</strong>ir birds andinsects. <strong>The</strong>ir Mammalia are <strong>of</strong> Papuan character, <strong>with</strong> someforeign intermixture. Two genera <strong>of</strong> <strong>the</strong> New Guinea marsupials,Belideus and Cuscus, abound ; and we have also <strong>the</strong> widespreadSus. But besides <strong>the</strong>se, we find no less than five genera<strong>of</strong> placental Mammals quite foreign to <strong>the</strong> Papuan or Australianfaunas. <strong>The</strong>se are 1. Cynopi<strong>the</strong>cus nigrescens, found only in <strong>the</strong>small island <strong>of</strong> Batchian, and probably introduced from Celebes,where <strong>the</strong> same ape occurs. 2. Viverra tangalunga, a commonIndo-Malayan species <strong>of</strong> civet, probably introduced. 3. Cervushippelaphus, var.Moluccensis, a deer abundant in all <strong>the</strong> islands,very close to a Javan species and almost certainly introducedby man, perhaps very long ago. 4. Babirusa alfurus, <strong>the</strong>babirusa, found only in <strong>the</strong> island <strong>of</strong> Bouru, and perhaps originallyintroduced from Celebes. 5. Sorex sp., small shrews.With <strong>the</strong> exception <strong>of</strong> <strong>the</strong> last, all <strong>the</strong>se species are <strong>animals</strong>habitually domesticated and kept in confinement by <strong>the</strong> Malays ;and when we consider that none <strong>of</strong> <strong>the</strong> smaller Mammalia <strong>of</strong> Javaand Borneo, numbering at least fifty different species, are found


——418 ZOOLOGICAL GEOGRAPHY. [part. hi.in any <strong>of</strong> <strong>the</strong> Moluccas, we can hardly suppose that such large<strong>animals</strong> as <strong>the</strong> deer and ape, could have reached <strong>the</strong>m bynatural means. <strong>The</strong>re is every reason to believe, <strong>the</strong>refore, that<strong>the</strong> indigenous Mammalia <strong>of</strong> <strong>the</strong> Moluccas are wholly <strong>of</strong> Papuanstock, and very limited in number.<strong>The</strong> birds are much more varied and interesting. About 200species <strong>of</strong> land-birds are now known, belonging to 85 genera. Of<strong>the</strong> species about 15 are Indo-Malayan, 32 Papuan, and about140 peculiar. Of <strong>the</strong> genera only two are peculiar, Semioptera,a paradise bird, and Lycocorax, a singular form <strong>of</strong> Corvidse ;but<strong>the</strong>re is also a peculiar rail-like wader, Habroptila. One genus,Basilornis, is found only in Ceram and Celebes ;ano<strong>the</strong>r, Scythrops,is Australian, and perhaps a migrant. About 30 genera arecharacteristic Papuan types, and 37 o<strong>the</strong>rs, <strong>of</strong> more or lesswiderange, are found in New Guinea and were <strong>the</strong>refore probablyderived <strong>the</strong>nce. <strong>The</strong>re remains a group <strong>of</strong> birds which are notfound in New Guinea, and are ei<strong>the</strong>r Palasarctic or Oriental.<strong>The</strong>se are 13 in number as follows :1. Monticola. 8. Corydalla.2. Acrocephalus. 9. Hydrornis.3. Cisticola. 10. Batrachostomus.4 Hypolais. 11. Loriculus.5. Criniger. 12. Treron.6. Butalis. 13. Neopus.7. Budytes.Of <strong>the</strong>se <strong>the</strong> Monticola, found only in Gilolo, appears tobe astraggler or migrant from <strong>the</strong> Philippine islands. Acrocephalus,<strong>of</strong> which four species occur, is a wide-spread group ; one <strong>of</strong><strong>the</strong> Moluccan birds is an Australian and ano<strong>the</strong>r a North-Asianspecies, which perhaps indicates that <strong>the</strong>re has long been somemigration southward from island to island, across <strong>the</strong> Moluccas.Cisticola is a genus <strong>of</strong> very wide range, extending to Australia.Hypolais is probably a modified form <strong>of</strong> a Chinese or Javanesespecies. Criniger is a pure Indo-Malay form, representedhere by three fine species. Butalis is a Chinese species, nodoubt straggling southward. Budytes and Corydalla are widespreadOriental and Palsearctic species or slight modifications <strong>of</strong><strong>the</strong>m. Hydrornis is a Malayan form <strong>of</strong> Pittidse. Batrachostomusis a distinct representative <strong>of</strong> a purely Indo-Malay genus. Lori-


—;CHAP. xaii.J THE AUSTRALIAN REGION. 419cuius is Malayan, and especially Philippine, but itreaches as faras Mysol. Treron is here at its eastern limit, and is representedin Bouru and Ceram by one <strong>of</strong> <strong>the</strong> most beautiful species.Ncopus, a Malayan eagle, is said to occur in <strong>the</strong> Moluccas. Wefind <strong>the</strong>n only three characteristic Indo-Malay • types in <strong>the</strong>Moluccas,Griniger, Batrachostomus, and Treron.All are representedby distinct and well marked species, indicating a somewhatremote period since <strong>the</strong>ir ancestors entered <strong>the</strong> districtbut allare birds <strong>of</strong> considerable powers <strong>of</strong> flight, so that a verylittle extension <strong>of</strong> <strong>the</strong> islands in a south-westerly directionwould afford <strong>the</strong> means <strong>of</strong> transmission, but this could not wellhave been by way <strong>of</strong> Celebes, because <strong>the</strong> two former genera areunknown in that island.It is evident, <strong>the</strong>refore, that <strong>the</strong> Moluccas are wholly Papuanin <strong>the</strong>ir zoology;yet <strong>the</strong>y are no less clearly derivative, and musthave obtained <strong>the</strong>ir original immigrants under conditions thatrendered a full representation <strong>of</strong> <strong>the</strong> fauna impossible. Suchremarkable and dominant types as<strong>the</strong> eleven genera <strong>of</strong> Paradiseidse,<strong>with</strong> Cracticus, Rectcs,Todopsis, Machccrirhynchus, Ge?ygone,Dacelo, Podargus, Cyclopsitta, Microglossum., Nasiterna, Chalcopsitta,and Goura,—all characteristic Papuan groups, found inalmost all <strong>the</strong> islands and most <strong>of</strong> <strong>the</strong>m very abundant, aretotally absent from <strong>the</strong> Moluccas. Taking this, in conjunction<strong>with</strong> <strong>the</strong>yetabsence <strong>of</strong> <strong>the</strong> two genera <strong>of</strong> Papuan kangaroos and<strong>the</strong> o<strong>the</strong>r smaller groups <strong>of</strong> marsupials, and we must beconvinced that <strong>the</strong> Moluccas cannot be mere fragments <strong>of</strong> <strong>the</strong>old Papuan land, or <strong>the</strong>y would certainly, in some one oro<strong>the</strong>r<strong>of</strong> <strong>the</strong>ir large and fertile islands, have preserved a more completerepresentation <strong>of</strong> <strong>the</strong> parent fauna. Most <strong>of</strong> <strong>the</strong> Mol^canbirds are very distinct from <strong>the</strong> allied species <strong>of</strong> New Guineaand this would imply that <strong>the</strong> entrance <strong>of</strong> <strong>the</strong> original formstook place at a remote period. <strong>The</strong> two peculiar genera <strong>with</strong>clearly Papuan affinities, show <strong>the</strong> same thing.<strong>The</strong> cassowary,found only in <strong>the</strong> large island <strong>of</strong> Ceram and distinct from anyPapuan species, would however seem to have required a landconnection for its introduction, almost as much as any <strong>of</strong> <strong>the</strong>larger mammalia.


420 ZOOLOGICAL GEOGRAPHY. [part hi.Taking all <strong>the</strong> facts into consideration, I would suggest as <strong>the</strong>most probable explanation, that if <strong>the</strong> Moluccas ever formed part<strong>of</strong> <strong>the</strong> main Papuan land, <strong>the</strong>y were separated at an early date,and subsequently so greatly submerged as to destroy a largeproportion <strong>of</strong> <strong>the</strong>ir fauna. <strong>The</strong>y have since risen, and haveprobably been larger than at present, and ra<strong>the</strong>r more closelyapproximated to <strong>the</strong> parent land, whence <strong>the</strong>y received a considerableimmigration <strong>of</strong> such <strong>animals</strong> as were adapted to crossnarrow seas. This gave <strong>the</strong>m several Papuan forms, but stillleft <strong>the</strong>m <strong>with</strong>out a number <strong>of</strong> <strong>the</strong> types more especially confinedto <strong>the</strong> forest depths, or powerful enough to combat <strong>the</strong>gales which <strong>of</strong>ten blow weaker flyers out to sea. Most <strong>of</strong> <strong>the</strong>birds whose absence from <strong>the</strong> Moluccas is so conspicuous belongto one or o<strong>the</strong>r <strong>of</strong> <strong>the</strong>se classes.Among <strong>the</strong> most characteristic birds <strong>of</strong> <strong>the</strong> Moluccas are <strong>the</strong>handsome crimson lories <strong>of</strong> <strong>the</strong> genera Lorius and Eos. <strong>The</strong>seare found in every island (but not in Celebes or <strong>the</strong> Timorgroup) ;and a fine species <strong>of</strong> Eos, peculiar to <strong>the</strong> small islands <strong>of</strong>Siau and Sanguir, just north <strong>of</strong> Celebes, obliges us to place<strong>the</strong>se <strong>with</strong> <strong>the</strong> Moluccas instead <strong>of</strong> <strong>with</strong> <strong>the</strong> former island, towhich <strong>the</strong>y seem most naturally to belong.<strong>The</strong> crimson parrots<strong>of</strong> <strong>the</strong> genus Eclectus are almost equally characteristic <strong>of</strong> <strong>the</strong>Moluccas, and add greatly to <strong>the</strong> brilliancy <strong>of</strong> <strong>the</strong> ornithology <strong>of</strong><strong>the</strong>se favoured islands.Reptiles.—<strong>The</strong> Reptiles, so far as known, appear to agree in<strong>the</strong>ir <strong>distribution</strong> <strong>with</strong> <strong>the</strong> o<strong>the</strong>r vertebrates. In some smallcollections from Ceram <strong>the</strong>re were no less than six <strong>of</strong> <strong>the</strong> generapeculiar to <strong>the</strong> Australian region, and which were before onlykn<strong>of</strong>n from Australia itself. <strong>The</strong>se are, <strong>of</strong> snakes, Liasis andEnygrus, genera <strong>of</strong> Pythonidae ;<strong>with</strong> Diemenia and Acanthophis(Elapidae) ; <strong>of</strong> lizards, Cyclodus, a genus <strong>of</strong> Scincidae ; and <strong>of</strong>Amphibia, a tree-frog <strong>of</strong> <strong>the</strong> genus Pelodryas.Insects— Peculiarities <strong>of</strong> <strong>the</strong> Moluccan Fauna.—In insects <strong>the</strong>Moluccas are hardly, if at all, inferior to New Guinea itself.<strong>The</strong>islands abound in grand Papilios <strong>of</strong> <strong>the</strong> largest size and extremebeauty ;and it is a very remarkable fact, that when <strong>the</strong> closelyalliedspecies <strong>of</strong> <strong>the</strong> Moluccas and New Guinea arecompared,


;chap. xiii. J• THEAUSTRALIAN REGION. 421<strong>the</strong> former are almost always <strong>the</strong> largest.As examples may bementioned, Ornithoptera priamus and 0. helena <strong>of</strong> <strong>the</strong> Moluccas,both larger than <strong>the</strong> varieties (or species) <strong>of</strong> Papua ; Papilioulysses and deiplwbus <strong>of</strong> Amboyna, usually larger than <strong>the</strong>irallies in New Guinea ; Hestia idea, <strong>the</strong> largest species <strong>of</strong> <strong>the</strong>genus ; Diadema pandarus and Charaxes euryalus, both largerthan any o<strong>the</strong>r species <strong>of</strong> <strong>the</strong> same genera in <strong>the</strong> whole archipelago.It is to be noted also, that in <strong>the</strong> Moluccas, <strong>the</strong> verylargest specimens or races seem always toisland <strong>of</strong> Amboyna ;come from <strong>the</strong> smalleven those <strong>of</strong> Ceram, <strong>the</strong> much larger islandto which it is a satellite, being almost always <strong>of</strong> less dimensions.Among Coleoptera, <strong>the</strong> Moluccas produce Euchirus longimanus,one <strong>of</strong> <strong>the</strong> largest and most remarkable <strong>of</strong> <strong>the</strong> LamellicornesSphingnotus dunningi, <strong>the</strong> largest <strong>of</strong> <strong>the</strong> Austro-Malayan Tmesisterninas;a Sphemira, <strong>the</strong> largest and handsomest <strong>of</strong> an extensivegenus ; an unusually large Schizorhina (Cetoniidse) ; andsome <strong>of</strong> <strong>the</strong> most remarkable and longest-horned Anthotribidoe.Even in birds <strong>the</strong> same law may be seen at work,—in <strong>the</strong> Tanysipteranais <strong>of</strong> Ceram, which has a larger tailthan any o<strong>the</strong>r in<strong>the</strong> genus ;in Centropus goliath <strong>of</strong> Gilolo, being <strong>the</strong> largest andlongest-tailed species ; in Hydrornis maximus <strong>of</strong> Gilolo, <strong>the</strong>largest and perhaps <strong>the</strong> most elegantly and conspicuouslycoloured <strong>of</strong> all <strong>the</strong> Pittidse ; in Platycercus amboinensis, beingpre-eminent in its ample blue tail ; in <strong>the</strong> two Moluccan loriesand Eos rubra, being more conspicuously red than <strong>the</strong> allied NewGuinea species ;and in Mcgapodius wallacei <strong>of</strong> Bouru, being <strong>the</strong>only species <strong>of</strong> <strong>the</strong> genus conspicuously marked and banded.All <strong>the</strong>se examples, <strong>of</strong> larger size, <strong>of</strong> longer tails or o<strong>the</strong>rappendages, and <strong>of</strong> more conspicuous colouring, are probablyindications <strong>of</strong> a less severe struggle for existence in <strong>the</strong>se islandsthan in <strong>the</strong> larger tract <strong>of</strong> New Guinea, <strong>with</strong> a more abundantand more varied fauna ; and this may apply even to <strong>the</strong> smallerislands, as compared <strong>with</strong> <strong>the</strong> larger in <strong>the</strong> immediate vicinity.<strong>The</strong> limited number <strong>of</strong> forms in <strong>the</strong> small islands compared<strong>with</strong> a similar area in <strong>the</strong> parent land, implies, perhaps, lesscompetition and less danger ; and thus allows, where all o<strong>the</strong>rconditions are favourable, an unchecked and continuous de-


422 ZOOLOGICAL GEOGEAPHY. [part hi.velopment in size, form, and colour, until <strong>the</strong>y become positivelyinjurious. This law may not improbably apply to <strong>the</strong> NewGuinea fauna itself, as compared <strong>with</strong> that <strong>of</strong> Borneo or anyo<strong>the</strong>r similar country ; and some <strong>of</strong> its peculiarities (such as itswonderful paradise-birds) may be due to long isolation, and consequentfreedom from <strong>the</strong> influence <strong>of</strong> any competing forms.<strong>The</strong> difference between <strong>the</strong> very sober colours <strong>of</strong> <strong>the</strong> Coleoptera,and in a less degree <strong>of</strong> <strong>the</strong> birds, <strong>of</strong> Borneo, as compared <strong>with</strong><strong>the</strong>ir brilliancy in New Guinea, always struck me most forcibly,and was long <strong>with</strong>out any, even conjectural, explanation.It isnot <strong>the</strong> place here to go fur<strong>the</strong>r into this most curious andinteresting subject. <strong>The</strong> reader who wishes for additional factsto aid him in forming an opinion, should consult Mr. Darwin'sDescent <strong>of</strong> Man, chapters x. to xv. ; and my own Contributionsto <strong>the</strong> <strong>The</strong>ory <strong>of</strong> Natural Selection, chapters iii.and iv.Timor Group.— Mammalia.—In <strong>the</strong> group <strong>of</strong> islandsJava and Australia, from Lombok tobetweenTimor inclusive, we find aset <strong>of</strong> mammals similar to those <strong>of</strong> <strong>the</strong> Moluccas, but some <strong>of</strong><strong>the</strong>m different species. A wide-spread species <strong>of</strong> Cuscus represents<strong>the</strong> Papuan element. A Sorex and a peculiar species <strong>of</strong>wild pig, we may also accept as indigenous. Three o<strong>the</strong>rs havealmost certainly been introduced. <strong>The</strong>se are, (1.) Macacus cynomolgus,<strong>the</strong> very commonest Malay monkey, which may havecrossed <strong>the</strong> narrow straits from island to island between Javaand Timor, though it seems much more probable that itwas introducedby Malays, who constantly capture and rear <strong>the</strong> young<strong>of</strong> this species. (2.) Cervus timoriensis, a deer, said to be a distinctspecies, inhabits Timor, but it is probably only a variety <strong>of</strong><strong>the</strong> Cervus hippelaphus <strong>of</strong> Java. This animal is, however, muchmore likely to have crossed <strong>the</strong> sea than <strong>the</strong> monkey. (3.) Paradoxurusfasciatus, takes <strong>the</strong> place <strong>of</strong> Viverra tangalunga in <strong>the</strong>Moluccas, both common and wide-spread civets which are <strong>of</strong>tenkept in confinement by <strong>the</strong> Malays. <strong>The</strong> Feli-s megalotis, longsupposed to be a native <strong>of</strong> Timor, has been ascertained by Mr.Elliot to belong to a different country altoge<strong>the</strong>r.Birds.—<strong>The</strong> birds are much more interesting, since <strong>the</strong>y are


:chap, xiii.] THE AUSTRALIAN REGION. 423sufficiently numerous to allow us to determine <strong>the</strong>ir <strong>relations</strong>,and trace <strong>the</strong>ir origin, <strong>with</strong> unusual precision. <strong>The</strong>re are 96genera and 160 species ot land-birds known to inhabit this group<strong>of</strong> islands ;and on a careful analysis, <strong>the</strong>y are found to be almostequally related to <strong>the</strong> Australian and Oriental regions, 30 generabeing distinctly traceable to <strong>the</strong> former, and <strong>the</strong> same number to<strong>the</strong> latter. <strong>The</strong>ir connection <strong>with</strong> <strong>the</strong> Moluccas is shown by <strong>the</strong>presence <strong>of</strong> <strong>the</strong> genera Mimeta, Gc<strong>of</strong>froyus, Cacatua, Ptilopus, andlanthcenas, toge<strong>the</strong>r <strong>with</strong> Megapodius and Cerchneis representedby Moluccan species. Turaccena shows a connection <strong>with</strong>Celebes, and Scops is represented by a Celebesian species. <strong>The</strong>connection <strong>with</strong> Australia is shown by <strong>the</strong> genera Sphceco<strong>the</strong>ra,Gerygone, Myiagra, Pardalotus, Gliciphila, Amadina, and Aprosmictus; while Milvus, Hypotriorchis, Eudynamis, and Eurystomus,are represented by Australian species. O<strong>the</strong>r genera confinedto or characteristic <strong>of</strong> <strong>the</strong> Australian region, are Rhipidura,Monarcha, Artamus, Campcpkaga, Pachycephalia, Philemon, Ptilotis,and Myzomela.We now come to <strong>the</strong> Indo-Malay or Javan element representedby <strong>the</strong> following genera1. Turdus (T.) 11. Oriolus. 21. Yungipicus.2. Geocichla (T.) 12. Pericrocotus. 22. Merops.3. Zoo<strong>the</strong>ra. 13. Cyornis (T.) 23. Pelargopsis.4. Megalurus (T.) 14. Hypothyinis. 24. Ceyx.5. Orthotomus. 15. Tchitrea. 25. Loriculus.6. Pratincola (T.) 16. Lanius (T.) 26. Treron (T.)7. Oreicola (T.) 17. Anthreptes. 27. lotreron (s.g. <strong>of</strong> Ptilopus).8. Drymocataphus (T.) 18. Eulabes. 28. Chalcophaps (T.)9. Parus. 19. Estrilda (T.) 29. Gallus (T.)10. Pycnonotus. 20. Erythrura (T.) 30. Strix.Such genera as Merops and Strix, which are as much Australianas Oriental, are inserted here because <strong>the</strong>y are represented byJavan species.<strong>The</strong> list is considerably swelled by genera whichhave reached Lombok across <strong>the</strong> narrow strait from Baly, buthave passed no fur<strong>the</strong>r. Such are Zoo<strong>the</strong>ra, Orthotomus, Pycnonotus,Pericrocotus and Strix. A much larger number (12) stopshort at Flores, leaving only 13, indicated in <strong>the</strong> list by (T) after<strong>the</strong>ir names, which reach Timor. It is evident, <strong>the</strong>refore, that<strong>the</strong>se islandshave been stocked from three chief sources,—<strong>the</strong>


:424 ZOOLOGICAL GEOGRAPHY. [part hi.Moluccas (<strong>with</strong> New Guinea and Celebes,) Australia, and Java.<strong>The</strong> Moluccan forms may well have arrived as stragglers fromisland to island, aided by whatever facilitiesby lands now submerged.have been affordedMost <strong>of</strong> <strong>the</strong> remainder have been derivedei<strong>the</strong>r from Australia or from Java ; and as <strong>the</strong>ir <strong>relations</strong>to <strong>the</strong>se islands are very interesting, <strong>the</strong>y must be discussed<strong>with</strong> some detail.Origin <strong>of</strong> <strong>the</strong> Timorese Fauna. — We must first note, that 80species, or exactly one-half <strong>of</strong> <strong>the</strong> land-birds<strong>of</strong> <strong>the</strong> islands, arepeculiar and mostly very distinct, intimating that <strong>the</strong> immigrationcommenced long enough back to allow <strong>of</strong> much specificmodification. <strong>The</strong>re is also one peculiar genus <strong>of</strong> kingfishers,Cariclonax, found only in Lombok and Flores, and more alliedto Australian than to Oriental types. <strong>The</strong> fine white-bandedpigeons (s. g. Leucotreron) are also almost peculiar ;one o<strong>the</strong>rless typical species only being known, a native <strong>of</strong> N. Celebes.In order to compare <strong>the</strong>species <strong>with</strong> regard to <strong>the</strong>ir origin, wemust first take away those <strong>of</strong> wide <strong>distribution</strong>special indications can be obtained.from which noIn this case 49 <strong>of</strong> <strong>the</strong> landbirdsmust be deducted, leaving 111 species which affordgoodmaterials for comparison. <strong>The</strong>se, when traced to <strong>the</strong>ir origin,show that 62 came from some part <strong>of</strong> <strong>the</strong> Australian region, 49from Java or <strong>the</strong> Oriental region. But if we divide <strong>the</strong>m intotwo groups, <strong>the</strong> one containing <strong>the</strong> species identical <strong>with</strong> those<strong>of</strong> <strong>the</strong> Australian or Oriental regions, <strong>the</strong> o<strong>the</strong>r containing alliedor representative species peculiar to <strong>the</strong> islands, we have <strong>the</strong> followingresultSpecies common to <strong>the</strong> Timorese Islands and <strong>the</strong> Oriental Region 30Peculiar Timorese species allied to those <strong>of</strong> <strong>the</strong>. Oriental Region 19Total 49Species common to <strong>the</strong> Timorese Islands and <strong>the</strong> AustralianEegion 18Peculiar Timorese species allied to those <strong>of</strong> <strong>the</strong> Australian Region 44Total , 62This table is very important, as indicating that <strong>the</strong> connection


chap, xiii.] THE AUSTRALIAN REGION. 425<strong>with</strong> Australia was probably earlier than that <strong>with</strong> Java ; since<strong>the</strong> majority <strong>of</strong> <strong>the</strong> Australian species have become modified,while <strong>the</strong> majority <strong>of</strong> <strong>the</strong> Oriental species have remained unchanged.This is due, no doubt, in part to <strong>the</strong> continued immigration<strong>of</strong> fresh individuals from Java, after that from Australia,<strong>the</strong> Moluccas and New Guinea had almost wholly ceased.must also notice <strong>the</strong> very small proportion <strong>of</strong> <strong>the</strong> genera,Weei<strong>the</strong>r<strong>of</strong> Australia or Java, that have found <strong>the</strong>ir way into <strong>the</strong>se islands,many <strong>of</strong> <strong>the</strong> largest and most wide-spread groups in both countriesbeing altoge<strong>the</strong>r absent.Taking <strong>the</strong>se facts into consideration,it is pretty clear that <strong>the</strong>re has been no close and longcontinuedapproximation <strong>of</strong> <strong>the</strong>se islands to any part <strong>of</strong> <strong>the</strong>Australian region ; and it is also probable that <strong>the</strong>y were fairlystocked <strong>with</strong> such Australian groups as <strong>the</strong>y possess before <strong>the</strong>immigration from Java commenced, or a largernumber <strong>of</strong> characteristicOriental forms would have been able to have established<strong>the</strong>mselves.On looking at our map, we find that a shallow submerged bankextends from Australia to <strong>with</strong>in about twenty miles <strong>of</strong> <strong>the</strong> coast<strong>of</strong> Timor; and this is probably an indication that <strong>the</strong> twocountries were once only so far apart.This would have allowed<strong>the</strong> purely Australian types to enter, as <strong>the</strong>y are not numerous;<strong>the</strong>re being about 6 Australian species,and 10 or 12 representatives<strong>of</strong> Australian species, in Timor.All <strong>the</strong> rest may have beenderived from <strong>the</strong> Moluccas or New Guinea, being mostly widespreadgenera <strong>of</strong> <strong>the</strong> Australian region; and <strong>the</strong> extension <strong>of</strong>Papua in a south-west direction towards Java (which was suggestedas a means <strong>of</strong> providing New Guinea <strong>with</strong> peculiar Indo-Malay types not found in any o<strong>the</strong>r part <strong>of</strong> <strong>the</strong> region) mayhave probably served to supply Timor and Flores <strong>with</strong> <strong>the</strong> mass<strong>of</strong> <strong>the</strong>ir<strong>the</strong> sea.Austro-Malayan genera across a narrow strait or arm <strong>of</strong>Lombok, Baly, and Sumbawa were probably not <strong>the</strong>nin existence, or nothing more than small \olcanic cones risingout <strong>of</strong> <strong>the</strong> sea, thus leaving a distance <strong>of</strong> 300 miles betweenFlores and Java. Subsequently <strong>the</strong>y grew into islands, which<strong>of</strong>fered an easy passage for a number <strong>of</strong> Indo-Malay generainto such scantily stocked territories as Flores and Timor. <strong>The</strong>Vol. I.—29


—426 ZOOLOGICAL GEOGRAPHY. [part hi.north coast <strong>of</strong> Australia <strong>the</strong>n sank, cutting <strong>of</strong>f <strong>the</strong> supply fromthat country ;and this left <strong>the</strong> Timorese group in <strong>the</strong> position itnow occupies.<strong>The</strong> reptiles and fishes <strong>of</strong> this group are too little known toenable us to make any useful comparison.Insects.—<strong>The</strong> insects, though not numerous, present many finespecies, some quite unlike any o<strong>the</strong>rs in <strong>the</strong> Archipelago.arePapilio liris, Pieris lecta,Cirrochroa lamarckii and C. leschenaultiiamong butterflies.Such<strong>The</strong> Coleoptera are comparatively littleknown, but in <strong>the</strong> insects generally <strong>the</strong> Indo-Malay element predominates.This may have arisen from <strong>the</strong> peculiar vegetationand arid climate not being suitable to <strong>the</strong> Papuan insects.WhyAustralian forms did not establish <strong>the</strong>mselves we cannot conjecture;but <strong>the</strong> field appears to have been open to immigrants fromJava, <strong>the</strong> climate and vegetation <strong>of</strong> which island at itseastern extremityapproximates to that <strong>of</strong> <strong>the</strong> Timorese group.<strong>The</strong> insectsare, however, so peculiarly modified as to imply a very great antiquity,and this is also indicated by a group <strong>of</strong> Sylviine birds hereclassedunder Oreicola, but some <strong>of</strong> which probably form disfinctgenera. <strong>The</strong>re may, perhaps, have been an earlier and a laterapproximation to Java, which, <strong>with</strong> <strong>the</strong> o<strong>the</strong>r changes indicated,would account for most <strong>of</strong> <strong>the</strong> factspresented by <strong>the</strong> fauna <strong>of</strong><strong>the</strong>se islands. One deduction is, at all events, clear :<strong>the</strong> extremepaucity <strong>of</strong> indigenous mammals along <strong>with</strong> <strong>the</strong> absence <strong>of</strong>so many groups <strong>of</strong> birds, renders it certain that <strong>the</strong> Timoreseislands did not derive <strong>the</strong>ir animal life by means <strong>of</strong> an actualunion <strong>with</strong> any <strong>of</strong> <strong>the</strong> large islands ei<strong>the</strong>r <strong>of</strong> <strong>the</strong> Australian or<strong>the</strong> Oriental regions.Celebes Group.—We now come to <strong>the</strong> Island <strong>of</strong> Celebes, inmany respects <strong>the</strong> most remarkable and interesting in <strong>the</strong> wholeregion, or perhaps on <strong>the</strong> globe, since no o<strong>the</strong>r island seems topresent so many curious problems for solution. We shall <strong>the</strong>reforegive a somewhat full account <strong>of</strong> its peculiar fauna, andendeavour to elucidate some <strong>of</strong> <strong>the</strong> causes to which its zoologicalisolation may be attributed.Mammalia.—<strong>The</strong> following is <strong>the</strong> list <strong>of</strong> <strong>the</strong> mammalia <strong>of</strong>


chap, xiii.] THE AUSTRALIAN REGION. 427Celebes as far as at present known, though many small speciesmay yet be discovered.1. Cynopi<strong>the</strong>cus nigrescens. 7. Barbirusa alfurns.2. Tarsius spectrum. 8. Sciurus (5 peculiar sp.)3. Viverra tangaluuga. 9. Mus (2 peculiar sp.)4. Cervus hippelaplms. 10. discus (2 peculiar sp.)5. Auoa depressicornis. Also 7 species <strong>of</strong> bats, <strong>of</strong>6. Sus celebensis. which 5 are peculiar.<strong>The</strong> first— a large black ape—is itself an anomaly, since it isnot closely allied to any o<strong>the</strong>r form <strong>of</strong> quadrumana. Its flatprojecting muzzle, large superciliary crests and maxillary ridges,<strong>with</strong> <strong>the</strong> form and appearance <strong>of</strong> its teeth, separate it altoge<strong>the</strong>rfrom <strong>the</strong> genus Macacus, as represented in <strong>the</strong> Indo-Malay islands,and ally it closely to <strong>the</strong> baboons <strong>of</strong> Africa. We 1have alreadyseen reason to suppose that it has been carried to Batchian,and <strong>the</strong>re is some doubt about <strong>the</strong> allied species or variety (C.niger) <strong>of</strong> <strong>the</strong> Philippines being really indigenous <strong>the</strong>re; in whichcase this interesting form will remain absolutely confined to Celebes.(2.) <strong>The</strong> tarsier is a truly Malayan species, but it is said tooccur in a small island at <strong>the</strong> nor<strong>the</strong>rn extremity <strong>of</strong> Celebes andon some <strong>of</strong> <strong>the</strong> Philippine islands.introduced <strong>the</strong>re.It might possibly have been(3) and (4)—a civet and a deer—are, almost certainly,as in <strong>the</strong> Moluccas, introduced species. (5.) Anoa depressicornis.This is one <strong>of</strong> <strong>the</strong> peculiar Celebesian types; a small straighthornedwild-bull, anatomically allied to <strong>the</strong> buffaloes, and somewhatresembling <strong>the</strong> bovine antelopes <strong>of</strong> Africa, but having nonear allies in <strong>the</strong> Oriental region. (6.) Sus Celebensis ;a peculiarspecies <strong>of</strong> wild-pig. (7.) Babirusa alfurus ; ano<strong>the</strong>r remarkabletype, having no near allies. It differs in its dentition from <strong>the</strong>typical Suidae, and seems to approach <strong>the</strong> African Phacochceridse.<strong>The</strong> manner in which <strong>the</strong> canines <strong>of</strong> <strong>the</strong> upper jaw are reversed,and grow directly upwards in a spiral curve over <strong>the</strong> eyes, isunique among mammalia. (8.)Five squirrels inhabit Celebes, andall are peculiar species. (9.) <strong>The</strong>se>are forest rats <strong>of</strong> <strong>the</strong> sub-genusGymnomys, allied to Australian species. 10. Cuscus. This typical1<strong>The</strong> general form <strong>of</strong> <strong>the</strong> skull agrees best <strong>with</strong> that <strong>of</strong> Cynocephalusmormon, <strong>the</strong> largest and most typical <strong>of</strong> <strong>the</strong> African baboons ;while <strong>the</strong>position <strong>of</strong> <strong>the</strong> nostrils brings it nearer <strong>the</strong> macaques.


