The Geographical and Ecological Distribution of Arboreal Psocoptera

Annu. Rev. Entomol. 1985.30:175-196. Downloaded from arjournals.annualreviews.org 

by Mr. Bas van Berkum on 10/10/07. For personal use only. 

Annual Reviews 

www.annualreviews.org/aronline 

Ann. Rev. EntomoL 1985.30:175-96 

Copyright © 1985 by Annual Reviews Inc. All rights reserved 

THE GEOGRAPHICAL AND 

ECOLOGICAL DISTRIBUTION 

OF ARBOREAL PSOCOPTERA 

Ian W. B. Thornton 

Department of Zoology, La Trobe University, Bundoora, Victoria, Australia 3083 

INTRODUCTION 

The order Psocoptera (the psocids) consists of some 36 families with perhaps 

5,000 species (some 3,000 described), and is generally considered to be the 

monophyletic sister group of the order Phthiraptera within the superorder 

Psocodea (32, 38, 69, 70). Fossils are known from the Permian, and most early 

fossils are generally placed in a separate suborder, Permopsocida. The earliest 

psocids are from the Kansas Lower Permian (75), and by the Oligocene most 

are referable to recent families and genera. The general biology and classification 

of the order has been the subject of several recent reviews (30, 62, 75, 

102). 

Some psocids occur in ground litter for a part or whole of their life cycle; 

others are found on rocks and in the nests of birds, rodents, and termites. A few 

species have been found in the feathers and fur of living birds and mammals. A 

number appear to live on herbs and grasses, and a few in moss, whereas others 

are found in caves (particularly Prionoglaridae and Psyllipsocidae). Several 

species are found in domestic habitats and use a wide range of foods that occur 

there. 

In this paper I attempt to review the present state of knowledge on the 

geographical and ecological distribution of those Psocoptera that inhabit trees 

and shrubs. The literature reviewed is largely confined to those publications 

appearing since 1964; the bibliography by Smithers (72) may serve as a source 

for earlier references. 

0066-4170/85/0101-0175502.00 

175





176 THORNTON 

Ecological Role 

FEEDING AND BIOMASS Although usually designated as scavengers or detritivores 

when ecological roles are assigned to the various components of 

ecosystems, arboreal Psocoptera are in fact grazers, and their pastures are the 

microepiphytes (fungi, algae, and lichens) that grow on the bark and. leaves 

trees and shrubs and in the litter. Some also take pollen grains. The peculiarities 

of psocid mouthparts and the massive clypeal muscles may in the future be 

shown to be specifically related to this grazing type of feeding. The ecological 

importance of psocopterans in forest ecosystems has until recently been either 

neglected or assumed to be small. However, considerable information is now 

available on their role in temperate and, to a lesser extent, tropical forest 

ecosystems as a major component of the guild of microepiphyte feeders. 

In temperate forests the primary productivity contribution of microepiphytes 

may be comparable to that of the herb layer (92), and grazing psocids may make 

up a significant part of the biomass. Densities of over 4000/m2 on the bark of 

larch (Larix decidua), equivalent to over 6000/m2 of land surface, have been 

recorded in Yorkshire, England, during late summer (16). The peak biomas s 

two species of Mesopsocus was about 1.2g/m2 of ~ bark surface, or about 2g/m 

of ground surface. This is comparable to the biomass of various kinds of big 

game animals such as antelopes feeding on grasslands, and at peak times the 

grazing of microepiphytes by psocids can be very intense. 

In the tropics, psocid densities are much lower. However, in an altitudinal 

transect of canopy arthropods from sea level to 2400 m subtropical Hawaii 

(26), psocids were frequent and abundant on the two dominant trees examined 

and outnumbered all other taxa combined at mid and high elevations. Clearly, 

psocid populations can be important primary consumers and saprophages. 

PREDATION " In Britain a polyphagous mite of the genus Anystis is frequently 

recorded as a predator of psocids. In Yorkshire it may be responsible for much 

of the high mortality of first instar larvae of Philotarsus picicornis besides 

causing significant egg mortality in Mesopsocus species (23). 

In southern England, spiders, mites, opilionids, neuropteran larvae, anthocorids, 

earwigs, and carabids have been reported to attack egg batches of 

foliicolous psocids. Some 26 predators have been recorded (15, 99): the most 

important are spiders (by far), mites, opilionids, neuropteran larvae, and 

coccinellid beetle. Of the three major groups of epiphyte herbivore:s on larch, 

the majority of predators select Psocoptera and Homoptera rather than the 

numerically more abundant Collembola. 

Ants (Crematogaster sp.) in Brazil have been seen at night attacking and 

carrying off archipsocids that live under large communal sheet-webs. In Florida 

archipsocids are preyed upon by Argentine ants and reduviid bugs.





DISTRIBUTION OF ARBOREAL PSOCOPTERA 177 

In Hawaii, larvae of the moth genus Eupithecia are ambush predators of 

psocids (57) and will feed on them in the laboratory. This guild of predaceous 

Lepidoptera is endemic to the Hawaiian Islands, an area that also contains 

several endemic complexes of psocopterans. These psocids constitute a considerable 

proportion of available prey for such predators. Larvae of endemic 

Neuroptera in Hawaii also prey on psocids, which are thought to have been 

their principal prey before the introduction of aphids. 

Sphecid wasps in Europe are known to take psocids occasionally; the 

stem-nesting Rhopalum claviceps, however, appears to specialize in psocids as 

prey. In France, R. claviceps is reported to attack 16 species, and many nests 

are wholly provisioned with psocids. In a Luxembourg garden, 25 species were 

found in 23 nests (68). From 1980 to 1983, N. Schneider (personal communication) 

examined 125 nests in Luxembourg and in 86 that were provisioned, 

all the prey were psocids (of 28 species), with an average of 24 per 

cell (270 cells). The wasp does not appear to discriminate between species 

between winged or apterous adults and nymphs. Apart from R. claviceps, 

arthropod predators appear to take psocids only as part of a wider spectrum of 

prey. 

Psocopterans are the hosts of hymenopteran egg and larval parasites, some of 

which seem to specialize on them as hosts. Some species even appear to show 

host-specificity within the order Psocoptera. 

Mymarid parasites of the genus Alaptus have been reared from psocids in 

many parts of the world, and it has been suggested that the geographical 

distribution of Alaptus coincides with that of arboreal psocids (62). In England, 

there is evidence of habitat-based host specificity, with Alaptus life cycles 

closely correlated with those of their hosts. In both England and Jamaica, 

parasitization has been related to the type of covering (feces, silk, or both) 

the egg masses of different psocid species, even when they occur on the same 

trees (23, 93). 

Braconids of the genera Euphoriella and Leiophron parasitize psocid 

nymphs in the Holarctic Region and in Central America. They have not been 

recorded elsewhere, and New, who has dissected large numbers of psocid 

nymphs of several genera in South America and Australia, has speculated that 

braconid parasitization either does not occur in those continents or is extremely 

localized (62). 

Several papers cite circumstantial evidence Of bird predation on psocids, and 

others from various parts of the world actually record predation by insectivorous 

birds (12, 13, 64, 85, 95, 98). In aviary experiments in England, more dark 

than light morphs of apterous females ofMesopsocus unipunctatus were taken 

by great tits (Parus major) from larch branches with a covering of epiphytes 

than from bare branches; Popescu (65) suggested that visual selection by birds





178 THORNTON 

could maintain the color polymorphism in this industrial melanic psocid, 

provided that there was competition for "cryptic" resting sites. 

Various forms of crypsis have been noted occasionally in other psocids, and 

avian predation may have provided selection pressure for this adaptation. 

Dorsal projections on the abdomen of apterous females of Hexacyrtoma, 

Gibbopsocus, and Camelopsocus resemble thorns and leaf scar swellings, and 

species of Calopsocus and Calocaecilius resemble coccinellid beetles in 

appearance and movement. Nymphs of some Psocidae cover "glandular hairs" 

with adherent debris, and others make web shelters into which debris is 

incorporated. Many winged forms, such as myopsocids, have intricately patterned 

wings resembling their mosaic background. In Brazil, epipsocids and 

archipsocids are partially cryptozoic by day and active at night, and ptiloneurids 

tend to be nocturnal. In their region of sympatry two North American 

species of Tapinella have a strikingly similar body pattern, and Mockford (47) 

has suggested that it may be advantageous for species to share a warning 

pattern. 

GEOGRAPHICAL DISTRIBUTION 

Although Psocoptera have been found in the sub-Antarctic islands, Tierra del 

Fuego, Finnish Lapland, and in Outer Mongolia, few species are found in these 

extreme environments. For example, Meinander (36), who collected extensively 

in northem Finland, found only four species. In contrast, diversity is 

great in the tropics; Broadhead collected 295 species in Panama, and in 

Trinidad 116 species have been recorded from mango alone (17). 

