Memoirs of the Museum of Victoria 56(2):339-347 (1997) 28 February 1997 https://doi.org/10.24199/j.mmv.1997.56.22 TROPICAL RAINFOREST MYGALOMORPH IN THE AUSTRALIAN DESERT: THE IRONY OF AN ADAPTIVE LEGACY

Barbara York Main

Department of , University of Western . Nedlands, WA 6009. Australia

Abstract

Main, B.Y., 1997. Tropical rainforest mygalomorph spiders in the Australian desert: the irony of an adaptive legacy. Memoirs of the Museum of Victoria 56(2): 339-347.

The semiarid regions of Australia have a high generic diversity of mygalomorph spiders . Several genera are postulated as being "tropical" rainforest genera. Three genera. Conothele, Selenocosmia and Cethegus have been selected for discussion. Persistence of these genera in the arid region is postulated as being due to a combination of natural restriction to relic habitats or those with verisimilitude with rainforest habitats and retention of behavioural

attributes which fortuitously fit them to persist. Aspects requiring conservation attention are perceived to be tourism and too frequent fires.

Introduction seasonal or monsoon rainforest forms small or large pockets in the Kimberley of northern West- The belief in the richness and diversity of life in ern Australia (McKenzie, 1991, Figure 3) and rainforests, particularly tropical rainforests, the Northern Territory. Dry vine forests in forms part of our inherited mythology as inland and some coastal areas of mid Queens- reflected in literature and art. No less in biologi- land are also generally included in broad state- cal science, where the diversity of tropical rain- ments about northern or tropical rainforests. forest life provides themes for theory, documen- Recently, some zoologists have challenged the tation and estimation: why so rich and how rich? "assumption that in the tropics is

From the middle of the last century with explo- vastly higher than .... in the temperate zones" ration in the tropics by adventurers and scien- (Platnick, 1991) and botanists likewise recog- tists, namely H. W. Bates, Charles Darwin and nise high diversity outside tropical rainforests,

Alfred Wallace, (and I must mention here the e.g., as in south-western largely overlooked observations of the dominant (Lamont et al., 1977). In looking at mygalo- figure of mid to late nineteenth century morph trapdoor spiders it is apparent, and per- systematics, Eugene Simon) fascination with haps surprising to those still hungover with the this phenomenon has developed into one of the spell generated by the rainforest mythology, that the driving themes of evolutionary biology as the semiarid regions of Australia are possibly the currently studied — why so rich — what is the richest areas at least in terms of generic diversity mechanism providing this richness and how is it and behavioural scope. On a broad geographic maintained? In the last part of this century, with front, Main (1982) listed 17 of "the 37 or more anxiety about diminution of the biological bank, mygalomorph genera in Australia" as occurring some biologists have become obsessed with esti- in semiarid and arid regions of Australia. At a mating the richness and diversity of life i.e. the smaller scale, a recent study by Main (1996) at array of life at the species level — how many Durokoppin in the semiarid Wheatbelt region of species (Ewing, 1983; Monteith, 1990). The con- Western Australia demonstrated occurrence of comitant ideal of documenting this diversity i.e. at least 25 species in 13 or 14 genera from six naming and describing the species and so mak- families in a small area of less than a square km ing a reality of the estimations, is sadly lack- within a reserve of remnant vegetation of 1030 ing. hectares. It is doubtful whether such taxonomic In Australia, rainforests (or "closed forests" in diversity can be matched in rainforest habitats. some terminologies, e.g., Specht 1981) comprise Wishart (1993) noted occurrence of eight myga- a small percentage of the landmass. Tropical lomorph species in a 95 x 55 m subtropical rainforests are confined to small areas of the rainforest remnant at Gerringong in New South northeast coastal region (Specht, 1981, Figure 2; Wales. Davies, Gray and other collectors appar- ently did not find a comparable taxonomic Webb and Tracey, 1981, Figure 1 ) while wet/dry