—;428 ZOOLOGICAL GEOGRAPHY. [past hi.Australian form is represented in Celebes by two peculiarspecies.Leaving out <strong>the</strong> Indo-Malay species, which may probably havebeen introduced by man, and are at all events comparativelyrecent immigrants, and <strong>the</strong> wild pig, a genus which ranges over<strong>the</strong> whole archipelago and which has <strong>the</strong>refore little significance,we find two genera which have come from <strong>the</strong> Australian side,— discus and Mus ; and four from <strong>the</strong> Oriental side, Cynopi<strong>the</strong>cus,Anoa, Bahirusa, and Sciurus. But Sciurus alone correspondsto discus, as a genus still inhabiting <strong>the</strong> adjacent islands<strong>the</strong> o<strong>the</strong>r three being not only peculiar to Celebes, but incapable<strong>of</strong> being affiliated to any specially Oriental group. We seem, <strong>the</strong>n,to have indications <strong>of</strong> two distinct periods ; one very ancient,when <strong>the</strong> ancestors <strong>of</strong> <strong>the</strong> three peculiar genera roamed over someunknown continent <strong>of</strong> which Celebes formed, perhaps, an outlyingportion ;—ano<strong>the</strong>r more recent, when from one side <strong>the</strong>re enteredSciurus, and from <strong>the</strong> o<strong>the</strong>r discus. But we must rememberthat <strong>the</strong> Moluccas to <strong>the</strong> east, possess scarcely any indigenousmammals except Cuscus ; whereas Borneo and Java on <strong>the</strong> west,have nearly 50 distinct genera. It is evident <strong>the</strong>n, that <strong>the</strong>facilities for immigration must have been much less <strong>with</strong> <strong>the</strong>Oriental than <strong>with</strong> <strong>the</strong> Australian region,and we may be prettycertain that at this later period <strong>the</strong>re was no land connection<strong>with</strong> <strong>the</strong> Indo-Malay islands, or some o<strong>the</strong>r <strong>animals</strong> than squirrelswould certainly have entered. Let us now see what light isthrown upon <strong>the</strong> subject by <strong>the</strong> birds.Birds.—<strong>The</strong> total number <strong>of</strong> birds known to inhabitCelebesis 205, belonging to about 150 genera. We may leave out <strong>of</strong>consideration <strong>the</strong> wading and aquatic birds, most <strong>of</strong> which arewide-ranging species.<strong>of</strong> land-birds,<strong>The</strong>re remain 123 genera and 152 species<strong>of</strong> which 9 genera and 66 species are absolutelyconfined to <strong>the</strong> island, while 20 more are found also in <strong>the</strong> Sulaor Sanguir Islands, so that we may take 86 to be <strong>the</strong> number <strong>of</strong>peculiar Celebes species. Lord Walden, from whose excellentpaper on <strong>the</strong> birds <strong>of</strong> Celebes (Trans. Zool. Soc. vol. viii. p. 23)most <strong>of</strong> <strong>the</strong>se figures are obtained, estimates, that <strong>of</strong> <strong>the</strong> specieswhich are not peculiar to Celebes, 55 are <strong>of</strong> Oriental and 22 <strong>of</strong>


chap, xiii.] THE AUSTRALIAN REGION. 429Australian origin, <strong>the</strong> remainder being common toboth regions.This shows a preponderant recent immigration from <strong>the</strong> Westand North, which is not to be wondered at when we look at <strong>the</strong>long coast line <strong>of</strong> Java, Borneo, and <strong>the</strong> Philippine islands, <strong>with</strong>an abundant and varied bird population, on <strong>the</strong> one side, and<strong>the</strong> small scattered islands <strong>of</strong> <strong>the</strong> Moluccas, <strong>with</strong> a comparativelyscanty bird-fauna, on <strong>the</strong> o<strong>the</strong>r.But, adopting <strong>the</strong> method here usually followed, let us lookat <strong>the</strong> <strong>relations</strong> <strong>of</strong> <strong>the</strong> genera found in Celebes, omitting for<strong>the</strong> present those which are peculiar to it. I divide <strong>the</strong>se generainto two series :—those which are found in Borneo or Java butnot in <strong>the</strong> Moluccas, and those which inhabit <strong>the</strong> Moluccas andnot Borneo or Java; <strong>the</strong>se being <strong>the</strong> respective sources fromwhich, primd facie, <strong>the</strong> species <strong>of</strong> <strong>the</strong>se genera must have beenderived. Genera which range widely into both <strong>the</strong>se districtsare rejected, as teaching us nothing <strong>of</strong> <strong>the</strong> origin <strong>of</strong> <strong>the</strong> Celebesianfauna. In a few cases, sub-genera which show a decidedeastern or western origin, are given.Genera derived from Borneo and Java.1. Geocichla. 9. Nectarophila. 17. Hydrocissa.2. Pratincola (sp.) 10. Anthreptes (sp.) 18. -Cranorrhinus.3. Trichastoma. 11. Munia (sp.) 19. Lyncornis.4. Oriolus (sp.) 12. Acrido<strong>the</strong>res. 20. Treron (sp.)5. Cyornis 13. Yuugipicus. 21. Gallus (sp.)6. Hypothymis. 14. Mulleripicus. 22. Spilornis.7. Hylocharis. 15. Rhamphococcyx. 23. Butastur.8. iEthopyga. 16. Hierococcyx. 24. Pernis.Genera derived from <strong>the</strong> Moluccas or Timor.1. Graucalus (sp.) 6. Tanygnathus. 11. Myristicivora (s. g.)2. Chalcostetha. 7. Trichoglossus. 12. Ducula (s. g.)3. Myzoraela. 8. Scythrops (sp.) 13. Zoncenas (s. g.)4. Munia (sp.) 9. Turaccena, 14. Lamproteron (s. g.)5. Cacatua (sp.) 10. Reinwardtcenas (sp.) 15. Megapodius.<strong>The</strong>se tables show a decided preponderance <strong>of</strong> Orientalover Australian forms. But we must remember that <strong>the</strong> immediatelyadjacent lands from whence <strong>the</strong> supply was derived, is


430 ZOOLOGICAL GEOGRAPHY. [part III.very much richer in <strong>the</strong> one case than in <strong>the</strong> o<strong>the</strong>r. <strong>The</strong> 24o-enera derived from Borneo and Java are only about one fourth<strong>of</strong> <strong>the</strong> characteristic genera <strong>of</strong> those islands; while <strong>the</strong> 15Moluccan and Timorese geuera are fully one third <strong>of</strong> <strong>the</strong>ircharacteristic types. <strong>The</strong> proportion derived from <strong>the</strong> Australian,is greater than that derived from <strong>the</strong> Oriental side.We shall exhibit this perhaps more clearly, by giving a list <strong>of</strong><strong>the</strong> important groups <strong>of</strong> each set <strong>of</strong> islands which are absentfrom Celebes.Important Families <strong>of</strong> Java and Borneoabsent from Celebes.Important Families <strong>of</strong> <strong>the</strong> Moluccasabsent from Celebes.1. Eurylsemidse.2. Timaliidse.3. Phyllornithidae.4. Pycnonotida?5. Laniidjfi.6. Megalseraidae.7. Trogonidse.8. Phasianida?.1. Meliphagidae.Additional important genera <strong>of</strong> Java orBorneo absent from Celebes.1. Orthotomus.2. Copsychus.3. Enicurus.4. Tchitrea.5. Pericrocotus.6. Irena.7. Platylophus.8. Dendrocitta.9. Eulabes.10. Hemicercus.11. Chrysocolaptes.12. Tiga.13. Microptemus.14. Batrachostomus.15. Palseornis.16. Rollulus.Important genera <strong>of</strong> <strong>the</strong> Moluccasabsent from Celebes.1. Mimeta.2. Monarcha.3. Rhipidura.4. Pachycephala.5. Lycocorax.6. Alcyone.7. Tanysiptera.8. Ge<strong>of</strong>froyus.9. Eclectus.10. Platycercus.11. Eos.12. Lorius.If we reckon <strong>the</strong> absent families to be each represented byonly two important genera, we shall find <strong>the</strong> deficiency on <strong>the</strong>Oriental side much <strong>the</strong> greatest;yet those on <strong>the</strong> side <strong>of</strong> <strong>the</strong>Moluccas are sufficiently remarkable.<strong>The</strong> Meliphagidse are notindeed absolutely wanting, since a Myzomela has now beenfound in Celebes ;but all its larger and more powerful formswhich range over almost <strong>the</strong> entire region, are absent. Thismay be balanced by <strong>the</strong> absence <strong>of</strong> <strong>the</strong> excessively abundantTimaliidffi <strong>of</strong> <strong>the</strong> Indo-Malay islands, which arerepresented by


;chap, xiil.] THE AUSTRALIAN REGION. 431only a single species ; and by <strong>the</strong> powerfdl Phasianidae, representedonly by <strong>the</strong> common Malay jungle fowl,perhaps introduced.<strong>The</strong> entire absence <strong>of</strong> Pycnonotidse is a very anomalousfact, since one <strong>of</strong> <strong>the</strong> largest genera, Criniger, is well representedin several islands <strong>of</strong> <strong>the</strong> Moluccas, and one has even been foundin <strong>the</strong> Togian islands in <strong>the</strong> great nor<strong>the</strong>rn inlet <strong>of</strong> Celebesbut yet it passes over Celebes itself. Ceyx, a genus <strong>of</strong> smallkingfishers, is a parallel case, since it isfound everywhere fromIndia to New Guinea, leaving out only Celebes ; but this comesamong those curiosities <strong>of</strong> <strong>the</strong> Celebesian fauna which we shallnotice fur<strong>the</strong>r on.In <strong>the</strong> list <strong>of</strong> genera derived from Borneo orJava, no less than 6 are represented by identical species (indicatedby sp. after <strong>the</strong> name) ; while in <strong>the</strong> Moluccan list 5are thus identical. <strong>The</strong>se must be taken to indicate, ei<strong>the</strong>r that<strong>the</strong> genus is a recent introduction, or that stragglers still occasionallyenter, crossing <strong>the</strong> breed, and thus preventing specificmodification. In ei<strong>the</strong>r case <strong>the</strong>y depend on <strong>the</strong> existing state<strong>of</strong> things, and throw no light on <strong>the</strong> different <strong>distribution</strong> <strong>of</strong>land and sea which aided or checked migration in former timesand <strong>the</strong>y <strong>the</strong>refore to some extent diminish <strong>the</strong> weight <strong>of</strong> <strong>the</strong>Indo-Malay affinity, as measured by <strong>the</strong> <strong>relations</strong> <strong>of</strong> <strong>the</strong> peculiarspecies <strong>of</strong> Celebes.From our examination <strong>of</strong> <strong>the</strong> evidence thus far,—that is, takingaccount firstly,<strong>of</strong> <strong>the</strong> species, and, secondly, <strong>of</strong> <strong>the</strong> genera, whichare common to Celebes and <strong>the</strong> groups <strong>of</strong> islands between whichit is situated, we must admit that <strong>the</strong> connexion seems ra<strong>the</strong>r<strong>with</strong> <strong>the</strong> Oriental than <strong>with</strong> <strong>the</strong> Australian region; but when wetake into account <strong>the</strong> proportion <strong>of</strong> <strong>the</strong> genera and speciespresent,to those which are absent, and giving some weight togreater extent <strong>of</strong> coast line on <strong>the</strong> Indo-Malay side, we seemjustified in stating that <strong>the</strong> Austro-Malay element is<strong>the</strong>ra<strong>the</strong>r <strong>the</strong>most fully represented. This result applies both to birds andmammals ; and it leads us to <strong>the</strong> belief, that during <strong>the</strong> epoch <strong>of</strong>existing species and genera, Celebes has never been united <strong>with</strong>any extensive tract <strong>of</strong> land ei<strong>the</strong>r on <strong>the</strong> Indo-Malay or Austro-Malay side, but has received immigrants from both during a verylong period, <strong>the</strong> facilities for immigration having been ra<strong>the</strong>r <strong>the</strong>


:.432 ZOOLOGICAL GEOGRAPHY. [part ill.greatest on <strong>the</strong> Austro-Malay or Australian side. We have nowto consider what fur<strong>the</strong>r light can be thrown on <strong>the</strong> subject by<strong>the</strong> consideration <strong>of</strong> <strong>the</strong> peculiar genera <strong>of</strong> Celebes, and <strong>of</strong> thosecuriosities or anomalies <strong>of</strong> <strong>distribution</strong> to which we havereferred.Nine genera <strong>of</strong> birds are altoge<strong>the</strong>r peculiar to Celebes ;threemore are found only in one o<strong>the</strong>r island, and seem to be typicallyCelebesian ; while one is found in <strong>the</strong> Sula islands (whichbelongs to <strong>the</strong> Celebes group) and probably exists in Celebesalso. <strong>The</strong> following is a list <strong>of</strong> <strong>the</strong>se 13 genera1. Artamides... (Campephagidse) 8. Monachalcyon (Alcedinidae)2. titreptocitta .. (Corvidse) 9. Cittura_... „3. Charitomis.. „ 10. Ceycopsis ... „4. Gazzola, (s. g.) „ 11. Meropogon .. (Meropidse)5. Basilomis .. (Sturnidse) 12. Prioniturus (Psittacidas)6. Enochs ... „ 13. Megacephalon (Megapodiidae)7. Scissirostrum „Of <strong>the</strong> above, Artamides, Monachalcyon,Cittura, and Megacephalon,are modifications <strong>of</strong> types characteristic <strong>of</strong> <strong>the</strong> Australianregion.All are peculiar to Celebes except Cittura, found also in<strong>the</strong> Sanguir islands to <strong>the</strong> northward, but which seems to belongto <strong>the</strong> Moluccan group. Strcptocitta, Charitomis, and Gazzola,are peculiar types <strong>of</strong> Corvidre; <strong>the</strong> two former allied to <strong>the</strong>magpies, <strong>the</strong> latter to <strong>the</strong> jackdaws.Charitomis is known onlyfrom <strong>the</strong> Sula islands east <strong>of</strong> Celebes, and is closely related toStrcptocitta. <strong>The</strong>re is nothing comparable to <strong>the</strong>se three groupsin any <strong>of</strong> <strong>the</strong> Malay islands, and <strong>the</strong>y seem to have <strong>relations</strong>ra<strong>the</strong>r <strong>with</strong> <strong>the</strong> Corvidse <strong>of</strong> <strong>the</strong> old-world nor<strong>the</strong>rn continent.Basilomis, Enodcs, and Scissirostrum, are remarkable forms <strong>of</strong>Sturnidse.Basilomis has a beautiful compressed crest, which in<strong>the</strong> allied species found in Ceram is elongated behind. Enodcshas remarkable red superciliary streaks, but seems allied toCalornis. Scissirostrum seems also allied to Calomis in generalstructure, but has a very peculiarly formed bill and nostrils.Wecan hardly say whe<strong>the</strong>r <strong>the</strong>se three forms show more affinity toOriental or to Australian types, but <strong>the</strong>y add to <strong>the</strong> weight <strong>of</strong>evidence as to <strong>the</strong> great antiquity and isolation <strong>of</strong> <strong>the</strong> Celebesianfauna. Scissirostrum has been classed <strong>with</strong> Euryceros, a Mada-


chap, xiii.] THE AUSTRALIAN REGION. 433gascar bird, and <strong>with</strong> Buphaga, an African genus ;but <strong>the</strong> peculiarbeak and nostrils approximate more to Crarticus and itsallies, <strong>of</strong> <strong>the</strong> Australian region, which should probably form adistinct family. Ceycopsis is undoubtedly intermediate between<strong>the</strong> Malayan Ccyx and <strong>the</strong> African Ispidina, and is <strong>the</strong>reforeespeciallyinteresting. Meropogon is a remarkable form <strong>of</strong> beeeater,allied to <strong>the</strong> Indo-Malayan Nydiornis.raquet-tailed parrots) <strong>of</strong> which two species inhabitone <strong>the</strong> Philippines, appears to be allied toGe<strong>of</strong>froyus.We must finally notice aPrioniturus (<strong>the</strong>Celebes, and<strong>the</strong> Austro-Malayanfew genera found in Celebes, whosenearest allies are not in <strong>the</strong> surrounding islands, and which thusafford illustrations <strong>of</strong> discontinuous <strong>distribution</strong>. <strong>The</strong> most remarkable,perhaps, is Coracias, <strong>of</strong> which a fine species inhabitsCelebes ;while <strong>the</strong> genus is quite unknown in <strong>the</strong> Indo-Malaysub-region, and does not appear again till we reach Burmah andIndia ; and <strong>the</strong> species has no closer affinity for Indian than forAfrican forms. Myialestes, a small yellow flycatcher, is ano<strong>the</strong>rexmple; its nearest ally (M. cinereocapilla) being a common Indianbird, but unknown in <strong>the</strong> Malay islands. <strong>The</strong> Celebesian birddescribed by me as Prionochilus aureolimbatus, is probably athird case <strong>of</strong> discontinuous <strong>distribution</strong>, if (as a more carefulexamination seems to show) it is not a Prionochilus, but congeneric<strong>with</strong> Pachyglossa, a bird only found in <strong>the</strong> Himalayas.<strong>The</strong> fine pigeon, Carpophaga forsteni, belongs to a group found in<strong>the</strong> Philippines, Australia, and New Zealand ; but <strong>the</strong> Celebesspecies is very distinct from all <strong>the</strong> o<strong>the</strong>rs, and seems, if anything,more allied to that <strong>of</strong> New Zealand.<strong>The</strong> Sula islands (Sula-mangola, Sula-taliabo, and Sula-besi)lie midway between Celebes and <strong>the</strong> Moluccas, being 80 milesfrom <strong>the</strong> nearest part <strong>of</strong> Celebes, <strong>with</strong> several interveningislands, and 40 miles from Bouru, all open sea. <strong>The</strong>ir birdsshow, as might be expected, a blending <strong>of</strong> <strong>the</strong> two faunas, but<strong>with</strong> a decided preponderance <strong>of</strong> that <strong>of</strong> Celebes. Out <strong>of</strong> 43land birds which have been collected in <strong>the</strong>se islands, we maydeduct 6 as <strong>of</strong> wide range and no significance. Of <strong>the</strong> 37 remaining,21 are Celebesian species, and 4 are new species but


—434 ZOOLOGICAL GEOGRAPHY. [part hi.allied to those <strong>of</strong> Celebes ; while <strong>the</strong>re are 10 Moluccan speciesand 2 new species allied to those <strong>of</strong> <strong>the</strong> Moluccas.that no less than 3 Moluccan genera, quite unknown initself, occur here,It is curiousCelebesMonarcha, Pachycephalia, and Criniger; but all<strong>the</strong>se, as well as several o<strong>the</strong>r <strong>of</strong> <strong>the</strong> Moluccan birds, are ra<strong>the</strong>rweak flyers, and such as are likely to have been carried acrossby strong winds. Of <strong>the</strong> genera, 23 are from Celebes, 10 from<strong>the</strong> Moluccas.<strong>The</strong>se facts show, that <strong>the</strong> Sula islands form part<strong>of</strong> <strong>the</strong> Celebes group, although <strong>the</strong>y have received an infusion <strong>of</strong>Moluccan forms, which will perhaps in time spread to <strong>the</strong> mainisland, and diminish <strong>the</strong> remarkable individuality that now characterisesits fauna.Insects.—Of <strong>the</strong> reptiles and fishes <strong>of</strong> Celebes we have notsufficient information to draw any satisfactory conclusions. I<strong>the</strong>refore pass to <strong>the</strong> insects <strong>of</strong> which something more isknown.<strong>The</strong> Butterflies <strong>of</strong> Celebes are not very numerous, less than200 species in all having been collected ; but a very large proportion<strong>of</strong> <strong>the</strong>m, probably three-fourths <strong>of</strong> <strong>the</strong> whole, are peculiar.<strong>The</strong>re is only one peculiar genus, Amcchania, allied to Ze<strong>the</strong>ra (agroup confined to <strong>the</strong> Philippine Islands), <strong>with</strong> which it shouldperhaps be united. Most <strong>of</strong> <strong>the</strong> genera are <strong>of</strong> wide <strong>distribution</strong>in <strong>the</strong> archipelago, or are especially Malayan, only two trulyAustralian genera, Elodina and Acropthalmia, reaching Celebes.On <strong>the</strong> o<strong>the</strong>r hand, 7 peculiar Oriental genera are found in Celebes,but not fur<strong>the</strong>r east, viz., Clerome, Adolias, Euripus, Apatura,Limenitis, lolaus, and Leptocircus. <strong>The</strong>re are also several indications<strong>of</strong> a direct affinity <strong>with</strong> <strong>the</strong> continent ra<strong>the</strong>r than <strong>with</strong>Malaya, as in <strong>the</strong> cases already enumerated among birds. Afine butterfly, yet unnamed, almostexactly resembles Dichorragiancsimachus, a Himalayan species. Euripus rdbustus is closelyallied to E. hali<strong>the</strong>rses <strong>of</strong> N. India ;<strong>the</strong>re are no less than o species<strong>of</strong> Limenitis, all quite unlike those found in o<strong>the</strong>r parts <strong>of</strong> <strong>the</strong>archipelago. <strong>The</strong> butterflies <strong>of</strong> Celebes are remarkably distinguishedfrom all o<strong>the</strong>rs in <strong>the</strong> East, by peculiarities <strong>of</strong> form, size,and colour, which run through groups <strong>of</strong> species belonging todifferent genera. Many Papilionidse and Pieridoe, and some


—chap, xiil] THE AUSTRALIAN REGION. 435Nymphalidse, have <strong>the</strong> anterior wings elongated, <strong>with</strong> <strong>the</strong> apex<strong>of</strong>ten acute, and, what is especially remarkable, an abrupt bendor shoulder near <strong>the</strong> base <strong>of</strong> <strong>the</strong> wing. (See Malay Archipelago,3rd Ed. p. 281, woodcut.) No less than 13 species <strong>of</strong> Papilio, 10Pieridse, and 4 or 5 Nymphalidse, are thus distinguished from<strong>the</strong>ir nearest allies in <strong>the</strong> surrounding islands or in India. Insize again, a large number <strong>of</strong> Celebesian butterflies stand preeminentover <strong>the</strong>ir allies. <strong>The</strong> fine Papilios adamantius, hlumei,and gigon— are perfect giants by <strong>the</strong> side <strong>of</strong> <strong>the</strong> closely-alliedforms <strong>of</strong> Java ;while P. androcles is <strong>the</strong> largest and longest-tailed,<strong>of</strong> all <strong>the</strong> true swallow-tailed group <strong>of</strong> <strong>the</strong> Old World.AmongNymphalidse, <strong>the</strong> species <strong>of</strong> Ehinopalpa and Euripus, peculiar toCelebes, are immensely larger than <strong>the</strong>ir nearest allies ; andseveral <strong>of</strong> <strong>the</strong> Pieridse are also decidedly larger, though in a lessmarked degree. In colour, many <strong>of</strong> <strong>the</strong> Celebesian butterflies differfrom <strong>the</strong> nearest allied species ; so that <strong>the</strong>y acquire a singularity<strong>of</strong> aspect which marks <strong>the</strong>m <strong>of</strong>f from <strong>the</strong> rest <strong>of</strong> <strong>the</strong> group.<strong>The</strong> most curious case is that <strong>of</strong> three butterflies, belongingto three distinct genera (Ccthosia myrina, Messaras mceonides,and Atella celebensis) all having a delicate violet or lilac gloss inlines or patches, which is wholly wanting in every allied species<strong>of</strong> <strong>the</strong> surrounding islands. <strong>The</strong>se numerous peculiarities <strong>of</strong>Celebesian butterflies are very extraordinary ; and imply isolationfrom surrounding lands, almost as much as do <strong>the</strong>strange forms<strong>of</strong> mammals and birds, which more prominently characterise thisinteresting island.Of <strong>the</strong> Coleoptera we know much less,but a few interestingfacts may be noted. <strong>The</strong>re are a number <strong>of</strong> fine species <strong>of</strong>Cicindela, some <strong>of</strong> peculiar forms ;and one Odontochila, a SouthAmerican genus; while Collyris reaches Celebes from <strong>the</strong>Oriental region. In Carabidse it has one peculiar genus, Dicraspeda; and a species <strong>of</strong> <strong>the</strong> fine Australian genus Catadromus.In Lucanidae it has <strong>the</strong> Oriental genus, Odontolabris. In Cetoniidaeit has a peculiar genus, Sternoplus, and several fine Cetonice;but <strong>the</strong> characteristic Malayan genus, Zomaptera, found in everyo<strong>the</strong>r island <strong>of</strong> <strong>the</strong> archipelago from Sumatra to New Guinea,is absent—an analogous fact to <strong>the</strong> case <strong>of</strong> Ceyx among birds.