Several species of psocids, most of which are domestic, are cosmopolitan in 

distribution, and others have wide tropical ranges. However, many regional 

faunas can be identified and many groups have limited distributions. Probably 

the least known faunas are those of southern Africa and tropical South America; 

and the Oriental region (notably peninsular India), the Indochina region, and 

Wallacea (the region between Wallace’s and Weber’s lines that is a transitional 

zone between the Oriental and Australian regions) have been underc, ollected. In 

contrast, the faunas of Europe and North America are fairly well known, but 

even here discoveries of considerable biogeographical interest have; been made 

in recent years. 

Distribution patterns may be related to plate tectonic events in only a few 

cases. The small family Sphaeropsocidae consists of the genera Sphaeropsocus, 

which is known only from oligocene amber, Sphaeropsocopsi,~r, which has 

five species in Chile and one in each of southern Argentina, Tasmania, and 

southern Africa, and Badonnelia, which has four species in Chile and one 

found in domestic situations in Europe. Apart from the European species, this 

family has a typical Gondwanian relict distribution and is the only known 

example of this type of distribution in the order Psocoptera (4, 5).






Another small family whose distribution can be explained, at least tentatively, 

by plate movements is the Asiopsocidae. The three constituent genera occur 

in Mongolia, Mexico, Arizona, and Spain (Asiopsocus), the neotropics, southern 

Florida, and tropical Africa (Notiopsocus), and Brazil (Pronotiopsocus). 

This pattern suggests a very old, relict distribution. Character-state analysis 

showed Pronotiopsocus to be the sister group of the Notiopsocus-Asiopsocus 

line, of which Asiopsocus is the northern and Notiopsocus the southern vicariant. 

Mockford (52) suggested that the first phylogenetic event occurred prior 

to the breakup of Pangaea, and the second was coincident with it. The absence 

of Notiopsocus from Madagascar, India, and Australia may be because this 

evolutionary line reached Africa from South America too late to occur on the 

plate containing Antarctica, India, and Australia, which separated from Africa- 

Madagascar 205-160 Myr BP, and too late also to occur on Madagascar, which 

separated from Africa 172-90 Myr BP. Mockford (52) suggested that after the 

Pleistocene emergence of the Panama isthmus, Notiopsocus moved north into 

Mexico and southern Florida. 

The distribution of the larger family Philotarsidae may also be related to plate 

movements (81, 83). One subfamily, the Zelandopsocinae, with 65 species 

three genera, is confined to the Australian plate and its fragments; although 

well represented in New Caledonia and New Zealand, it is absent from the 

Outer Melanesian Arc. Its sister group, the Philotarsinae, with 85 extant 

species, consists of two tribes--Philotarsini (Philotarsus and Haplophallus) 

and Aaroniellini (Aaroniella, Latrobiella, and Tarsophallus). Haplophallus 

has a gondwanan distribution (Africa, Australia, South America) with limited 

extensions, and it is absent from the Holarctic; Philotarsus is basically Holarctic. 

Of the Aaroniellini, Aaroniella is largely tropical, virtually absent from the 

Palaearctic and Africa, and has only one species in Australia. In contrast, its 

sister group, Latrobiella, occurs on the Australian plate and its fragments, on 

the Bismarcks and Solomons, and in southern South America. Tarsophallus is 

known only from the highlands of East Africa (two species) (55) and, although 

no females are available, it appears to be the African sister group of Aaroniella 

and Latrobiella. 

Europe 

The number of species known from France has increased from 71 to 97 in the 

last 40 years, and 4 new genera have been discovered in Europe during the last 

decade. These include a litter-dwelling amphipsocid from southern France, 

which is only the second representative of this family to be known from Europe; 

a new genus of the Mesopsocidae; a new electrentomid genus with a beetle-like 

habit, which is the first Troctopsocidae from outside the neotropics; and a new 

genus of the Pseudocaeciliidae, a family previously unknown from Europe and 

best represented in the Oriental and Australian regions. The last genus, with





180 THORNTON 

three-segmented tarsi, is unique in the family; it is primitive in several respects 

but has apomorphous characters in common with Elipsocidae. These recent 

discoveries are of biogeographical and phylogenetic interest and underscore the 

incompleteness of knowledge, even of such a well-investigated are, a. 

Recent work on European Elipsocidae (29) has shown that Hemineura dispar 

occurs in Scandinavia, Estonia, and Germany and at high altitudes in Switzerland 

and France and in Moravia. Badonnel regarded the species as a postglacial 

relict; its southern limit is in the mountains of southern France where two 

congeneric species also occur, H. bigoti and H. hispanica, the latter also 

known from Spain,. Whereas dispar evidently has two generations per year, at 

least in Moravia, the two Mediterranean species are univoltine winter species. 

Both are parthenogenetic, with apterous females, but they differ from one 

another in aspects of biology and behavior. It is possible that all three species 

are glacial relicts, since dispar alone has survived in the north and extended to 

high locations in the south of its range; the other two persist only in the 

northwest Mediterranean area where they have a long resting stage in the egg 

and a short active generation in winter. 

The Atlantic islands have also recently received considerable attention (35, 

37). The Canary Islands, which are nearest to the mainland, have 28 species (10 

endemic), followed by Madeira, with 19 (4 endemic). The Azores have 

species (4 endemic), and only 3 species (2 endemic) are known from the 

Verde Islands. Atlantopsocus (Psocidae), a genus that occurs in the Atlantic 

islands, comprises three species and is known from Ireland, Morocco, the 

Canaries, Madeira, and the Azores. Its closest relative appears to be Camelopsocus 

(54), of which five species are known from western North America. 

Mesopsocus, which is well represented in northwest Africa, extends only to the 

Canaries, where one species is endemic; this genus, in which most species have 

flightless females, is also absent from Madagascar (below). Twelve psocopteran 

species (four endemic) are known from Saint Helena (7); three of these 

cosmopolitan and four occur in Northwest Africa and the Atlantic islands 

mentioned above, possibly introduced by Portuguese maritime activity. Of the 

endemics, three have closely similar counterparts in West and Central Africa, 

and one is evidently an older endemic of unknown affinities. 

Ethiopian Region 

Coverage of Africa has been patchy; almost 80% of the species are known only 

from the type locality. The known fauna totals 493 species, and its relationships 

with those of Madagascar and the Mascarene islands have recently been 

considered. In West Africa 160 species are known, in East Africa 155 (20, 21), 

and in southern Africa 239 species are recorded from Angola (9) and 90 from 

other areas. Africa is rich in a number of families while others that are well 

represented elsewhere, such as Lachesillidae and Philotarsidae, are absent or 

have only few species.






The family Mesopsocidae appears to have its center of diversity in Africa; it 

is absent from Neotropical and Australian regions and Madagascar and only 

weakly represented in the Nearctic and Oriental regions (21). The genera 

Hexacyrtoma and Labocoria are African, and Psoculus occurs in Europe and 

Africa. The largest genus, Mesopsocus (40 species, 29 described), has 

species in the Ethiopian region, 14 in the Palaearctic (three also occurring in the 

Nearctic), and one in the Oriental region. 

Of three East African mountain massifs, Mount Kenya, the Aberdares, and 

the Empakaai region, the first two, which are closest to one another, were 

found to have the most similar psocopteran faunas and each had 19% endemicity, 

whereas the endemicity of the more isolated Empakaai region was 41%. 

There is a parallel here with geographical archipelagos (80). A closely related 

group of four species of Mesopsocus, all with apterous females, occurs at high 

elevations on mounts Kenya and Kilimanjaro (one species), Mount Kenya and 

the Aberdares (one), and Empakaai (two). A fifth species with apterous 

females is evidently restricted to the shores of Lake Naivasha, and a sixth is 

found on Mount Kenya, the Aberdares, and Empakaai. In the last case, the 

differentiation of an Empakaai species appears to have been impeded by the 

presence (unusual in Mesopsocus) of winged females (21). 

In West Africa the psocid fauna of the 250 km wide Togo-Benin gap, which 

separates the rain forests to the west from those of Nigeria and Central Africa to 

the east, has affinities with eastern rather than western regions (101). Six of the 

28 species collected from the gap link it with the eastern regions; no Ivory Coast 

or Guinea species occurred in the collections. Of the 12 new species found in 

the Gap, only 2 have affinities with western species, and both are widespread; 

the most similar counterparts of the other 10 species occur to the east. More 

collecting to the west is needed to confirm the isolation of the western forest 

bloc species from those of the main body of equatorial rain forest, a finding 

which would lend support to the hypothesis that the lowland forest isolate to the 

west was confined to refugia during arid glacial periods. 