339 340 B. Y. MAIN

richness in eastern Australian rainforests during Taxonomic diversity of Australian various rainforest surveys, to that found at Durokoppin (see Monteith, and Davies, 1991). There are ten familes of Mygalomorphae cur- Nor did the Bellenden Ker survey in north rently recognized from Australia (Raven, produce such a diversity (Monteith 1985a). Recent taxonomic revisions have and Davies, 1991). Nevertheless it is known brought the number of mygalomorph genera of from the taxonomic works of Raven (Raven, mainland Australia and offshore islands and 1994 and earlier papers), museum collections, Tasmania to 43 according to Main ( 1 985a) and my own collection and personal observations with those recorded or described since then that there is indeed a rich mygalomorph fauna in (Main, 1985b, 1985c, 1986, 1991c; Raven, the eastern Australian rainforests including the 1986, 1988. 1994; Churchill and Raven, 1992) tropics. Main (1976) also noted 12 species in a plus some reinstated, and one found not to occur short transect across the sclerophyll/rainforest in Australia (Raven, 1994), there are now 57 boundary to the Nothofagus habitat in Laming- named genera considered valid while there are ton National Park. With later refinements in the still several recognised but unnamed genera this number of species and genera (pers. obs.). would be higher but possibly not as rich as at the Durokoppin site. Surveys in the monsoon rain- forests of neither Kakadu (Kikkawa and Mon- Tropical rainforest and desert — terminology teith. 1980) nor the Rimberley (Main, 1991a) and distribution indicated a taxonomic richness to that observed in the semiarid. Before discussing the occurrence of rainforest genera in the "desert" some definition of the Main (199 lb) noted 25 genera of the "forty or boundaries of the two habitat regions in the pres- so" named mygalomorph genera as occurring in ent context is necessary. I use the term rainforest "rainforests". However, this number included to mean predominantly closed "wet" forest but genera occurring in southern or temperate rain- also to include seasonal i.e. monsoon forest as in forest while the present discussion is concerned the Kimberlcy and Northern Territory and vine only with those found in tropical rainforests

thickets as in Queensland . Thus "tropical rain- (although of course some of those genera may forest" as used here equates roughly to "closed occur in temperate rainforests as well). The pres- forest" of Specht (1981, Fig.2). ent study notes 26 genera occurring in trop- Northern "open forest woodland" of Specht ical rainforests (Table 1). Most of these occur (1981, Fig. 4) actually embraces many patches of also in tall, open forest (mesophytic or sclero- rainforest as documented in McKenzie et al phyll forest) but a few arc confined to tropical rainforest, (1991) and patches and gallery forest in the e.g., Masteria and . A few Northern Territory. Thus tropical rainforest in others (e.g., Kiama, Amiralolhele, Carrai, small or large areas is mostly found within the and Pk-sioilwlc ) are also confined to rain- rainfall isohyets of 750mm per annum and forest but including southern rainforests and above (sec Nix, Nothofagus. 1981, Figure 10). "Desert" and semiarid/arid is loosely defined The theme of this paper concerns those genera here to encompass that huge interior of Australia which are regarded as being primarily "tropical with less than 500 mm rainfall per annum, rainforest genera" but which also extend into receding to less than 250 mm and 1 50 mm (sec semiarid regions. In that such genera are Nix, 1981, Figure 10). The southern boundary adapted to humid or seasonally humid forest accepted in the present context is not as restric- situations it may appear anomalous that some tive as that defined as the 250 mm isohyet by species occur in semiarid/arid regions. I shall Williams and Calaby (1985) and followed by now attempt to show that it is the very nature of Morton et al 1 ( 995, Figure 3. 1 ). However, there- the spiders' adaptations to tropical rainforest are many refugia associated with topographic- habitas which ironically enables them to live features throughout the whole of the low rainfall in the desert region. Furthermore I argue (con- areas of Australia — refugia which can be trary to some of my earlier interpretations) regarded as harbouring biotic leftovers of an that the spiders, rather than having invaded earlier "wetter" climatic regime. Striking the desert, have been stranded there following examples are found in the valleys of the Central the retreat of their original tropical rainforest Australian mountains, the gorges of the Hamers- landscapes. Iey Ranges in Western Australia and around low TROPICAL RAINFOREST SPIDERS IN THE AUSTRALIAN DESERT 341 hills and granite outcroppings on the subdued taxonomic papers but some natural history landscapes of the Western Australian Plateau works) of Churchill and Raven (1992), Gray and Eyre Peninsula in South Australia. Morton (1987,1992), Main (1985b, 1985c, 1986, 1991a, et al. (1995) list 74 refugia in the semiarid and 1991c, 1995, 1996), Raven (1984a, 1984b. arid region. 1985b, 1986, 1993, 1994) and to some extent Even outside these obvious refugial areas, from data with museum collections and finally within the "desert" there are numerous micro- from my own observations and field records.

