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list and offered an analysis of our obligative spidersto allow comparison with such animals of othergroups.Recognition of troglobites is not so precise ascould be wished and the certainty of this specialstatus is a matter requiring much more informationthan mere presence of a blind taxon in a cave. Thereare some spiders with attributes of troglobites thatdo not live in caves. Blind spiders are found in termitenests in Africa and others with greatly reducedor obsolete eyes live in ant nests or under moist humusof tropical and temperate forests on many continents.Obsolescence of the eyes is not necessarily ameasure of length of cave life or even indication ofit. A few cavernicoles with essentially normal eyeshave not been taken outside of caves; a prime exampleis Phanetta suhterranea (Emerton), a cavernicolewith small eyes studied by me on specimensfrom 141 caves ranging from Pennsylvania and Illinoissouth into Alabama, with so far no epigeanrecords, but which live deep within caves often underlitter and rocks. This taxon is listed arbitrarily asa troglophile. By contrast, in other instances I haveassumed a more generous attitude in identifyingtroglobites and list a number with degenerated eyesin trivial remnants that are seemingly limited tosingle caves or cave systems. Spider troglobitesshare most of the special features of cave animals ofother groups. The integument is pallid or whitishbecause of loss of pigment and often is less stronglysclerotized than in surface forms. The eyes showvarying degrees of regression from mere loss ofpigment to complete disappearance of all externalvestiges. This occurs in regular pattern in spiderswith loss first of the anterior median pair and thengeneral reduction and gradual disappearance of thethree remaining diads. The troglobite status is rarelyreached without strong modification of the eyes:most of our obligative cavernicoles are eyeless ornearly so. Finally, in spiders and many other groupsthe appendages tend to become long and much thinnerthan in surface forms.Spider troglobites have been derived from cryptozoictaxa of both primitive and specialized groups.Most are sedentary and spin webs in crevices oncave walls and ceilings or in or under detritus on thefloors. Within these webs spiders have minimizedneed for sight and substitute an increasingly keenchemotactic sense centered in the hairs and setae oftheir appendages. Such sedentary types move to newstations only to satisfy food and climate needs. Allof our American spider troglobites and most fromworld centers come from spiders of this type. Thefew hunting spiders that attain troglobite rank havelikewise de-emphasized sight as a life requirement.A prime example is an eyeless troglobitic wolf spider,Adelocosa anops Gertsch, which lives in KoloaCave, Kauai, Hawaii. Also in Hawaii occurs Lycosahowarthi Gertsch, a yellowish, essentially blindtroglobitic species with vestigial eyes, known fromseveral caves.In 1985 Barr and Holsinger published a basic paperentitled "Speciation in Cave Faunas" in which amap was included of the major limestone areas ofthe United States, in which most of our cavernicolespiders are to be found. Spider troglobites are nowknown from the three principle areas as follows: 1.The Appalachian Valley and Western Plateaus; 2.The Texas Region; and 3. The cave areas of CentralCalifornia. Absence of troglobites from variouslimestone nuclei outside of the cited districts reflectsmore than a failure of intensive collecting and indicatesunlikely presence of troglobites. From none ofthe caves in Utah, Idaho, Arizona, and New Mexicohave come spider troglobites in spite of intensivecollecting by arachnologists. Carlsbad Caverns, oneof our great caves, thus far lacks truly derivativetypes. It is likely that additional troglobites may befound in areas south of these three regions but fewwill be found much to the north of the indicatedlimits.Spider Troglobites of AppalachiaWithin this broad province are found manycaves, including such giants as Manunoth and manyothers famous for their faunas. The great wealth ofcave habitats of this region early became an explOrativearea for speleologists who found spidertroglobites relatively numerous: at least 18 are soclaimed and they belong to eight genera of fourfamilies. Especially numerous are taxa of the familiesLinyphiidae and Nesticidae and include 14 ofthe known troglobites. The first known Americanspider troglobite, Anthrobia mammouthia, a smallwhitish spider without trace of eyes, was describedby Tellkampf in 1844 from Manunoth Cave in Kentucky;it is still the only species of its genus. Severaladditional cavernicoles were added by various studentsso that in 1960 Nicholas was able to list ninepresumed obligative taxa from the United States:five of them do not now qualify for that status.American cave spiders are now known from quitelarge collections from many localities, so it is nowpossible to present a favorable picture of our faunaon the basis of new publications and much unpublisheddata in my possession. Our fauna showssimilarity to the European one, with many of the76