The Madagascar fauna has all the features of insularity; 166 of the 197 

species are endemic (84%) as are 6 of 37 genera (16%) (3, 6). Endemic species 

complexes exist in eight genera. The species flocks of Thylacella, Amphientomum, 

Ctenopsocus, and Blastopsocidus are each thought to derive from a 

single founder, whereas those of Myopsocus and Ptycta represent two lines (3). 

Explosive speciation is also evident in Caecilius (26 species, 24 endemic) and 

Amphipsocus (29 species, all endemic). Radiation is greatest in the amphipsocid 

subtribe Amphipsocina, which has three endemic genera. Notable absences 

of groups well represented in Africa include the genera Belapha, Fulleborniella, 

Ectopsocopsis, Myopsocus, the families Elipsocidae and Mesopsocidae, 

and the subfamily Cerastipsocinae of the Psocidae. Nevertheless, links with 

Africa are evident. The genera Paracaecilius, Epipsocopsis, Harpezoneura, 

and Blastopsocidus are only known from Africa and Madagascar, and there are





182 THORNTON 

seven cases of species pairs with a member of each occurring in Africa and 

Madagascar; other species pairs appear to represent a more recent disjunction 

within Madagascar itself. 

The Mascarene Islands, Reunion and Mauritius, together have a fauna of 44 

species, half of which is endemic to the group. The islands have know faunas of 

34 (12 endemic) and 16 (6 endemic) species respectively, with 6 species 

common (4 endemic) (2, 10, 94). Three genera endemic--lsophanopsis 

(Caeciliidae) on Reunion and Mockfordiella (Caeciliidae) and Mascaropsocus 

(Peripsocidae) on both islands. The fauna has remarkably little affinity with 

that of Madagascar. For example, the relationships of the Reunion endemics 

with Madagascar are few and probably represent very old phylogenetic events; 

present conditions are unfavorable for faunal exchange. 

North America and the Neotropics 

About a score of North American species have Holarctic distributions. Some 

may be true relicts, but about half are probably the result of recent introductions 

(49, 50). 

Lachesilla, one of the largest genera in the order with almost 300 recognized 

species (27; A. N. Garcia-Aldrete personal communication), has its chief 

representation in North America (over 120 species endemic). Of the 18 species 

groups recognized, all are represented in the Americas and all but two in North 

and Central America. Only three groups extend beyond the New World to 

Africa (all), Europe (one), and Madagascar (one). 

Systematic studies of the cerastipsocine Psocidae suggest an early New 

World radiation with African and Oriental groups as derivatives (51). 

example of a monophyletic group confined to Central and North America is 

Kaestneriella (Peripsocidae) (56). 

Recent work on the North American fauna has concentrated on detailed 

continental distributions of species (25), species pairs (48), and species groups 

(14, 40, 41, 43-45, 53). Many of these distributions may be related to the 

effects of glacial changes (disruptions of previously continuous ranges, Pleistocene 

refugia, and early Tertiary invasions from the south). 

In view of the massive representation of Lachesilla on the continent it is 

remarkable that only four species occur in Cuba, only one of which is unknown 

elsewhere. Of the 68 psocids known from Cuba (8, 46), 35 (51%) are presumed 

endemics. About half have close relatives in the Caribbean area or the adjacent 

mainland, and in the Psocidae the genera Blaste and Indiopsocus have closely 

related groups of species that are probably autochthonous. There is no endemic 

genus. Most of the widespread tropical species occur also in Africa and were 

possibly introduced to the American tropics during the slave trade. 

Jamaica has also been surveyed (93), and 83 species are known./kgain, there






are no endemic genera, and again remarkably few species (three, two endemic) 

of Lachesilla occur there. Several genera have speciated in the highlands, and 

55 species (66%) are presumed endemics. Fourteen species are neotropical, 

five African, and nine cosmopolitan or tropicopolitan. There are few species in 

common with Cuba. 

In both Cuba and Jamaica there are rather few affinities with Mexico, and 

whereas links with the Floridean paxt of North America have been detected in 

the case of Cuba, South America has been suggested as a main source area for 

Jamaica. The very weak representation of Lachesilla and weak affinities with 

Central America argue against a land connection to the west, and the hurricane 

patterns in the Caribbean are in accord with the Floridean and South American 

affinities suggested. A further 40-50 species are known from other Caribbean 

islands (11, 24, 42, 43), over half of which are endemic to individual islands; 

they include 6 of the 10 species of the Caribbean genus Spurostigma, 3 of the 

endemic genus Troctopsocopsis, and a largely Caribbean group of Caecilius. 

Additional collections, both in the Caribbean and mainland areas to the south, 

are needed before hypotheses of Caribbean psocid biogeography can be formulated. 

Groups well represented in the neotropics include the families Pachytroctidae, 

Archipsocidae, Epipsocidae, and Liposcelidae and the amphipsocid genus 

Polypsocus. The Neurostigmatidae, Spurostigmatidae, Thyrosophorinae (Psocidae), 

and the genera Steleops and Dasydemella are confined to the region 

while the Troctopsocidae, Dolabellopsocidae, and Ptiloneuridae each have but 

a single species outside South America (in Europe, India-China, and Africa 

respectively). Endemic genera include Spurostigma, Isthmopsocus, and 

Dolabellopsocus. Some otherwise rather widespread families are absent (Psilopsocidae, 

Mesopsocidae) and others are relatively weakly represented (Lepidopsocidae, 

Amphientomidae, Ectopsocidae, Elipsocidae, Myopsocidae, Philotarsidae, 

and Stenopsocidae). It is thus possible to characterize the neotropical 

psocopteran fauna in these general terms. 

The known Galapagos fauna (90) comprises 40 species, and where affinities 

can be determined they appear to be largely with the above region. Two species 

of Indiopsocus are of particular interest. I. dentatus is related to species from 

the Caribbean, the gulf coast of Mexico, and southern Florida (E. L. Mockford, 

personal communication); it occurs at low and high altitudes and is on almost 

all Galapagos islands. I. acraeus appears to be most closely related to I. 

expansus, known from the Colombian Andes at 2750 m. In contrast to I. 

dentatus it is a high altitude stenotope, found only on high islands at elevations 

above 500 m. In both species, differences between island populations and 

between isolates on volcanoes of a single island have been recognized, and 

peripheral isolates are more differentiated than those on more centrally located, 

less isolated islands or volcanoes (I. W. B. Thornton, unpublished). Correla-





184 THORNTON 

tion between differentiation and isolation is greater for the stenotopic acraeus 

than for the eurytopic dentatus, and it appears that both are actively speciating 

on the archipelago. In view of the known geological history (31),. dispersal 

rather than fragmentation events appear to have been the likely precursors of 

differentiation both between archipelago populations and between the continental 

and Galapagos species. A related species, I. texanus, was an early and 

regular colonizer of Florida mangrove islets defaunated by tent fumigation 

(71). 

Temperate South America 

The temperate South American fauna is rather better known than that of the 

tropics, particularly west of the Andes. Many groups that are typical of the 

tropics are lacking, and the characteristic genus Ptenopsila is endemic to the 

region. The temperate fauna is less diverse; only a few species are both frequent 

and abundant, a situation similar to that in northern latitudes. The Elipsocidae 

appear to have radiated particularly in Chile, where the genera Eolachesilla and 

Roesleria are endemic and Nothopsocus is endemic to Chile and the Juan 

Fernandez Islands. Although Chile may be regarded as an ecological island 

with the Atacama desert to the north and mountains to the east, the Andean 

barrier declines in significance in the south as the range becomes lower and 

eventually runs into the sea. The Valdivian forests possess a rich and varied 

fauna; the more southern Magellanic forests, dominated by Nothofagus, are 

impoverished, with an attenuated fauna (63). 

The arboreal psocid fauna of Argentina appears to include southward incursives, 

and many species there do not occur in Chile. However, in the south, at 

least two of the four most abundant Chilean species are trans-Andean. The 

Embidopsocinae and Pachytroctidae, which are well represented in Brazil and 

Argentina, have not been found in Chile, and Sphaeropsocidae appear to be 

rare east of the Andes yet have nine species in Chile (4). In other genera several 

species pairs with a member on each side of the Andes have been noted (1). 

Of the Juan Fernandez group, only Robinson Crusoe (Mas a Tierra) has been 

surveyed (88). The nine known species comprise an endemic complex of five 

Nothopsocus species, a species widely distributed it/Chile, and three with an 

extensive range and probably recently introduced. Remarkably, on the Chilean 

mainland Nothopsocus is represented by only two species, which are neither 

widely distributed nor common. In contrast, the genus Drymopsocus (also 

Elipsocidae), which has four widely distributed species in Chile, was not found 

on Robinson Crusoe. Possibly, ecological exclusion is responsible for these 

contrasting patterns. The families Caeciliidae and Philotarsidae, both common 

in Chile, also appear to be absent from Robinson Crusoe. The Galapagos and 

Robinson Crusoe faunas are strikingly different; elipsocids are not known from 

the Galapagos, and only Peripsocus nitens, abundant on the Chilean mainland, 

is probably common to the two.