436 ZOOLOGICAL GEOGRAPHY. [part hi.In Buprestidae, <strong>the</strong> principal Austro-Malay genus, Sambus, isfound here ; while Sponsor, a genus 8 species <strong>of</strong> which inhabitMauritius, has one species here and one in New Guinea. InLongicorns <strong>the</strong>re are four peculiar genera, Comusia, Pytholia,Bityle, and Ombrosaga ; but <strong>the</strong> most important features are <strong>the</strong>occurrence <strong>of</strong> <strong>the</strong> o<strong>the</strong>rwise purely Indo-Malayan genera Agelasta,Nyctimene, and Asta<strong>the</strong>s ; and <strong>of</strong> <strong>the</strong> purely Austro-Malayan Arrhenotus, Trysimia, Xenolea, Amblymora, Diallus,and JEgocidnus.<strong>The</strong> remaining genera range over both portions<strong>of</strong> <strong>the</strong> archipelago. In <strong>the</strong> extensive family <strong>of</strong> Curculionidsewe can only notice <strong>the</strong> elegant genus, Celebia, allied to Eupholus,which, owing to its abundance and beauty, isfeature in <strong>the</strong> entomology <strong>of</strong> <strong>the</strong> island.a conspicuousOrigin <strong>of</strong> <strong>the</strong> fauna <strong>of</strong> Celebes.—We have now to consider,briefly, what past changes <strong>of</strong> physical geography are indicatedby <strong>the</strong> curious assemblage <strong>of</strong> facts here adduced. We haveevidently, in Celebes, a remnant <strong>of</strong> an exceedingly ancient land,which has undergone many and varied revolutions; and <strong>the</strong>stock <strong>of</strong> ancient forms which it contains must be taken account<strong>of</strong>, when we speculate on <strong>the</strong> causes that have so curiouslylimited more recent immigrations. Going back to <strong>the</strong> arrival<strong>of</strong> those genera which are represented in Celebes by peculiarspecies, and taking first <strong>the</strong> Austro-Malay genera, we findamong <strong>the</strong>m such groups as Zoncenas (s.g.), Phlogcenas, Leucotreron(s.g.), and Turaccena, which are not found in <strong>the</strong> Moluccasat all ; and Myzomela, found in Timor and Banda, but not inCeram or Bourn, which are nearest to Celebes. This, combined<strong>with</strong> <strong>the</strong> curious absence <strong>of</strong> so many <strong>of</strong> <strong>the</strong> commonest Moluccangenera, leads to <strong>the</strong> conclusion that <strong>the</strong> Austro-Malay immigra-•tion took place by way <strong>of</strong> Timor and <strong>the</strong> sou<strong>the</strong>rn part<strong>of</strong> NewGuinea. It will be remembered, that to account for <strong>the</strong> Indo-Malayan forms in New Guinea, we suggested an extension <strong>of</strong>that country in a westerly direction just north <strong>of</strong> Timor. Nowthis is exactly what we require, to account for <strong>the</strong> stocking<strong>of</strong> Celebes <strong>with</strong> <strong>the</strong> Australian forms it possesses. At this timeBorneo, did not approach so near, and itwas at a somewhat laterperiod that <strong>the</strong> last great Indo-Malay migration set in; but


chap, xiii.] THE AUSTRALIAN REGION. 437finding <strong>the</strong> country already fairly stocked, comparatively fewgroups were able to establish <strong>the</strong>mselves.Going back a little far<strong>the</strong>r, we come to <strong>the</strong> entrance <strong>of</strong> thosefew birds and insects which belong to India or Indo-China ;andthis probably occurred at <strong>the</strong> same time as that continentalextension southward, which we found was required to account fora similar phenomenon in Java. Celebes, being more remote,received only a few stragglers. We have now to go muchfar<strong>the</strong>r back, to <strong>the</strong> time when <strong>the</strong> ancestors <strong>of</strong> <strong>the</strong> peculiarCelebesian genera entered <strong>the</strong> country, and here our conjecturesmust necessarily be less defined.On <strong>the</strong> Australian side we have to account for Megacephalon,and <strong>the</strong> o<strong>the</strong>r genera <strong>of</strong> purely Papuan type. It may perhapsbe sufficient to say, that we do not yet know that <strong>the</strong>se genera,or some very close allies, do not still exist in New Guinea ; inwhich case <strong>the</strong>y may well have entered at <strong>the</strong> same time <strong>with</strong><strong>the</strong> species, already referred to. If, on <strong>the</strong> o<strong>the</strong>r hand, <strong>the</strong>y arereally as isolated as <strong>the</strong>y appear to be, <strong>the</strong>y representan earliercommunication, ei<strong>the</strong>r by an approximation <strong>of</strong> <strong>the</strong> two islandsover <strong>the</strong> space now occupied by <strong>the</strong> Moluccas ; or, what is perhapsmore probable, through a former extension <strong>of</strong> <strong>the</strong> Moluccas,which have since undergone so much subsidence, as to lead to<strong>the</strong> extinction <strong>of</strong> a large proportion <strong>of</strong> <strong>the</strong>ir ancient fauna.<strong>The</strong> wide-spread volcanic action, and especially <strong>the</strong> prevalence <strong>of</strong>raised coral-reefs in almost all <strong>the</strong> islands, render this lastsupposition very probable.On <strong>the</strong> Oriental side <strong>the</strong> difficulty is greater ; for here we find,what seem to be clear indications <strong>of</strong> a connection <strong>with</strong> Africa, aswell as <strong>with</strong> Continental Asia, at some immensely remote epoch.Cynopi<strong>the</strong>cus, Babirusa, and Anoa; Ceijcopsis, Streptocitta, and Gazzola(s. g.), and perhaps Scissirostrum, may be well explained asdescendants <strong>of</strong> ancestral types in<strong>the</strong>ir respective groups, whichalso gave rise to <strong>the</strong> special forms <strong>of</strong> Africa on <strong>the</strong> one hand, and <strong>of</strong>Asia on <strong>the</strong> o<strong>the</strong>r.For this immigration we must suppose, that ata period before <strong>the</strong> formation <strong>of</strong> <strong>the</strong> present Indo-Malay Islands,a great tract <strong>of</strong> land extended in anorth-westerly direction, tillit met <strong>the</strong> old Asiatic continent. This may have been before


438 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>the</strong> Himalayas had risen to any great height, and when a largepart <strong>of</strong> what are now <strong>the</strong> cold plateaus <strong>of</strong> Central Asia mayhave teemed <strong>with</strong> life, some forms <strong>of</strong> which are preserved inAfrica, some in Malaya, and a few in Celebes. Here mayhave lived <strong>the</strong> common ancestor <strong>of</strong> Stis, Babimsa, and Phacochosrus; as well as <strong>of</strong> Cynopi<strong>the</strong>cus, Cynocephalus, and Macacus ;<strong>of</strong> Anoa and Bubalus; <strong>of</strong> Scissirostrum and Euryceros ; <strong>of</strong> Ceyx,Ceycopsis, and Ispidina. Such an origin accounts, too, for <strong>the</strong>presence <strong>of</strong> <strong>the</strong> North-Indian forms in Celebes ;and it <strong>of</strong>fers lessdifficulties than a direct connection <strong>with</strong> continental Africa, whichonce appeared to be <strong>the</strong> only solution <strong>of</strong> <strong>the</strong> problem. If thissouth-eastward extension <strong>of</strong> Asia occurred at <strong>the</strong> same time as<strong>the</strong> north-eastward extension <strong>of</strong> South Africa and Madagascar,<strong>the</strong> two early continents may have approached each o<strong>the</strong>r sufficientlyto have allowed <strong>of</strong> some interchange <strong>of</strong> forms : Tarsiusmay be <strong>the</strong> descendant <strong>of</strong> some Lemurine animal that <strong>the</strong>nentered <strong>the</strong> Malayan area, while <strong>the</strong> progenitors <strong>of</strong> Cryptoproctamay <strong>the</strong>n have passed from Asia to Madagascar.It is true that we here reach <strong>the</strong> extremest limits <strong>of</strong> speculation; but when we have before us such singular phenomena asare presented by <strong>the</strong> fauna <strong>of</strong> <strong>the</strong> island <strong>of</strong> Celebes, we can hardlyhelp endeavouring to picture to our imaginations by what pastchanges <strong>of</strong> land and sea (in <strong>the</strong>mselves not improbable) <strong>the</strong> actualcondition <strong>of</strong> things may have been brought about.II. Australia and Tasmania, or <strong>the</strong> Australian Sub-region.A general sketch <strong>of</strong> Australian zoology having been given in <strong>the</strong>earlier part <strong>of</strong> this chapter, it will not be necessary to occupy muchtime on this sub-region, which isas remarkably homogeneous as<strong>the</strong> one we have just left is heterogeneous. Although much <strong>of</strong><strong>the</strong> nor<strong>the</strong>rn part <strong>of</strong> Australia is <strong>with</strong>in <strong>the</strong> tropics, while Victoriaand Tasmania are situated from 36° to 43° south latitude,<strong>the</strong>re is no striking change in <strong>the</strong> character <strong>of</strong> <strong>the</strong> faunathroughout <strong>the</strong> continent ;a number <strong>of</strong> important genera extendingover <strong>the</strong> whole country, and giving a very uniform characterto its zoology. <strong>The</strong> eastern parts, including <strong>the</strong> colonies <strong>of</strong> NewSouth Wales and Queensland, are undoubtedly <strong>the</strong> richest, several


;chap, xiii.] THE AUSTRALIAN REGION. 439peculiar types being found only here. <strong>The</strong> sou<strong>the</strong>rn portion issomewhat poorer, and has very few peculiar forms ; and Tasmaniabeing isolated is poorer still, yet its zoology has much resemblanceto that <strong>of</strong> Victoria, from which country it has evidentlynot been very long separated. <strong>The</strong> north, as far as yet known, ischaracterised by hardly any peculiar forms, but by <strong>the</strong> occurrence<strong>of</strong> a number <strong>of</strong> Papuan types, which have evidently been derivedfrom New Guinea.Mammalia.—<strong>The</strong> Australian sub-region contains about 160species <strong>of</strong> Mammalia, <strong>of</strong> which 3 are Monotremata, 102 Marsupials,23 Chiroptera, 1 Carnivora (<strong>the</strong> native dog, probably notindigenous), and 31 Muridae. <strong>The</strong> north is characterised by aspecies <strong>of</strong> <strong>the</strong> Austro-Malayan genus Cuscus.Phascolarctos (<strong>the</strong>koala, or native bear) is found only in <strong>the</strong> eastern districts;Phascolomys (<strong>the</strong> wombat) in <strong>the</strong> south-east and TasmaniaPetaurista (a peculiar form <strong>of</strong> flying opossum), in <strong>the</strong> east.Thylacinus(<strong>the</strong> zebra-wolf), and Sarcophilus (<strong>the</strong> " native devil "),two carnivorous marsupials, are confined to Tasmania. WestAustralia, <strong>the</strong> most isolated and peculiar region botanically,alone possesses <strong>the</strong> curious littlehoney-eating Tarsipes, and <strong>the</strong>Peragalea, or native rabbit. <strong>The</strong> remarkable Myrmecobius, asmall ant-eating marsupial, is found in <strong>the</strong> west and south;and Onychogalea, a genus <strong>of</strong> kangaroos, in West and CentralAustralia. All <strong>the</strong> o<strong>the</strong>r genera have a wider <strong>distribution</strong>, aswill be seen by a reference to <strong>the</strong> list at <strong>the</strong> end <strong>of</strong> thischapter.Plate XL A Scene in Tasmania, <strong>with</strong> Characteristic Mammalia.—As some <strong>of</strong> <strong>the</strong> most remarkable Mammalia <strong>of</strong> <strong>the</strong> Australianregion are now found only in Tasmania, we have chosen thisisland for <strong>the</strong> scene <strong>of</strong> our first illustration <strong>of</strong> <strong>the</strong> fauna <strong>of</strong> <strong>the</strong>Australian sub-region. <strong>The</strong> pair <strong>of</strong> large striped <strong>animals</strong> arezebra-wolves {Thylacinus cynocephalus), <strong>the</strong> largest and most destructive<strong>of</strong> <strong>the</strong> carnivorous marsupials.<strong>The</strong>se creatures used tobe tolerably plentiful in Tasmania, where <strong>the</strong>y are alone found.<strong>The</strong>y are also called "native tigers," or " native hyaenas;" and bein^destructive to sheep, <strong>the</strong>y have been destroyed by <strong>the</strong> farmersand will doubtless soon be exterminated.In <strong>the</strong> foreground on-'e*


:440 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>the</strong> left is a bandicoot (Peramclcs gunnii). <strong>The</strong>se are delicate little<strong>animals</strong> allied to <strong>the</strong> kangaroos ; and <strong>the</strong>y are found in all parts <strong>of</strong>Australia, and Tasmania, to which latter country this speciesis confined. On <strong>the</strong> right is <strong>the</strong> wombat (Phascolomys wombat),a root-eating marsupial, <strong>with</strong> large incisor teeth like those <strong>of</strong> ourrodents. <strong>The</strong>y inhabit south-east Australia and Tasmania. In<strong>the</strong> foreground is <strong>the</strong> porcupine ant-eater (Echidna sctosa), belongingto a distinct order <strong>of</strong> mammalia, Monotremata, <strong>of</strong> which <strong>the</strong>only o<strong>the</strong>r member is <strong>the</strong> duck-billed OrnitJwrhynchus. <strong>The</strong>se<strong>animals</strong> are, however, more nearly allied to <strong>the</strong> marsupials, thanto <strong>the</strong> insectivora or edentata <strong>of</strong> <strong>the</strong> rest <strong>of</strong> <strong>the</strong> world, which insome respects <strong>the</strong>y resemble. An allied species (Echidna hystrix)inhabits south-east Australia.Birds.—Australia (<strong>with</strong> Tasmania) possesses about 630 species<strong>of</strong> birds, <strong>of</strong> which 485 are land-birds. Not more than about onetwentieth<strong>of</strong> <strong>the</strong>se are found elsewhere, so that it has a largerproportion <strong>of</strong> endemic species than any o<strong>the</strong>r sub-region on <strong>the</strong>globe. <strong>The</strong>se birds are divided among <strong>the</strong> several orders asfollowsPasseres


—chap, xiii.] THE AUSTRALIAN. REGION. 441to Tasmania ; which island appears to possess no peculiargenus <strong>of</strong> birds except Eudyptes, one <strong>of</strong> <strong>the</strong> penguins. WestAustralia has no wholly peculiar genus except Geopsittacus, acurious form <strong>of</strong> ground parroquet; <strong>the</strong> singular Atrichia, firstfound here, having been discovered in <strong>the</strong> east. In NorthAustralia, Emblcma (Ploceidce) is <strong>the</strong> only peculiar Australiangenus, but several Austro-Malayan and Papuan genera enter,as, Syma and Tanysiptera (Alcedinidse) ;Machcerihynchus(Muscicapidse) ; Calornis (Sturnidae) ; Manucodia, Ptilorhis, andjElurcedus (Paradiseidse) ; Megapodius ; and Casuarius. <strong>The</strong> presence<strong>of</strong> a species <strong>of</strong> bustard (Eupodotis) in Australia, is verycurious, its nearest allies being in <strong>the</strong> plains <strong>of</strong> India and Africa.Among waders <strong>the</strong> genus Tribonyx, a thick-legged bird somewhatresembling <strong>the</strong> Notornis <strong>of</strong> New Zealand, though not closelyallied to it, is <strong>the</strong> most remarkable. <strong>The</strong> district where <strong>the</strong>typical Australian forms most abound is undoubtedly <strong>the</strong> easternside <strong>of</strong> <strong>the</strong> island. <strong>The</strong> north and south are both somewhatpoorer, <strong>the</strong> west much poorer, although it possesses a few verypeculiar forms, especially among Mammalia. Tasmania is <strong>the</strong>poorest <strong>of</strong> all,a considerable number <strong>of</strong> genera being here wanting;but, except <strong>the</strong> two peculiar carnivorous marsupials, itpossesses nothing to mark it <strong>of</strong>f zoologically from <strong>the</strong> adjacentparts <strong>of</strong> <strong>the</strong> main land. It is probable that its insular climate,more moist and less variable than that <strong>of</strong> Australia, may not besuitable to some <strong>of</strong> <strong>the</strong> absent forms ; while o<strong>the</strong>rs may requiremore space and more varied conditions, than are <strong>of</strong>fered by acomparatively small island.<strong>The</strong> remaining classes <strong>of</strong> <strong>animals</strong> have been already discussedin our sketch <strong>of</strong> <strong>the</strong> region as a whole (p. 396).Plate XII Illustrating <strong>the</strong> Fauna <strong>of</strong> Australia.—In thisplate we take New South "Wales as our locality, and representchiefly, <strong>the</strong> more remarkable Australian types <strong>of</strong> birds. <strong>The</strong>most conspicuous figure is <strong>the</strong> wonderful lyre-bird (Menura superba),<strong>the</strong> elegant plumage <strong>of</strong> whose tail is altoge<strong>the</strong>r unique in<strong>the</strong> whole class <strong>of</strong> birds. <strong>The</strong> unadorned bird is <strong>the</strong> female. In<strong>the</strong> centre is <strong>the</strong> emu (Dromceus novm-hollandice) ,<strong>the</strong> representativein Australia, <strong>of</strong> <strong>the</strong> ostrich in Africa and America, but be-Yol. I.—30


442 ZOOLOGICAL GEOGKAPHY. [part in.longing to a different family, <strong>the</strong> Casurariidaa* To <strong>the</strong> right area pair <strong>of</strong> crested pigeons {Ocyphaps lophotes), one <strong>of</strong> <strong>the</strong> many singularforms <strong>of</strong> <strong>the</strong> pigeon family to which <strong>the</strong> Australian regiongives birth. In every o<strong>the</strong>r part <strong>of</strong> <strong>the</strong> globe pigeons aresmooth-headed birds, but here <strong>the</strong>y have developed three distinctforms <strong>of</strong> crest, as seen in this bird, <strong>the</strong> crowned pigeonfigured in Plate X., and <strong>the</strong> double-crested pigeon (Lopholcemusantardicus). <strong>The</strong> large bird on <strong>the</strong> tree is one <strong>of</strong> <strong>the</strong> Australianfrog-mou<strong>the</strong>d goat-suckers (Podargits strigoides), which arecalled in <strong>the</strong> colony " More-pork," from <strong>the</strong>ir peculiar cry. <strong>The</strong>ydo not capture <strong>the</strong>ir prey on <strong>the</strong> wing like true goat-suckers, buthunt about <strong>the</strong> branches <strong>of</strong> trees at dusk, for large insects, andalso for unfledged birds.A large kangaroo {Macropus giganteus)is seen in <strong>the</strong> distance ;and passing through <strong>the</strong> air, a flyingopossum (Petaurus sciureus), a beautiful modification <strong>of</strong> a marsupial,so as to resemble in form and habits <strong>the</strong> flying squirrels<strong>of</strong> <strong>the</strong> nor<strong>the</strong>rn hemisphere.III. Hie Pacific Islands, or Polynesian Sub-region.Although <strong>the</strong> area <strong>of</strong> this sub-region is so vast, and <strong>the</strong>number <strong>of</strong> islands it contains almost innumerable, <strong>the</strong>re is aconsiderable amount <strong>of</strong> uniformity in its forms <strong>of</strong> animal life.From <strong>the</strong> Ladrone islands on <strong>the</strong> west, to <strong>the</strong> Marquesas on <strong>the</strong>east, a distance <strong>of</strong> more than 5,000 miles, <strong>the</strong> same characteristicgenera <strong>of</strong> birds prevail ; and this is <strong>the</strong> only class <strong>of</strong> <strong>animals</strong> onwhich we can depend, mammalia being quite absent, and reptilesvery scarce.<strong>The</strong> Sandwich Islands, however, form an exceptionto this uniformity; and, as far as we yet know, <strong>the</strong>y are sopeculiar that <strong>the</strong>y ought, perhaps, to form a separatesub-region.<strong>The</strong>y are, however, <strong>geographical</strong>ly a part <strong>of</strong> Polynesia; and amore careful investigation <strong>of</strong> <strong>the</strong>ir natural history may showmore points <strong>of</strong> agreement <strong>with</strong> <strong>the</strong> o<strong>the</strong>r islands. It is <strong>the</strong>reforea matter <strong>of</strong> convenience, at present, to keep <strong>the</strong>m in <strong>the</strong> Polynesiansub-region, which may be divided into Polynesia properand <strong>the</strong>Sandwich Islands.Polynesia proper consists <strong>of</strong> a number <strong>of</strong> groups <strong>of</strong> islands <strong>of</strong>some importance, and a host <strong>of</strong> smaller intermediate islets.


ECS3"SE- 13O•*!2Oaat»rpitr1HH>>oS•s.&>r-


———chap, xin.] THE AUSTRALIAN REGION. 443-For <strong>the</strong> purpose <strong>of</strong> zoological comparison, we may class<strong>the</strong>m infour main divisions. 1. <strong>The</strong> Ladrone and Caroline Islands;2. New Caledonia and <strong>the</strong> New Hebrides ; 3. <strong>The</strong> Fiji, Tonga,and Samoa Islands; 4. <strong>The</strong> Society, and Marquesas Islands.<strong>The</strong> typical Polynesian fauna is most developed in <strong>the</strong> thirddivision ; and it will be well to describe this first, and <strong>the</strong>n showhow <strong>the</strong> o<strong>the</strong>r islands diverge from it, and approximate o<strong>the</strong>rsub-regions.Fiji, Tonga, and Samoa Islands.—<strong>The</strong> land-birds inhabiting<strong>the</strong>se islands belong to 41 genera, <strong>of</strong> which 17 are characteristic<strong>of</strong> <strong>the</strong> Australian region, and 9 more peculiarly Polynesian.<strong>The</strong> characteristic Australian genera are <strong>the</strong> following : Petroica(Sylviidae) ; Lalage (Campephagidae) ; Monarcha, Myiagra, Rhipidura(Muscicapidse) ; Pachyccphala (Pachycephalidae) ; Eectes(Laniida?) ; Myzomela, Ptilotis, Anthochmra (Meliphagidae) ; Amadina,Eythrura, (Ploceidse) ; Artamus (Artamidse) ; Lorius (Trichoglossidre); Ptilopus, Phlogainas (Columbidse) ; Megapodius(Megapodiidse).<strong>The</strong> peculiar Polynesian genera are :Tatare,Lamprolia(Sylviidae) ;Aplonis, Sturnodes (Sturnidae) ; Todiramphus(Alcedinidffi) ;Pyrhulopsis, Cyanoramphus, (Platycercidae) ; Coriphilus(Trichoglossidae) ;Didunculus (Didunculidae).<strong>The</strong> wide-spread genera are Turdus, Zosterops,Hirundo, Halcyon,Collocalia, Eudynamis Cuculus, Ianthoenas, Carpopliaga,Turtur, Halimdus, Astur, Circus, Strix, Asio. <strong>The</strong> aquatic birdsare fifteen in number, all wide-spread species except one—a form<strong>of</strong> moor-hen (Gallinulidse), which has been constituted a newgenus Pareudiastes.Society, and Marquesas Islands.—Here, <strong>the</strong> number <strong>of</strong> genera <strong>of</strong>land-birds has considerably diminished, amounting only to 16in all. <strong>The</strong> characteristic Australian genera are 5 ;Monarcha,Anthoclucra, Trichoglossus, Ptilopus, and Phlogcenas. <strong>The</strong> Polynesiangenera are 4 ;Tatare, Todiramphus, Cyanoramphus,Coriphilus, and one recently described genus, Scrresius, an extraordinaryform <strong>of</strong> large fruit pigeon, here classed under Carpopliaga.<strong>The</strong>se remote groups have thus all <strong>the</strong> character <strong>of</strong>Oceanic islands, even as regards <strong>the</strong> rest <strong>of</strong> Polynesia, since <strong>the</strong>y


——;444 ZOOLOGICAL GEOGKAPHY. [part m.possess hardly anything, but what <strong>the</strong>y might have receivedby immigration over a wide extent <strong>of</strong> ocean.Zadrone, and Caroline Islands.—<strong>The</strong>se extensive groups <strong>of</strong>small islands are very imperfectly known, yet a considerablenumber <strong>of</strong> birds have been obtained. <strong>The</strong>y possess twopeculiar Polynesian genera, Tatare and Sturnodes ; one peculiarsub-genus, Psammathia (here included under Acrocephalus);and ten <strong>of</strong> <strong>the</strong> typicalAustralian genera found in Polynesia,Lalage, Monarcha, Myiagra, Rhipidura, Myzomela, Erythrura,Artamtts, Phlogcenas, Ptilopus, and Megapodius, as well as <strong>the</strong>Papuan genus Pedes, and <strong>the</strong> Malayan Calornis ; —so that <strong>the</strong>ycan be certainly placed in <strong>the</strong> sub-region. Genera which donot occur in <strong>the</strong> o<strong>the</strong>r Polynesian islands are, Acrocephalus, (s.g.Psammathia) originally derived perhaps from <strong>the</strong> Philippinesand Caprimulgus, a peculiar species, allied to one from Japan.New Caledonia, and <strong>the</strong> New Hebrides.—Although <strong>the</strong>se islandsseem best placed <strong>with</strong> Polynesia, yet <strong>the</strong>y form a transition toAustralia proper, and to <strong>the</strong> Papuan group. <strong>The</strong>y possess 30genera <strong>of</strong> land-birds, 18 <strong>of</strong> which are typical <strong>of</strong> <strong>the</strong> Australianregion ; but wdiile 13 are also Polynesian, <strong>the</strong>re are 5 which donot pass fur<strong>the</strong>r east. <strong>The</strong>se are Acanthiza, Eopsaltria, Gliciphila,Philemon, and Ianthoenas.<strong>The</strong> peculiar Polynesian genus,Aplonis, <strong>of</strong> which three species inhabit New Caledonia, link it to<strong>the</strong> o<strong>the</strong>r portions <strong>of</strong> <strong>the</strong> sub-region. <strong>The</strong> following are <strong>the</strong>genera at present known from New Caledonia :Turdus, Acanthiza,Campephaga, Lalage, Myiagra, Rhipidura, Pachycephala,Eopsaltria, Corvus, Physocorax (s.g. <strong>of</strong> Corvus, allied to <strong>the</strong> jackdaws),Glicphila, Anthochaira, Philemon, Zosterops, Erythrura,Aplonis, Artamtts, Cuculus, Halcyon, Collocalia, Cyanoramphus,Trichoglossus, Ptilopus, Carpophaga, Macropygia, Ianthoenas,Chalcophaps, Haliastur, Accipiter. <strong>The</strong> curious Phinochetusjubatus, forming <strong>the</strong> type <strong>of</strong> a distinct family <strong>of</strong> birds(Rhinochetidee),allied to <strong>the</strong> herons, is only known from New Caledonia.It thus appears, that not more than about 50 genera and 150species <strong>of</strong> land-birds, are known from <strong>the</strong> vast number <strong>of</strong> islandsthat are scattered over <strong>the</strong> Central Pacific, and it is not probable


—;chap, xiu.] THE AUSTRALIAN REGION. 445that <strong>the</strong> number will be very largely increased. Some <strong>of</strong> <strong>the</strong>species, as <strong>the</strong> Eudynamis taitensis and Tatare longirostris, rangeover 40° <strong>of</strong> longitude, from <strong>the</strong> Fiji Islands to <strong>the</strong> Marquesas. Ino<strong>the</strong>r genera, as Cyanommplms and Ptilopus, each importantisland or group <strong>of</strong> islands, has its peculiar species. <strong>The</strong> connection<strong>of</strong> all <strong>the</strong>se islands <strong>with</strong> each o<strong>the</strong>r, on <strong>the</strong> one hand, and<strong>the</strong>ir close relation to <strong>the</strong> Australian region, on <strong>the</strong> o<strong>the</strong>r, areequally apparent ; but we have no sufficient materials for speculating<strong>with</strong> any success, on <strong>the</strong> long series <strong>of</strong> changes that havebrought about <strong>the</strong>ir existing condition, as regards <strong>the</strong>ir peculiarforms <strong>of</strong> animal life.Sandwich Islands.—Thissomewhat extensive group <strong>of</strong> largeislands, is only known to contain 11 genera and 18 species <strong>of</strong>indigenous land-birds ; and even <strong>of</strong> this small number, two birds<strong>of</strong> prey are wide ranging species, which may well have reached <strong>the</strong>islands during <strong>the</strong>ir present isolated condition. <strong>The</strong>se latter are,Strix delicatula, an owl spread over Australia and <strong>the</strong> Pacific;and Asio accipitrinus, a species which has reached <strong>the</strong> Galapagosfrom S. America, and <strong>the</strong>nce perhaps <strong>the</strong> Sandwich Islands.<strong>the</strong> remaining 8 genera, one is a crow (Corvus hawaiensis), andOfano<strong>the</strong>r a fishing eagle (Pandion solitarius), <strong>of</strong> peculiar species;leaving 7 genera, which are all (according to Mr. Sclater) peculiar.First we have Chasiempis, a genus <strong>of</strong> Muscicapidse, containingtwo species (which may however belong to distinct genera) ;andas <strong>the</strong> entire family is unknown on <strong>the</strong> American continent<strong>the</strong>se birds must almost certainly be allied to some <strong>of</strong> <strong>the</strong>numerous Muscicapine forms <strong>of</strong> <strong>the</strong> Australian region. Nextwe have <strong>the</strong> purely Australian family Meliphagida?, representedby two genera, Moho, an isolated form, and Chwtoptila, a genusestablished by Mr. Sclater for a bird before classed in Entomyza,an Australian group. <strong>The</strong> four remaining genera are believedby Mr. Sclater to belong to one group, <strong>the</strong> Drepanididse, altoge<strong>the</strong>rconfined to <strong>the</strong> Sandwich Islands. Two <strong>of</strong> <strong>the</strong>m, Drepanis andHemignathus, <strong>with</strong> three species each, are undoubtedly allied<strong>the</strong> o<strong>the</strong>r two, Loxops and Psittirostra, have usually been classedas finches. <strong>The</strong> former seem to approach <strong>the</strong> Dicaeidse; and allresemble this group in <strong>the</strong>ir coloration.


446 ZOOLOGICAL GEOGRAPHY. [part hi.<strong>The</strong> aquatic birds and waders all belong to wide-spreadgenera, and only one or two are peculiar species."<strong>The</strong> Sandwich Islands thus possess a larger proportion <strong>of</strong>peculiar genera and species<strong>of</strong> land-birds than any o<strong>the</strong>r group<strong>of</strong> islands, and <strong>the</strong>y are even more strikingly characterised bywhat seems to be a peculiar family. <strong>The</strong> only o<strong>the</strong>r class <strong>of</strong>terrestrial <strong>animals</strong> at all adequately represented on <strong>the</strong>se islands,are <strong>the</strong> land shells ;and here too we find a peculiar family, subfamily,or genus (Achatinella or Achatinellidse)consisting <strong>of</strong> anumber <strong>of</strong> genera, or sub-genera,— according to <strong>the</strong> divergent views<strong>of</strong> modern conchologists,—and nearly 300 species. <strong>The</strong> Eev. J.T. Gulick, who has made a special <strong>study</strong> <strong>of</strong> <strong>the</strong>se shells on <strong>the</strong>spot, considers that <strong>the</strong>re are 10 genera, some <strong>of</strong> which are confinedto single islands. <strong>The</strong> species are so restricted that <strong>the</strong>iraverage range is not more than five or six square miles, whilesome areconfined to a tract <strong>of</strong> only two square miles in extent,and very few range over an entire island. Some species areconfined to <strong>the</strong> mountain ridges, o<strong>the</strong>rs to <strong>the</strong> valleys ;and eachridge or valley possesses its peculiar species. Considerablymore than half <strong>the</strong> species occur in <strong>the</strong> island <strong>of</strong> Oahu, where<strong>the</strong>re is a good deal <strong>of</strong> forest. Very few shells belonging too<strong>the</strong>r groups occur, and <strong>the</strong>y are all small and obscure ; <strong>the</strong>Achatinellse almost monopolising <strong>the</strong> entire archipelago.Remarks on <strong>the</strong> probable past history <strong>of</strong> <strong>the</strong> Sandwich Islands.—<strong>The</strong> existence <strong>of</strong> <strong>the</strong>se peculiar groups <strong>of</strong> birds and landshellsin so remote a group <strong>of</strong> volcanic islands, clearly indicatesthat <strong>the</strong>y are but <strong>the</strong> relics <strong>of</strong> a more extensive land ; and <strong>the</strong>reefs and islets that stretch for more than 1,000 miles in a westnorth-westdirection, may be <strong>the</strong> remains <strong>of</strong> a country oncesufficiently extensive to develope <strong>the</strong>se and many o<strong>the</strong>r, nowextinct, forms <strong>of</strong> life. 1Some light may perhaps be thrown on <strong>the</strong> past history <strong>of</strong> <strong>the</strong>1A new genus <strong>of</strong> Beetles (Apterocyclus) <strong>of</strong> <strong>the</strong> family Lucanidse, hasrecently been described from <strong>the</strong> Sandwich Islands, and it is said to be mostnearly related to a group inhabiting Chih,—an indication ei<strong>the</strong>r <strong>of</strong> <strong>the</strong> greatantiquity <strong>of</strong> <strong>the</strong> fauna, or <strong>of</strong> <strong>the</strong> varied accidental migrations from which ithas had its origin.