habitats which present, in microcosm, habitats I have selected for the present discussion the that have verisimilitude with tropical rainfor- three genera Conothele (Ctcnizidae). Selenocos- ests. These persistent, small and large, scattered mia (Theraphosidac) and Cethegus () and isolated refugia preserve certain taxa which which have in common a distribution right

by their overall distribution suggest a tropical across the north i.e. throughout the tropics and rainforest origin. However, it is not just that primarily in rainforest and which extends widely such taxa may have been left in situ as the con- through other southern habitats within the gen- tinent has become dryer and wet forest habitats eral "desert" region but they are excluded from have shrunk. There are also behavioural and life the extreme southwest and southeast of the con- history factors peculiar to the relic genera which tinent. This distribution suggests a tropical ori- favour their persistence. gin and possibly a relatively recent entry into Furthermore the weather patterns, as well as Australia, especially for Conothele and Seleno- the vegetation cover and topography of rainfor- cosmia both of which also occur to the north of est and "desert" refuges play a big part in main- Australia, a hypothesis already espoused by taining the genera in their peripheral range. Main (1981, 1982). Cethegus in having partly Northern rainforest and open forests and wood- and perhaps primarily a tropical and subtropical land regions are dominated by summer rain rainforest distribution (Main, 1960; Raven,

(Nix, 1981, see Figure 1); the semiarid and 1984c) can similarly be regarded as being a desert, south of the tropics by winter rain. recent denizen of the semiarid. (Neme- However, as well as winter rain which may be siidae) and Missulena (Actinopodidae) occur very irregular, the whole of the vast inland also widely throughout the continent including the experiences some summer rain associated with tropical north. Aname also occurs in Tasmania isolated thunderstorms and cyclones or mon- from where however Missulena is absent. The soonal rains which occasionally extend well distribution of ldiommata () may south of the tropics. These unpredictable rain parallel that of the three selected genera and events now profoundly affect the persistence of some of its behavioural attributes may similarly "tropical" elements of the mygalomorph fauna account for its broad habitat inclusion. The in desert regions. is currently under review by Raven

( 1 994). The remaining genera have more restric-

ted ranges (Table 1 ). Tropical rainforest genera — distribution and All those genera occurring in rainforest appear characteristics to be dependent on a moist, shaded habitat and

It is now pertinent to state that of the 57 cur- species of the three genera also occurring in rently accepted mygalomorph genera in Aus- "desert" areas selected for discussion are restric- tralia, 37 occur in rainforests and of these 26 ted to more or less permanently moist habitats as

occur in tropical rainforests (Table 1 ). Of the lat- I will show below. ter, many genera also occur in open forest and some have species in drier habitats ranging from Behavioural and life style characteristics of heath and desert. The following woodland to rainforest "deserticoles" seven genera are found only in rainforests: Kiama, Australothele, Carrai. Masteha, Plesio- Most genera which exhibit strong adaptations to desert living are regarded as old. autochthonous thele, Migas and Sason . Of these, Mastena and Sason are restricted to northern i.e. tropical Australian genera (Main, 1981 ). The three rainforest. Data on rainforest and other relevant "rainforest" genera under discussion persist in habitat distributions to arrive at the information the desert firstly by avoiding the desert environ- are situated in refugia however min- summarised in Table I is derived from Main ment (they (1985a) where bibliographic sources are given iscule), and secondly by retaining a suite of life style attributes appropriate e.g. taxonomic literature associated with species behavioural and descriptions and later publications (mainly to their original rainforest habitat and which 342 B. Y. MAIN

now predispose or fortuitously fit them to sur- cal features which reduce water loss and various vive in generalized arid areas. (In view of the combinations of behavioural characteristics controversy surrounding such terms as pre- such as (1) avoidance of environmental con-

adaptation and related contrivances, I am reluc- ditions through fossorial habits (2) sedentary life tant to add to the confusion by using similar style e.g. burrow site fidelity (3) specialised for- words like "a predisposition" but nevertheless aging strategies and seasonal feeding by non- dare to coin the term "fortuitous adap- aestivating segments of the population (4) tation"!). coincidence of reproductive behaviour and dis- Main (1982) summarised the desert adap- persion ofjuveniles with rainy periods (however tations of mygalomorphs to include morphologi- erratic) (5) extreme longevity of females (6)

Table 1. The 26 genera which occur in tropical rainforest and the extent of their distribution i.e. to include southern areas and whether absent from the southwest and southeast corners of Australia. * = 14 genera which extend into "desert" (arid/semiarid). P = present; sw,

southwest, se , south east.