Links between the southern South American fauna and that of the Australian 

region are few--the Sphaeropsocidae and the genera Haplophallus (Philotarsidae) 

and Drymopsocus are examples--and the most typical species of temperate 

South America, Ptenopsila delicatella, has no analogue in temperate 

Australia. 

Australia and New Zealand 

Tropical Australia’s fauna includes a few representatives of basically Oriental- 

Papuan-Melanesian groups, such as the Calopsocidae, Psilopsocidae, subgroups 

of the Myopsocidae, Psocidae, and Amphipsocidae, and other more 

widespread tropical groups, e.g. Archipsocidae, Hemipsocidae, and Epipsocidae, 

which are absent from temperate regions of the continent. In the south the 

Elipsocidae (Propsocinae) and Philotarsidae are characteristic. Few strong 

affinities with islands of the western Pacific have been discerned. The philotarsid 

subfamily Zelandopsocinae is endemic to Australia and fragments of the 

Australian plate, and for this distribution a vicariance explanation is sufficient 

and appropriate. The elipsocid subfamily Propsocinae also appears to have its 

headquarters in this region: the widespread Propsocus occurs in Australia, 

Tasmania, South Africa, Chile, and Hawaii, Pentacladus in Australia, Antarctopsocus 

on Marion Island and the Crozet Islands, and Spilopsocus in Australia, 

Lord Howe Island, New Zealand, and Campbell Island. 

In general, the New Zealand fauna (73) appears to be a reduced Australian 

fauna (74). Here biological interest centers on the genus Austropsocus (Philotarsidae), 

which in this area exhibits a tendency towards flightlessness. Most 

species of the main islands have flightless females, while the species on the 

Chathams and that on the Campbell Plateau islands and Macquarie are flightless 

in both sexes. Except in the case of Macquarie, the presence of related 

flightless island species can be sufficiently explained by fragmentation events; 

the species common to Macquarie and the Campbell Plateau islands, which are 

geologically unrelated, is more difficult to account for (81). 

Oriental Region 

Knowledge of this fauna, crucial to an interpretation of Melanesian zoogeography, 

is insufficient. Recent expeditions somewhat augmented the basic work of 

Enderlein on this region, but more work is needed. As an example, 10 days of 

collecting on Bali and Lombok recently raised the number of species known 

from Indonesia from 140 to over 200. 

A few typically Oriental groups have been recognized. The family Calopsocidae 

extends from the Oriental region to northern Australia and the Solomons, 

with endemic genera in New Guinea (89). The Oriental group of Epipsocidae 

more closely related to African than tropical South American species; Epipsocopsis 

has distinct African and Oriental species groups, and Hinduipsocus is 

endemic to India.





186 THORNTON 

Although New Guinea has been inadequately surveyed, there are indications 

that it has been an important link between the greater Sunda islands and the 

islands of the Outer Melanesian Arc in the case of the Pseudocaeciliidae, 

Peripsocidae, Ectopsocidae, a subgroup of the Myopsocidae, and Calopsocus 

(Calopsocidae) and Epipsocopsis (Epipsocidae) (77, 89). Considerable evolution 

has occurred in New Guinea itself with the production of seven endemic 

genera. There is also speculation that differentiation of philotarsid genera may 

have occurred in the area, where several unusual forms are found (77, 86). 

Relatively recent contributions from the New Guinea area to the faunas of 

northeast Australia in the above families and the Stenopsocidae, and to the 

fauna of east Australia in the Myopsocidae, Stenopsocidae, and Psilopsocidae 

are indicated. 

Biogeographical studies on the archipelagos of the southwestern Pacific are 

still incomplete, although specialist collections have been made as far as 

Tonga. It is already clear that no single hypothesis can explain existing 

patterns, which vary from family to family and within some families. Within 

the Philotarsidae, for example, vicaxiance and dispersal explanations have been 

offered for different sections of the family (81,83). The subfamily Zelandopsocinae, 

strictly confined to the Inner Melanesian Arc, has a species swarm on 

New Caledonia, and links between New Zealand and New Guinea. are recognizable. 

In contrast, the Outer Arc has been colonized by the sister subfamily 

Aaroniellinae, probably by dispersal. The Calopsocidae and Epipsocidae also 

occur on the Outer Arc, and in the Pseudocaeciliidae, Psocidae, Lepidopsocidae, 

and Myopsocidae there has been radiation in Fiji (82). The Fijian fauna (81 

species) is 60% endemic, and in general an Oriental-Melanesian origin is 

indicated. Tonga’s fauna is an attenuation of that of Fiji. 

In the Psocidae also, a variety of patterns can be seen. Amphigerontiines 

extend only as far as Norfolk Island and New Zealand, and the genus Psococerastis 

does not appear to have penetrated either the Inner or Outer Melanesian 

Arcs. Of the Psocinae, Ptycta, the only genus of the family in Micronesia 

(six species), occurs in Australia, has three species in the Bismarcks, and 

weakly represented along the Outer Melanesian Arc as far as Tonga, but, 

remarkably, has not yet been found in New Guinea. Its range in this region 

appears to complement that of the related Copostigma, a genus that is unknown 

from Australia, has five species in New Guinea, is absent from the Bismarcks 

and Solomons, but has four species in the New Hebrides and seven in Fiji. 

Clematoscenea, like Copostigtna, is absent from Australia and well represented 

in New Guinea (eight species), but through the Melanesian Arcs their 

ranges are mutually exclusive. 

Ninety species are known from Micronesia, 54 of which occur in the 

southern Mafianas. Endemism is highest (over 30%) on Truk, the southern 

Marianas, and Palaus; Yap and Kusaie, with faunas of a size similar to that of






Truk, show relatively little endemism, whereas Ponape shows a moderate 

degree. Apparently there is no endemism in the northern Marianas, which are 

younger and more impoverished than the southern Marianas, nor in the Marshalls, 

Gilberts, or Caroline atolls, which are very low and very young. 

Oriental species make up about 15% of the fauna of each island group, and 

there are a few New Guinea species (87). 

In the Hawaiian archipelago there is a remarkable development of two 

endemic genera of elipsocids, Kilauella (ca 145 spp.) and Palistreptus (20 

spp.) and the genus Ptycta (Psocidae) (51 spp.), making up over 80% of 

fauna. Endemicity is greatest on Kauai, the oldest and most isolated island of 

the group. In the case of the genus Ptycta, the "Maui complex," which is a 

group of four islands subjected to anastomoses and subdivisions during glacial 

cycles, appears to have been the site of a major burst of speciation. The 

endemic elipsocids have affinities with the Propsocinae of the southern hemisphere 

rather than the Elipsocinae. The remaining 42 species represent 20 

genera of 12 families (84, 85). 

From the above it will be clear that we are not yet in a position to summarize 

definitively the biogeography of Psocoptera on a word scale. The lace curtain 

of hard data has all kinds of holes in it, although in a few places interesting 

patterns can be seen. 

ECOLOGICAL DISTRIBUTION 

Information on the distribution of arboreal psocids within geographical areas 

has been accumulating in recent years. In general, their occurrence appears to 

be related to the distribution and nature of available food on tree surfaces. 

Habitat 

ALTITUDE A reduction in diversity with increasing altitude has been demonstrated 

in Switzerland: only 14 of 55 species occur over 1900 m (upper 

subalpine zone) and all of these species occur in the lower, montane zone 

(1100-1500 m) (33). In the Engadine valley the richest psocid fauna was found 

in montane pine forest (34). Of course, vegetation type must also be considered 

along with temperature, moisture, and food availability (which are probably all 

interrelated) in determining the altitudinal distributions of psocids. 

Mount Kenya has relatively clear-cut altitudinal vegetative zones that are 

determined by changes in temperature, humidity, and rainfall. The Podocarpus 

zone (2440-3120 m) was found to support 28 psocid species, and 12 were 

confined to this zone (21). Only two species were found in the extremely wet 

Hagenia zone (3050-3200 m), and both occurred at lower elevations. One 

these was very rare; the other appears to be stenotopic and is found only





188 THORNTON 

between 2440 and 3200 m. Mesopsocus montanus, found in the Erica zone 

(3320-3660 m), occurs at similar heights on Kilimanjaro. 