—;chap, xiii.] THE AUSTRALIAN REGION. 447Sandwich Islands, by <strong>the</strong> peculiar plants which are found on<strong>the</strong>ir mountains. <strong>The</strong> peak <strong>of</strong> Teneriffe produces no Alpineplants <strong>of</strong> European type, and this has been considered to provethat it has been always isolated ; whereas <strong>the</strong> occurrence <strong>of</strong> NorthTemperate forms on <strong>the</strong> mountains <strong>of</strong> Java, accords <strong>with</strong> o<strong>the</strong>r evidence<strong>of</strong> this island having once formed part <strong>of</strong> <strong>the</strong> Asiatic continent.Now on <strong>the</strong> higher summits <strong>of</strong> <strong>the</strong> Sandwich Islands,nearly 30 genera <strong>of</strong> Arctic and North Temperate flowering plantshave been found. Many <strong>of</strong> <strong>the</strong>se occur also in <strong>the</strong> South Temperatezone, in Australia or New Zealand ; but <strong>the</strong>re are o<strong>the</strong>rs whichseem plainly to point to a former connection <strong>with</strong> some NorthTemperate land, probably California, as a number <strong>of</strong> islets arescattered in <strong>the</strong> ocean between <strong>the</strong> two countries. <strong>The</strong> mostinteresting genera are <strong>the</strong> following :Silene, which is whollyNorth Temperate, except that it occurs in S. Africa ;Vicia,also North Temperate, and in South Temperate America;Fragaria, <strong>with</strong> a similar <strong>distribution</strong>; Aster, widely spreadin America, o<strong>the</strong>rwise North Temperate only ; Vaccinium,wholly confined to <strong>the</strong> nor<strong>the</strong>rn hemisphere, in cold andtemperate climates. None <strong>of</strong> <strong>the</strong>se are found in Australia orNew Zealand ;and <strong>the</strong>ir presence in <strong>the</strong> Sandwich Islandsseems clearly to indicate a former approximation to NorthTemperate America, although <strong>the</strong> absence <strong>of</strong> any Americanforms <strong>of</strong> vertebrata renders it certain that no actual land connectionever took place.Recent soundings have shown, that <strong>the</strong> Sandwich Islandsrise from a sea which is 3,000 fathoms or 18,000 feet deepwhile <strong>the</strong>re is a depth <strong>of</strong> at least 2,000 fathoms all across toCalifornia on one side, and to Japan on <strong>the</strong> o<strong>the</strong>r. Between<strong>the</strong> Fiji Islands, New Caledonia, <strong>the</strong> Solomon Islands, andAustralia, <strong>the</strong> depth isabout 1,300 fathoms, and between Sydneyand New Zealand 2,600 fathoms ; showing, in every case, ageneral accordance between <strong>the</strong> depth <strong>of</strong> sea and <strong>the</strong> approximation<strong>of</strong> <strong>the</strong> several faunas. In a few more years, when it isto be hoped we shall know <strong>the</strong> contour <strong>of</strong> <strong>the</strong> sea-bottom betterthan that <strong>of</strong> <strong>the</strong> continents, we shall be able to arrive at moredefinite and trustworthy conclusions as to <strong>the</strong> probable changes


—.448 ZOOLOGICAL GEOGRAPHY. [PART III.<strong>of</strong> land and sea by which <strong>the</strong> phenomena <strong>of</strong> animal <strong>distribution</strong>in <strong>the</strong> Pacific have been brought about.Reptiles <strong>of</strong> <strong>the</strong> Polynesian Sub-region.—<strong>The</strong> researches <strong>of</strong>Mr. Darwin on Coral Islands, proved, that large areas in <strong>the</strong>Pacific Ocean have been recently subsiding; but <strong>the</strong> peculiarforms <strong>of</strong> life which <strong>the</strong>y present, no less clearly indicate <strong>the</strong>former existence <strong>of</strong> some extensive lands. <strong>The</strong> total absence <strong>of</strong>Mammalia, however, shows ei<strong>the</strong>r that <strong>the</strong>se lands never formedpart <strong>of</strong> <strong>the</strong> Australian or Papuan continents, or if<strong>the</strong>y did, that<strong>the</strong>y have been since subjected to such an amount <strong>of</strong> subsidenceas to exterminate most <strong>of</strong> <strong>the</strong>ir higher terrestrial forms <strong>of</strong> life.It is a remarkable circumstance, that although Mammalia (exceptbats) are wanting, <strong>the</strong>re are a considerable number <strong>of</strong> reptilesrano-in« over <strong>the</strong> whole sub-region. Lizards are <strong>the</strong> mostnumerous, five families and fourteen genera being represented,as follows :1. Cryptoblepharus2. Ablepharus ...3. Lygosoma4. Mabouya5. Euprepes.6. Dactyloperus ...7. Doryura8. Gehyra9. Amydosaurus .10. Heteronota11. (hrrdophus ...Brachylophus . .12.13. Lophura14. Chioroscartes(Gyninopthalnridse)n(Scincidgej»(Geckotidse)(Geckotidse)(Iguanidse)(Agaiuidse)Fiji Islands.All <strong>the</strong> islands.Pelew Islands, New Caledonia.Samoa Islands.Pacific Islands.Sandwich Islands.Pacific Islands.Fiji Islands.Tahiti.Fiji Islands.New Caledonia.Fiji Islands.Pelew Islands.Fiji Islands.<strong>The</strong> first five are wide-spread genera, represented mostly bypeculiar species ; but sometimes <strong>the</strong> species <strong>the</strong>mselves have awide range, as in <strong>the</strong> case <strong>of</strong> Ablepharus pazcilopleurus, which(according to Dr. Gun<strong>the</strong>r) is found in Timor, Australia, NewCaledonia, Savage Island (one <strong>of</strong> <strong>the</strong> Samoa group), and <strong>the</strong>Sandwich Islands! Gehyra and Heteronota are Australiangenera ;while Lophura has reached <strong>the</strong> Pelew Islands from <strong>the</strong>Moluccas. <strong>The</strong> remainder (printed in italics), are peculiar genera;Brachylophus being especially interesting as an example <strong>of</strong> an


chap, xiii.] THE AUSTRALIAN REGION. 449o<strong>the</strong>rwise peculiar American family, occurring so far acrossPacific.<strong>the</strong>Snakes are much less abundant, only four genera being represented,one <strong>of</strong> <strong>the</strong>m marine. <strong>The</strong>y are, Anoplodipsas, a peculiargenus <strong>of</strong> Amblycephalidse from New Caledonia; Enygrus, agenus <strong>of</strong> Pythonidse from <strong>the</strong> Fiji Islands ; Ogmodon, a peculiargenus <strong>of</strong> Elapidae, also from <strong>the</strong> Fiji Islands, but ranging toPapua and <strong>the</strong> Moluccas ;and Platurus, a wide-spread genus <strong>of</strong>sea-snakes (Hydrophidre). In <strong>the</strong> more remote Sandwich andSociety Islands <strong>the</strong>re appear to be no snakes. This accords<strong>with</strong> our conclusion that lizards have some special means <strong>of</strong>dispersal over <strong>the</strong> ocean which detracts from <strong>the</strong>ir value asindicating zoo-<strong>geographical</strong> affinities ;which is fur<strong>the</strong>r provedby <strong>the</strong> marvellous range <strong>of</strong> a single species (referred to above)from Australia to<strong>the</strong> Sandwich Islands.A species <strong>of</strong> Hyla is said to inhabit <strong>the</strong> New Hebrides, andseveral species <strong>of</strong> Platymantis (tree-frogs) are found in <strong>the</strong> FijiIslands ; but o<strong>the</strong>rwise <strong>the</strong> Amphibians appear to be unrepresentedin <strong>the</strong> sub-region, though <strong>the</strong>y will most likely be foundin so large an island as New Caledonia.From <strong>the</strong> foregoing sketch, it appears, that although <strong>the</strong>reptiles present some special features, <strong>the</strong>y agree on <strong>the</strong> whole<strong>with</strong> <strong>the</strong> birds, in showing, that <strong>the</strong> islands <strong>of</strong> Polynesiaall belong to <strong>the</strong> Australian region, and that in <strong>the</strong> Fiji Islandsis to be found <strong>the</strong> fullest development <strong>of</strong> <strong>the</strong>ir peculiarfauna.IV. New Zealand Sub-region.<strong>The</strong> islands <strong>of</strong> New Zealand are more completely oceanicthan any o<strong>the</strong>r extensive tract <strong>of</strong> land, being about 1,200miles from Australia and nearly <strong>the</strong> same distance from NewCaledonia and <strong>the</strong> Friendly Isles. <strong>The</strong>re are, however, severalislets scattered around, whose productions show that <strong>the</strong>ybelong to <strong>the</strong> same sub-region ;—<strong>the</strong> principal being, NorfolkIsland, Lord Howe's Island, and <strong>the</strong> Kermadec Isles, on <strong>the</strong>north ; Chatham Island on <strong>the</strong> east ; <strong>the</strong> Auckland and MacquarieIsles on <strong>the</strong> south ;—and if <strong>the</strong>se were once joined to


—450 ZOOLOGICAL GEOGRAPHY. [part hi.New Zealand, <strong>the</strong>re would have been formed an island-continentnot much inferior in extent to Australia itself.New Zealand is wholly situated in <strong>the</strong> warmer portion <strong>of</strong><strong>the</strong> Temperate zone, and enjoys an exceptionally mild andequable climate. It has abundant moisture, and thus comes<strong>with</strong>in <strong>the</strong> limits <strong>of</strong> <strong>the</strong> South-Temperate forest zone ; and thisleads to its productions <strong>of</strong>ten resembling those <strong>of</strong> <strong>the</strong> tropical,but moist and wooded, islands <strong>of</strong> <strong>the</strong>Pacific, ra<strong>the</strong>r than those<strong>of</strong> <strong>the</strong> temperate, but arid and scantily wooded plains <strong>of</strong> Australia.<strong>The</strong> two islands <strong>of</strong> New Zealand are about <strong>the</strong> sameextent (approximately) as <strong>the</strong> British Isles, but <strong>the</strong> difference in<strong>the</strong> general features <strong>of</strong> <strong>the</strong>ir natural history is very great.<strong>The</strong>reare, in <strong>the</strong> former, no mammalia, less than half as many birds,very few reptiles and fresh-water fishes, and an excessive andmost unintelligible poverty <strong>of</strong> insects; yet, considering <strong>the</strong>situation <strong>of</strong> <strong>the</strong> islands and <strong>the</strong>ir evidently long-continuedisolation, <strong>the</strong> wonder ra<strong>the</strong>r is that <strong>the</strong>ir fauna is so variedand interesting as it is found to be. Our knowledge <strong>of</strong> thisfauna, though no doubt far from complete, is sufficientlyample ; and it will be well to give a pretty full account <strong>of</strong>it, in order to see what conclusions may be drawn as to itsorigin.Mammalia.—<strong>The</strong> only mammals positivelyknown as indigenousto New Zealand are two bats, both peculiar to it,Scotophiliatuberculatus and Mystadna tuberculata. <strong>The</strong> former isallied to Australian forms ;<strong>the</strong> latter is more interesting, asbeing a peculiar genus <strong>of</strong> <strong>the</strong> family Noctilionidae, which doesnot exist in Australia ; and in having decided resemblances to<strong>the</strong> Phyllostomidse <strong>of</strong> South America, so that it may almost beconsidered to be a connecting link between <strong>the</strong> two families.forest rat is said to have once abounded on <strong>the</strong> islands, and tohave been used for food by <strong>the</strong> natives ; but <strong>the</strong>re is much doubtas to what it really was, and whe<strong>the</strong>r it was not an introducedspecies. <strong>The</strong> seals are wide-spread antarctic forms which haveno <strong>geographical</strong> significance.Birds.—About 145 species <strong>of</strong> birds are natives <strong>of</strong> New Zealand,<strong>of</strong> which 88 are waders or aquatics, leaving 57 land-birds belong-A


—:;chap, xni.] THE AUSTEALIAN REGION. 451ing to 34 genera. Of this latter number, 16, or nearly half xarepeculiar ; and <strong>the</strong>re are also 5 peculiar genera <strong>of</strong> waders andaquatic birds, making 21 in all. Of <strong>the</strong> remaining genera <strong>of</strong>land-birds, four are cosmopolite or <strong>of</strong> very wide range, while <strong>the</strong>remainder are characteristic <strong>of</strong> <strong>the</strong> Australian region. <strong>The</strong>following is a list <strong>of</strong> <strong>the</strong> Australian genera found in NewZealand : Sphenmacus, Gerygone, Orthonyx (Sylviidse) ; Graucalus(Campephagidse) ; Rhipidura (Muscicapidse) ; Anthochwra (Meliphagidae); Zosterops (Dicaeida?) ; Cyanoramphus (Platycercidse)Carpophaga (Columbidse) ; Hieracidea (Falconidse) ; Tribonyx(Rallidae). Besides <strong>the</strong>se <strong>the</strong>re are several genera <strong>of</strong> widerange, as follows :Anthus (Motacillidse) ; Hirundo (Hirundinidse); Chrysococcyx, Eadynamis (Cuculidse) ; Halcyon (Alcedinidae); Coturnix (Tetraonidse) ; Circus (Falconidae) ;A<strong>the</strong>ne(Strigidae).Most <strong>of</strong> <strong>the</strong> above genera are represented by peculiar NewZealand species, but in several cases <strong>the</strong> species are identical<strong>with</strong> those <strong>of</strong> Australia, as in <strong>the</strong> following : Anthochcera carunculata,Zosterops lateralis, Hirundo nigricans, and Chrysococcyxlucidus ;also oneWe now come toare <strong>of</strong> especialEudynamis taitensis—which isinterestPolynesian.<strong>the</strong> genera peculiar to New Zealand, whichList <strong>of</strong> Geneea <strong>of</strong> Birds Peculiar to New Zealand.Family and Genus.Sylviid^e.1. Myiomoira2. Miro


452 ZOOLOGICAL GEOGRAPHY. [PART III.Family and Genus.STURNID.E.10. Creadion11. Heterolocha ...12. Callseas


—chap, xiii.] THE AUSTRALIAN REGION. 453part <strong>of</strong> Polynesia, distances <strong>of</strong> more than 1,000 miles ! <strong>The</strong>sefacts seem, however, to have been accepted on insufficient evidenceand to be in <strong>the</strong>mselves extremely improbable. It is observedthat <strong>the</strong> cuckoos appear annually in certain districts and againdisappear ;but <strong>the</strong>ir course does not seem to have been traced,still less have <strong>the</strong>y ever been actually seen arriving or departingacross <strong>the</strong> ocean. In a country which has still such wide tracts<strong>of</strong> unsettled land, it is very possible that <strong>the</strong> birds in questionmay only move from one part <strong>of</strong> <strong>the</strong> islands to ano<strong>the</strong>r.Islets <strong>of</strong> <strong>the</strong> New Zealand Sub-region.We will here notice <strong>the</strong> smaller islands belonging to <strong>the</strong> subregion,as it ischiefly <strong>the</strong>ir birds that possess any interest.Norfolk Island.—<strong>The</strong> land-birds recorded from this islandamount to 15 species, <strong>of</strong> which 8 are Australian, viz. : Climacterisscandens, Symmorphus leucopygius, Zosterops tenuirostris andZ. albogularis, Halcyon sanctus, Platycercus pennanti, Garpophagaspadicea, Phapspicata and P. chalcoptera. Of <strong>the</strong> peculiarspecies three belong to Australian genera ; Petroica, Gerygone, andRhipidura ; one to a cosmopolitan genus, Turdus. So far <strong>the</strong>affinity seems to be all Australian, and <strong>the</strong>re remain only threebirds which ally this island to New Zealand,Gyanoramphus rayneri, and Notornis alba.Nestor productus,<strong>The</strong> former inhabited<strong>the</strong> small Phillip Island (close to Norfolk Island) but is now extinct.Being a typical New Zealand genus, quite incapable <strong>of</strong> flyingacross <strong>the</strong> sea, its presence necessitates some former connexionbetween <strong>the</strong> two islands, and it is <strong>the</strong>refore perhaps <strong>of</strong> moreweight than all <strong>the</strong> Australian genera and species, which are birdscapable <strong>of</strong> long flights. <strong>The</strong> Cyanoramphus is allied to a NewZealand broad-tailed parroquet. <strong>The</strong> Notornis alba is extinct,but two specimens exist in museums, and it is even a strongercase than <strong>the</strong> Nestor, as showing a former approximation or union<strong>of</strong> this island <strong>with</strong> New Zealand. A beautiful figure <strong>of</strong> thisbird is given in <strong>the</strong> Ibis for 1873.Lord Howe's Island.—This small island, situated half-waybetween Australia and Norfolk Island, is interesting, as containinga peculiar species <strong>of</strong> <strong>the</strong> New Zealand genus Ocydromus, or


—454 ZOOLOGICAL GEOGRAPHY. [part hi.wood-hen (0. sylvestris).<strong>The</strong>re is also a peculiar thrush, Turdusvinitinctus. Its o<strong>the</strong>r birds are wholly <strong>of</strong> Australian types, andmost <strong>of</strong> <strong>the</strong>m probably Australian species. <strong>The</strong> following havebeen observed, and no doubt constitute nearly its whole indigenousbird fauna.Acanthiza sp., Bhipidura sp., Pachycephala gutturalis,Zosterops strennuus and Z. tephropleurus, Strepera sp.,Halcyon sp., and Chalcophaga chrysochlora.<strong>The</strong> two species <strong>of</strong>Zosterops are peculiar. <strong>The</strong> Ocydromus is important enough toally this island to New Zealand ra<strong>the</strong>r than to Australia ;and if<strong>the</strong> white bird seen <strong>the</strong>re is, as supposed, <strong>the</strong> Notornis alba whichis extinct in Norfolk Island, <strong>the</strong> connection will be renderedstill more clear.Chatham Islands.—<strong>The</strong>se small islands, 450 miles east <strong>of</strong> NewZealand, possess about 40 species <strong>of</strong> birds, <strong>of</strong> which 13 are landbirds.All but one belong to New Zealand genera, and all butfive are New Zealand species. <strong>The</strong> following are <strong>the</strong> genera <strong>of</strong><strong>the</strong> land-birds : Sphenoeacus, Gerygone, Myiomoira, Bhipidura,Zosterops, Anthus, Pros<strong>the</strong>madera, Anthornis, Chrysococcyx, Cyanoramphus,Carpophaga, Circus. <strong>The</strong> peculiar species are Anthornismelanoccphala, Myiomoira, difenbachi and M. traversi,Bhipidura flabellifera, and a peculiar rail incapable <strong>of</strong> flight,named by Captain Hutton Cabalus modestus. It is stated that<strong>the</strong> Zosterops differs from that <strong>of</strong> New Zealand, and is also amigrant ; and it is <strong>the</strong>refore believed to come every year fromAustralia, passing over New Zealand, a distance <strong>of</strong> nearly 1,700miles ! Fur<strong>the</strong>rinvestigation will perhaps discover some o<strong>the</strong>rexplanation <strong>of</strong> <strong>the</strong> facts. It is also stated, that <strong>the</strong> pigeon andone <strong>of</strong> <strong>the</strong> small birds (? Gerygone or Zosterops) have arrived at<strong>the</strong> islands <strong>with</strong>in <strong>the</strong> last eight years. <strong>The</strong> natives fur<strong>the</strong>r declare,that both <strong>the</strong> Stringops and Apteryx once inhabited <strong>the</strong>islands, but were exterminated about <strong>the</strong> year 1835.<strong>The</strong> Auckland Islands.—<strong>The</strong>se are situated nearly 300 milessouth <strong>of</strong> New Zealand, and possess six land-birds, <strong>of</strong> which threeare peculiar,Anthus auchlandicus, Cyanoramphus aucklandicus,and C. malherbii, <strong>the</strong> o<strong>the</strong>rs being New Zealand species <strong>of</strong>Myiomoira, Pros<strong>the</strong>madera, and Anthornis. It is remarkablethat two peculiar parrots <strong>of</strong> <strong>the</strong> same genus should inhabit <strong>the</strong>se


;chap, xiii.] THE AUSTRALIAN REGION. 455small islands ; but such localities seem favourable to <strong>the</strong> Platycercidae,for ano<strong>the</strong>r peculiar species isfound in <strong>the</strong> remote MacquarieIslands, more than 400 miles far<strong>the</strong>r south. A peculiarspecies and genus <strong>of</strong> ducks, Nesonetta aucklandica, is also foundhere, and as far as yet known, nowhere else. A species <strong>of</strong><strong>the</strong> nor<strong>the</strong>rn genus Mergus isbeen recently obtained by Baron von Hiigel.also found on <strong>the</strong>se islands, and hasPlate XIII. Illustrating <strong>the</strong> peculiar Ornithology <strong>of</strong> New Zealand.—Ourartist has here depicted a group <strong>of</strong> <strong>the</strong> most remarkableand characteristic <strong>of</strong> <strong>the</strong> New Zealand birds.In <strong>the</strong> middleforeground is <strong>the</strong> Owl-parrot or Kakapoe (Stringops habroptilus),a nocturnal burrowing parrot, that feeds on fern-shoots, roots,berries, and occasionally lizards ; that climbs but does not flyand that has an owl-like mottled plumage and facial disc.<strong>The</strong> wings however are not rudimentary, but fully developed;and it seems to be only <strong>the</strong> muscles that have become uselessfor want <strong>of</strong> exercise. This would imply, that <strong>the</strong>se birds havenot long been inhabitants <strong>of</strong> New Zealand only, but were developedin o<strong>the</strong>r countries (perhaps Australia) where <strong>the</strong>ir wingswere <strong>of</strong> use to <strong>the</strong>m.Beyond <strong>the</strong> Kakapoe are a pair <strong>of</strong> <strong>the</strong> large rails, Notornismantelli; heavy birds <strong>with</strong> short wings quite useless forflight, and <strong>with</strong> massive feet and bill <strong>of</strong> a red colour. On<strong>the</strong> right is a pair <strong>of</strong> Kiwis {Apteryx australis), one <strong>of</strong> <strong>the</strong>queerest and most unbird-like <strong>of</strong> living birds.It has very smalland rudimentary wings, entirely concealed by <strong>the</strong> hair-likeplumage, and no tail. It is nocturnal, feeding chiefly on worms,which it extracts from s<strong>of</strong>t earth by means <strong>of</strong> its long bill. <strong>The</strong>genus Apteryx forms a distinct family <strong>of</strong> birds, <strong>of</strong> which fourspecies are now known, besides some which are extinct. <strong>The</strong>yare allied to <strong>the</strong> Cassowary and to <strong>the</strong> gigantic extinct Dinornis.On <strong>the</strong> wing are a pair <strong>of</strong> Crook-billed Plovers {Anarhynchusfrontalis), remarkable for being <strong>the</strong> only birds known whichhave <strong>the</strong> bill bent sideways. This was at first thought to be amalformation ;but it is now proved to be a constant character <strong>of</strong><strong>the</strong> species, as it exists even in <strong>the</strong> young chicks;yet <strong>the</strong> purposeserved by such an anomalous structure is not yet discovered.


406 ZOOLOGICAL GEOGRAPHY. [part hi,No country on <strong>the</strong> globe can <strong>of</strong>fer such an extraordinary set <strong>of</strong>birds as are here depicted.Reptiles.—<strong>The</strong>se consistno land-snakes and only one frog.almost wholly <strong>of</strong> lizards, <strong>the</strong>re beingTwelve species <strong>of</strong> lizards areknown, belonging to three genera, one <strong>of</strong> which is peculiar, asare all <strong>the</strong> species.Hinulia, <strong>with</strong> two species, and Mocoa, <strong>with</strong>four species (one <strong>of</strong> which extends to <strong>the</strong> Chatham Islands),belong to <strong>the</strong> Scincidse ; both are very wide-spread genera andoccur in Australia. <strong>The</strong> peculiar genus Naultinns, <strong>with</strong> six species,belongs to <strong>the</strong> Geckotidse, a family spread over <strong>the</strong> whole world.<strong>The</strong> most extraordinary and interesting reptile <strong>of</strong> New Zealandis, however, <strong>the</strong> Hatteria punctata, a lizard-like animalliving in holes, and found in small islands on <strong>the</strong> north-eastcoast, and more rarely on <strong>the</strong> main land.It is somewhat intermediatein structure between lizards and crocodiles, and also hasbird-like characters in <strong>the</strong> form <strong>of</strong>. its ribs. It constitutes, notonly a distinct family, Rhyncocephalidse, but a separate order <strong>of</strong>reptiles, Khyncocephalina. It is quite isolated from all o<strong>the</strong>rmembers <strong>of</strong> <strong>the</strong> class ; and is probably a slightly modified representative<strong>of</strong> an ancient and generalised form, which has beensuperseded in larger areas by <strong>the</strong> more specialized lizards andsaurians.<strong>The</strong> only representatives <strong>of</strong> <strong>the</strong> Ophidia are two sea-snakes<strong>of</strong> Australian and Polynesian species, and <strong>of</strong> no <strong>geographical</strong>interest.Amphibia.—<strong>The</strong> solitary frog indigenous to New Zealand,belongs to a peculiar genus, Liopelma, and to <strong>the</strong> family Bomburatoridse,o<strong>the</strong>rwise confined to Europe and temperate SouthAmerica.Fresh-water Fishes.—<strong>The</strong>re are, according to Captain Hutton,15 species <strong>of</strong> fresh-water fish in New Zealand, belonging to 7genera ; six species, and one genus (Retropinna), being peculiar.Betropinna richardsoni belongs to <strong>the</strong> Salmonidae, and is <strong>the</strong>only example <strong>of</strong> that family occurring in <strong>the</strong> Sou<strong>the</strong>rn hemisphere,where it is confined to New Zealand and <strong>the</strong> ChathamIslands.<strong>The</strong> wide <strong>distribution</strong> <strong>of</strong> Galaxias attennatus—from <strong>the</strong>


chap, xin.] THE AUSTRALIAN REGION. 457Chatham Islands to South America—has already been noticed \while ano<strong>the</strong>r species,G. fasciatus, is found in <strong>the</strong> Chatham andAuckland Isles as well as New Zealand. A second genuspeculiar to New Zealand, Neochanna, allied to Galaxias, hasrecently been described. Prototroctes oxyrhynchus is allied to anAustralian species, but belongs to a family (Haplochitonidse)which is o<strong>the</strong>rwise South American. An eel, Anguilla latirostris,is found in Europe, China, and <strong>the</strong> West Indies, as well as inNew Zealand ! while <strong>the</strong> genus Agonostoma ranges to Australia,Celebes, Mauritius, and Central America.Insects.—<strong>The</strong> great poverty <strong>of</strong> this class iswell shown by <strong>the</strong>fact, that only eleven species <strong>of</strong> butterflies are known to inhabitNew Zealand. Of <strong>the</strong>se, six are peculiar, and one, Argyrophenga(Satyridse), is a peculiar genus allied to <strong>the</strong> Nor<strong>the</strong>rn genusErebia. <strong>The</strong> rest are ei<strong>the</strong>r <strong>of</strong> wide range, as Pyrameis carduiand Diadema bolina ; or Australian, as Hamdyaas zoilus ;whileone, Danais erippus, is American, but has also occurred in Australia,and is no doubt a recent introduction into both countries.Only one Sphinx is recorded, and no o<strong>the</strong>r species <strong>of</strong> <strong>the</strong> Sphinginaexcept <strong>the</strong> British currant-moth, JEgeria tipuliformis,doubtless imported. Coleoptera are better represented, nearly300 species having been described, all or nearly all being peculiar.<strong>The</strong>se belong to about 150 genera, <strong>of</strong> which more than 50are peculiar. No less than 14 peculiar genera belong to <strong>the</strong>Carabidse,mostly consisting <strong>of</strong> one or two species, but Demetridahas 3, and Metaglymma 8 species. O<strong>the</strong>r important genera areDicrochile, Homalosoma, Mecodema, and Scopodes, all in common<strong>with</strong> Australia. Mecodema and Metaglymma are <strong>the</strong> largestgenera. Even <strong>the</strong> Auckland Islands have two small genera<strong>of</strong> Carabidse found nowhere else.Cicindelidse are represented in New Zealand by 6 species <strong>of</strong>Cicindela, and 1 <strong>of</strong> Dystiysidera, a genus peculiar to <strong>the</strong> Australianregion.<strong>The</strong> Lucanidse are represented by two peculiar genera, Dendrdblaxand Oxyomus ; two Australian genera, Lissotes andCeratognathus ;and by <strong>the</strong> almost cosmopolite Dorcus.<strong>The</strong> Scarabeidse consist <strong>of</strong> ten species only, belonging to fourVol. I.—31


458 ZOOLOGICAL GEOGRAPHY. [part hi.genera, two <strong>of</strong> which are peculiar (Odontria and Stethaspis) ;andtwo Australian (Pericoptus and Calonota). <strong>The</strong>re are noCetoniidse.<strong>The</strong>re is only one Buprestid, belonging to <strong>the</strong> Australian genusCisseis. <strong>The</strong> Elateridae, (about a dozen species,) belong mostlyto Australian genera, but two, Metablax and Ochosternus, arepeculiar.<strong>The</strong>re are 30 species <strong>of</strong> Curculionidse, belonging to 22 genera.Of <strong>the</strong> genera, 12 are peculiar ; 1 is common to New Zealandand New Caledonia ;restare widely distributed.Longicorns are, next to5 belong to <strong>the</strong> Australian region, and <strong>the</strong>Carabidse, <strong>the</strong> most numerous family,<strong>the</strong>re being, according to Mr. Bates (Ann. Nat. Hist., 1874), about35 genera, <strong>of</strong> which 26 are peculiar or highly characteristic,and 7 <strong>of</strong> <strong>the</strong> o<strong>the</strong>rs Australian.<strong>The</strong> largest and most characteristicgenera are JEmona and Xyloteles, both being peculiar toNew Zealand ; few <strong>of</strong> <strong>the</strong> remainder having more than one ortwo species. Demonax extends to <strong>the</strong> Moluccas and S. E.Asia.A dozen <strong>of</strong> <strong>the</strong> genera have no near <strong>relations</strong> <strong>with</strong> those<strong>of</strong> any o<strong>the</strong>r country.Phytophaga are remarkably scarce, only two species <strong>of</strong>Colaspis being recorded ; and <strong>the</strong>re is only a single species <strong>of</strong>Coccinella.<strong>The</strong> o<strong>the</strong>r orders <strong>of</strong> Insects appear to be equally deficient.Hymenoptera are veiy poorly represented, only a score <strong>of</strong> speciesbeing yet known ; but two <strong>of</strong> <strong>the</strong> genera are peculiar, as are all<strong>the</strong> species. <strong>The</strong> Neuroptera and Heteroptera are also veryscarce, and several <strong>of</strong> <strong>the</strong> species are wide-spread forms <strong>of</strong> <strong>the</strong>Australian region. <strong>The</strong> few species <strong>of</strong> Homoptera are allpeculiar. <strong>The</strong> Myriapoda afford some interesting facts. <strong>The</strong>reare nine or ten species, all peculiar. One genus, Lithobius,ranges over <strong>the</strong> nor<strong>the</strong>rn hemisphere as far south as Singapore,and probably through <strong>the</strong> Malay Archipelago, but is not foundin Australia. Henicops occurs elsewhere only in Tasmaniaand Chili. Cryptops, only in <strong>the</strong> north temperate zone ; whiletwo o<strong>the</strong>rs, Cermatia and Cormocephalus, both occur inAustralia.