same families and genera, but also carries its distinctiveAmerican flavor.The list of names and authors that follows refrainsfrom duplicative listing available in our standardbibliographic journals. For the sake of completeness,undoubted new species taxa, some beingstudied elsewhere, are included for general interestand tentative completeness.List of the Spider Troglobites of AppalachiaFamily LinyphiidaeAnthrobia mammouthia Tellkampf, 1844. Kentucky,some bordering states.Porhomma cavernicolus (Keyserling), 1886.Widespread from Illinois, Arkansas, Virginia, toAlabama.lslandiana cavealis Ivie, 1965. Kentucky.lslandiana muma Ivie, 1965. Virginia.lslandiana speophila Ivie, 1965. West Virginia.Smilax, new species. Iowa, Wisconsin.Oreonetides, new species. Virginia.Family NesticidaeNesticus dilutus Gertsch, 1984. Tennessee.Nesticus furtivus Gertsch, 1984. Tennessee.Nesticus valentinei Gertsch, 1984. Tennessee.Nesticus barrowsi Gertsch, 1984. Tennessee.Nesticus jonesi Gertsch, 1984. Alabama.Nesticus stygius Gertsch, 1984. Tennessee.Nesticus georgia Gertsch, 1984. Georgia.Family LeptonetidaeAppaleptoneta georgia (Gertsch), 1974. Georgia.Family DictynidaeCicurina (Cicurina), new species. Georgia.Cicurina (Cicurina), new species. Alabama.Cicurina (Cicurella), new species. Alabama.Spider Troglobites of CaliforniaThe small cluster of spots in the middle of California(Barr and Holsinger, 1985, p. 313) identifiesrich cave localities in the foothills of the Sierran andCascadian Mountain Ranges. All the troglobitesbelong to different genera than those of Appalachiaand Texas. Most California caves are quite small butintensive collecting by Richard Graham, DarrellUbick, and a growing group of interested studentshave indicated presence of a rmmmum of 20troglobites belonging to five families. The familiesand seven genera are all strongly represented in thewhole region by numerous epigean species. TheCalifornia region has a very large fauna of crypticagelenines (these now assigned to two special families),so it is not surprising that at least nine speciesrepresenting three distinct genera have producedtroglobites. Missing from this list is any troglobiusspecies of Cicurina, which has produced a multiplicityof eyeless taxa in Texas and is also representedby three taxa in the Appalachia area.It need not be surprising that only one of thelisted troglobites has been formally named but therichness of the fauna in the cited area marks it as aspecial project. The presence of the genus Telema(its single eyeless species known only from a fewcaves in the French Pyrenees), earlier assigned tothe American genus Usofila, is now known fromCalifornia by numerous eyed species and severaleyeless ones.The listed undescribed taxa of the Californian regionare being studied by several students.List of the Spider Troglobites of CaliforniaFamily NesticidaeNesticus potterius (Chamberlin), 1933. ShastaCounty.Family LinyphiidaeLinyphiinae, new species. Amador County.Linyphiinae, new species. Tulare County.Family LeptonetidaeCalileptoneta, new species. Trinity County.Family CybaeidaeCybaeus, new species. Shasta County.Cybaeozyga, new species. Shasta County.Cybaeozyga, new species. Shasta County.Cybaeozyga, new species. Shasta County.Family DictynidaeBlabomma, new species. Shasta County.Blabomma, new species. Calaveras County.Blabomma, new species. Calaveras County.Blabomma, new species. Tulare County.Blabomma, new species. Calaveras County.Blabomma, new species. Amador County.77
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PREFACEThe present volume is the se
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TABLE OF CONTENTSHOLSINGER, JOHN R.
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the state of Coahuila in northern M
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Fig. 2.-Anesia welboumi, new specie
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\\. \ - -' ..........---~\ I// --..
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Mexiweckelia hardeni, new speciesFi
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2 sets of 1 or 2 setae each; dactyl
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AFig. 7.-Holsingerius smaragdinus,
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have as many setae on the inner pla
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Antenna 1 about 33 % length of body
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Of biogeographic interest for the h
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Bowman, T .E. 1992. Two subterranea
Page 35 and 36: A note by Scott Harden that accompa
Page 37 and 38: ~~..~ ~~ I))\\ "-,'.=bFig. 3.-Speoc
Page 39 and 40: unarmed, except in pleopod 2, which
Page 41 and 42: Cokendolpher, LC., and l.R. Reddell
Page 43 and 44: lobes. The larger setae vary greatl
Page 45 and 46: the relationships of the order and
Page 47 and 48: zomids their absence is considered
Page 49 and 50: some species could be either split,
Page 51 and 52: have small pores over the surface o
Page 53 and 54: inhabits tropical deciduous forest
Page 55 and 56: huitvnolotitlensis from A. stygius
Page 57 and 58: 8-10, figs. 5-7; Rowland, 1973c:136
Page 59 and 60: (in row) and one pair large posteri
Page 61 and 62: (0.34); tarsus 0.64 (0.58); total 4
Page 63 and 64: Abdomen: Tergite I with two pairs a
Page 65 and 66: setae near posterior margin. stemit
Page 67: setae, and ten ventral setae. Stemi
Page 71 and 72: 1,980 m elev., 26 Dec. 1986 (T. Tre
Page 73 and 74: asitarsal-tarsal proportions: 15:4:
Page 75 and 76: and one pair setae at base of proce
Page 77 and 78: Cephalothorax: Propeltidium 1.66 mm
Page 79 and 80: Male adult unknown.Immature paratyp
Page 81 and 82: IV: trochanter 1.20 (1.10); femur 3
Page 83 and 84: Cokendolpher, 1.C. 1981. The order
Page 85: Gertsch, W.J. 1992. Distribution pa
Page 89 and 90: species from the United States and
Page 91 and 92: the posterior median pair short and
Page 93 and 94: great many species of North America
Page 95 and 96: Key to the Eyed Females1. Eight eye
Page 97 and 98: 39111012Figs. 1-12.-Ventral and dor
Page 99 and 100: 1419 23Figs. 13-24.-Ventral and dor
Page 101 and 102: 27 293334Figs. 25-36.-Ventral and d
Page 103 and 104: Figs. 37-48.-Ventral and retrolater
Page 105 and 106: Cicurina blanco, new speciesFigs. 7
Page 107 and 108: Type-data.-Female holotype from ins
Page 109 and 110: Description.-Female holotype: Lengt
Page 113 and 114: lengths: first femur 2 rom, fourth
Page 115 and 116: Cicurina pablo, new speciesFigs. 10
Page 119 and 120: canal in nearly vertical posItion;
Page 121 and 122: Cicurina vespera, new speciesFigs.