The effect of altitude on psocid distribution can perhaps best be examined by 

studying the fauna of a single tree species that has a sufficiently wide altitudinal 

range. On mango in Jamaica an increase in diversity of 3.3 species per 300 m 

increase in altitude from 150 to 1220 m was found (100). Over this range more 

high-altitude stenotopes were added as species were replaced by close relatives 

at successively higher, less disturbed habitats. About 95% of the high altitude 

stenotopes were endemic, in contrast to 50% of those from regions below about 

600 m. Such confinement of endemics to native highland vegetation is common 

on many tropical islands. As in the Hagenia zone of Mount Kenya, psocids 

were extremely sparse in Jamaica from 1200 to 2300 m; at these altitudinal 

ranges, humidities are extremely high and bryophytes and hymenophyllous 

ferns replace the lichens of the lower sclerophyll forest. A marked reduction in 

psocid numbers and diversity is also characteristic of mountain tops with wet 

conditions and epiphytic mosses in many parts of the Pacific. On mango in 

Trinidad the psocid fauna was more diverse, but the same general trends were 

apparent over the range of altitudes studied (0 to 610 m) (19). 

In Yorkshire, England, the fauna of larch was studied from near sea level to 

400 m (16). In contrast to the Caribbean studies, no change in species numbers 

or species replacement was found, but density increased markedly with altitude. 

Mesopsocus unipunctatus was found to occur at both low and high 

elevations (above 260 m), whereas its close relative M. immunis, believed to be 

at the northern end of its range, was confined to lower altitudes. Greater winter 

loss of egg batches and greater sensitivity to low temperatures in regard to 

oviposition were suggested as possible critical factors in deternfining the 

absence of M. immunis from the higher forests (23). Elipsocus mclachlani, a 

facultative lichen feeder, was found only above 200 m, but the six other 

common species were eurytopic. 

On Kauai Island, Hawaiian Islands, I collected some 34 species from Acacia 

koa from 500 to 1200 m on the Kokee massif over a period of a few days. Of 

these, seven could be considered eurytopic (occurring from 650 to 1200 m) six 

were only taken from 800 to 950 m, nine from 800 to 1200 m (four most 

common at 1200 m), and nine only on the Kokee plateau at about 1200 m. Eight 

of the high altitude stenotopes were endemics, including all five species of the 

endemic genus Palistreptus found in this transect. 

TREES AS HABITATS There is evidence that the psocid faunal spectrum on 

trees varies considerably according to the type and species of tree, even in the 

same locality. 

Tree "preferences" In Europe, a broad division between associations on 

coniferous and broad-leaved trees is generally accepted. Although similar in






psocid diversity, these types of tree generally differ in species spectrum, and 

"conifer species" and "broad-leaved species" of psocids can be identified. 

In North America there is evidence (40) that natural species groups 

Caecilius occur predominantly on particular vegetation types, such as grasses, 

sedges, palms, conifers, and broad-leaved trees. 

In Brazil, a "preference" for the dead fronds arid leaf bases of palms has 

been noted for some predominantly litter-frequenting epipsocids (60), and 

three closely related species of Lachesilla appear to be extremely closely associated 

with Syagrus palms (59, 61). Various other species are probably palmicolous 

in other parts of the world and, in Caecilius, may be more closely 

related to those frequenting grasses and sedges than to those frequenting other 

trees (40). 

In a 16-ha area near Sydney, Australia, Smithers (76) found that the different 

vegetation types supported different associations of psocopterans. Of the 43 

species present, three were found predominantly on Casuarina spp., three on 

bracken, two on Maclura pomifera. One was restricted to a sandstone shrub 

layer, and five to a small patch of depauperate rain forest. 

Within the general categories of coniferous and broad-leaved trees, "preferences" 

for particular tree species have been noted (22, 30, 33, 58, 67, 91). For 

example, of three closly related Elipsocus species at Malham, Yorkshire, E. 

mclachlani predominated on larch, E. westwoodi on hawthorn, and E. hyalinus 

on sallow and spruce. In Jamaica, weekly samples taken from Juniperus and 

Podocarpus at 1200 m over a period of 10 months revealed that seven of the 

eight most common psocids predominated on one or the other tree species. A 

striking preference for juniper was seen in Peripsocus juniperi (3787 individuals, 

to 43 on Podocarpus) while the reverse preference was clear in 

Caecilius pallidobrunneus (72:2714) and H emipsocus roseus (4:1197). These 

trees offer very different substrates; for example, the needles of Podocarpus are 

broad and flattened, in contrast to those of juniper. Turner (91) suggested food 

requirements, differential protection from rain, and the structural features of 

the needles, which on Podocarpus may favor normally leaf-inhabiting species 

such as H. roseus, as factors that may affect the observed preferences. 

New (58) showed that the natural movement of Betula and Populus leaves in 

southern England inhibits settling by foliicolous psocid species such as Caeciliusflavidus. 

Such species lay eggs on leaves, which become widely scattered 

in autumn, and overwinter in leaf litter where the spring generation is passed. 

The adults then fly up to the trees. Settlement at this time is thus a crucial 

completion of the life cycle. Surface texture of leaves, which varies between 

tree species, was also shown to affect their suitability as oviposition sites for 

three psocids, and the distribution and size of leaf hairs apparently affect the 

availability of food to young psocid nymphs. The amount and kind of food 

carried on leaves of different trees differs: some trees develop distinctive crops 

of microepiphytes as the season progresses.





190 THORNTON 

Vertical stratification Systematic collections of psocopterans at various 

heights above ground have been made in the rain forests of Zaire, Sulawesi, 

Brunei, New Guinea, and Panama. In the Zaire (79) and Panama and Brunei 

(78) studies, there was a concentration of flying psocids in the upper canopy, 

but more at 1 m above ground than at 10 m. In Panama, collections from light 

traps that were run for long periods on Barro Colorado Island and in montane 

forest, both in the canopy and near the ground, showed little vertical difference 

in psocid abundance and no exclusively canopy or understory species (17). 

However, 19 species were significantly more and 23 significantly less numerous 

in the canopy than near the ground. 

Partitioning of tree substrates Some species, for example, of Loensia, 

Trichadenotecnum, Cymatopsocus, Palistreptus, Cerobasis, and Reuterella, 

seem to be particularly associated with tree trunks, and bracket fungi are known 

to support associations of psocids, particularly liposcelids. Other psocids, such 

as embidopsocines, are predominantly subcortical. The main partitioning of the 

substrates afforded by trees, however, is reflected in the recognition, particularly 

in Europe, of predominantly corticolous or foliicolous species (e.g. 33, 

58, 57). Such a dichotomy is believed to be related to presumed adaptations of 

the tarsal claw and empodium for negotiating the smooth and rough surfaces of 

leaves and bark respectively. Psocidae, Myopsocidae, Philotarsidae, Pseudocaeciliidae, 

and northern hemisphere Peripsocidae are generally regarded as 

bark frequenters, and Caeciliidae, Stenopsocidae, Hemipsocidae, Lachesillidae, 

Ectopsocidae, and (in Australia) Peripsocidae, as foliage frequenters. 

There is now evidence from studies in Trinidad that foliicolous and corticolous 

species are also differentiated in the tropics (19). Of 22 species present 

mango and Citrus in Trinidad, 7 were restricted to leaves, 12 to bark, and only 3 

occurred on both. 

The foliicolous habit on deciduous trees has been associated with a redistribution 

of eggs at autumn leaf-fall, leaf litter dormancy, and a redispersal of 

adults onto trees the following year. Such dispersal flights are unknown in 

corticolous species and evidently do not occur in those foliicolous species that 

live predominantly on deciduous trees without a widely dispersed leaf fall (58). 

In general, conifer psocids are corticolous, including those on the deciduous 

larch. New has suggested that in those species frequenting evergreen conifer 

needles, on which the microflora is similar to that on bark, the foliicolous habit 

is at an early stage of development; foliicoly may have first evolved in species 

that exploited the highly specialized and reproductively potent microflora of 

leaves. 

When offered as substrate an experimental compound larch twig comprising 

dead twig with lichen and living twig with Desmococcus, E. westwoodi 

nymphs and adults preferred the live twig and E. mclachlani nymph~,; preferred






the dead, whereas adults showed little preference (22). In northern England 

three common species (E. westwoodi, Atnphigerontia contaminata, and Peripsocus 

phaeopterus) predominate on living twigs of larch, and a common 

species of each genus (E. rnclachlani, A. bifasciata, and P. didymis) predominates 

on dead twigs. E. mclachlani and Reuterella helvimaculata are virtually 

absent from living twigs. Several of the remaining species show a "preference" 

for one type of twig (16). Such preferences have also been noted in Switzerland 

and Brazil (on palms). In England the preferences have been related, except 

the case of Peripsocus species (18), to feeding. E. westwoodi, for example, 

feeds on Desmococcus even when lichen is available, and E. mclachlani is a 

facultative lichen feeder (16, 22). 