—chap, xiii.] THE AUSTRALIAN REGION. 459Land-Shells.—Of <strong>the</strong>se, 114 species are known, 97 beingpeculiar. Three species <strong>of</strong> Helix are also found in Australia,and five more in various tropical islands <strong>of</strong> <strong>the</strong> Pacific. Nanina,Lymncea, and Assiminea, are found in Polynesia or Malaya,but not in Australia. Amphibola is an Australian genus, as isJanella. Testacella and Limax belong to <strong>the</strong> Palsearctic region.From <strong>the</strong> Chatham Islands, 82 species <strong>of</strong> shells are known,all being New Zealand species, except nine, which arepeculiar.Tlie Ancient Fauna <strong>of</strong> New Zealand.—One <strong>of</strong> <strong>the</strong> most remarkablefeatures <strong>of</strong> <strong>the</strong> New Zealand fauna, is <strong>the</strong> existence,till quite recent times, <strong>of</strong> an extensive group <strong>of</strong> wingless birds,—calledMoas by <strong>the</strong> natives—many <strong>of</strong> <strong>the</strong>m <strong>of</strong> gigantic size,and which evidently occupied <strong>the</strong> place which, in o<strong>the</strong>r countries,is filled by <strong>the</strong> mammalia. <strong>The</strong> most recent account <strong>of</strong> <strong>the</strong>sesingular remains, is that by Dr. Haast, who, from a <strong>study</strong> <strong>of</strong><strong>the</strong> extensive series <strong>of</strong> specimens in <strong>the</strong> Canterbury museum,believes, that <strong>the</strong>y belong to two families, distinguished byimportant differences <strong>of</strong> structure,and constitute four genera,Dinornis and Miornis, forming <strong>the</strong> family Dinornithidse;Palapteryx and Euryapteryx, forming <strong>the</strong> family Palapterygidae.<strong>The</strong>se were mostly larger birds than <strong>the</strong> living Apteryx, andsome <strong>of</strong> <strong>the</strong>m much larger even than <strong>the</strong> African ostrich, andwere more allied to <strong>the</strong> Casuariidse and Struthionidae than to<strong>the</strong> Apterygidse. No less than eleven species <strong>of</strong> <strong>the</strong>se birdshave been discovered; all are <strong>of</strong> recent geological date, and<strong>the</strong>re are indications that some <strong>of</strong> <strong>the</strong>m may have been inexistence less than a century ago, and were really exterminatedby man. Eemains have been found (<strong>of</strong> apparently<strong>the</strong> same recent date) <strong>of</strong> species <strong>of</strong> Apteryx, Stringops, Ocydromus,and many o<strong>the</strong>r living forms, as well as <strong>of</strong> Harpagornis,a large bird <strong>of</strong> prey, and Gnemiornis, a gigantic goose. Bodies<strong>of</strong> <strong>the</strong> Hatteria 'punctata have also been found along <strong>with</strong> those<strong>of</strong> <strong>the</strong> Moa, showing that this remarkable reptile was once moreabundant on <strong>the</strong> main islands than it is now.TJie Origin <strong>of</strong> <strong>the</strong> New Zealand Fauna.—Having now given


460 ZOOLOGICAL GEOGRAPHY. [part hi.an outline sketch <strong>of</strong> <strong>the</strong> main features <strong>of</strong> <strong>the</strong> New Zealandfauna and <strong>of</strong> its <strong>relations</strong> <strong>with</strong> o<strong>the</strong>r regions, we may considerwhat conclusions are fairly deducible from <strong>the</strong> facts. As <strong>the</strong>outlying Norfolk, Chatham, and Lord Howe's Islands, are allinhabited (or have recently been so) by birds <strong>of</strong> New Zealandtype or even identical species, almost incapable <strong>of</strong> flight, we mayinfer that <strong>the</strong>se islands show us <strong>the</strong> former minimum extent <strong>of</strong> <strong>the</strong>land-area in which <strong>the</strong> peculiar forms which characterise <strong>the</strong>sub-region were developed. If we include <strong>the</strong> Auckland andMacquarie Islands to <strong>the</strong> south, we shall have a territory <strong>of</strong> notmuch less extent than Australia, and separated from it by perhapsseveral hundred miles <strong>of</strong> ocean. Some such ancient landmust have existed to allow <strong>of</strong> <strong>the</strong> development and specialization<strong>of</strong> so many peculiar forms <strong>of</strong> birds, and it probably remained<strong>with</strong> but 'slight modifications for a considerable geological period.During all this time it would interchange many <strong>of</strong> its forms <strong>of</strong>life <strong>with</strong> Australia, and <strong>the</strong>re would arise that amount <strong>of</strong> identity<strong>of</strong> genera between <strong>the</strong> two countries winch we find to exist.extension southwards, perhaps considerably beyond <strong>the</strong> Macquaries,would bring it <strong>with</strong>in <strong>the</strong> range <strong>of</strong> floating ice duringcolder epochs, and <strong>with</strong>in easy reach <strong>of</strong> <strong>the</strong> antarctic continentduring <strong>the</strong> warm periods ;<strong>of</strong> genera and species <strong>with</strong>Itsand thus would arise that interchangeSouth America, which forms one <strong>of</strong><strong>the</strong> characteristic features <strong>of</strong> <strong>the</strong> natural history <strong>of</strong> New Zealand.Captain F. W. Hutton (to whose interesting paper on <strong>the</strong>Geographical <strong>relations</strong> <strong>of</strong> <strong>the</strong> New Zealand Fauna we areindebted for some <strong>of</strong> our facts) insists upon <strong>the</strong> necessity <strong>of</strong>former land-connections in various directions, and especially<strong>of</strong> an early sou<strong>the</strong>rn continental period, when New Zealand,Australia, Sou<strong>the</strong>rn Africa, and South America, were united.Thus he would account for <strong>the</strong> existence <strong>of</strong> Struthious birdsin all <strong>the</strong>se countries, and for <strong>the</strong> various o<strong>the</strong>r groups <strong>of</strong>birds, reptiles, fishes, or insects which have no obvious means<strong>of</strong> traversing <strong>the</strong> ocean,—and this union must have occurredbefore mammalia existed in any <strong>of</strong> <strong>the</strong>se countries. Butsuch a supposition is quite unnecessary, if we consider thatall wingless land-birds and some water-birds (as <strong>the</strong> Gare-fowl


chap, xni.] THE AUSTRALIAN REGION. 461and Steamer Duck) are probably cases <strong>of</strong> abortion <strong>of</strong> uselessorgans, and that <strong>the</strong> common ancestors <strong>of</strong> <strong>the</strong> variousforms <strong>of</strong> Struthiones may have been capable <strong>of</strong> a moderatedegree <strong>of</strong> flight ; or <strong>the</strong>y may have originated in <strong>the</strong> nor<strong>the</strong>rnhemisphere, as already explained in Chap. XI. p. 287.<strong>The</strong> existence<strong>of</strong> two, if not three, distinct families <strong>of</strong> <strong>the</strong>se birds in NewZealand, proves that <strong>the</strong> original type was here isolated at avery early date, and being wholly free from <strong>the</strong> competition <strong>of</strong>mammalia, became more differentiated than elsewhere. <strong>The</strong>Hatteria is probably coeval <strong>with</strong> <strong>the</strong>se early forms, and is <strong>the</strong>only relic <strong>of</strong> a whole order <strong>of</strong> reptiles, which once perhapsranged far over <strong>the</strong> globe.Still less does any o<strong>the</strong>r form <strong>of</strong> animal inhabiting New Zealand,require a land connection <strong>with</strong> distant countries to accountfor its presence. With <strong>the</strong> example before us <strong>of</strong> <strong>the</strong> Bermudasand Azores, to which a great variety <strong>of</strong> birds fly annually over vastdistances, and even <strong>of</strong> <strong>the</strong> recent arrival <strong>of</strong> new birds in NewZealand and Chatham Island, we may be sure that <strong>the</strong> ancestors<strong>of</strong> every New Zealand bird could easily have reached itsshoresduring <strong>the</strong> countless ages which elapsed while <strong>the</strong> Dinornis andApteryx were developing. <strong>The</strong> wonderful range <strong>of</strong> some <strong>of</strong> <strong>the</strong>existing species <strong>of</strong> lizards and fresh-water fish, asalready given,proves that <strong>the</strong>y too possess means <strong>of</strong> dispersal which havesufficed to spread <strong>the</strong>m, <strong>with</strong>in a comparatively recent period,over countries separated by thousands <strong>of</strong> miles <strong>of</strong> ocean; and <strong>the</strong>fact that a group like <strong>the</strong> snakes, so widely distributed and forwhich <strong>the</strong> climate <strong>of</strong> New Zealand is so well adapted, does notexist <strong>the</strong>re, is an additional pro<strong>of</strong> that land connection had nothingto do <strong>with</strong> <strong>the</strong> introduction <strong>of</strong> <strong>the</strong> existing fauna. We havealready (p. 398), discussed in some detail <strong>the</strong> various modes inwhich <strong>the</strong> dispersal <strong>of</strong> <strong>animals</strong> in <strong>the</strong> sou<strong>the</strong>rn hemisphere hasbeen effected ;and in accordance <strong>with</strong> <strong>the</strong> principles <strong>the</strong>re established,we conclude, that <strong>the</strong> New Zealand fauna, living andextinct, demonstrates <strong>the</strong> existence <strong>of</strong> an extensive tract<strong>of</strong> landin <strong>the</strong> vicinity <strong>of</strong> Australia, Polynesia, and <strong>the</strong> Antarctic continent,<strong>with</strong>out having been once actually connected <strong>with</strong> ei<strong>the</strong>r<strong>of</strong> <strong>the</strong>se countries, since <strong>the</strong> period when mammalia had peopled


462 ZOOLOGICAL GEOGRAPHY. [part in.all <strong>the</strong> great continents. That event certainly dates back toSecondary, if not to Palaeozoic, times, because so dominant agroup must soon have spread over <strong>the</strong> whole continuous landarea<strong>of</strong> <strong>the</strong> globe. We have no reason for believing that birdswere an earlier development ; and certainly cannot, <strong>with</strong> anyprobability, place <strong>the</strong> origin <strong>of</strong> <strong>the</strong> Struthiones before that <strong>of</strong>Mammals.Causes <strong>of</strong> <strong>the</strong> Poverty <strong>of</strong> Insect-life in New Zealand : its Influenceon <strong>the</strong> Character <strong>of</strong> <strong>the</strong> Flora.—<strong>The</strong> extreme paucity <strong>of</strong> insectsin New Zealand, to which we have already alluded, seemsto call for some attempt at explanation. No o<strong>the</strong>r country in <strong>the</strong>•world, in which <strong>the</strong> conditions are equally favourable for insectlife,and which has ei<strong>the</strong>r been connected <strong>with</strong>, or is in proximityto, any <strong>of</strong> <strong>the</strong> large masses <strong>of</strong> land, presents a similar phenomenon.<strong>The</strong> only approach to it is in <strong>the</strong> Galapagos, and insome <strong>of</strong> <strong>the</strong> islands <strong>of</strong> <strong>the</strong> Pacific ; and in each <strong>of</strong> <strong>the</strong>se cases <strong>the</strong>absence <strong>of</strong> mammals leads us to infer, that no connection <strong>with</strong> acontinent has ever taken place. Yet <strong>the</strong> fauna <strong>of</strong> New Zealandevidently dates back to a remote geological epoch, and it seemsstrange that an abundance <strong>of</strong> indigenous insects have not beendeveloped, especially when we consider <strong>the</strong> vastantiquity thatmost <strong>of</strong> <strong>the</strong> orders and families, and many <strong>of</strong> <strong>the</strong> genera, <strong>of</strong> insectspossess (see p. 166), and that <strong>the</strong>y must always have reached <strong>the</strong>country in greater numbers and variety than any <strong>of</strong> <strong>the</strong> higher<strong>animals</strong>. <strong>The</strong> undoubted fact that such an indigenous insectfaunahas not arisen, would <strong>the</strong>refore lead us to conclude, thatinsects find <strong>the</strong> conditions requisite for <strong>the</strong>ir development only in<strong>the</strong> great continental masses <strong>of</strong> land, instrict adaptation to, anddependance on, a varied fauna and flora <strong>of</strong> ever-increasing richnessand complexity. A small number <strong>of</strong> widely-separated forms, introducedinto a country where <strong>the</strong> fauna and flora are alike scantyand unrelated to <strong>the</strong>m, seem to have little tendency to varyand branch out into that vast network <strong>of</strong> insect-life whichenriches all <strong>the</strong> great continents and <strong>the</strong>ir once connectedislands.It is a striking confirmation on a large scale,<strong>of</strong> Mr. Darwin'sbeautiful <strong>the</strong>ory—that <strong>the</strong> gay colours <strong>of</strong> flowers have mostly, or


—chap, xiii.] THE AUSTRALIAN REGION. 4G3perhaps, wholly been produced, in order to attract insects which aidin <strong>the</strong>ir fertilization—that in New Zealand, where insectsare sostrikingly deficient in variety, <strong>the</strong> flora should be almost as strikinglydeficient in gaily-coloured blossoms. Of course <strong>the</strong>re are someexceptions, butas a whole, green,. inconspicuous, and imperfectflowers prevail, to an extent not to be equalled in any o<strong>the</strong>r part<strong>of</strong> <strong>the</strong> globe ;and affording a marvellous contrast to <strong>the</strong> generalbrilliancy <strong>of</strong> Australian flowers, combined <strong>with</strong> <strong>the</strong> abundanceand variety <strong>of</strong> its insect-life. We must remember, too, that <strong>the</strong>few gay or conspicuous flowering-plants possessedby New Zealand,are almost all <strong>of</strong> Australian, South American, orEuropeangenera; <strong>the</strong> peculiar New Zealand or Antarctic genera beingalmost wholly <strong>with</strong>out conspicuous flowers. In <strong>the</strong> tropicalGalapagos <strong>the</strong> same thing occurs. Mr. Darwin notices <strong>the</strong>wretched weedy appearance <strong>of</strong> <strong>the</strong> vegetation ; and states thatit was some time before he discovered that most <strong>of</strong> <strong>the</strong> plantswere in flower at <strong>the</strong> time <strong>of</strong> his visit ! And <strong>the</strong> insect-life wascorrespondingly deficient, consisting mainly <strong>of</strong> a few terrestrialbeetles.<strong>The</strong> poverty <strong>of</strong> insect-life in New Zealand must, <strong>the</strong>refore, bea very ancient feature <strong>of</strong> <strong>the</strong> country ; and it furnishes an additionalargument against <strong>the</strong> <strong>the</strong>ory <strong>of</strong> land-connection <strong>with</strong>, oreven any near approach to, ei<strong>the</strong>r Australia, South Africa, orSouth America. For in that case numbers <strong>of</strong> winged insectswould certainly have entered, and <strong>the</strong>flowers would <strong>the</strong>n, as inevery o<strong>the</strong>r part <strong>of</strong> <strong>the</strong> world, have been rendered attractive to<strong>the</strong>m by <strong>the</strong> development <strong>of</strong> coloured petals ; and this characteronce acquired would long maintain itself, even if <strong>the</strong> insects had,from some unknown cause, subsequently disappeared.After <strong>the</strong> preceding paragraphs were written, it occurred to me,that if this reasoning were correct, New Zealand plants ought tobe also deficient in scented flowers ; because it is a part <strong>of</strong> <strong>the</strong>same <strong>the</strong>ory, that <strong>the</strong> odours <strong>of</strong> flowers have, like <strong>the</strong>ir colours,been developed to attract <strong>the</strong> insects required to aid in <strong>the</strong>ir fertilization.I <strong>the</strong>refore at once applied to my friend Dr. Hooker,as <strong>the</strong> highest authority on New Zealand botany ;simply askingwhe<strong>the</strong>r <strong>the</strong>re was any such observed deficiency. His reply was:


464 ZOOLOGICAL GEOGRAPHY. [part hi" New Zealand plants are remarkably scentless, both in regard to<strong>the</strong> rarity <strong>of</strong> scented flowers, <strong>of</strong> leaves <strong>with</strong> immersed glandscontaining essential oils, and <strong>of</strong> glandular hairs." <strong>The</strong>re are afew exceptional cases, but <strong>the</strong>se seem even more rare than mightbe expected, so that <strong>the</strong> confirmation <strong>of</strong> <strong>the</strong>ory is very complete.<strong>The</strong> circumstance that aromatic leaves are also very scarce, suggests<strong>the</strong> idea that <strong>the</strong>se, too, serve as an attraction to insects.Aromatic plants abound most in arid countries, and on Alpineheights ; both localities where winged insects are comparativelyscarce, and where it may be necessary to attract <strong>the</strong>m in everypossible way.Dr. Hooker also informs, me that since his Introductionto <strong>the</strong> New Zealand Flora was written, many plants <strong>with</strong>handsome flowers have been discovered, especially among <strong>the</strong>Raoiunculi, shrubby Veronicas, and herbaceous Composite. <strong>The</strong>two former, however, are genera <strong>of</strong> wide range, which may haveoriginated in New Zealand by <strong>the</strong> introduction <strong>of</strong> plants <strong>with</strong>handsome flowers, which <strong>the</strong> few indigenous insects would beattracted by, and thus prevent <strong>the</strong> loss <strong>of</strong> <strong>the</strong>ir gay corollas ; sothat <strong>the</strong>se discoveries will not much affect <strong>the</strong> general character<strong>of</strong> <strong>the</strong> flora, and its very curious bearing on <strong>the</strong> past history <strong>of</strong><strong>the</strong> islands through <strong>the</strong> <strong>relations</strong> <strong>of</strong> flowers and insects.In judging <strong>of</strong> <strong>the</strong> relationhere supposed to exist, it must beremembered, that if <strong>the</strong> New Zealand insects have been introducedfrom <strong>the</strong> surrounding countries by chance immigrations atdistant intervals, <strong>the</strong>n, as we go back into <strong>the</strong> past <strong>the</strong> insectfauna will become poorer and poorer, and still more inadequatethan at present to lead to <strong>the</strong> development <strong>of</strong> attractive flowersand odours.This quite harmonizes <strong>with</strong> <strong>the</strong> fact <strong>of</strong> <strong>the</strong> ancientindigenous flora being so remarkably scentless and inconspicuous,while a few <strong>of</strong> <strong>the</strong> more recently introduced genera <strong>of</strong>plants have retained <strong>the</strong>ir floral attractions.Concluding Remarks on <strong>the</strong> Early History <strong>of</strong> <strong>the</strong> AustralianRegion.We have already discussed in some detail, <strong>the</strong> various <strong>relations</strong><strong>of</strong> <strong>the</strong> Australian sub-regions to <strong>the</strong> surrounding Eegions, and <strong>the</strong><strong>geographical</strong> changes that appear to have taken place. A very


chap, xiu.] THE AUSTRALIAN REGION. 465few observations will <strong>the</strong>refore suffice, on <strong>the</strong> supposed earlyhistory <strong>of</strong> <strong>the</strong> Australian region as a whole.It was probably far back in <strong>the</strong> Secondary period, that someportion <strong>of</strong> <strong>the</strong> Australian region was in actual connection <strong>with</strong><strong>the</strong> nor<strong>the</strong>rn continent, and became stocked <strong>with</strong> ancestral forms<strong>of</strong> Marsupials ; but from that time till now <strong>the</strong>re seems to havebeen no fur<strong>the</strong>r land connection, and <strong>the</strong> Australian lands have<strong>the</strong>nceforward gone on developing <strong>the</strong> Marsupial and Monotrematetypes, into <strong>the</strong> various living and extinct races we now find<strong>the</strong>re. During some portion <strong>of</strong> <strong>the</strong> Tertiary epoch Australia probablycomprised much <strong>of</strong> its existing area, toge<strong>the</strong>r <strong>with</strong> Papuaand <strong>the</strong> Solomon Islands, and perhaps extended as far east as <strong>the</strong>Fiji Islands ; while it might also have had a considerable extensionto <strong>the</strong> south and west.Some light has recently been thrownon this subject by Pr<strong>of</strong>essor McCoy's researches on <strong>the</strong> Palaeontology<strong>of</strong> Victoria. He finds abundant marine fossils <strong>of</strong>Eocene and Miocene age, many <strong>of</strong> which are strikingly similarto those <strong>of</strong> Europe at <strong>the</strong> same period. Among <strong>the</strong>se are Cetaceans<strong>of</strong> <strong>the</strong> genus Squaloclon ;European species <strong>of</strong> Plagiostomousfishes ; mollusea and corals closely resembling those <strong>of</strong>Europe and North America <strong>of</strong> <strong>the</strong> same age,—such as numerousVolutes closely allied to those <strong>of</strong> <strong>the</strong> Eocene beds <strong>of</strong> <strong>the</strong> Isle<strong>of</strong>Wight, and <strong>the</strong> genus Dentalium in great abundance, almost orquite identical <strong>with</strong> European tertiary species. Along <strong>with</strong><strong>the</strong>se, are found some living species, but always such as nowlive far<strong>the</strong>r north in tropical seas. <strong>The</strong> Cretaceous and Mesozoicmarine fossils are equally close to those <strong>of</strong> Europe.<strong>The</strong> whole <strong>of</strong> <strong>the</strong>se remains demonstrate that, as in <strong>the</strong>nor<strong>the</strong>rn so in <strong>the</strong> sou<strong>the</strong>rn hemisphere, a much warmer climateprevailed in <strong>the</strong> Eocene and Miocene periods than at <strong>the</strong> presenttime. This is a most important result, and one which stronglysupports Mr. Belt's view, before referred to, that <strong>the</strong> warmerclimates in past geological epochs, and especially that <strong>of</strong> <strong>the</strong>Miocene as compared <strong>with</strong> our own, was caused by a diminution <strong>of</strong><strong>the</strong> obliquity <strong>of</strong> <strong>the</strong> ecliptic, leading to a much greater uniformity<strong>of</strong> <strong>the</strong> seasons for a considerable distance from <strong>the</strong> equator, andgreatly reducing <strong>the</strong> polar area <strong>with</strong>in which <strong>the</strong> sun would ever


4GG ZOOLOGICAL GEOGRAPHY. [part hi.disappear during an entire rotation <strong>of</strong> <strong>the</strong> earth. During sucha period, tropical forms <strong>of</strong> marine <strong>animals</strong> would have been ableto spread north and south, into what are now cool latitudes ;andidentical genera, and even species, might <strong>the</strong>n have ranged along<strong>the</strong> sou<strong>the</strong>rn shores <strong>of</strong> <strong>the</strong> old Palsearctic continent, fromBritainto <strong>the</strong> Bay <strong>of</strong> Bengal, and southward along <strong>the</strong> Malayan coaststo Australia.Numerous Miocene plant-beds have alsobeen found in Victoria,saidcontaining abundance <strong>of</strong> Dicotyledonous leaves, which aregenerally to resemble those <strong>of</strong> <strong>the</strong> Asiatic flora, and <strong>of</strong> <strong>the</strong>Miocene plant-beds <strong>of</strong> <strong>the</strong> Khine.It is to be hoped <strong>the</strong>se bedswill be more closely examined for remains <strong>of</strong> insects, land-shells,and vertebrates, and that <strong>the</strong> plants will be carefully preserved andcritically studied ; for here probably lies hidden <strong>the</strong> key, thatwill solve much <strong>of</strong> <strong>the</strong> mystery that<strong>of</strong> <strong>the</strong> Australian fauna.attaches to <strong>the</strong> past history


chap, xiii.] THE AUSTRALIAN REGION. 467TABLES OF DISTRIBUTIONIn drawing up <strong>the</strong>se tables, showing <strong>the</strong> <strong>distribution</strong> <strong>of</strong> <strong>the</strong>various classes <strong>of</strong> <strong>animals</strong> in <strong>the</strong> Australian region, <strong>the</strong> followingsources <strong>of</strong> information have been relied on, in addition to <strong>the</strong>general treatises, monographs, and catalogues used in compiling<strong>the</strong> 4th Part <strong>of</strong> this work.Mammalia.—Gould, Mammals <strong>of</strong> Australia ;Waterhouse onMarsupials ;Dr. J. E. Gray's List <strong>of</strong> Mammalia <strong>of</strong> New Guinea;Midler, Temminck and Schlegel on Mammals <strong>of</strong> <strong>the</strong> Moluccas;papers by Dr. Gray ;and personal observation by <strong>the</strong> Author.JKrtfo.—Gould's Birds <strong>of</strong> Australia; Buller's Birds <strong>of</strong> NewZealand ; G. R. Gray's Lists <strong>of</strong> Birds <strong>of</strong> Moluccas, &c. ;Hartlauband Finsch on Birds <strong>of</strong> Pacific Islands ;Sclater on Birds<strong>of</strong> Sandwich Islands ;papers by Haast, Hutton, Meyer, Salvin,Schlegel, Sclater, Travers, Lord Walden and <strong>the</strong> Author.Reptiles.— Krefft, Catalogue <strong>of</strong> Snakes ; Gun<strong>the</strong>r, List <strong>of</strong>Lizards in Voyage <strong>of</strong> Erebus and Terror (1875) ; and numerouspapers.


.468 ZOOLOGICAL GEOGRAPHY. [part III.TABLE I.FAMILIES OF ANIMALS INHABITING THE AUSTRALIAN REGION.Explanation.Names in italics show families which are peculiar to <strong>the</strong> region.Names inclosed thus ( ) show families which only just enter <strong>the</strong> region, and arenot considered properly to belong to it.Numbers correspond to <strong>the</strong> series <strong>of</strong> numbers to <strong>the</strong> families in Part IV.Sub-regions.Order and Family.©


...CHAP. XIII.] THE AUSTRALIAN REGION. 460Sub-regions.Order and Family.x .23~-Range beyond <strong>the</strong> Region.81. Phalangistidce82. PhascolomyidceMONOTREMATA83. Ornithorhynchidce..84. EchidnidceBIRDS.Passeres.1. Turdidae2. Sylviidse3. Timaliidae ...5. Cinclidae8. Certhiidae9. Sittidae10. Paridce13. Pycnonotidse . .14. Oriolidae15. Campephagidae16. Dicruridae17. Muscicapidae...18. Pachycephalidce19. Laniidai20. Corridae21. Paradiseidce ...22. M'eli-phagidce .23. Nectariniidae24. Dicaeidae2 5. Drepanididce .30. Hirundinidae .34. Ploceidae35. Sturnidae36. Artamida?37. Alaudidae.38. Motacillidee ...47. Pittidae49. Menuridce50. Atrichiidce ...CosmopoliteCosmopoliteOriental familyOriental familyOriental and EthiopianOriental and EthiopianOriental and Ethiopian<strong>The</strong> Old WorldAlmost peculiar to region<strong>The</strong> Old WorldCosmopoliteOriental and EthiopianOriental and EthiopianCosmopoliteOriental. Ethiopian<strong>The</strong> Old' WorldOriental<strong>The</strong> Old World and N. America<strong>The</strong> Old WorldOriental, EthiopianPeculiar to AustraliaPeculiar to AustraliaPlCARI^!.51. Picidse58. Cnculidae62. Coraciidae63. Meropidae67. Alcedinidae ...68. Bucerotidae ...71. Podargidae ...73. Capriroulgidas74. CypselidaeAll o<strong>the</strong>r regionsCosmopoliteOriental and EthiopianOriental and EthiopianCosmopoliteOriental and EthiopianOrientalCosmopoliteCosmopolite


470 ZOOLOGICAL GEOGRAPHY. [part hi.


iCHAP. XIII.] THE AUSTRALIAN REGION. 471Sub-regions.Order and Family.en i—3 —


.472 ZOOLOGICAL GEOGRAPHY. [PART 111Sub-regions.Order and Family.OS 3 —3 Range beyond <strong>the</strong> Region.14. Alytidae15. Pelodryadae ...16. Hylidae17. Polypedatidae18. Ranidae19. DiseoglossidaeFISHES (FRESH-WATER).ACANTHOPTERYGII.11. Trachinidae ...35. Labyrinthici ...37. A<strong>the</strong>rinidae ...38. Mugillidae ...Anacanthini.53. Gadopsidce ...Physostomi.59. Siluridse61. Haplochitonidse65. Salmonidae ...67. Galaxidae78. Ostegolossidae85. (Symbranchidae)Dipnoi.92. SirenoideiINSECTS. LEPI-DOPTERA (PART).Durini (Butterflies).1. Danaidse2. Satyridae3. Elymniidae ...4. Morphidae6. Acraeidae8. Nymphalidae . .9. Liby<strong>the</strong>idae ...10. Nemeobeidas ..13. Lycaenidae14. Pieridae15. Papilionidae ...16. HesperidaeSphingidea.17. Zygaenidae18. Castniidae19. Agaristidae ...20. Uraniidae23. Sphingidae ...All regions but OrientalNeotropicalAll regions but EthiopianAll <strong>the</strong> regionsAlmost cosmopoliteAll regions but NearcticPatagonia (? marine)Oriental, S. AfricaEurope, AmericaEthiopian, NeotropicalAll warm regionsTemperate S. AmericaPalaearctic, NearcticTemperate S. AmericaAll tropical regionsOriental, NeotropicalEthiopian, NeotropicalAll warm regions, and to CanadaCosmopoliteOriental, EthiopianOriental, NeotropicalAll tropical regionsCosmopoliteAll <strong>the</strong> o<strong>the</strong>r regionsAll o<strong>the</strong>r regions but NearcticCosmopoliteCosmopoliteCosmopoliteCosmopoliteCosmopoliteNeotropicalOriental, EthiopianAll tropical regionsCosmopolite


.CHAP. XIII.] THE AUSTRALIAN REGION. 473TABLE II.GENERA OF TERRESTRIAL MAMMALIA AND BIRDS INHABITING THEA USTRALIAN REGION.Explanation.Names in italics show genera peculiar to <strong>the</strong> regionNames enclosed thus ( ) show genera which just enter <strong>the</strong> region, but are not consideredproperly to belong to it.Genera truly belonging to <strong>the</strong> region are numbered consecutively.MAMMALIA.Order, Family, andGenus.q a).'3O 0>Range <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.PRIMATES.Cynopi<strong>the</strong>cidje.(Macacus1. Cynopi<strong>the</strong>cus ...Lombok to Timor)Celebes and BatchianOriental genusPhilippines ?Lemurid-e.(Tarsius ,Celebes)Indo-Malayan genusCHIROPTERA.Pteropid^e.2. Pteropus3. Xantharpyia ...4. Cynopterus5. Macroglossus ..6. Harpyia7. Hypoderma8. Notopteris15111111<strong>The</strong> whole reg. except New Zeal.Moluccas and TimorMorty IslandCelebes, Moluccas, TimorCelebes and MoluccasCelebes, Moluccas, and TimorFiji IslandsTropics <strong>of</strong> E. Hemisp.Oriental, S. PalaearcticOrientalIndo-MalayaPhilippinesRhinolophid^;.9. Rhinolophus ..10. Hipposideros ..11. Phyllorhina ..12. AseUia13. MegadermaMoluccas, Timor, AustraliaMoluccas and Aru IslandsMoluccas and TimorAmboynaTernate"Warmer pts. <strong>of</strong> E. Hem is.OrientalIndo-MalayaIndo-MalayaOriental, EthiopianVESPERTILIONIDiE.14. Scotophilus ..15. Vespertilio16. Miniopteris17. Taphozous18. Plecotus19. Nyctophilus ..Vol. I.—32Moluccas, Timor, AustraliaAustraliaMoluccas, Timor, and AustraliaCelebes, Moluccas, N. AustraliaTimorAustralia and TasmaniaOrientalCosmopoliteIndo-Malaya, S. AfricaOrien.,Ethiop., Neotrop.N. India, S. PalaearcticIndia


474 ZOOLOGICAL GEOGEAPHY. [PART 111.Order, Family, andGenus.


CHAP. XIII.] THE AUSTRALIAN REGION. 475Order, Family, andGenus.