Tropical Rainforest Widespread East/Aust Absent only se/sw

Actinopodidae * Missulena + * Conothele + Dipluridae * Cethegus +

Namirea -f Masteria +ne Qld — Hexathelidae * Hadronyche +& Eyre P P se * Arbanitis P Cataxia +Qld only — Homogona + Pse

Migas + P se * Aname + P * Chenistonia + — P * Kwonkan +w half Aust — Psw Xamiat us + Namea + *?Yilgamia +w half Aust Theraphosidae * Sclenocosmia + Barychelidae * Idiommata + Psw * Synothele + WA/SA Psw Trittamc -i-ne Sason +ne Zophorame +ne * * Ozycrypta +ne/cent Tungari +ne +n/trop TROPICAL RAINFOREST SPIDERS IN THE AUSTRALIAN DESERT 343

capacity to fast for long periods (7) iteroparous similar sites in Western Australia and various reproduction and (8) limited dispersal capacity mountain blocks ("ranges") in Central Aus- in most species . tralia. Although rocky, these sites due to their At the same time these attributes were given as geological structure encourage seepage into soil explaining the restricted geographic ranges of interstices. This dampness combined with shade many such species which " are tied to particular albeit of shrubs and tussocks provides a habitat habitat types categorized by soil/vegetation that mimics in microcosm an earlier rainforest attributes". Conversely it was pointed out that habitat, of which indeed such sites are relics. Conothele has a wide geographic range "assisted (2) Elsewhere in open country of the dry by its aerial dispersal" and that "the large, interior the spiders are found in association with aggressive and relativley mobile Selenocosmia rock tumbles around low hills, and in clay and stirlingi has been able to colonize unstable habi- alluvial soils in small depressions of drainage tats of the interior". lines. Thus they occur in either relic microhabi- It is these contrary attributes, plus additional tats (valleys of mountain ranges which also ones but all reminiscent of their rainforest heri- retain ancient flora such as Livistona ) or in habi- tage, of Conothele and Selenocosmia and shared tats with verisimilitude. by Cethegus, which I wish to emphasise here as (3) The burrow of Conothele in most sites is being the "fortuitous adaptations" enabling relatively shallow, and lined with a stocking-like them to live in the desert region. In other words tube of tough silk that adheres to a plaster wall they live in the desert but are not the of and with a tightly-fitting, cap-like silk/soil door. desert. The base of the burrow unless in the process of In considering the "fortuitous adaptations" of being deepened is usually fully lined. Thus the Conothele, Selenocosmia and Cethegus the burrow acts like a sealed flask — preventing salient ones are probably associated with the fol- flooding during immersion in wet tropical sites. lowing factors: Secondarily this secure tube in flood prone areas 1. Persistence in relic microhabitats within the in the otherwise dry inland similarly protects the arid region; spiders and conversely ameliorates environmen- 2. Adoption of microhabitats with verisimili- tal dessicating conditions. tude with rainforest habitats; (4) Juveniles disperse aerially (although it is 3. Burrow/nest structure; assumed not over great distances) which assists 4. Dispersion method of juveniles; dissemination in rugged terrain or habitats with 5. Mobility i.e. capacity to relocate nest/burrow patchy shrubbery or rocky sites. The method is site; advantageous in steep sided, rocky gorges as well 6. Foraging behaviour; and as open woodland with discontinuous favour- 7. Reproductive behaviour and phenology. able microhabitats. I will now discuss each of the genera in turn to show the relevance of these factors to persistence (5) In unstable creek banks, as in the rainforest of the respective taxa as deserticoles. (such as in the Kimberley and many northern Australian sites) and steep slopes in open forest Conothele e.g., north Queensland and New Guinea, if the The genus has a wide distribution from burrows are washed over by sheet flooding or dislodged the spiders are securely Burma, various island groups through New even partly Guinea and Australia (Roewer, 1942; Main, cocooned because of the sealed stocking struc- ture of the nest and the spiders can relocate or 1981; Raven, 1985). It occurs generally in rain- reestablish after being buffeted by sudden eros- forest and open forest in humid, tropical conditions. regions. Nests are usually in the ground but some ive rainforest species are arboreal and make small (6) Spiders are typical sit-and-wait predators cocoon-like tubes with trap doors in bark, thus and do not fully emerge from the burrow when avoiding inundation in very wet habitats (Main, capturing prey. This possibly limits them to 1993). Spiders are moderate sized, with a high-prey habitats. However the habitats they smooth shiny cuticle, very spiny anterior legs occupy would also appear to favour invertebrate and apart from courting males and dispersing density. Spiders do not store rejectamenta in the juveniles spiders never leave the nest. basally sealed burrow but eject it at the surface (I) In semiarid and arid regions in Australia (personal observations) which again implies a spiders occur in shaded cliff faces of mountain certain transitoriness unlike the extreme seden- valleys as in the Hamersley Ranges and other tary nature of most arid adapted species which )