Distribution in Relation to Food 

In the Yorkshire Pennines the microepiphyte food available to psocid grazers 

on larch bark consists of lichen, Desmococcus, and fungal spores. Seven of the 

common species feed on Desmococcus rather than lichen, and two prefer 

lichen. Of the latter, R. helvimaculata feeds on the whole lichen; E. mclachlani 

only on the apothecia. Both these species are most abundant at higher altitudes. 

In nature, the alga-preferring specie§ also feed on fungal spores. Thus in 

northern England the microepiphyte grazers fall into two categories: Desmococcus 

and fungal spore feeders, and lichen feeders, and their altitudinal 

distributions and tree preferences are related to the availability of these foods. 

In contrast to corticolous species, some of which prefer a certain type of food 

in the presence of others, foliicolous species on broad-leaved trees in England 

graze indiscriminately on the available food (58). However, both the amount 

and proportion of foods available on leaves differ between tree species and vary 

throughout the summer. The food spectrum affects psocid fecundity and may 

thus affect the mix of psocid grazers. On larch bark also there is a successional 

development as the substrate branch ages: the lichen Lecanora increases, 

epiphyte density rises, and photosynthetic potential decreases. These changes 

may have an effect on the distribution of psocids (96). 

Most of the psocids on conifers at 1200 m in Jamaica take small quantities of 

all food components available (sooty moulds, powdery lichens, Phycopeltis 

sp., pollen grains, bark flakes), and Turner believes that the absence of the 

feeding dichotomy so clearly seen on larch in England may be because the loose 

associations of algae and fungi do not produce lichen acids (91). Only 2 of 

montane tropical forest psocids in East Africa were tentatively regarded as 

feeders on what appeared to be powdery lichens (20, 21). Several feeders 

fungus (sooty moulds and honeydew moulds) were not recorded as taking algal 

cells, and two were unusual in taking few or no spores. 

The altitudinal diversity of psocopterans on mango in Jamaica has been 

related to indices of microepiphyte diversity (100). Both the variety and





192 THORNTON 

quantity of microepiphytes increase with altitude, thus allowing the coexistence 

of a greater number of species (although modification of lowlan,ds by man 

is also an important factor). There have been no specialist studies of the vertical 

distribution of psocid food on trees, but Turner (97) found significan~Ily greater 

epiphyte densities above 1.5 m than nearer to the ground. 

Several species of some eight families are known to be specialist exploiters 

of dying and dead vegetation and in fact can be collected by first: breaking 

branches and then returning later to collect a sample. Ectopsocus briggsi is a 

typical example. Turner (96) noted that this small species, which feeds on the 

spores and hyphae of decomposer fungi, had the greatest biomass of any psocid 

in the late summer in one of the three years of his study, yet was otherwise rare. 

He ascribed this to unusually moist conditions in late summer of that year, 

following a summer drought, that favored the development of fungi on dead 

plant material. Unpredictable climatic phenomena such as this as well as 

random local disturbances to the canopy will have an important effect on the 

distribution of such opportunistic species. 

HUMIDITY Humidity clearly plays a key role in the life of psocids and in 

aboreal species its effects may be related to food supply. Liposcelis species are 

capable of absorbing water vapor from an atmosphere down to 58% relative 

humidity (R.H.); below 58% R.H. water loss leads to death. Rudolph (66) 

demonstrated this process in 22 psocid species of diverse habitats and representing 

all major groups. The rates of both water loss and uptake were shown to 

be markedly high for arthropods, and the uptake-to-loss ratio, a measure of the 

efficiency of the water balance mechanism, was among the highest of all 

arthropods studied. This ability appears to be characteristic of the order, 

unrelated to specific environmental conditions. Differences in critical equilibrium 

humidities (58-85%) and water loss and uptake rates were not correlated 

with habitat, apart from the observation that domestic and nidicolous species 

have lower critical values (58-76%) than arboreal species (76-85%). Intensive 

and extensive studies of the microhabitats of species that differ greatly in 

uptake/loss ratios are now needed. 

The mechanism’s operation was shown to be highly dependent on food 

supply, particularly in arboreal species in which food storage ability is evidently 

poor and the physiological mechanism is upset unless the insects can feed at 

short intervals. The indications are that above about 75% R.H., food supply, 

rather than relative humidity itself, may be the limiting factor. 

AIR POLLUTION AND PSOCID DISTRIBUTION Gilbert (28) has den~tonstrated 

the indirect effects (through microepiphytes) of air pollution on the distribution 

of corticolous psocids in the Newcastle area of the industrial north of England. 

He showed a very high correlation between the diversity of the psocid fauna of






the trunks of ash (Fraxinus excelsior) and distance from the pollution focus in 

the town center. The lichen feeders R, helvimaculata, E. mclachlani, and 

Loensia fasciata were absent from town sample sites, and the Desmococcus 

feeders M. unipunctatus, E. hyalinus, and E. westwoodi were common at rural 

sites, abundant in the perimeter zone, and also virtually absent from the town. 

These distributions were closely related to the distribution of the autotrophs 

lichen and Desmococcus. A. bifasciata was exceptional, evidently able to 

survive in the town center on bark flakes, detritus, and fungal spores. 

The density and distribution of Loensia fasciata were also found to be 

inversely correlated with SOz emission load in a study of this species in the 

vicinity of a German power plant (39). In a recent study in Luxembourg this 

was confirmed, but the related species Loensia variegata was most dense in 

heavily polluted areas and, along with Cerobasis guestfalicus, appeared to be a 

good indicator of heavy pollution. In sharp contrast, Pseudopsocus rostocki 

was absent from areas with more than 80 Ixg SO2/m3 (68). 

Popescu identified an "industrial component" involving lichen cover and 

SOa level that accounted for most of the variation in frequency of the melanic 

morph of M. unipunctatus and suggested that lichen was detrimental to the 

crypsis of melanics (65). Thus the variation in the distribution of epiphytes as 

result of pollution may affect psocopterans not only directly with respect to 

food, but also indirectly through its effect on predation. 

Literature Cited 

1. Badonnel, A. 1963. Psocopt~res terricoles, 

lapidicoles et corticoles du Chili. 

Biol. Am. Aust. 2:291-338 

2. Badonnel, A. 1966. Sur quelques Psocopt~res 

des ~les Mascareignes. Bull. 

Soc. Entomol. Fr. 71:234-38 

3. Badonnel, A. 1967. Insectes Psocoptrres. 

Faune Madagascar 23:1-238 

4. Badonnel, A. 1971. Psocopt~res 6daphiques 

du Chili (3e note). Bull. Mus. Nat. 

Hist. Nat., Zool. 3e ser., 1:1-38 

5. Badonnel, A. 1971. Sphaeropsocopsis 

reisi n.sp., premier reprrsentant africain 

connu de la famille des Sphaeropsocidae 

(Psocoptera, Nanopsocetae), avec complrments 

h la faune des Psocoptrres 

angolais. Publ. Cult. Cia. Diamantes 

Angola 84:13-28 

6. Badonnel, A. 1976. Complrments ~t 

l’rtude des Psocopt~res de Madagascar. 

Bull. Mus, Nat. Hist. Nat., Zool. 3e tats des Expdditions Biosp~ologiques 

Cubano-Roumaines ~ Cuba, ed. T. 

Orghidan, A. N. Jimrnez, V. Decou, ~. 

Negrea, N. V. Bayrs, pp. 339-44. 

Bucharest: Acad. Republ. Soc. Rumania 

9. Badonnel. A. 1977. Psocoptbres de l’Angola: 

V. Publ. Cult. Cia. Diamantes 

Angola 89:103--52 

10. Badonnel, A. 1981. Psocopt~res de l’~le 

de La Rrunion. 2 

ser., 

410(287):1143-98 

7. Badonnel, A. 1976. La faune terrestre de 

l’lle de Sainte-Hrl~ne. 16 Psocoptera. 

Ann. Mus. R. Afr. Cent., Ser. 8, 

215:206-30 

8. Badonnel, A. 1977. Psocoptbrescavernicotes 

de Cuba (Premiere note). In R~sul 

e note. Bull. Mus. Nat. 

Hist. Nat., Zool. 4e ser., 3:663-74 

11. Badonnel, A. 1981. Psocopt~res /~6ophiles 

des Petites Antilles. Rev. Ecol. 

Biol. Sol 18(3):425-34 

12. Baldwin, P. A. 1953. Annual cycle, environment, 

and evolution in the Hawaiian 

honeycreepers (Aves: Drepaniidae). 

Univ. Calif. Publ. Zool. 52(4):285-398 

13. Betts, M. M. 1956. A list of insects taken 

by titmice in the forest of Dean (Gloueestershire). 

Entomol. Mon. Mag. 92:68-71 

14. Betz, B. W. 1983. Systematics of the 

Trichadenotecnum alexanderae species 

complex (Psocoptera: Psocidae) based 

an investigation of modes of reproduction 

and morphology. Can. Entomol. 115: 

1329-54 

15. Broadhead, E. 1958. Some records of





194 THORNTON 

animals preying upon psocids. Entomol. 