476 ZOOLOGICAL GEOGRAPHY. [part III.Order, Family, and


CHAP. XIII.] THE AUSTRALIAN REGION. 477Order, Family, andGenus.


478 ZOOLOGICAL GEOGRAPHY. [PART III.


CHAP. XIII.] THE AUSTRALIAN REGION. 479Order, Family, andGenus.


1480 ZOOLOGICAL GEOGRAPHY. [far :Order, Family, andGenus.r


.(HAP. XIII.] THE AUSTRALIAN REGION. 481Order, Family, andGenus.. cRange <strong>with</strong>in <strong>the</strong> Region.Range beyond <strong>the</strong> Region.CuCULIDiE.167. Rhamphococcyx168. Centropus169. Cueulus170. Calicchthrus ..171. Cacomantis ...172. Chrysococcyx(Hierococcyx...173. Eudynamis ...174. ScythropsCoraciid^:.1135110516CelebesAustro-Malaya and AustraliaAustro-Malaya and AustraliaPapuan IslandsAustro-Malaya and AustraliaAustro-Malaya to Fiji Islandsand N. ZealandCelebes)<strong>The</strong> whole region ; excl. SandwichIslandsCelebes, Moluccas, and AustraliaOriental, EthiopianPale, Orien., EthiopianOrientalOriental, EthiopianOriental genusOriental(Coracias175. Eurystomus ...Celebes)Austro-Malaya aud AustraliaOriental and EthiopianOriental and EthiopianMeropid^;.176. Meropogon177. Merops.CelebesAustro-Malaya and AustraliaPale, Orien.,EthiopianAlCEDINIDjE.178. Alcedo179. Alcyone180. Pelargopsis ...181. Ceyx182. Ceycopsis183. Syma184. Halcyon185. Todirhamphus186. Dacelo187. Monachalcyon188. Caridonax189. Tanysiptcra ...190. Cittura191. MelidoraBtJCEROTID-E.4627121936111421Celebes to New IrelandBatchian to TasmaniaCelebes, FloresCelebes to New GuineaCelebesPapuan Islands and N. Australia<strong>The</strong> whole region ; excl. SandwichIslandsCentral Pacific and Sandwich Ids.Papuan Islands and AustraliaCelebesLombok and FloresBatchian to N. Guinea and N.AustraliaCelebes and Sanguir IslandsNew GuineaPale, Orien.,PhilippinesOrientalOrientalOriental, EthiopianEthiopian192. Hydrocissa? ...193. Calao194. Cranorrhinus ?CelebesMoluccas to Solomon IslandsCelebesOrientalMalayanMalayanPODARGID.E.195. Podargus196. Batrachostomus197. uEgothdes ...1025Papuan Islands to TasmaniaMoluccasPapuaD Islands to TasmaniaOrientalCAPRIMULGID.E.198. Caprimulgus ..Lombok to Australia, N. Guineato Pelew IslandsPale, Ethiopian, Orien.


482 ZOOLOGICAL GEOGRAPHY. [part hi.


CHAP. XIII.] THE AUSTRALIAN REGION. 483Order, Family, andGenus.O 0)*5tRange <strong>with</strong>in <strong>the</strong> Region.Range bej-ond <strong>the</strong> Region.Stringopmle.235. StringopsN. Zealand, Chatham Islands ?COLUMB.E.Columbidje,236. Treron237. Ptilopus238. Carpophaga ...239. Ianthaenas240. Leitcomelcena...241. Lopholcemus ...242. Geopelia.243. Macropygia ...244. Turacczna245. Reinwardto&nas246. Turtur247. Ocyphaps248. Petrophassa ...249. Chalcophaps ..250. Trugon251. Henicophaps ...252. Phaps253. Leucosarcia ...254. Geophaps255. Lophophaps ...256. Catenas257. Otidiphaps ...258. Phlogcenas ...259. GouraDlDUNCULIDiE.260. Didwnculus ...5040611563121141131231173Celebes, Bouru, and Ceram,Floresand Timor<strong>The</strong> whole region ; excl. N.Zealand<strong>The</strong> whole regionGilolo, Timor, Papuan Ids. toSamoan IslandsAustraliaAustraliaLombok to TasmaniaAustro-Malaya, AustraliaCelebes, Timor, Solomon Ids.Celebes to New GuineaAustro-MalayaAustraliaN. W. AustraliaAustro-Malaya, AustraliaN. GuineaPapuan IslandsAustralia and TasmaniaAustraliaAustraliaAustraliaAustro-MalayaN. GuineaCelebes, N. Guinea to MadagascarPapuan IslandsSamoan IslandsOriental,Indo-MalayaEthiopianAn-OrientalJapan, Philippines,daman IslandsMalaya, ChinaIndo-MalayaPalsearc, Orien., Ethiop.OrientalIndo-MalayaPhilippine IslandsGALLING.TETRAONIDiE.261. CoturnixPhasianice.(GallusTurnicid^;.262. TurnixCelebes, Timor, Australia, N.ZealandCelebes to Timor)Celebes & Moluccas to TasmaniaPalaearc, Orien., Ethiop.Oriental genusPalsearc, Orien., Ethiop.MEGAPODIID.E.263. Talegallus ...264. Megacephalon265. Lipoa266. Megapodius ...1112Papuan Islands and AustraliaCelebesS. AustraliaCelebes to Austral. & Samoan Ids.Philippines, Nicobar Ids*


484 ZOOLOGICAL GEOGRAPHY. [part III.Order, Family, and .Genus.


CHAP. XIII.] THE AUSTRALIAN REGION. 485Order, Family, andGenus.


INDEX TO VOL. I.


INDEX TO VOL. I.Note.—In this Index <strong>the</strong> names in Italics all refer to fossil genera or familiesmentioned in Part II. <strong>The</strong> systematic names <strong>of</strong> genera and families occurring inalmost every page <strong>of</strong> Part III. are not given, as <strong>the</strong>y would unnecessarily swell<strong>the</strong> Index ; but <strong>the</strong>y can be readily referred to by <strong>the</strong> Class or Order, or by <strong>the</strong>Geographical Division (Region or Sub-region) under which <strong>the</strong>y occur. <strong>The</strong>ywill, however, all be found in <strong>the</strong> General Index, <strong>with</strong> a reference to <strong>the</strong> page (inVol. II., Part IV.) where a systematic account <strong>of</strong> <strong>the</strong>ir <strong>distribution</strong> is given.Aardvark <strong>of</strong> East Africa, figure <strong>of</strong>, 261Accipitres, European Eocene, 163Accipitres, classification <strong>of</strong>, 97range <strong>of</strong> Palsarctie genera <strong>of</strong>, 248range <strong>of</strong> Ethiopian genera <strong>of</strong>, 312range <strong>of</strong> Oriental genera <strong>of</strong>, 385range <strong>of</strong> Australian genera <strong>of</strong>, 486Acera<strong>the</strong>riwn, European Miocene, 119N. American Tertiary, 136Achcenodon, N. American Tertiary, 138Aco<strong>the</strong>rium, European Eocene, 126Adapts, European Eocene, 125/Elurogale, European Eocene, 125sEpyornis, <strong>of</strong> Madagascar, 164/Eshna, from <strong>the</strong> Lias, 167Agnopterus, European Eocene, 163Agriochcerus, N. American Tertiary, 138Agrion, from <strong>the</strong> Lias, 167Alcephalus, Indian Miocene, 122Aldabra Islands, land-tortoises <strong>of</strong>, 289Ahtornis, N. American Eocene, 163Algeria, Post-Pliocene deposits and caves <strong>of</strong>,111Allen. Mr. J. A., on Zoological regions, 61objections to his system <strong>of</strong> circumpolarzones, 67objections to his zoo-<strong>geographical</strong> nomenclature,68Altai mountains, fossils in caves, 111Amblyrhiza, Pliocene <strong>of</strong> Antilles, 148America, recent separation <strong>of</strong> North andSouth, 40extinct mammalia <strong>of</strong>, 129North, Post-Pliocene fauna <strong>of</strong>, 129Amomys, N. American Tertiary, 134Amphechinus, European Miocene, 117Amphibia, means <strong>of</strong> dispersal <strong>of</strong>, 28classification <strong>of</strong>, 100peculiar to Palaearctic region, 186<strong>of</strong> Central Europe, 196<strong>of</strong> <strong>the</strong> Mediterranean sub-region, 205<strong>of</strong> Siberian sub-region, 220Vol. L—33Amphibia, <strong>of</strong> <strong>the</strong> Manchurian sub-region, 226table <strong>of</strong> Paltearctic families <strong>of</strong>, 237<strong>of</strong> <strong>the</strong> Ethiopian region, 255<strong>of</strong> West Africa, 264South African, 268<strong>of</strong> Madagascar, 280table <strong>of</strong> Ethiopian families <strong>of</strong>, 298<strong>of</strong> <strong>the</strong> Oriental region, 317<strong>of</strong> <strong>the</strong> Indian sub-region, 326<strong>of</strong> Ceylon, 327<strong>of</strong> Indo-Chinese sub-region, 331<strong>of</strong> Indo-Malay sub- region, 340table <strong>of</strong> Oriental families <strong>of</strong>, 369<strong>of</strong> <strong>the</strong> Australian region, 397resemblances <strong>of</strong> Australian and SouthAmerican, 400<strong>of</strong> New Guinea, 416<strong>of</strong> New Zealand, 457Ampkibos, Indian Miocene, 122Amphicjion, European Miocene, 118Indian Miocene, 121N. American Tertiary, 134Amphinuricidai, European Miocene, 119Amphimoschus, European Miocene, 120Amphisorex, European Miocene, 118Amphitragulns, European Miocene, 120Anastoma, European Tertiary, 169Anchilophus. European Eocene, 125Anchippodus, N. American Eocene, 139Anchippus, N. American Tertiary, 135Ancki<strong>the</strong>ruhe, N. American Tertiary, 135Anchi<strong>the</strong>rium, European Miocene, 119European Eocene, 125N American Tertiary, 135Ancient fauna <strong>of</strong> New Zealand, 459Ancylo<strong>the</strong>rium, Miocene <strong>of</strong> Greece, 116European Miocene, 121Andaman Islands, zoology <strong>of</strong>. 333probable past history <strong>of</strong>, 334Andreas, European Miocene, 165Animal kingdom, primary divisions <strong>of</strong>, 85Animals, development <strong>of</strong>, affecting <strong>distribution</strong>,7dispersal and migration <strong>of</strong>, 10


490 INDEX.Animals, rapid multiplication <strong>of</strong>, 10Anisacodon, N. American Tertiary, 137Anoa <strong>of</strong> Celebes, peculiarities <strong>of</strong>, 428Anoplo<strong>the</strong>riidce, European Miocene, 119Anoplo<strong>the</strong>rium, European Miocene, 1 19European Eocene, 126S. American Eocene, 148Anseres, arrangement <strong>of</strong>, 98peculiar Palasarctic genera, 250peculiar Ethiopian genera <strong>of</strong>, 313peculiar Australian genera <strong>of</strong>, 487Antelopes in <strong>the</strong> Indian Miocene deposits, 122birthplace and migrations <strong>of</strong>, 155Paliearctic, 182Antelo<strong>the</strong>rhim, Indian Miocene, 122Anthracothcrid.ee, N. American Tertiary, 137Anthraeo<strong>the</strong>rium, European Miocene, 119Antiacodon, N. American Tertiary, 133Antilles, Pliocene Mammalia <strong>of</strong>, 148Antilope, Post-Pliocene, 112in Brazilian caves, 144Antiquity <strong>of</strong> <strong>the</strong> genera <strong>of</strong> insects, 166<strong>of</strong> <strong>the</strong> genera <strong>of</strong> land and freshwatershells, 168Aphanapteryx <strong>of</strong> Mauritius, 164Aphelo<strong>the</strong>rium, European Eocene, 125Aquila, European Miocene, 161Archcenpteryx, Bavarian Oolite, 163Arctic zone not a separate region, 68Arctocyon, European Eocene, 125Arctodus, N. American Post-Pliocene, 130Arctomys, European Pliocene, 113Arcto<strong>the</strong>rium in Brazilian caves, 144S. American Pliocene, 146Argus pheasant, figure <strong>of</strong>, 339peculiarity in display <strong>of</strong> plumage, andconfirmation <strong>of</strong> Mr. Darwin's views,340Artiodactyla, European Eocene, 126N. American Tertiary, 137S. American Pliocene, 146Arvicola, European Pliocene, 113in Brazilian caves, 145S. American Pliocene, 147S. American Eocene, 148Auchena, N. American Post-Pliocene, 130Auckland Islands, birds <strong>of</strong>, 455Australia, physical features <strong>of</strong>, 387Australia and S. America, supposed land connectionbetween, 398Australian region, description <strong>of</strong>, 387zoological characteristics <strong>of</strong>, 390mammalia <strong>of</strong>, 390birds <strong>of</strong>, 391reptiles <strong>of</strong>, 396amphibia <strong>of</strong>, 397fresh-water fish <strong>of</strong>, 397summary <strong>of</strong> vertebrata <strong>of</strong>, 397supposed land-connection <strong>of</strong> <strong>with</strong> 8. America,398insects <strong>of</strong>, 403lepidoptera <strong>of</strong>, 404coleoptera <strong>of</strong>, 405land shells <strong>of</strong>, 407sub-regions <strong>of</strong>, 408early history <strong>of</strong>, 465Australian sub-region, mammalia <strong>of</strong>, 438illustration <strong>of</strong> mammalia <strong>of</strong>, 439birds <strong>of</strong>, 440illustration <strong>of</strong> fauna <strong>of</strong>, 441Austro- Malayan sub-region, physical features<strong>of</strong>, 388zoology <strong>of</strong>, 409Aye-aye, figure <strong>of</strong>, 278Azores, visited by European birds, 17birds <strong>of</strong>, 207Azores, butterflies <strong>of</strong>, 207beetles <strong>of</strong>, 207, 209peculiarly modified birds <strong>of</strong>, 207stragglers to, 208how stocked <strong>with</strong> animal life, 208BBabirusa <strong>of</strong> Celebes, peculiarities <strong>of</strong>, 42bBadger, figure <strong>of</strong>, 195Balaam, European Pliocene, 112Balwnodon, European Pliocene, 112Baly, Mr, on Phytophaga <strong>of</strong> Japan, 230Banca, its peculiar species and solution <strong>of</strong> aproblem in <strong>distribution</strong>, 356Barriers, as affecting <strong>distribution</strong>, 6permanence <strong>of</strong>, as affecting <strong>distribution</strong>, 7to <strong>the</strong> dispersal <strong>of</strong> birds, 17Bates, Mr., on Carabidae<strong>of</strong> Japan, 228on Longicorns <strong>of</strong> Japan, 230Bathmodon, N. American Tertiary, 13€Bathrodon, N. American Tertiary, 133Batrachia, Tertiary, 165Bats, powers <strong>of</strong> flight <strong>of</strong>, 15classification <strong>of</strong>, 87<strong>of</strong> New Zealand, 450Bears, probable cause <strong>of</strong> absence <strong>of</strong>, fromtropical Africa, 291Beaver, N. American Tertiary, 140Beetles, families selected for <strong>study</strong>, 103from <strong>the</strong> Lias, 167<strong>of</strong> Azores, 207<strong>of</strong> Japan, 228Belemnoziphius, European Pliocene, 112Belt, Mr. , his <strong>the</strong>ory <strong>of</strong> a great Siberian lakeduring <strong>the</strong> glacial epoch, 218on change <strong>of</strong> climate caused by diminution<strong>of</strong> obliquity <strong>of</strong> ecliptic, 466Birds, means <strong>of</strong> dispersal <strong>of</strong>, 15dispersal <strong>of</strong> by winds, 16American, found in Europe, 16reaching <strong>the</strong> Azores, 17barriers to dispersal <strong>of</strong>, 17limited by forests, 17classification <strong>of</strong>, 93Miocene <strong>of</strong> Greece, 116extinct, 160fossil <strong>of</strong> Palrearetic region, 161European <strong>of</strong> Miocene period, 161Eocene <strong>of</strong> Europe, 162<strong>relations</strong> <strong>of</strong>, 162extinct <strong>of</strong> North America, 163recently extinct in New Zealand, 164Cretaceous <strong>of</strong> N. America, 164remains <strong>of</strong> in Brazilian caves, 164recently extinct in Madagascar and <strong>the</strong>Mascarene Islands, 164cosmopolitan groups <strong>of</strong>, 176numerous genera, Palaearetic, 1S3<strong>of</strong> <strong>the</strong> European sub-region, 193nor<strong>the</strong>rn range <strong>of</strong> in Europe, 193<strong>of</strong> <strong>the</strong> zone <strong>of</strong> pine forests, 194<strong>of</strong> Iceland, 198<strong>of</strong> <strong>the</strong> Mediterranean sub-region, 203<strong>of</strong> Malta, 206 (note)<strong>of</strong> Azores, 207<strong>of</strong> <strong>the</strong> Cape Verd Islands, 215<strong>of</strong> Siberian sub-region, 219Oriental found in Siberia, 219extreme nor<strong>the</strong>rn Asiatic, 219<strong>of</strong> nor<strong>the</strong>rn Asiatic forests, 220<strong>of</strong> <strong>the</strong> Manchurian sub-region, 223Pal^arctic genera <strong>of</strong>, in <strong>the</strong> Manchuriansub-region, 224


AmericanINDEX.491Birds, Oriental genera <strong>of</strong>, in <strong>the</strong> Manchuriansub-region, 224characteristic <strong>of</strong>golia, 226N.W. China and Mon-table <strong>of</strong> Palsearctic families <strong>of</strong>, 235<strong>of</strong> West Africa, 243list <strong>of</strong> Paliearctic genera <strong>of</strong>, 243<strong>of</strong> <strong>the</strong> Ethiopian region, 253<strong>of</strong> <strong>the</strong> East African sub-region, 260S. African, 267genera <strong>of</strong>, peculiar to Madagascar, 275common to Madagascar and Oriental orEthiopian regions, 276species common to Madagascar and Africaor Asia, 277table <strong>of</strong> Ethiopian families <strong>of</strong>, 295table <strong>of</strong> Ethiopian genera <strong>of</strong>, 306<strong>of</strong> <strong>the</strong> Oriental region, 316<strong>of</strong> <strong>the</strong> Indian sub-region, 323Oriental genera <strong>of</strong> in Central India, 324Pala^arctic and Ethiopian genera inCentral India, 325<strong>of</strong> Ceylon, 327<strong>of</strong> Indo-Chinese subr egion, 330<strong>of</strong> Indo-Malayan sub-region, 337illustration <strong>of</strong> peculiar Malayan, 339<strong>of</strong> <strong>the</strong> Philippine Islands, 346table <strong>of</strong> Oriental families <strong>of</strong>, 366table <strong>of</strong> Oriental genera <strong>of</strong>, 375<strong>of</strong> Australian region, 391specially organized Australian families <strong>of</strong>,392<strong>of</strong> <strong>the</strong> Papuan Islands, 410peculiarities <strong>of</strong>, 413brilliant colours <strong>of</strong>, 413remarkable forms <strong>of</strong>, 414<strong>of</strong> <strong>the</strong> Moluccas, 418peculiarities <strong>of</strong>, 421<strong>of</strong> Timor group, 423<strong>of</strong> Celebes. 42S<strong>of</strong> Australia, 440<strong>of</strong> New Zealand, 451peculiar to New Zealand, 452<strong>of</strong> Norfolk Island, 453<strong>of</strong> Lord Howe's Island, 453<strong>of</strong> <strong>the</strong> Chatham Islands, 454<strong>of</strong> <strong>the</strong> Auckland Islands, 455table <strong>of</strong> families <strong>of</strong> Australian, 471table <strong>of</strong> genera <strong>of</strong> Australian, 478Black ape <strong>of</strong> Celebes, 427Blanford, Mr. W. T., on <strong>the</strong> " Indian"region, 60on <strong>relations</strong> <strong>of</strong> Indian sub-region <strong>with</strong>Africa, 321Blapsirtium, Oolitic insect, 167Blyth, Mr., on zoological regions, 60on <strong>the</strong> <strong>relations</strong> <strong>of</strong> Indian sub-region <strong>with</strong>Africa, 321Borneo, probable recent changes in, 357Bos, Post-Pliocene, 112Indian Miocene, 122Bourbon, zoology <strong>of</strong>, 280reptiles <strong>of</strong>, 281Bovidce, European Miocene, 120Brachymys, European Miocene, 120Bruma<strong>the</strong>rivm, Miocene <strong>of</strong> Perim Island, 122Brazilian cave-fauna, 143remarks on, 145Breyeria borinensis, carboniferous insect, 168Britain, peculiar species in, 197British Isles, zoology <strong>of</strong>, 197Broad-bill, Malayan, figure <strong>of</strong>. 340Bronto<strong>the</strong>rida', N. Tertiary, 137Bronto<strong>the</strong>rinm, X. American Tertiary, 137B-ubo, European Miocene, 162Bulimus, Eocene, 169Buncehiras, N. American Tertiary, 134Buprestidium, Oolitic insect, 167Butterflies, arrangement <strong>of</strong>, 103Pal«arctic, 187<strong>of</strong> Central Europe, 196<strong>of</strong> <strong>the</strong> Mediterranean sub-region, ^05<strong>of</strong> Azores, 207peculiar to Siberian sub-region, 220<strong>of</strong> Japan and North China, 227<strong>of</strong> <strong>the</strong> Ethiopian region, 255number <strong>of</strong> Ethiopian species, 256<strong>of</strong> Indo-Malay sub-region, 342<strong>of</strong> <strong>the</strong> Australian region, 404<strong>of</strong> <strong>the</strong> Austro-Malay sub-region, 404<strong>of</strong> <strong>the</strong> Moluccas, 419<strong>of</strong> Celebes, peculiarities <strong>of</strong>, 434<strong>of</strong> New Zealand, 457Cadurco<strong>the</strong>rium, European Eocene, 125Cmlodon, in Brazilian caves, 145Ccelogenys, in Brazilian caves, 144Cosnopi<strong>the</strong>cus, European Eocene, 124Caino<strong>the</strong>rium, European Miocene, 120European Eocene, 126Calamodon, N. American Eocene, 139Callithrix in Brazilian caves, 184Canaries, birds <strong>of</strong>, 208beetles <strong>of</strong>, 209Canidw, European Miocene, 118European Eocene, 125N. American Tertiary, 134remarkable S. African, 267Canis, European Pliocene, 112Post-Pliocene, 112European Miocene, 118Indian Miocene, 121European Eocene, 125N. American Post-Pliocene, T29N. American Tertiary, 134, 135in Brazilian caves, 144S. American Pliocene. 146Camel, fossil in Indian Miocene, 122birth-place and migrations <strong>of</strong>, 155Fahearctic, 182Camelida; essentially extra-tropical, 112N. American Tertiary, 138Camelopardalis, Miocene <strong>of</strong> Greece, 116Indian Miocene, 122Cameto<strong>the</strong>rium, S. American Pliocene, 147Cape <strong>of</strong> Good Hope, peculiar flora and fauna<strong>of</strong>, 266Cape Verd Islands, zoology <strong>of</strong>, 214Cape-hare, S. African, 267Cardiodus; S. American Pliocene, 147Cariama, Brazilian caves, 164Carnivore <strong>of</strong> European Pliocene, 112Miocene <strong>of</strong> Greece, 115European Miocene, 118Indian Miocene, 121European Eocene, 125N. American Post- Pliocene, 129N. American Tertiary. 134<strong>of</strong> Brazilian caves, 144S. American Pliocene, 146Carnivora, classification <strong>of</strong>, 88antiquity <strong>of</strong>, 153<strong>of</strong> <strong>the</strong> Pala'arctie region, 182list <strong>of</strong> Palsearctic genera <strong>of</strong>, 240list <strong>of</strong> Ethiopian genera <strong>of</strong>, 302range <strong>of</strong> Oriental genera <strong>of</strong>, 373list <strong>of</strong> Australian genera <strong>of</strong>, 476Caroline Islands, birds <strong>of</strong>, 444Carterodon in Brazilian caves, 145


492 INDEX.Cams, and Gerstaeker on classification <strong>of</strong><strong>animals</strong>, 85Pr<strong>of</strong>essor, on classification <strong>of</strong> <strong>the</strong> Cetacea,88Castor, European Pliocene, 113European Miocene, 120Casoryx, N. American Tertiary, 138Cathartes, Brazilian caves, 164Cave-fauna <strong>of</strong> Brazil, 143Cavia, European Miocene, 121in Brazilian caves, 144S. American Pliocene, 147Cebochcerus, European Eocene, 126Cebus in Brazilian caves, 144Celebes, physical features <strong>of</strong>, 389mammalia <strong>of</strong>, 426birds <strong>of</strong>, 428insects <strong>of</strong>, 434origin <strong>of</strong> fauna <strong>of</strong>, 436Centetidm, European Miocene, 118Ceratodus, remarkable Australian fish, 397Cercolabes in Brazilian caves, 145Cercopi<strong>the</strong>cus in European Pliocene, 112Cervidce, European Miocene, 120birthplace and migrations <strong>of</strong>, 155Cervus, European Pliocene, 113Indian Pliocene and Miocene, 122N. American Post-Pliocene, 130N. American Tertiary, 138in Brazilian caves, 144S. American Pliocene, 147Cetacea, European Pliocene, 112European Miocene, 119N. American Post-Pliocene, 130N. American Tertiary, 140Cetacea, classification <strong>of</strong>, 89range <strong>of</strong> Oriental genus, 374Ceylon and Malaya, resemblance <strong>of</strong> insects <strong>of</strong>,327Ceylonese sub-region, 326mammalia <strong>of</strong>, 327birds <strong>of</strong>, 327reptiles <strong>of</strong>, 327amphibia <strong>of</strong>, 327insects <strong>of</strong>, 327past history <strong>of</strong>, as indicated by its fauna,328Chalicomys, European Pliocene, 113Chalico<strong>the</strong>rium, European Miocene, 119Indian Miocene, 122fossil in N. China, 123Chamaleo, N. American Eocene, 165Chamois, figure <strong>of</strong>, 195Chatham Islands, birds <strong>of</strong>, 454Chelonia, classification <strong>of</strong>, 100Chelydra, European Pliocene, 165Chevrotain <strong>of</strong> Malaya, figure <strong>of</strong>, 336Chili should not be placed in <strong>the</strong> Palseaor Nearctic regions, 63China, fossil mammals in, resembling those <strong>of</strong>Indian and European Miocene, 362North, mammalia <strong>of</strong>, 222Chinchillida: in Brazilian caves, 145S. American Pliocene, 147Pliocene <strong>of</strong> Antilles, 148Chiroptera, classification <strong>of</strong>, 87list <strong>of</strong> Palsearctic genera <strong>of</strong>, 239list <strong>of</strong> Ethiopian genera <strong>of</strong>, 300range <strong>of</strong> Oriental genera <strong>of</strong>, 371list <strong>of</strong> Australian genera <strong>of</strong>, 475Chiroptera, European Eocene, 125in Brazilian caves, 144Chlamydolherium in Brazilian caves, 145Chmromorus, European Miocene, 119Chceropotiimvs, European Eocene, 126Chairo<strong>the</strong>riwn, Indian Miocene, 122Choneziphivs, European Pliocene, 112Chough, Alpine, figure <strong>of</strong>, 195Circuuipolar zones, objections to system <strong>of</strong>, 67Classification as affecting <strong>the</strong> <strong>study</strong> <strong>of</strong> <strong>distribution</strong>,83Clausilia, Eocene, 169Climate, as a limit to <strong>the</strong> range <strong>of</strong> mammalia,11gradual change <strong>of</strong>, before <strong>the</strong> glacial epoch,41Coleoptera. families selected for <strong>study</strong>, 103Palsearctic, 188number <strong>of</strong> Palsearctic species, 189<strong>of</strong> Central Europe, 196<strong>of</strong> <strong>the</strong> M editerranean sub-region, 205<strong>of</strong> <strong>the</strong> Cape Verd Islands, 215<strong>of</strong> <strong>the</strong> Ethiopian region, 256S. African, 268<strong>of</strong> Madagascar, 282, 283<strong>of</strong> <strong>the</strong> Oriental region, 319<strong>of</strong> Indo-Malay sub-region, 342<strong>of</strong> <strong>the</strong> Australian region, 405affinity <strong>of</strong> Australian and South American406, 407<strong>of</strong> Celebes, 435<strong>of</strong> New Zealand, 457Collocalia, European Miocene, 161Colobus, European Miocene, 117Colonoceras, N. American Tertiary, 136Colossochelys <strong>of</strong> Indian Miocene, 123,U65Columbse, classification <strong>of</strong>, 96range <strong>of</strong> Palsearctic genera <strong>of</strong>, 248range <strong>of</strong> Ethiopian genera <strong>of</strong>, 311range <strong>of</strong> Oriental genera <strong>of</strong>, 384range <strong>of</strong> Australian genera <strong>of</strong>, 485Comoro islands, zoology <strong>of</strong>, 281Continents, <strong>distribution</strong> <strong>of</strong>, 37recent changes <strong>of</strong>, 38Continental extension in Mesozoic times, 156Corvus, European Miocene, 161Coryphndon, European Eocene, 126Cosmopolitan groups enumerated, 175Cricetodon, European Miocene, 120Cricetus, European Tliocene, 113Crocodiles, Eocene, 165Crocodilia, classification <strong>of</strong>, 100Crook-billed plovers <strong>of</strong> New Zealand, 456Crotch, Mr., on beetles <strong>of</strong> <strong>the</strong> Azores, 209Crowned-pigeon, figure <strong>of</strong>, 415Cryptornis, European Eocene, 163Ctenomys, S. American Pliocene, 147Cuba, extinct mammalia <strong>of</strong>, 148Curculionidium, Oolitic insect, 167Cyclostoma, Eocene, 169Cyllo sepulta, European Cretaceous, 167Cynoelurvs, in Brazilian caves, 144Cynopi<strong>the</strong>cus <strong>of</strong> Celebes, affinities <strong>of</strong>, 427Cyo<strong>the</strong>rium, European Eocene, 125D.Daptophilus, N. American Tertiary, 134Darwin, Mr., his explanation <strong>of</strong> <strong>the</strong> cause <strong>of</strong><strong>the</strong> abundance <strong>of</strong> apterous insects inMsdeira, 211on <strong>the</strong> relation <strong>of</strong> flowers and insects, 463Dasyprocta, European Miocene, 121in Brazilian eaves, 144Dasypus, in Brazilian caves, 145S. American Pliocene, 147Dasyurus, Australian Post-Tertiary. 157David, Pere, his researches in China and Thibet,221, 222on birds <strong>of</strong> N. China, 226