344 B. Y. MAIN

are more vulnerable to sudden disturbance of around the burrow entrance. In addition they

habitat (and habitually store rejectamenta). emerge to hunt or actively chase prey . (7) Little is known about the reproductive (7) Spiders are mostly summer breeding, behaviour of the genus other than that wander- which in the monsoon forests at least ties in with ing of males and juvenile dispersion is linked the "wet". In the arid region spiders are more with rainy periods (at whatever season). opportunistic, males taking advantage of irregu- lar summer rains to wander. Longevity of females enables individual spiders to forego breeding "Selenocosmia" during drought years while persisting as a population (a common strategy of arid The genus occurs from to New Guinea adapted mygalomorphs (Main, 1976, 1978)). and Australia (Roewer. 1942). However the family representatives of Theraphosidae in Aus- Cethegus tralia require taxonomic revision and the gen- Commonly called "curtain web spiders", eric category "Selenocosmia" is loosely applied these spiders have long spinnerets (associated in the present context. Schmidt (1995) trans- with their profuse web building). They are mod- ferred the Australian species of Selenocosmia to erately sized, hairy, with relatively long legs Phlogius in which genus early species were orig- which tend to turn backwards at the tips. inally placed. The spiders are very large and Although they are web weavers, they are remark- hairy with long legs. Spiders make very deep ably agile on the ground and can move very burrows. quickly. In rainforests the spiders make diffuse silk tubes amongst rocks, in logs or on irregular ( 1 ), (2) and (3) While it is difficult to identify cliff faces. In monsoon forest they frequently "desert" sites of a relictual nature, most habitats occur in the rocky river beds. These are inun- (throughout the inland in various States) cer- dated during the "wet" when the spiders are tainly have verisimilitude in that they are possibly washed out or enclosed in their floccu- lowlying, wet or seasonally wet, in relatively lent silk and temporarily submerged. unstable situations and may be flooded during There is no data to suggest that spiders leave their the rainy season. Because the spiders are already burrows and climb trees or vegetation priorXo the onset of adjusted to inundation of burrows (e.g. in inundation as occurs with some South seasonal "swamps" or "bogs" in the Kimberley American Ischnothelines (Hofer, 1 990) but this is another inundation in flood prone, arid flats is not inimi- possibility. However, their cal. It is possible that the very deep burrows taxonomic affinity with this group suggests also a behavioural maintain air pockets. Also the furriness of the flexi- bility which may include spiders' bodies encourages formation of an a predisposition to transient web siting. enclosing air bubble thus spiders may be immersed without drowning. The earliest (1) Like Conothele, Cethegus often occurs in records of the habitat of the "barking" or "whis- rocky cliff faces which determine seepage of tling" spider, Selenocosmia stir/ingi is in the moisture to the surface or in cracks in "relict" classic account of the spider in The Report ofThe habitats in gorges and valleys in the arid Horn Expedition (Spencer, 1896) where the region. spiders are noted as occurring in "grassy flats (2) and (3) A notable behaviour in open, semi- amongst low hills". These would be the swales arid habitats is that spiders site the burrow and between the ranges and sand dunes, low lying curtain web against a supporting butt of a shrub areas subject to sheet flooding and temporary or small tree which secondarily provides the boggy conditions following intermittent rain. benefit of harvesting water into the burrow They in this way resemble seasonal wet/dry which ramifies the soil amongst the roots. Water depressions where other species occur in the harvesting by trees and shrubs in arid areas is a Kimberley and north Queensland (pers. obs.) well known phenomenon (Slatyer, 1 965; Nulsen (4) Little is known about juvenile dispersion et al„ 1986). other than that it is assumed spiderlings scatter (4 ) Juveniles are aerially dispersed (Main, freely on the ground (Kotzman, 1 986). 1995 and unpublished records). (5) Spiders appear to be readily mobile and (5) Spiders appear to relocate nest sites in there is some evidence (Kotzman, 1986 and unstable rainforest habitats (inferred behaviour) pers. obs.) that spiders move sites at least in arid (Main, 1 993) and certainly relocate (due to rain habitats. damaged webs and burrows) after rain in semi- (6) Spiders may lay an entrapping mesh of web arid habitats. The relocation of numerous nests TROPICAL RAINFOREST SPIDERS IN THE AUSTRALIAN DESERT 345 has been documented at Durokoppin (Main, "deserticoles", by virtue of retaining their rain- 1993, pers. obs., unpublished records). forest life style and repertoire of adaptations (6) Spiders ensnare prey in their webs and take (transmuted as "fortuitous adaptations") are a variety of crawling and flying insects and other masquerading as "deserticoles". . The web probably ensnares a higher In the light of this the implications for conser- prey take than that available to typical sit-and- vation management are obvious. Especial care wait trapdoor spiders. needs to be directed to preserving intact relic- (7) As with other genera reproductive behav- sites in gorges and less easily identifiable sites iour and dispersion is opportunistic depending with "verisimilitude". In that rainforest habitats on seasonal or irregular rain. However there is are generally not tire prone, relic habitats within some evidence (pers. records) that males the arid zone need to be protected to prevent an develop more rapidly than most mygalomorph edge erosion by fires. With the present infil-