Mon. Mag. 94:68-69 

16. Broadhead, E. 1958. The psocid fauna of 

larch trees in northern Englandnan ecological 

study of mixed species populations 

exploiting a common resource. J. 

Anita. Ecol. 27:217-63 

17. Broadhead, E. 1983. The assessment of 

faunal diversity and guild size in tropical 

forests with particular reference to the 

Psocoptera. In Tropical Rain Forest: 

Ecology and Management, ed. S. L. Sutton, 

T. C. Whitmore, A. C. Chadwick, 

pp. 107-19. Oxford/Boston: Blackwell 

18. Broadhead, E., Datta, B. 1960. The taxonomy 

and ecology of British species of 

Peripsocus Hagen (Corrodentia, Pseudocaeciliidae). 

Trans. Soc. Br. Entomol. 

14:131-46 

19. Broadhead, E., Evans, I-I. A. 1979. The 

diversity and ecology of the Psocoptera in 

tropical forests. Acta IV Symp. Int. Ecol. 

Trop., Panama, 1977, 1:185-96 

20. Broadhead, E., Richards, A. M. 1980, 

The Peripsocidae and Psocidae (Psocoptera) 

of East Africa. Syst. Entomol. 5: 

357-97 

21. Broadhead, E., Richards, A. M. 1982. 

The Psocoptera of East Africa---a taxonomic 

and ecological survey. Biol. J. 

Linn. Soc. 17:137-216 

22. Broadhead, E., Thornton, I. W. B. 1955. 

An ecological study of three closely related 

psocid species. Oikos 6:1-50 

23. Broadhead, E., Wapshere, A. J. 1966. 

Mesopsocus populations on larch in England-the 

distribution and dynamics of 

two closely-related coexisting species of 

Psocoptera sharing the same food resources. 

Ecol. Monogr. 36:327-88 

24. Eertmoed, G. E. 1973. The phenetic relationships 

of the Epipsocetae (Psocoptera): 

the higher taxa and the species of 

two new families. Trans. Am. Entomol. 

$oc. Philadelphia 99:373414 

25. Eertmoed, G. E. 1978. Embryonic diapause 

in the psocid Peripsocus quadrifasciatus: 

photoperiod, temperature, ontogeny 

and geographic variation. Physiol. 

Entomol. 3:197-206 

26. Gagnr, W. C. 1979. Canopy-associated 

arthropods in Acacia koa and Metrosideros 

tree communities along an altitudinal 

transect on Hawaii Island. Pac. Insects 

21(1):56-82 

27. Garcia-Aldrete, A. N. 1974. A classification 

above species level of the genus 

Lachesilla Westwood (Psocoptera: 

Lachesillidae). Folia Entomol. Mex. 27: 

1-88 

28. Gilbert, O. L. 1971. Some indirect 

effects of air pollution on bark-living invertebrates. 

J. Appl. Ecol. 8:77-84 

29. Guillaumont, F. 1977. Sur la biologic et 

l’rcologie des esp~ces du genre Hemineura 

Tetens en Provence occidentale 

(Psocopt~res, Elipsocidae). Ecol. 

Mediterr. 3:55-65 

30. Gunther, K. K. 1974. Staublause, Psocoptera. 

Die Tierwelt Deul!schlands 61. 

Jena: G. Fischer. 314 pp. 

31. Hey, R. 1977. Tectonic ewalution of the 

Cocos-Nazca spreading centre. Geol. 

Soc. Am. Bull. 88(10):140,~1-20 

32. Kim, K. C., Ludwig, H. W. 1982. Parallel 

evolution, cladistics, and classification 

of parasitic Psocodea. Ann. Entomol. 

Soc. Am. 75(5):537-48 

33. Lienhard, C. 1977. Die Psocopteren des 

Schweizerischen Nationalparks und seiner 

Umgebung (Insecta: Psocoptera). 

Ergeb. Wiss. Unters. Schweiz. Nationalpark 

14(75):417-551 

34. Lienhard, C. 1980. Oekologische Untersachungen 

im Unterengadin. D2. Psocopteren 

(Insecta: Psocoptera). Ergeb. 

Wiss. Unters. Schweiz. Nationalpark 

12(8): 16-33 

35. Lienhard, C. 1983. Surquelquespsocopt~res 

de Mad~re avec c16 de drtermination 

pour les esp~ces de Trichopsocus 

Kolbe de la rrgion palrarctique occiden- 

Funchal tale (Insecta: 67:1-12Psocoptera). 

Bocagiana 

36. Meinander, M. 1972. The invertebrate 

fauna of the Kilpisjarvi area, Finnish 

Lapland. 6. Psocoptera. Acta. Soc. 

Fauna Flora Fenn. 80:65-66 

37. Meinander, M. 1973. The Psocoptera of 

the Canary Islands. Not. Entomol. 53: 

141-58 

38. Meinander, M., I-Ialkka, O., Srderlund, 

V. 1974. Chromosomal evolution in the 

Psocoptera. Not. Entomol. 14:81-84 

39. Mey, W., Tietze, F. 1979. Zur Indikation 

von Luftverunreinigungen mittels 

Psocoptera. Hercynia 16(41):417-19 

40. Moekford, E. L. 1965. The genus Caecilius 

(Psocoptera: Caeciliidae). Part 

Species groups and the North American 

species of the flavidus group. Trans. Am. 

Entomol. Soc. P hilade lphh~ 91:121-66 

41. Mockford, E. L. 1966. The genus Caecilius 

(Psocoptera: Caeciliidae). Part II. 

Revision of the species groups, and the 

North American species of thefasciatus, 

confluens and africanus groups. Trans. 

Am. Entomol. Soc. Philadelphia 92:133- 

72 

42. Mockford, E. L. 1967. The electrentomoid 

psocids (Psocoptera). Psyche 74: 

118-65 

43. Mockford, E. L. 1969. The genus Caecilius 

(Psocoptera: Caecilliidae). Part III. 

The North American species of the alcinus, 

caligonus and subflavus groups.





Trans. Am. Entomol. Soc. Philadelphia 

95:77-151 

44. Mockford, E. L. 1971. Peripsocus species 

of the alboguttatus group (Psocoptera: 

Peripsocidae). J. NY Entomol. Soc. 

79:89-115 

45. Mockford, E. L. 1974. TheEchmepteryx 

hageni complex (Psocoptera: Lepidopsocidae) 

in Florida. Fla. Entomol. 57(3): 

255-67 

46. Mockford, E. L. 1974. Records and descriptions 

of Cuban Psocoptera. Entomol. 

Am. 48:103-215 

47. Mockford, E. L. 1975. A new species of 

Tapinella from Mexico and notes on 

variation in Tapinella maculata (Psocoptera: 

Pachytroctidae). Folia Entomol. 

Mex. 31-32:101-15 

48. Mockford, E. L. 1979. Diagnoses, distribution 

and comparative life history 

notes on Aaroniella maculosa (Aaron) 

and A. eertmoedi n.sp. (Psocoptera: Philotarsidae). 

Great Lakes Entomol. 12: 

35-44 

49. Mockford, E. L. 1979. Psocoptera. In 

Canada and its Insect Fauna, ed. H. V. 

Danks, Mem. Entornol. Soc. Can. 108: 

324-26 

50. Mockford, E. L. 1980. Identification of 

Elipsocus species of western North 

America, with descriptions of two new 

species (Psocoptera: Elipsocidae). Pan- 

Pac. Entomol. 56:241-59 

51. Mockford, E. L. 1981. Systematics of 

new world genera of Cerastipsocini and 

species of Psococerastis Pearman (Psocoptera: 

Psocidae: Cerastipsocinae). 

Trans. Am. Entomol. Soc. Philadelphia 

107:249-98 


Asiopsocidae (Psocoptera) including 

Pronotiopsocus amazonicus n.gen, n. sp. 

Fla. Entomol. 66(2):241-49 


the Blaste postica complex with descriptions 

of three new species (Psocoptera: 

Psocidae). In press 

54. Mockford, E. L. 1984. A systematic 

study of genus Camelopsocus with descriptions 

of three new species (Psocoptera, 

Psocidae). In press 

55. Mockford, E. L., Broadhead, E. 1982. A 

new genus and two new species of Philotarsidae 

(Insecta: Psocoptera) from East 

Africa. J. Nat. Hist. 16:431-37 

56. Moekford, E. L., Wong, S. K. 1969. The 

genus Kaestneriella (Psocoptera: Peripsocidae). 