INDEX. 493Deer, fossil in N. American Tertiary formations,138Palsearctic, 182probable cause <strong>of</strong> absence <strong>of</strong> from tropicalAfrica, 291Delphinus, European Pliocene, 112Dendrocyg na, European Miocene, 162Desman <strong>of</strong> S. Russia, figure <strong>of</strong>, 219Dicera<strong>the</strong>rium, N. American Tertiary, 137Dichobune, European Eocene, 12'iDicotyles, N. American Post- Pliocene, 130N. American Tertiary, 137in Brazilian caves, 144S. American Pliocene, 146birthplace and migrations <strong>of</strong>, 155Dicrocerus, European Miocene, 120Didelphys, European Eocene, 126N. American Post-Pliocene, 130in Brazilian caves, 145Didida:, 164Dinocerata, N. American Tertiary, 139Dinaceras, N. American Eocene, 139Dinoruis, allied form in European Eocene, 163<strong>of</strong> New Zealand and Australia, 164Dinoriiithidcc <strong>of</strong> New Zealand, 164Dino<strong>the</strong>riim, Miocene <strong>of</strong> Greece, 116European Miocene, 120Miocene <strong>of</strong> Perim Island, 123Dinyctis, N. American Tertiary, 134Dinylus, European Miocene, 117Diplacodon,^. American Tertiary, 136Diprotodon, Australian Post-Tertiary, 157Dispersal <strong>of</strong> <strong>animals</strong>, 10<strong>of</strong> mamma' ia, 10<strong>of</strong> reptiles and amphibia, 28Distribution, affected by climate, 5affected by physical features, 5contrasts <strong>of</strong>, in similar climates, 5similarities <strong>of</strong>, in diverse climates, 6barriers as affecting, 6<strong>study</strong> <strong>of</strong>, dependent on a good classification,83<strong>of</strong> <strong>animals</strong> an adjunct to geology, 8<strong>of</strong> <strong>animals</strong> requires certain preliminarystudies, S<strong>of</strong> <strong>animals</strong> dependent on physical geography,35<strong>of</strong> <strong>animals</strong>, as affected by <strong>the</strong> glacial epoch.40<strong>of</strong> <strong>animals</strong>, as affected by changes <strong>of</strong>vegetation, 43<strong>of</strong> <strong>animals</strong>, as affected bv organic changes,44<strong>of</strong> <strong>animals</strong>, hypo<strong>the</strong>tical illustration <strong>of</strong>,46<strong>of</strong> <strong>animals</strong>, complexity <strong>of</strong> <strong>the</strong> causes affecting<strong>the</strong>, 49<strong>of</strong> <strong>animals</strong>, problems in, 51<strong>of</strong> plants, as affected by <strong>the</strong> glacial epoch,42Dodo <strong>of</strong> Mauritius, 282Dolichopterus, European Miocene, 162Dommina, N. American Tertiary, 134Dorca<strong>the</strong>rium, European Miocene, 120Dremo<strong>the</strong>.rium, Miocene <strong>of</strong> Greece, 116European Miocene, 120Dresser, Mr. H. E., on nor<strong>the</strong>rn range <strong>of</strong>European birds, 193Droma<strong>the</strong>ritim, N, American Triassic, 134oldest American mammal, 160Drnngo-shrike, Malayan figtne o p , 340Dryopntkecus European Miocene, 117E.East Africa, <strong>geographical</strong> features <strong>of</strong>, 258wide range <strong>of</strong> genera and species over, 259few special types in, 260East African sub-region, description <strong>of</strong>, 258genera and species ranging over <strong>the</strong> whole<strong>of</strong>, 259mammalia <strong>of</strong>, 260birds <strong>of</strong>, 260reptiles <strong>of</strong>, 260amphibia and fishes <strong>of</strong>, 260insects <strong>of</strong>, 260few peculiar types in, 260illustration <strong>of</strong> zoology <strong>of</strong>, 261East Australia, peculiar birds <strong>of</strong>, 440East Thibet, mammalia <strong>of</strong>, 222Eaton, Rev. A. E., on insects <strong>of</strong> KerguelenIsland, 211Echimyidiv, in Brazilian caves, 145Eckinogale, European Miocene, 118Ectognathus, N. American Eocene, 139Edentata, Miocene <strong>of</strong> Greece, 116European Miocene, 121N. American Post-Pliocene, 130N. American Pliocene, 140<strong>of</strong> Brazilian caves, 145S. American Pliocene, 147Edentata, classification <strong>of</strong>, 90probable birthplace <strong>of</strong>, 155range <strong>of</strong> Ethiopian genera <strong>of</strong>, 305range <strong>of</strong> Oriental genus rf, 375Elephants, fossil <strong>of</strong> Indian M\ >cene, 123fossil in N. American Post- Pliocene formations,130birthplace and migrations <strong>of</strong>, 155Elephant shrews, 8. African, 267Elephas, Post-Pliocene, 112fossil in N. China, 123N. American Tertiary, 138Elliot, Mr., his great work on <strong>the</strong> birds oparadise, 415Elomis, European Miocene, 162Elo<strong>the</strong>rium, N. American Tertiary, 137, 139Elwes, Mr., on birds <strong>of</strong> Peisia, 204on true <strong>relations</strong> <strong>of</strong> <strong>the</strong> birds <strong>of</strong> CentralIndia, 323Embasis, N. American Tertiary, 134Emu, figure <strong>of</strong>, 441Emys, Indian Miocene, 123Miocene and Eocene, 1 65Emydida, Indian Miocene, 123Enhydrion, Indian Miocene, 121Eobasileus, N. American Eocene, 139Eocene period, 124fauna <strong>of</strong> S. America, 148Ephemera, from <strong>the</strong> Lias, 167Bporeodon, N. American Tertiary, 138Equiiln; European Pliocene, 112Miocene <strong>of</strong> Greece, 115European Eocene, 125Eqmts, European Pliocene, 112Pust-Pliocene, 112Indian Miocene, 121N. American Post-Pliocene, 130N. American Tertiary, 135Brazilian caves, 144S. American Pliocene, 146Ereptodon, N. American Post- Pliocene, 130Erinaceus, European Miocene, 117Erythromachus <strong>of</strong> Rodriguez, 164Esthonyx, N. American Eocene, 139Ethiopian region should not include any part<strong>of</strong> India, 63defined, 7isubdivisions <strong>of</strong>, 73


494 INDEX.Ethiopian region, general features <strong>of</strong>, 251zoological characteristics <strong>of</strong>, 25?mammalia <strong>of</strong>, 253great speciality <strong>of</strong>, 253birds <strong>of</strong>, 253reptiles <strong>of</strong>, 254amphibia <strong>of</strong>, 255fresh-water fish <strong>of</strong>, 255summary <strong>of</strong> vertebrates <strong>of</strong>, 255insects <strong>of</strong>, 255coleoptera <strong>of</strong>, 256terrestrial mollusca <strong>of</strong>, 257sub-regions <strong>of</strong>, 258Atlantic islands <strong>of</strong>, 269<strong>the</strong> probable past history <strong>of</strong>, 285tables <strong>of</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong> <strong>of</strong>, 293Ewtnys, N. American Tertiary, 140Euphrachis, S. American Pliocene, 147Europe, recent changes in physical geography<strong>of</strong>, 39Miocene fauna <strong>of</strong> Central, 117Miocene fauna <strong>of</strong>, allied to existing fauna<strong>of</strong> tropical Asia and Africa, 124European sub-region, description <strong>of</strong>, 191forests <strong>of</strong>, 192mammalia <strong>of</strong>, 192birds <strong>of</strong>, 193reptiles and amphibia <strong>of</strong>, 195fresh-water fish <strong>of</strong>, 196insects <strong>of</strong>, 196islands <strong>of</strong>, 197Euryceros <strong>of</strong> Madagascar, figure <strong>of</strong>, 278Eurydon, in Brazilian caves, 145Eurii<strong>the</strong>rium, European Eocene, 126Eutatus, S. American Pliocene, 147Eutelodon, European Eocene, 126Eutemnodus, S. American Eocene, 148Extinct mammalian fauna <strong>of</strong> Europe, generalconsiderations on, 126mammalia <strong>of</strong> N. America and Europe,comparison <strong>of</strong>, 140mammalia <strong>of</strong> <strong>the</strong> Antilles, 148mammalia <strong>of</strong> Old and New Worlds,general remarks on, 148fauna <strong>of</strong> New Zealand, 459Extinction <strong>of</strong> large <strong>animals</strong>, causes <strong>of</strong>, 158Fauna <strong>of</strong> Japan, general character and affinities<strong>of</strong>, 230<strong>of</strong> Patearetic region, general conclusionsas to, 231extinct, <strong>of</strong> Madagascar and MascareneIslands, 282Malayan, probable origin <strong>of</strong>, 359Moluccan, peculiarities <strong>of</strong>, 419Timorese, origin <strong>of</strong>, 422<strong>of</strong> Celebes, origin <strong>of</strong>, 436<strong>of</strong> New Zealand, origin <strong>of</strong>, 460Felis spelcea, 110Felis, Miocene <strong>of</strong> Greece, 115European Miocene, 118Indian Miocene, 121N. American Post-Pliocene, 129in Brazilian caves, 144Fernando Po, zoological features <strong>of</strong>, 265Fiji, Tonga, and Samoa Islands, birds <strong>of</strong>, 443Fishes, means <strong>of</strong> dispersal <strong>of</strong>, 29classification <strong>of</strong>, 101cosmopolitan groups <strong>of</strong>, 176<strong>of</strong> <strong>the</strong> Palaearctic region, 186<strong>of</strong> <strong>the</strong> European sub-region, 196<strong>of</strong> <strong>the</strong> Mediterranean sub-region, 205<strong>of</strong> <strong>the</strong> Manchurian sub-region. 227Fishes.fresh -water,table <strong>of</strong> Palaearctic families<strong>of</strong>, 227<strong>of</strong> <strong>the</strong> Ethiopian region, 255<strong>of</strong> South Africa, 268fresh-water, table <strong>of</strong> Ethiopian families <strong>of</strong>,298fresh-water, <strong>of</strong> <strong>the</strong> Oriental region, 318<strong>of</strong> <strong>the</strong> Indo-Malay sub-region, 341fresh-water, table <strong>of</strong> Oriental families <strong>of</strong>,369fresh-water, <strong>of</strong> <strong>the</strong> Australian region, 397fresh-water,resemblance <strong>of</strong> Australian andS. American, 400how <strong>the</strong> transmission may have takenplace, 401fresh-water, <strong>of</strong> New Zealand, 457Flamingoes, European Miocene, 162Flora, <strong>of</strong> New Zealand, as influenced by scarcity<strong>of</strong> insects, 462fossil <strong>of</strong> Australia, 467Flower, Pr<strong>of</strong>essor, on classification <strong>of</strong> mammalia,85classification <strong>of</strong> carnivora, 87Flying Lemur, Malayan, figure <strong>of</strong>, 337Flying Opossum, figure <strong>of</strong>, 442Formosa, zoology <strong>of</strong>, 332Forests, essential to existence <strong>of</strong> many European<strong>animals</strong>, 192Siberian, greatest extent <strong>of</strong>, 216G.Galapagos, scarcity <strong>of</strong> insects in, 463Galecynus, in European Pliocene, 112Galera, N. American Post- Pliocene, 13Galeospalax, European Miocene, 118Galeo<strong>the</strong>rium, Post-Pliocene, 111Galethylax, European Eocene, 125Galictis, in Brazilian caves, 144Gallinse, classification <strong>of</strong>, 96range <strong>of</strong> Palaearctic genera <strong>of</strong>, 248range <strong>of</strong> Ethiopian genera <strong>of</strong>, 311range <strong>of</strong> Oriental genera <strong>of</strong>, 3S4range <strong>of</strong> Australian genera <strong>of</strong>, 485GoMus, Miocene <strong>of</strong> Greece, 116Gallus bravardi, European Pliocene, 161Gast


INDEX. 495Grallae, peculiar Oriental genera <strong>of</strong>, 386peculiar Australian genera <strong>of</strong>, 486Gray, Dr. J. E., on classification <strong>of</strong> Cetacea,88Greece, Upper Miocene deposits <strong>of</strong>, 115summary <strong>of</strong> Miocene fauna <strong>of</strong>, 116Groups peculiar to a region, how denned, 184Gulick, Rev. J. T., on Achatinellidas <strong>of</strong> <strong>the</strong>Sandwich Islands, 446Giin<strong>the</strong>r, Dr., his classification <strong>of</strong> reptiles, 98his classification <strong>of</strong> fishes, 101on gigantic tortoises <strong>of</strong> Galapagos and <strong>the</strong>Mascarene Islands, 289on range <strong>of</strong> Indian reptiles in <strong>the</strong> Himalayas,329H.Haast, Dr. , on extinct birds <strong>of</strong> New Zealand,460Habitat, definition <strong>of</strong>, 4Hainan, zoology <strong>of</strong>, 334Haleyomis, European Eocene, 103Hali<strong>the</strong>rium, European Pliocene, 112European Miocene, 119Hellado<strong>the</strong>rium, Miocene <strong>of</strong> Greece, 116European Miocene, 120Hatteria <strong>of</strong> New Zealand, 456Helictis, Himalayan, figure <strong>of</strong>, 331Helix, Eocene, 169Hemibos, Indian Miocene, 122Hemicyon, European Miocene, 118Herpeto<strong>the</strong>rium, N. American Tertiary, 134llesperomys, N. American Tertiary, 140in Brazilian caves, 145S. American Pliocene, 147Hesperomis, N. American Cretaceous, 164Heterodon, in Brazilian caves, 145Hexaprotodon, Indian Miocene, 122Hickman, Mr. John, on a cause <strong>of</strong> <strong>the</strong> extinction<strong>of</strong> large <strong>animals</strong>, 158Himalayas, altitude reached by various groupsin <strong>the</strong>, 329, 333Hipparion, European Pliocene, 112Miocene <strong>of</strong> Greece, 115European Miocene, 119N. American Post-Pliocene, 130N. American Tertiary, 135Hippopotamus, Post- Pliocene, 112Europe m Pliocene, 113Indian Pliocene, 122Hipposyus, N. American Tertiary, 133Hippo<strong>the</strong>ri-um, European Miocene, 119Indian Miocene, 122Hippotragus, European Miocene, 120Homalodonto<strong>the</strong>rium, S. American Pliocene,146Homalophvs, European Miocene, 161Homocamelus, N. American Tertiary, 138Honevsuckers, birds specially adapted to"Australia, 392Hooker, Dr. , on deficiency <strong>of</strong> odours in NewZealand plants, 464Hoplocetus, European Pliocene, 112Hoplophoneus, N. American Tertiary, 134Hor&es, fossil, in Indian Miocene, 121perfect series <strong>of</strong> ancestral, in N. America,136probable birthplace <strong>of</strong>, 154Hutton, Capt. F. W., on origin <strong>of</strong> New Zealandfauna, 461Huxley, Pr<strong>of</strong>essor, on zoological regions, 59division <strong>of</strong> animal kingdom by, 85Hycena, Post-Pliocene, 112Miocene <strong>of</strong> Greece, 115European Miocene, 118Hyoma, Indian Miocene, 121fossil in N. China, 123Hyrenwrctos in European Pliocene, 112European Miocene, 118Indian Miocene, 121S. American Pliocene, 146Hycenictis, Miocene <strong>of</strong> Greece, 115European Miocene, 118Hyrenidw, European Miocene, 118Hycenodon, European Miocene, 118European Eocene, 125N. American Tertiary, 134Hyomodontidoe, European Miocene, 118Hydrochcerus, N. American Post-Pliocene, 130Hydrornis, European Miocene, 162Hyohippus, N. American Tertiary, 135Hyomoschns, European Miocene, 120Hyopotamus, European Miocene, 119N. American Tertiary, 137Hyopsodus, N. AmericanTertiary, 133Hyo<strong>the</strong>rium, European Miocene, 119Hypertragulus, N. American Tertiary, 13SHypisodus, N. American Tertiary, 138Hi/psiprymnus, Australian Post-Tertiary, 157Hyrachyus, N. American Tertiary, 136Hyracodon, N. American Tertiary, 136Hyracoidea, classification <strong>of</strong>, 90Palaearetic, 242Ethiopian, 304Hyraco<strong>the</strong>rium, supposed, in European Eocene,125European Eocene, 126Hystrix, European Pliocene, 113Miocene <strong>of</strong> Greece, 116N. American Tertiary, 140Ibidipodui, European Miocene, 162Ibidorhynclms, figure <strong>of</strong>, 331Iceland, zoology <strong>of</strong>, 198Icthyornis, N. American Cretaceous, 164Icticyon in Brazilian caves, 144Icti<strong>the</strong>rium, Miocene <strong>of</strong> Greece, 115European Miocene, 118Ictops, N. American Tertiary, 133India, Miocene fauna <strong>of</strong>, allied to that <strong>of</strong>Europe, 123geological features <strong>of</strong>, 328Indian, sub-region, description <strong>of</strong>, 321supposed relation to Ethiopian region, 321mammalia <strong>of</strong>, 322birds <strong>of</strong>, 323reptiles and amphibia <strong>of</strong>, 326Indo-Chinese, sub-region, description <strong>of</strong>, 329zoological characteristics <strong>of</strong>, 330illustration <strong>of</strong>, 331reptiles <strong>of</strong>, 331amphibia <strong>of</strong>, 331insects <strong>of</strong>, 332islands belonging to, 333Indo-Malayan sub- region, description <strong>of</strong>, 334mammalia <strong>of</strong>, 336illustrations <strong>of</strong>, 336, 339birds <strong>of</strong>, 337remote <strong>geographical</strong> <strong>relations</strong> <strong>of</strong>, 339reptiles and amphibia <strong>of</strong>, 340fishes <strong>of</strong>, 341insects <strong>of</strong>, 341coleoptera <strong>of</strong>, 342terrestrial mollusca <strong>of</strong>, 343zoological <strong>relations</strong> <strong>of</strong> islands <strong>of</strong>, 345recent <strong>geographical</strong> changes in, 357probable origin <strong>of</strong> fauna <strong>of</strong>, 359Insects, means <strong>of</strong> dispersal <strong>of</strong>, 32


496 INDEX.Insects, tenacity <strong>of</strong> life <strong>of</strong>, 33adapted to special conditions, 33groups selected for <strong>the</strong> <strong>study</strong> <strong>of</strong> <strong>the</strong>ir<strong>geographical</strong> <strong>distribution</strong>, 102antiquity <strong>of</strong> <strong>the</strong> genera <strong>of</strong>, 166fossil <strong>of</strong> European Miocene, 166European Cretaceous, 167European Wealden, 167Palaeozoic, 168Palsearctic, 187<strong>of</strong> Central Europe, 196<strong>of</strong> <strong>the</strong> Mediterranean sub-region, 205<strong>of</strong> <strong>the</strong> Siberian sub-region, 220<strong>of</strong> <strong>the</strong> Manchurian sub-region, 227<strong>of</strong> <strong>the</strong> Ethiopian region, 255<strong>of</strong> <strong>the</strong> East African sub-region, 260<strong>of</strong> West Africa, 265S. African, 268<strong>of</strong> Madagascar, 282general remarks on, 284<strong>of</strong> tropical Africa and America, probablecause <strong>of</strong> similarities in, 291<strong>of</strong> Indo-Chinese sub-region, 332<strong>of</strong> <strong>the</strong> Oriental region, 318<strong>of</strong> Ceylon, 327<strong>of</strong> Indo-Malay sub-region, 341statistics <strong>of</strong> collecting in <strong>the</strong> variousislands <strong>of</strong> <strong>the</strong> Malay Archipelago, 343<strong>of</strong> <strong>the</strong> Australian region, 403<strong>of</strong> New Guinea, 417<strong>of</strong> <strong>the</strong> Moluccas, 420<strong>of</strong> Timor group, 426<strong>of</strong> Celebes, 454<strong>of</strong> New Zealand, 458scarcity <strong>of</strong>, in New Zealand, 462influence <strong>of</strong> on <strong>the</strong> flora, 463Insectivora, European Miocene, 117N. American Post-Pliocene, 129N. American Tertiary, 133Insectivora, classification <strong>of</strong>, 87<strong>of</strong> <strong>the</strong> Palsearctic region, 181<strong>of</strong> N. China and E. Thibet, 222range <strong>of</strong> Palsearctic genera <strong>of</strong>, 239<strong>of</strong> Madagascar, 273range <strong>of</strong> Ethiopian genera <strong>of</strong>, 301<strong>of</strong> <strong>the</strong> Oriental region, 315range <strong>of</strong> Oriental genera <strong>of</strong>, 372range <strong>of</strong> Australian genera <strong>of</strong>, 476Isacis, N. American Tertiary, 133Ischyromys, N. American Tertiary, 140Islands, N. European, zoology <strong>of</strong>, 197<strong>of</strong> <strong>the</strong> Mediterranean sub-region, 206<strong>of</strong> <strong>the</strong> West African sub-region, 265<strong>of</strong> Ethiopian region, 269Mascarene, 280<strong>of</strong> <strong>the</strong> Indo-Chinese sub-region, 333<strong>of</strong> Indo-Malay sub-region, 345Fiji, Tonga, and Samoa, 443Society and Marquesas. 444New Caledonia and New Hebrides, 445Sandwich, 446<strong>of</strong> New Zealand sub-region, 453Norfolk, 453Lord Howe's, 454Chatham, 454Auckland, 455Issiodromys, European Pliocene, 113Jacchus, in Brazilian caves, 144Japan and North China, physical features <strong>of</strong>.221sou<strong>the</strong>rn extremity <strong>of</strong> perhaps belongs to<strong>the</strong> Oriental region, 226Japan, general character <strong>of</strong> <strong>the</strong> fauna <strong>of</strong>, 230former land-connexions <strong>of</strong>, 231Java, mammalia <strong>of</strong>, 349productions <strong>of</strong>, well known, 350birds <strong>of</strong>, 351representative species <strong>of</strong> birds in, 352origin <strong>of</strong> <strong>the</strong> anomalous features <strong>of</strong> itsfauna, 352Sumatra and Borneo, <strong>the</strong>ir <strong>geographical</strong>contrasts and zoological peculiaritiesexplained, 357Junonia, European Miocene, 167Kakapoe, <strong>of</strong> New Zealand, 455Kangaroos, extinct in Australia, 157Kerguelen Island, apterous insects <strong>of</strong>, 211{note)Kerodon, in Brazilian caves, 144S. American Pliocene, 147King-fisher, racquet tailed, <strong>of</strong> New Guinea,figure <strong>of</strong>, 415Kiwi <strong>of</strong> New Zealand, 455Koodoo antelope, figure <strong>of</strong>, 261Lacertilia, classification <strong>of</strong>, 99Ladrone Islands, birds <strong>of</strong>, 444Lagomys, European Pliocene, 113European Miocene, 120Lagostomus, in Brazilian eaves, 145S. American Pliocene, 147Lake Baikal, seals <strong>of</strong>, 218Land and water, proportions <strong>of</strong>, 35Land and fresh-water shells,antiquity <strong>of</strong> <strong>the</strong>genera <strong>of</strong>, 168Land-shells, Palseozoic, 169Paliearctic, 190<strong>of</strong> Madeira, 209<strong>of</strong> <strong>the</strong> Cape Verd Islands, 215<strong>of</strong> <strong>the</strong> Ethiopian region, 257<strong>of</strong> W. Africa, 265<strong>of</strong> Madagascar and <strong>the</strong> Mascarene Islands,285<strong>of</strong> Indo-Malay sub-region, 344<strong>of</strong> <strong>the</strong> Australian region, 407<strong>of</strong> Sandwich Islands, 446<strong>of</strong> New Zealand, 459Lanius, European Miocene, 161Laopi<strong>the</strong>cits, \ .American Tertiary, 133Laor?iis, N. American Cretaceous, 164Lemuria, a hypo<strong>the</strong>tical land, 76Lamuravidce, N. American Tertiary, 133Lemuravus, N American Tertiary, 133Lemuridm, European Eocene, 124Lemuroidea, range <strong>of</strong> Ethiopian genera <strong>of</strong>, 300range <strong>of</strong> Oriental genera <strong>of</strong>, 371Lepictis, N. American Tertiary, 133Lepidoptera, cosmopolitan families <strong>of</strong>, 177table <strong>of</strong> Palsearctic families <strong>of</strong>, 23SS. African, 268table <strong>of</strong> Ethiopian families <strong>of</strong>, 299<strong>of</strong> <strong>the</strong> Oriental region, 318table <strong>of</strong> Oriental families <strong>of</strong>, 369<strong>of</strong> <strong>the</strong> Australian region, 404table <strong>of</strong> Australian families <strong>of</strong>, 472Leptarchus, N. American Tertiary, 135Leptauchenia, N. American Tertiary, 138Leptochosrus, N. American Tertiary, 137Leptodon, Miocene <strong>of</strong> Greece, 116Leptomeryx, N. American Tertiary, 138Leptoplilvs, European Miocene, 162


INDEX. 497Leptosomus, allied form in European Eocene,168Leptosomus <strong>of</strong> Madagascar, 27Sfigure <strong>of</strong>, 279Leptothcrium, in Brazilian caves, 144Lepus, in Brazilian caves, 145S. American Pliocene, 147Lestodon, S. American Pliocene, 147Lewis, Mr. George, his collection <strong>of</strong> Japaninsects, 228Lebellula, from <strong>the</strong> Lias, 167Lilljeborg, Pr<strong>of</strong>essor, on classification <strong>of</strong> <strong>the</strong>Rodentia, 90Limnwa, Eocene, 169European Secondary, 169Limnatornis, European Miocene, 161Limnoctjon, N. American Tertiary, 134Limnohyus, N. American Tertiary, 136Limnotneridce, N. American Tertiary, 133Limnolherivm, N. American Tertiary, 133Listriodon, European Miocene, 119Lithomys, European Miocene. 120Lithornis, European Eocene, 163Lizards, classification <strong>of</strong>, 99Tertiary, 165wide range <strong>of</strong> a species in Polynesia, 448Loncheres, in Brazilian caves, 145Lonchophorus, in Brazilian caves, 145Lophiodon, European Eocene, 125N. American Tertiary, 136Lophio<strong>the</strong>rium, N. American Tertiary, 136 .Lord Howe's Island, birds <strong>of</strong>, 453Loxomylus, Pliocene <strong>of</strong> Antilles, 148Lund, Dr., his researches in caves <strong>of</strong> Brazil, 143Lutra, European Miocene, 118Indian Miocene, 121Lyccena, Miocene <strong>of</strong> Greece, 115Lyrebird, figure <strong>of</strong>, 441M.Macacus, European Pliocene, 112Miocene <strong>of</strong> Greece, 115Indian Miocene, 121supposed in European Eocene, 125Machairodus, 110, 111Miocene <strong>of</strong> Greece, 115European Miocene, 118Indian Miocene, 121N. American Tertiary, 134in Brazilian caves, 144S. American Pliocene, 146Macranchenia, S. American Pliocene, 146Macro<strong>the</strong>rium, Miocene <strong>of</strong> Greece, 116European Miocene, 121Madagascar, extinct birds <strong>of</strong>, 164description <strong>of</strong>, 272mammalia <strong>of</strong>, 272birds <strong>of</strong>, 274reptiles <strong>of</strong>, 279amphibia <strong>of</strong>, 280extinct fauna <strong>of</strong>, 282general remarks on insect fauna <strong>of</strong>, 284Madeira, birds <strong>of</strong>, 20Sland shells <strong>of</strong>, 208beetles <strong>of</strong>, 210wingless insects numerous in, 211how stocked <strong>with</strong> <strong>animals</strong>, 213Malacca, Sumatra, and Borneo, zoologicalunity <strong>of</strong>, 353comparison <strong>of</strong> mammalia, 354<strong>of</strong> birds, 355Malagasy sub-region, description <strong>of</strong>, 272mammalia <strong>of</strong>, 272birds <strong>of</strong>, 274illustration <strong>of</strong> zoology <strong>of</strong>, 278Malagasy sub-region, reptiles <strong>of</strong>, 279amphibia <strong>of</strong>, 280extinct fauna <strong>of</strong>, 2S2, 289insects <strong>of</strong>, 282early history <strong>of</strong>, 2S6Malaya and Indo-Malaya, terms defined, 345(note)Malavan forms <strong>of</strong> life reappearing in West" Africa, 263fauna, probable origin <strong>of</strong>, 359resemblances to that <strong>of</strong> Madagascar andCeylon explained, 361Malta, Post-Pliocene fauna <strong>of</strong>, 114formerly joined to Africa, 201fossil elephants <strong>of</strong>, 201birds <strong>of</strong>, 206 (note)Mammalia, means <strong>of</strong> dispersal <strong>of</strong>, 10as limited by climate, 11as limited by rivers, 12how far limited by <strong>the</strong> sea, 13dispersed by ice-floes and drift-wood, 14means <strong>of</strong> dispersal <strong>of</strong> aquatic, 15<strong>of</strong> most importance in determining zoologicalregions, 57classification <strong>of</strong>, 85birthplace and migrations <strong>of</strong> some families<strong>of</strong>, 142, 153cosmopolitan groups <strong>of</strong>, 176<strong>of</strong> <strong>the</strong> Palsearetic region, 181<strong>of</strong> <strong>the</strong> European sub-region, 192<strong>of</strong> <strong>the</strong> Mediterranean sub-region, 202<strong>of</strong> <strong>the</strong> Siberian sub-region, 217characteristic <strong>of</strong> Western Tartary, 218<strong>of</strong> <strong>the</strong> Manchurian sub-region, 222Pal»arctio genera <strong>of</strong>, in <strong>the</strong> Manchuriansub-region, 222Oriental genera <strong>of</strong>, on borders <strong>of</strong> samesub-region. 223peculiar to Japan, 223characteristic <strong>of</strong> N. W. China and Mongolia,226table <strong>of</strong> Palaearetic families <strong>of</strong>, 234range <strong>of</strong> Pal*arctic genera <strong>of</strong>, 239<strong>of</strong> <strong>the</strong> Ethiopian region, 253absence <strong>of</strong> certain important groups, 253<strong>of</strong> <strong>the</strong> East African sub-region, 260<strong>of</strong> West Africa, 262<strong>of</strong> S. Africa, 267<strong>of</strong> Madagascar, 272table <strong>of</strong> Ethiopian families <strong>of</strong>, 294table <strong>of</strong> Ethiopian genera <strong>of</strong>, 300<strong>of</strong> <strong>the</strong> Oriental region, 315range <strong>of</strong> <strong>the</strong> genera inhabiting <strong>the</strong> Indiansub-region, 322<strong>of</strong> Ceylon, 327<strong>of</strong> <strong>the</strong> Indo-Chinese sub-region, 330<strong>of</strong> <strong>the</strong> Indo-Malayan sub-region, 336illustration <strong>of</strong> characteristic Malayan,336<strong>of</strong> <strong>the</strong> Philippine Islands, 345table <strong>of</strong> Oriental families <strong>of</strong>, 365table <strong>of</strong> Oriental genera <strong>of</strong>, 371<strong>of</strong> Australian region, 390<strong>of</strong> <strong>the</strong> Papuan Islands, 410<strong>of</strong> <strong>the</strong> Moluccas, 417<strong>of</strong> Timor group, 422<strong>of</strong> Celebes, 427<strong>of</strong> Australia, 439illustration <strong>of</strong>, 439<strong>of</strong> New Zealand, 450table <strong>of</strong> families <strong>of</strong> Australian, 470table <strong>of</strong> genera <strong>of</strong> Australian, 475Mammal, <strong>the</strong> most ancient American, 134Mammalia, extinct, <strong>of</strong> Old World, 107extinct, <strong>of</strong> historic period, 110extinct, comparative age <strong>of</strong> in Europe, 127