Page 123 and 124: procurved canal across sac; connect
Page 125 and 126: Cicurina caverna, new speciesFigs.
Page 127 and 128: with sac of similar size set in obl
Page 129 and 130: February 1964 (J. Reddell, D. McKen
Page 131 and 132: County: Diamond Cave, 16 August 196
Page 133 and 134: Roth, V.D. 1992. A new and first tr
Page 135 and 136: LITERATURE CITEDBarr, T.C. 1963. Ec
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Muchmore, W.B. 1992. Cavernicolous
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Species of Aphrastochthonius are kn
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FAMILY NEOBISIIDAE CHAMBERLINGenus
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trochanter 2.5 (2.6), femur 5.55 (5
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ottom of entrance pit, Ogle Cave (O
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tactile seta on tibia and basitarsu
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Missouri may be conspecific (unpubl
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Trichobothriotaxy of chela generall
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Female (figures given first for all
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transverse furrows; eyespots not ev
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Reddell and W. Russell); I female f
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and Acuminochernes, along with the
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Chamberlin, J.C. 1946. The genera a
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(0.36); chela (without pedicel) 2.0
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although not recently studied is no
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and stylar outgrowths, present in s
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in Phalangodes (et al.) is thick an
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pairs) are found in six species: T.
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elated species IS probably best int
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legs than expected (2.6-3.2). This
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~ 3.3 are synapomorphic); all trogl
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TAXONOMYTEXELLA Goodnight and Goodn
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14. BK absent (Figs. 177, 180). SA
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Figs. 8-11.-Texella bijUrcata (Brig
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Description.-Total body length, 1.5
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male examined closely has fewer set
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Figs. 26-29.-Texella kokoweej, new
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apical region which loses the apica
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Figs. 38-41.-Texella shoshone, new
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Distribution.-Known only from the t
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Figs. 52-55.-Texella brevistyla, ne
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Texellajungi, new speciesFigs. 60-7
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Figs. 66-69.-Texellajungi, new spec
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than S; SA with laterobasal carina
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~81//J/'/ ;'?/ ~~.........--~~I, II
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cylindrical, retina and cornea abse
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Figs. 93-96.-Texella cokendolpheri,
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SA with well developed prong and re
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Figs. 105-108.-Texelia mulaiki Good
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and McCarty Caves, which are known
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Color orange. Body of medium rugosi
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Notes.-The type locality was errone
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Figs. 128-131.-Taella reyesi, new s
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Figs. 136-139.-Texella reyesi, new
Page 227 and 228:
Page 229 and 230:
Page 231 and 232:
1989 (W. Elliott, J. Reddell, and M
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Table 3.-Continued.# locality sex S
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mesoapical; patella, 2 mesal; tibia
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Figs. 162-165.-Texella gmbbsi, new
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Figs. 166-169.-Texella diplospina,
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Figs. 174-177.-Texella renkesae, ma
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Figs. 178-18\.-Teulla spinoperca, n
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Distribution.-Known only from Fayet
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Figs. 190-193.-Texellafendi, new sp
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CLASSIFICAnONTexellabifurcata group
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Chandler, D.S. 1992. The Pselaphida
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Key to Species1. Abdominal segments
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stemite VI slightly impressed at ba
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vertexal carinae, and the laterally
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is associated with rotten woods (Ch
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small ventral carina near base, pro
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Grigarick, A.A., and R.O. Schuster.
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Cicurifla (Cicurella) holsiflgeri G
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