J. NY Entomol. Soc. 77(4): 

221-49 

57. Montgomery, S. L. 1982. Biogeography 

of the moth genus Eupithecia in Oceania 

and the evolution of ambush predation 

in Hawaiian caterpillars (Lepidoptera: 


Geometridae). Entomol. Gen. 8(1):27- 

34 

58. New, T. R. 1970. The relative abundance 

of some British Psoeoptera on different 

species of trees. J. Anim. Ecol. 39:521- 

40 

59. New, T. R. 1971. Species of Lachesilla 

associated with palm trees in central 

Brazil. J. Linn. Soc. London Zool. 50: 

431-47 

60. New, T. R. 1972. Some Epipsocetae 

(Psocoptera) from central Brazil. Trans. 

R. Entomol. Soc. London 123:455-97 

61. New, T. R. 1973. Local distribution of 

Psocoptera in the Mato Grosso, central 

Brazil. Pap. Avulsos Dep. Zool. Sao 

Paulo 27:115-44 

62. New, T. R. 1984. A Biology of Psocoptera. 

R. Entomol. Soc. London. In press 

63. New, T. R., Thornton, I. W. B. 1981. 

Psocoptera from central and southern 

Chile. Pac. Insects Monogr. 37:13(~ 

78 

64. Palmgren, P. 1932. Zur biologie von 

Regulus r. regulus (L.) und Parus atricapillus 

borealis Selys. Eine vergleichend 

ikologische utersuchung. Acta Zool. 

Fenn. 14:1-113 

65. Popescu, C. 1979. Natural selection in 

the industrial melanic psocid Mesopsocus 

unipunctatus (Mull.) (Insecta: Psocoptera) 

in northern England. Heredity 

42(2): 133-42 

66. Rudolph, D. 1982. Occurrence, properties 

and biological implications of the 

active uptake of water vapour from the 

atmosphere in Psocoptera. J. Insect 

Physiol. 28(2):111-21 

67. Schneider, N. 1979. Les psocopt~res du 

Grand-Duch6 de Luxembourg. II. 

Faunistique et 6cologie des esprces sylvicoles 

du Kalebierg. Bull. Ann. Soc. R. 

Beige Entomol. 115:197-208 

68. Schneider, N. 1982. Les psocopt~res du 

Grand-Duch6 de Luxembourg. III. 

Faunistique et 6cologie des esp~ces 

urbaines. Bull. Ann. Soc. R. Belge Entomol. 

118:131-44 

69. Seeger, W. 1975. Funktionsmorphologie 

an Spezialbildungen der Fuhlergeissel 

von Psocoptera und anderen Paraneoptera 

(Insecta); Psocodea als monophyletische 

Gruppe. Z. Morphol. Tiere 

81:137-59 

70. Seeger, W. 1979. Special structures in 

egg membranes and embryos of Psoeoptera, 

providing the monophyly of this 

order, with special regard to other Paraneoptera 

(Insecta). Stuttg. Beitr. Naturk. 

Ser. A 329:1-57 

71. Simberloff, D. 1976. Species turnover 

and equilibrium island biogeography, 

Science 194:572-78





196 THORNTON 

72. Smithers, C. N. 1965. A bibliography of 

the Psocoptera (Insecta). Aust. Zool. 13: 

137-209 

73. Smithers, C. N. 1969. The Psocoptera of 

New Zealand. Rec. Canterbury Mus. 

7(4):259-344 

74. Smithers, C. N. 1970. Some thoughts on 

trans-Tasman relationships in the Pso- 

D. 1972. Psocoptera. Insects Mieronesia 

8(4):45-144 

88. Thornton, I. W. B., New, T. R. 1981. 

Psocoptera from Robinson Crusoe Island, 

Juan Fernandez Archipelago. Pac. 

Insects Monogr. 37:179-91 

89. Thornton, I. W. B., Smithers, C. N. 

1984. The systematics of the Calopsocicoptera. 

N. Z. Entomol. 4(3):79-85 

75. Smithers, C. N. 1972. The classification 

and phylogeny of the Psocoptera. Mem. 

Aust. Mus. 14:1-351 

76. Smithers, C. N. 1977. The Psocoptera of 

Muogamarra Nature Reserve. Rec. Aust. 

Mus. 31(7):251-306 

77. Smithers, C. N., Thornton, I. W. B. 

1982. The role of New Guinea in the 

evolution and biogeography of some 

families of psocopteran insects. In 

Biogeography and Ecology of New 

Guinea, ed. J. L. Gressitt, 2:621-38. The 

Hague: Junk 

78. Sutton, S. L., Ash, C. P., Grundy, A. 

1983. The vertical distribution of flying 

insects in lowland rain-forests of Panama, 

Papua New Guinea and Brunei. J. 

Linn. Soc. London Zool. 78:287-97 

79. Sutton, S. L., Hudson, P. J. 1980. The 

vertical distribution of small flying indae, 

an Oriental and Melanesian family 

of Psocoptera. Syst. Entomol. 9(2):183- 

244 

90. Thornton, I. W. B., Woo, K. T. 1972. 

Psocoptera of the Galapagos Islands. 

Pac. Insects 15(I):1-57 

91. Turner, B. D. 1974. The population 

dynamics of tropical arboreal Psocoptera 

(Insecta) on two species of conifers in the 

Blue Mountains, Jamaica. J. Anim. Ecol. 

43:323-37 

92. Turner, B. D. 1975. Energy flow in 

arboreal epiphytic communities. An empirical 

model of net primary productivity 

in the alga Pleurococcus on larch trees. 

Oecologia 20:179-88 

93. Turner, B. D. 1975. The Psocoptera of 

Jamaica. Trans. R. Entomol. Soc. London 

126:533-609 

94. Turner, B. D. 1976. Psocoptera of the 

Mascarene Islands. Syst. Entomol. 1: 

sects in the lowland rainforest of Zaire. 

J. Linn. Soc. London Zool. 68:111- 

23 

80. Thornton, I. W. B. 1967. The measurement 

of isolation in archipelagos, and its 

relation to insular faunal size and endemism. 

Evolution 21(4):842-49 

81. Thornton, I. W. B. 1980. Plate tectonics 

and the distribution of the insect family 

Philotarsidae (order Psocoptera) in the 

southwest Pacific. Palaeogeogr. Palaeoclimatol. 

Palaeoecol. 31:251-66 

82. Thornton, I. W. B. 1981. Psocoptera of 

the Fiji Islands. Pac. Insects Monogr. 

37:1-105 

83. Thornton, I. W. B. 1981. The systematics, 

phylogeny and biogeography of the 

psocopteran family Philotarsidae. Syst. 

Entoraol. 6(4):413-52 

84. Thornton, I. W. B. 1981. Psocoptera of 

the Hawaiian Islands. I and II. Introduction 

and the nonendemic fauna. Pac. Insects 

23:1-49 

85. Thornton, I. W. B. 1984. Pscoptera of 

the Hawaiian Islands. lII. The endemic 

complex of Ptycta species; systematics, 

distribution and possible phylogeny. Int. 

J. Entomol. 26(1-2): 1-128 

86. Thornton, I, W. B. 1984. An unusual 

psocopteran from New Guinea, and its 

relationships within the Philotarsidae. 

Int. J. Entornol. In press 

87. Thornton, I. W. B., Lee, S. S., Chui, W. 

201-25 

95. Turner, B. D. 1979. Psocids as prey for 

Mascarene Swiftlets. Entomol. Mon. 

Mag. 113(1360):210 

96. Turner, B. D. t983. Annual respiration 

and production estimates for collembolan 

and psocopteran epiphyte herbivores on 

larch trees in southern England. Ecol. 

Entornol. 8:213-28 

97. Turner, B. D. 1983. Biomass, chlorophyll 

concentration and calorific content 

dynamics of epiphytes on larch trees in 

southern England, Proc. Leeds Philos. 

Lit. Soc. Sci. Sect. 9(3):61-76 

98. Turner, B. D. 1983. Psocoptera from 

Venezuela. A collection of psocids from 

the food boluses of swift.,;. Entomol. 

Scand. In press. 

99. Turner, B. D. 1984. Predation pressure 

on the arboreal epiphytic herbivores of 

larch trees in southern England. Ecol. 

Entomol. 9:In press. 

100. Turner, B. D., Broadhead, E. 1974. The 

diversity and distribution of psocid 

populations on Mangifera indica L. in 

Jamaica and their relationship to altitude 

and micro-epiphyte diversity. J. An#n. 

Ecol. 43:173-90 

101. Turner, B. D., Cheke, R. A. 1983. Psocoptera 

of the Togo-Benin Gap, West 

Africa. J. Nat. Hist. 17:379-~-04 

102. Weidner, H. 1972. Copeognatha (Psocodea). 

Hand. Zool. 4(2):1-94


by Mr. Bas van Berkum on 10/10/07. For personal use only.


by Mr. Bas van Berkum on 10/10/07. For personal use only.

The Geographical and Ecological Distribution of Arboreal Psocoptera

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?