498 INDEX.Mammalia, extinct, <strong>of</strong> <strong>the</strong> New World, 129extinct, <strong>of</strong> N. America and Europe, compared,141original birth-place <strong>of</strong> some families andgenera, 142, 153<strong>of</strong> <strong>the</strong> secondary period, 160Manatus, N. American Post-Pliocene, 130Manchurian sub-region, description <strong>of</strong>, 220mammalia <strong>of</strong>, 222birds <strong>of</strong>, 223reptiles and amphibia <strong>of</strong>, 227fresh-water fish <strong>of</strong>, 227insects <strong>of</strong>, 227coleoptera <strong>of</strong>, 228Marquesas Islands, birds <strong>of</strong>, 443Marsh, Mr., on unprovability <strong>of</strong> Asiatic andAfrican deserts, 200on camels and goats as destructive tovegetation, 200Marsupials, classification <strong>of</strong>, 91N. American Post-Pliocene, 130European Miocene, 121first migration to America, 155diversified forms <strong>of</strong>, 391<strong>of</strong> America prove no connexion <strong>with</strong>Australia, 399list <strong>of</strong> Australian genera <strong>of</strong>, 476Martes, N. American Tertiary, 135Mascarene Islands, zoology o'f, 280extinct fauna <strong>of</strong>, 282gigantic land-tortoises <strong>of</strong>, 289Mastodon, European Pliocene, 113Miocene <strong>of</strong> Greece, 116European Miocene, 120in Brazilian caves, 144S. American Pliocene, 147Indian Miocene, 123N. American Post-Pliocene, 130N. American Tertiary, 138Mauritius, zoology <strong>of</strong>, 280reptiles <strong>of</strong>, 2S1McCoy, 'Pr<strong>of</strong>essor, on Paleontology <strong>of</strong>Victoria, 466Mediterranean, recent changes in, 39sub-region, description <strong>of</strong>, 199mammalia <strong>of</strong>, 202birds <strong>of</strong>, 203reptiles and amphibia <strong>of</strong>, 204fresh-water fish <strong>of</strong>, 205insects <strong>of</strong>, 205islands <strong>of</strong>, 206sea not separating distinct faunas, 201Megacerops, N. American Tertiary, 137Megalomeryc, N. American Tertiary, 13SMegalociius, fossil in Cuba, 14SMegalonyx, N. American Post-Pliocene, 130in Brazilian caves, 145S. American Pliocene, 147Megalostoma, Eocene, 169Megamys, S. American Eocene, 148Megaspira, European Tertiary, 169Mega<strong>the</strong>ridn?, in Brazilian caves, 145Mega<strong>the</strong>rium, N. American Post-Pliocene 130in Brazilian caves, 145S. American Pliocene, 147Melania, European secondary, 169Meleagris, N. American Miocene, 163Mellivora, Indian Miocene, 121Melolonthidium, Oolitic insect, 167MenUco<strong>the</strong>rium, N. American Tertiary 13SMeno<strong>the</strong>riiim, N. American Tertiary, 133Mephitis, in Brazilian caves, 144Merychus, N. American Tertiary, 138Merychipws, N. American Tertiary, 135MerychocJuzrws, N. American Tertiary, 138Merycodus, N. American Tertiary, 13SMerycopotamus, Indian Miocene, 122Meryco<strong>the</strong>rium <strong>of</strong> Siberian drift,' 112Mesacodon, N. American Tertiary, 133Mesohippxis. N. American Tertiary, 135Mesonyx, N. American Tertiary, 134Mcsopi<strong>the</strong>cus, Miocene <strong>of</strong> Greece, 115Meyer, Dr. A. B., on reptiles and amphibia otNew Guinea, 415Microlestes, oldest European mammal, 160Micromeryx, European Miocene, 120Micmsyops, N. American Tertiary, 133Micro<strong>the</strong>rium, European Miocene, 120Middendorf, on extreme nor<strong>the</strong>rn birds, 219Migrating birds, in which region to be placed185Migration <strong>of</strong> <strong>animals</strong>, 10general phenomena <strong>of</strong>, 18<strong>of</strong> birds, 19<strong>of</strong> birds in Europe, 19probable origin <strong>of</strong>, 22<strong>of</strong> birds in India and China, 23<strong>of</strong> birds in N. America, 23changes in extent <strong>of</strong>, 24<strong>of</strong> birds in S. Temperate America, 25general remarks on, 25Milvus, European Miocene, 162Miocene fauna <strong>of</strong> <strong>the</strong> Old World, 114fauna <strong>of</strong> Greece, 115fauna <strong>of</strong> Greece, summary <strong>of</strong>, 116fauna <strong>of</strong> Central Europe,117deposits <strong>of</strong> Siwalik Hills, 121faunas <strong>of</strong> Europe and Asia, general observationson, 123Miohippus, N. American Tertiary, 135Mivart, Pr<strong>of</strong>essor, on classification <strong>of</strong> primates,86on classification <strong>of</strong> insectivora, 87on classification <strong>of</strong> amphibia, 101Moles almost wholly Palrearetic, 1S1Mole-rat, <strong>of</strong> W. Tartary, 218Mollusca, means <strong>of</strong> dispersal <strong>of</strong>, 30classification <strong>of</strong>, 104groups selected for <strong>study</strong>, 104Moluccas, zoology <strong>of</strong>, 417birds <strong>of</strong>, 419reptiles <strong>of</strong>, 420Insects <strong>of</strong>, 420peculiarities <strong>of</strong> fauna <strong>of</strong>, 421Monkeys on <strong>the</strong> high Himalayas, 12fossil in N. American Miocenein E. Thibet, 222abundance <strong>of</strong> in <strong>the</strong> Oriental region, 315Monotremata, classification <strong>of</strong>, 91list <strong>of</strong> Australian genera <strong>of</strong>, 477" More-pork" <strong>of</strong> Australia, figure <strong>of</strong>, 442Moro<strong>the</strong>rium, N. American Pliocene, 140MotaciUa, European Miocene, 161Mound -builders, peculiar Australian birds393Moupin, position and zoology <strong>of</strong>, 221Marida;. S. American Pliocene, 147Murray, Mr. Andrew, on zoological region, 60Mustela, Miocene <strong>of</strong> Greece, 115European Miocene. 118S. American Pliocene, 146Mii^telidre, in Brazilian caves. 144Mylodon, N. American Post-Pliocene, 130S. American Pliocene, 147Mi/ogalc, European Miocene, 118Mi/omorphus, fossil in CubaMuopotamus, in Brazilian caves, 145Myoxus, European Miocene, 120European Eocene, 126Mysarachne, European Miocene, 118Mi/sops, N. American Eocene, 140Myxophagus, N. American Post-Pliocene, 130


INDEX. 499Nannhyus, N. American Tertiary, 137Nos-ua, in Brazilian caves, 144Nearctic region, denned, 79subdivisions <strong>of</strong>, 80distinct from Pal*arctie, 79Necrornis, European Miocene, 161Neotropical region, denned, 78subdivisions <strong>of</strong>, 78<strong>relations</strong> <strong>of</strong> W. African sub-region <strong>with</strong>, 265Kesodon, S. American Pliocene, 147Newton, Pr<strong>of</strong>essor, on position <strong>of</strong> Menuridceand Atrickiidce, 95on birds <strong>of</strong> Iceland, 198New Caledonia, birds <strong>of</strong>, 444New Guinea, zoology <strong>of</strong>, 409mammalia <strong>of</strong>, 410birds <strong>of</strong>, 411peculiarities <strong>of</strong> its ornithology, 413illustration <strong>of</strong> ornithology <strong>of</strong>, 414reptiles ar.d amphibia <strong>of</strong>, 415insects <strong>of</strong>, 416New Zealand, objections to making a primaryzoological region, 62extinct birds <strong>of</strong>, 164sub-region, description <strong>of</strong>, 449compared <strong>with</strong> British Isles, 449mammalia <strong>of</strong>, 451islets <strong>of</strong>, 453illustration <strong>of</strong> ornithology <strong>of</strong>, 455reptiles <strong>of</strong>, 456amphibia <strong>of</strong>, 457fresh-water fish <strong>of</strong>, 457insects <strong>of</strong>, 458Longicorns <strong>of</strong>, 458Myriapoda <strong>of</strong>, 458land-shells <strong>of</strong>, 459ancient fauna <strong>of</strong>, 460origin <strong>of</strong> fauna <strong>of</strong>, 460poverty <strong>of</strong> insects in, 462<strong>relations</strong> <strong>of</strong> insect-fauna and flora, 472Nicobar Islands, <strong>the</strong>ir zoological <strong>relations</strong>, 332Nightingale, migration <strong>of</strong> <strong>the</strong>, 21Norfolk Island, birds <strong>of</strong>, 453North America, remarks on Post-Pliocenefauna <strong>of</strong>, 130Post-Pliocene fauna <strong>of</strong>, partly derived fromS. America, 131extinct birds <strong>of</strong>, 163North Africa, zoological <strong>relations</strong> <strong>of</strong>, 202Notharctos, N. American Tertiary, 133Notornis <strong>of</strong> New Zealand, 455Noto<strong>the</strong>rium, Australian Post-Tertiary, 157O.Ocho<strong>the</strong>rivm, in Brazilian caves, 145Octodontidce, S. American Pliocene, 147Ophidia, classification <strong>of</strong>, 99Opisthocomus, Brazilian caves, 164Opossum, extinct, in European Miocene, 121Oreodon, N. American Tertiary, 138Oreodontidce, N. American Tertiary, 138Oriental region, defined, 75subdivisions <strong>of</strong>, 75description <strong>of</strong>, 314zoological features <strong>of</strong>, 315mammalia <strong>of</strong>, 315birds <strong>of</strong>, 316reptiles <strong>of</strong>, 317amphibia <strong>of</strong>, 317fresh-water fishes <strong>of</strong>, 318summary <strong>of</strong> vertebrata, 31Sinsects <strong>of</strong>, 318sub-regions <strong>of</strong>, 321Oriental region, concluding remarks on, 362tables <strong>of</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong> <strong>of</strong>, 364Oriental <strong>relations</strong> <strong>of</strong> VV. African sub-region, 265Oriental and Patearctic faunas once identical,362Oriental and Ethiopian faunas, cause <strong>of</strong> <strong>the</strong>irresemblances, 363Orohippus, N. American Tertiary, 136Ostrich, Miocene <strong>of</strong> N. India, 162Otaria, European Miocene, 118Ovibos, N. American Post-Pliocene, 130Oxen, birthplace and migrations <strong>of</strong>, 155Palwarctie, 182Oxya-na, N. American Tertiary, 134Oxygomphits, European Miocene, 118Oxymycterus, in Brazilian caves, 145S. American Pliocene, 147Pachyana, N. American Tertiary, 134Pachynolophus, European Eocene, 126Pachy<strong>the</strong>rium, in Brazilian caves, 145Palaearctic region, defined, 71subdivisions <strong>of</strong>, 71general features <strong>of</strong>, 180zoological characteristics <strong>of</strong>, 181has few peculiar families, 181mammalia <strong>of</strong>, 181birds <strong>of</strong>, 182high degree <strong>of</strong> speciality <strong>of</strong>, 184reptiles and amphibia <strong>of</strong>, 186fresh-water fish <strong>of</strong>, 186summary <strong>of</strong> vertebrata <strong>of</strong>, 186insects <strong>of</strong>, 186coleoptera <strong>of</strong>, 187number <strong>of</strong> coleoptera <strong>of</strong>, 189land-shells <strong>of</strong>, 190sub-regions <strong>of</strong>, 190general conclusions on <strong>the</strong> fauna <strong>of</strong>, 231tables <strong>of</strong> <strong>distribution</strong> <strong>of</strong> <strong>animals</strong> <strong>of</strong>, 233Palceacodcn, N. American Tertiary, 133Palceetus, European Miocene, 162Palceyithalus, European Eocene, 162Palwtodus, European Miocene, 162Paheocastor, N. American Tertiary, 140Palceocercv.s, European Miocene, 162Palceochanis, European Miocene, 119Polwohierax, European Miocene, 162Palceolagus, N. American Tertiary, 140Palceolama,$. American Pliocene, 147Palceomephitis, European Miocene, 118Palceovieryx, European Miocene, 120Pc.l&omys, European Miocene, 121PalcBontina oolitica, Oolitic insect, 167Palaeontology, 107how best studied in its bearing on <strong>geographical</strong><strong>distribution</strong>, 168as an introduction to <strong>the</strong> <strong>study</strong> <strong>of</strong> <strong>geographical</strong><strong>distribution</strong>, concluding remarkson, 169Palmonyctis, European Eocene, 12SPalceoperdix, European Miocene, 161Palmoyforymis, European Miocene, 166Palaoreas, Miocene <strong>of</strong> Greece, 116Palcrortyx, European Miocene, 161Palceoryx, Miocene <strong>of</strong> Greece, 116Palceospalax, 111European Miocene, 117PaUrosyops, N. American Tertiary, 136Palceo<strong>the</strong>ridc", European Eocene, 125Palceo<strong>the</strong>rium, European Eocene, 125S. American Eocene, 148Palaeotragus, Miocene <strong>of</strong> Greece, 116Palceotringa, N. American Cretaceous, 164


!500 INDEX.Palapt


INDEX. 501Region, <strong>the</strong> best term for <strong>the</strong> primary zoologicaldivisions, 68Arctic, why not adopted, 69Palaearctic, defined, 71Palaearctic, subdivisions <strong>of</strong>, 71Ethiopian, defined, 73Ethiopian, subdivisions <strong>of</strong>, 73Oriental, defined, 75Oriental, subdivisions <strong>of</strong>, 75Australian, defined, 77Australian, subdivisions <strong>of</strong>, 77Neotropical, defined, 78Neotropical, subdivisions <strong>of</strong>, T8Nearctic, defined, 79Nearctic, distinct from Palaearctic, 79Nearctic. subdivisions <strong>of</strong>, 80Regions, zoological, 50zoological, how <strong>the</strong>y should be formed, 53zoological, may be defined by negative orpositive characters, 54zoological, by what class <strong>of</strong> <strong>animals</strong> bestdetermined, 56for each class <strong>of</strong> <strong>animals</strong>, not advisable,58zoological, proposed since 1857, 58zoological, Mr. Sclater's, 59zoological, discussion <strong>of</strong> those proposedby various authors, 61zoological, proportionate richness <strong>of</strong>, 64temperate and tropical, well marked illnor<strong>the</strong>rn hemisphere, 65and zones, table <strong>of</strong>, 66comparative richness <strong>of</strong>, 81and sub-regions, table <strong>of</strong>, 81order <strong>of</strong> succession <strong>of</strong> <strong>the</strong>, 173Representative species, 4Reptiles, means <strong>of</strong> dispersal <strong>of</strong>, 28classification <strong>of</strong>, 98Miocene <strong>of</strong> Greece, 116<strong>of</strong> Indian Miocene deposits, 123extinct Tertiary, 165cosmopolitan groups <strong>of</strong>, 176peculiar to Palaearctic region, 186<strong>of</strong> Central Europe, 195<strong>of</strong> <strong>the</strong> Mediteiranean sub-region, £04<strong>of</strong> Siberian sub-region, 220<strong>of</strong> <strong>the</strong> Manchurian sub-region, 227table <strong>of</strong> Palaearctic families <strong>of</strong>, £36<strong>of</strong> <strong>the</strong> Ethiopian region, 254<strong>of</strong> <strong>the</strong> East African sub-region, 260<strong>of</strong> West Africa, 264S. African, 268<strong>of</strong> Madagascar, 279table <strong>of</strong> Ethiopian families <strong>of</strong>, 297<strong>of</strong> <strong>the</strong> Oriental region, 317<strong>of</strong> <strong>the</strong> Indian sub-region, 326<strong>of</strong> Ceylon, 327<strong>of</strong> Indo-Chinese sub-region, 331<strong>of</strong> Indo-Malay sub-region, 340table <strong>of</strong> Oriental families <strong>of</strong>. 368<strong>of</strong> <strong>the</strong> Australian region, 396<strong>of</strong> New Guinea, 415<strong>of</strong> <strong>the</strong> Moluccas, 4£0<strong>of</strong> <strong>the</strong> Polynesian sub-region, 447<strong>of</strong> New Zealand, 456table <strong>of</strong> Australian families <strong>of</strong>, 472Rhea, in Brazilian caves, 164Rhinoceros, Post- Pliocene, 112European Pliocene, 113Miocene <strong>of</strong> Greece, 116Indian Miocene, 122fossil remain.' <strong>of</strong>, at 16,000 feet elevationin Thibet, 122fossil in N China, 123N. American Tertiary, 136Rhinoceros-hornbill, figure <strong>of</strong>, 339Rhinocerotidm, N. American Tertiary, 136River-hog, <strong>of</strong> West Africa, figure <strong>of</strong>, 264<strong>of</strong> Madagascar, figure <strong>of</strong>, 278Rivers, limiting <strong>the</strong> range <strong>of</strong> mammalia, 12limiting <strong>the</strong> range <strong>of</strong> birds, 17River-scene in West Africa, 264Kodentia, classification <strong>of</strong>, 90range <strong>of</strong> Palsarctic genera <strong>of</strong>, 242range <strong>of</strong> Ethiopian genera <strong>of</strong>, 304range <strong>of</strong> Oriental genera <strong>of</strong>, 374range <strong>of</strong> Australian genera <strong>of</strong>, 476Rodentia, European Pliocene, 113Miocene <strong>of</strong> Greece, 116European Miocene, 120European Eocene, 126N. American Post-Pliocene, 130N. American Tertiary, 139<strong>of</strong> Brazilian ca\es, 144S. American Pliocene, 147<strong>of</strong> S. American Eocene, 148Ruff, figure <strong>of</strong>, 1958.Sahara, a debatable land, 251Saiga, antelope <strong>of</strong> W. Tartary, 218Samoa Islands, birds <strong>of</strong>, 413Sandwich Islands, birds <strong>of</strong>, 445probable past history <strong>of</strong>, 446mountain plants <strong>of</strong>, 446depth <strong>of</strong> ocean around, 447Sand grouse, Pallas, <strong>of</strong> Mongolia, 226Suty rites Rei/nesii, .European Cretaceous insect,167Saunders, Mr. Edward, on <strong>the</strong> Buprestidae <strong>of</strong>Japan, 229Scelido<strong>the</strong>rium, in Brazilian caves, 145S. American Pliocene, 147SchistopUurum, S. American Pliocene, 147Schweinfurth, Dr., ou natural history <strong>of</strong> Centra]Africa, 252on limits <strong>of</strong> W. African sub-region, 262{note)Sciurus, European Miocene, 120European Eocene, 126Scivravvs, N. American Eocene, 140Sclater, Mr., on zoological regions, 59why his six regions are adopted, 63on birds <strong>of</strong> Sandwich Islands, 445Sea, as a barrier to mammalia, 13Seals, fossil in European Miocene, 118<strong>of</strong> Lake Baikal, 218Secondary formations, mammalian remainsin, 159Secretary bird <strong>of</strong> Africa, figure <strong>of</strong>, 261Semnopithccus, European Pliocene, 112Miocene <strong>of</strong> Greece, 115European Miocene, 117Indian Miocene, 121Semper, Dr., on Philippine mammalia, 345Serpentarius, European Miocene, 162Seychelle Islands, zoology <strong>of</strong>, 281amphibia <strong>of</strong>, 281'Sharp, Dr., on Japan beetles, 229Sharpe, Mr. R. B., his arrangement <strong>of</strong> Accipitres,97on birds <strong>of</strong> Cape Verd Islands, 215Sheep, Palaearctic, 182Siberia, climate <strong>of</strong>. 217Siberian sub-region, description <strong>of</strong>, 216mammalia nf, 217birds <strong>of</strong>, 219reptiles and amphibia <strong>of</strong>, 220insects <strong>of</strong>, 220Simocyon, Miocene <strong>of</strong> Greece, 115


American502 INDEX.Sinopa, N. American Tertiary, 134Sirenia, classification <strong>of</strong>, 89range <strong>of</strong> Ethiopian genera <strong>of</strong>, 303range <strong>of</strong> Oriental genus, 374range <strong>of</strong> Australian genus <strong>of</strong>, 476Sirenia, European Pliocene, 112European Miocene, 119Siva<strong>the</strong> riti m, Indian Miocene, 122Siwalik Hills, Miocene deposits <strong>of</strong>, 121Smith, Mr. Frederick, on Hynienoptera <strong>of</strong>Japan, 230Snake, at great elevation in Himalayas, 220Snakes, classification <strong>of</strong>, 99Eocene, 165large proportion <strong>of</strong> venomous species inAustralia, 396<strong>of</strong> New Zealand, 457Society Islands, birds <strong>of</strong>, 443Soricictis, European Miocene, 118Soricidcc, European Miocene, 118South African sub-region, description <strong>of</strong>, 266mammalia <strong>of</strong>, 267birds <strong>of</strong>, 267reptiles <strong>of</strong>, 268amphibia <strong>of</strong>, 268fresh-water fish <strong>of</strong>, 268butterflies <strong>of</strong>, 26Scoleoptera <strong>of</strong>, 268summary <strong>of</strong> its zoology, 269South America, fossil fauna <strong>of</strong>, 143Pliocene deposits <strong>of</strong>, 146supposed land connection <strong>with</strong> Australia,398South Australia, peculiar birds <strong>of</strong>, 441Species, representative, 4Speothos, in Brazilian caves, 144Spermo]ihihis, European Miocene, 120Sphenodon, in Brazilian caves, 145Sphinx, in European Oolite, 167St. Helena, zoological features <strong>of</strong>, 269coleoptera <strong>of</strong>, 270landshells <strong>of</strong>, 271St. Thomas' Island, birds <strong>of</strong>, 266Stations, definition <strong>of</strong>, 4Stene<strong>of</strong>iber, European Miocene, 120S<strong>the</strong>nurvs, Australian Post-Tertiary, 157Strix, European Miocene, 162Struthiones, arrangement <strong>of</strong>, 9Srange <strong>of</strong> Ethiopian genera <strong>of</strong>, 313range <strong>of</strong> Australian genera <strong>of</strong>, 487Struthious birds, probable origin <strong>of</strong>, 287Stylinodontida>, X. American Eocene, 139Stylinodontia,N . Eocene, 139Sub-regions, on what principle formed, SOPalaearctic, 191Ethiopian, 258Oriental, 321Australian, 40SSuidce, European Miocene, 119Sula Islands, fauna <strong>of</strong>, 433Sus, European Pliocene, 113Miocene <strong>of</strong> Greece, 116European Miocene, 119Indian Miocene, 122Swinhoe, Mr., on zoology <strong>of</strong> Formosa andHainan, 332Symborodon, N. American Tertiary, 137Synaphodus, European Miocene, 119Synoplotlierhim, N. American Tertiary, 134Tables <strong>of</strong> <strong>distribution</strong> <strong>of</strong> families and generaexplained, 177Talpa, European Miocene, 117Tapir, fossil in N. China, 123Tapirs, birthplace and migrations <strong>of</strong>, 154Tapir, Malayan, figure <strong>of</strong>, 3371'apiridcB, European Eocene, 125Tapirus, European Pliocene, 113Indian Miocene, 122in Brazilian caves, 144Tarsier, Malayan, figure <strong>of</strong>, 337Tasmania, comparative zoological poverty <strong>of</strong>,441Toxodon, European Miocene, 118TclmatobuLS, N. American Cretaceous, 1C4Telmatolestes, N. American Tertiary, 133Testudo, Miocene <strong>of</strong> Greece, 116Indian Miocene, 123Testudo, great antiquity <strong>of</strong> <strong>the</strong> genus, 289Tetrachus, European Miocene, 117Tetrao albus, in Italian caverns, 161Thalassictis, Miocene <strong>of</strong> Greece, 115European Miocene, 118<strong>The</strong>ridomys, European Miocene, 126European Eocene, 126S. American Eocene, 148Thinoliyns, N. American Tertiary, 137Thiuolestes, N.American Tertiary, 133Thylacinus, Australian Post-Tertiary, 157Thylacoleo, Australian Post-Tertiary, 157Tillodontia, N. American Eocene, 139Tillo<strong>the</strong>ridce, N. American Eocene, 139Tillo<strong>the</strong>rium, N. American Eocene, 139Timor, physical features <strong>of</strong>, 389group, mammalia <strong>of</strong>, 422birds <strong>of</strong>, 422origin <strong>of</strong> fauna <strong>of</strong>, 424insects <strong>of</strong>, 420Tinoeeras, N. American Eocene, 139Titinmmys, European Miocene, 121Titano<strong>the</strong>rivm, N. American Tertiary, 137Tomarctos, N. American 'tertiary, 135Tonga Islands, birds <strong>of</strong>, 443Tortoises, classification <strong>of</strong>, 100<strong>of</strong> Mascarene Islands and Galapagos, 289Touraco <strong>of</strong> W. Africa, figure <strong>of</strong>, 264Toxodon, S. American Pliocene, 13.ToxodontidK, S. American Pliocene, 147Trachy<strong>the</strong>rivm, European Miocene, 119Tragocerus, Miocene <strong>of</strong> Greece, 116European Miocene, 120Tragopan, Himalayan, figure <strong>of</strong>, 331Tree-shrew <strong>of</strong> Borneo, figure <strong>of</strong>, 337Tree-kangaroo, figure <strong>of</strong>, 415Trichechvs, N American Post-Pliocene, 130Trichoglossida>, birds specially adapted toAustralia, 393Trionyx, Indian Miocene, 123Miocene and Eocene, 165Tristan d'Acunha, zoology <strong>of</strong>, 271Tristram, Canon, summary <strong>of</strong> <strong>the</strong> birds <strong>of</strong>Palestine, 203Trogon, European Miocene, 161Troqon<strong>the</strong>rivm, Post-Pliocene <strong>of</strong> Europe. IllTrvcijelis, N. American Post-Pliocene, 129Tundras <strong>of</strong> Siberia, greatest extent <strong>of</strong>, 216Tnpaiida', European Miocene, 118Turner, Mr., on classification <strong>of</strong> Edentata, 90Tylodon, European Eocene, 125Typo<strong>the</strong>rium, S. American Pliocene, 147U.Uintacyon, N. American Tertiary, 134UintaOierium, N. American Eocene, 139Uintornis, N. American Eocene, 163Unio, European Secondary, 169Ungulata, classification <strong>of</strong>, 89


INDEX. 503I JJngulata, antiquity <strong>of</strong>, 151<strong>of</strong> <strong>the</strong> Palsearctic region, 182range <strong>of</strong> Pafcearctic genera <strong>of</strong>, 241range <strong>of</strong> Ethiopian genera <strong>of</strong>, 303range <strong>of</strong> Oriental genera <strong>of</strong>, 374range <strong>of</strong> Australian genera <strong>of</strong>, 476Ungulate, European Pliocene, 112Miocene <strong>of</strong> Greece, 115European Miocene, 119Indian Miocene, 121European Eocene, 125N. American Post-Pliocene, 130N. American Tertiary, 135<strong>of</strong> Brazilian caves, 144S. American Pliocene, 146Urania <strong>of</strong> Madagascar, 282Ursidce, N. American Tertiary, 135in Brazilian caves, 144Ursitaxus, Indian Miocene, 121Ursus, Post-Pliocene, 112Indian Miocene, 121Vanga <strong>of</strong> Madagascar, figure <strong>of</strong>, 278Varanus, Miocene <strong>of</strong> Greece, 116Indian Miocene, 123Vertebrata, summary <strong>of</strong> Palaearctic, 1S6summary <strong>of</strong> Ethiopian, 255summary <strong>of</strong> Oriental, 318summary <strong>of</strong> Australian, 397Vespertilio, European Eocene, 125Viperus, European Miocene, 1(55Vivcrra, European Pliocene, 112European Miocene, 118Viverridce, European Miocene, 118European Eocene, 125W.Walden, Viscount, on birds <strong>of</strong> PhilippineIslands, 346on birds <strong>of</strong> Celebes, 428Washakius, N. American Tertiary, 134Waterhouse, Mr. G. E., on classification olrodentia, 90on classification <strong>of</strong> marsupials, 91West African sub-region, description <strong>of</strong>, 262mammalia <strong>of</strong>, 262birds <strong>of</strong>, 262Oriental or Malayan element in, 263river scene <strong>with</strong> characteristic <strong>animals</strong>, 264reptiles <strong>of</strong>, 264amphibia <strong>of</strong>, 264Oriental and Neotropical <strong>relations</strong> <strong>of</strong>, 265insects <strong>of</strong>, 265land-shells <strong>of</strong>, 265islands <strong>of</strong>, 265West Australia, peculiar birds <strong>of</strong>, 441Whydah finch <strong>of</strong> W. Africa, figure <strong>of</strong>, 264Wollaston, Mr. T. V., on <strong>the</strong> coleoptera <strong>of</strong> <strong>the</strong>Atlantic Islands, 209 *on <strong>the</strong> wings <strong>of</strong> <strong>the</strong> Madeiran beetles, 211on <strong>the</strong> origin <strong>of</strong> <strong>the</strong> insect fauna <strong>of</strong> <strong>the</strong>Atlantic Islands, -J14on coleoptera <strong>of</strong> <strong>the</strong> Cape Verd Islands, 215on beetles <strong>of</strong> St. Helena, 270Xenurus, in Brazilian caves, 145Xiphodontidw, European Miocene, 119Z.Zeuglodontidce, N. American Tertiary, 140Zonites prisms. Palaeozoic, 169Zoological characteristics <strong>of</strong> Palaearctic region,181Ethiopian region, 252Oriental region, 315Australian region, 390Zoological regions, discussion on, 50END OF VOL. I.


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