species . Precocious development, as argued by tration of tourism (see also Morton et al.. 1995)

Main ( 1 99 1 ) would mean lower mortality due to that is oriented to once off, casual, sight seeing shortened exposure to environmental hazards visitations, the increasing danger of disturbance and thereby increased surety of reproduction in simply by people pressure to many biologically unstable, flood prone (tropical) and climatically resilient but physically brittle sites is inevitable. unpredictable (arid) habitats. And this is quite apart from the inimical effects of too frequent fires upsetting the habitats and faunal populations. Discussion The spiders cannot be the only potential vic- The three tropical rainforest genera Conothele, tims — but they are surely the silent advocates Cethegus and Selenocosmia have been shown for a whole suite of invertebrates. from the taxonomic literature, museum collec- tions and personal observations and collections Acknowledgments to be also widely distributed throughout Aus- tralia, including the low rainfall areas but exclus- I am grateful to the many colleagues who, over ive of the mesophytic southwest and south east the years, have presented me with relevant speci- forests and southern coastal regions. Evidence mens of the genera discussed here. I acknowl- respective from natural history observations and distri- edge the following curators and bution records substantiates the hypothesis that museums for making available relevant collec- these genera survive in the broad semiarid to tions, particularly of Conothele specimens: M.R. desert region, not by having evolved specific Gray, Australian Museum, Sydney ; R.J. Raven, Malipatil adaptations to the desert but by retaining adap- Queensland Museum, Brisbane; M. Territory Museums tations which primarily fitted them to the insta- and G. Brown, Northern and Art Galleries, Darwin; K. Walker and G. bility of wet (even if only seasonally wet) tropical Melbourne; M.S. habitats. Milledge, Museum of Victoria, Australian Perth. I At the same time the spiders do not appear to Harvey, Western Museum, the referee's comments. have expanded their habitat range but rather to also appreciated have become restricted to relic rainforest habi- tats or to "newer" microhabitats which exhibit References verisimilitude, at least seasonally, with wet trop- Churchill, T.B. and Raven, R.J., 1992. Systcmalics ol where Cethegus occurs in the ical habitats. Even the intertidal trapdoor spider genus Idioclis dry, open "desert" country of the Western Aus- (Mygalomorphae: Barychclidac) in the western tralian Goldfields. which appears to have little Pacific with a new genus from the northeast.

there are ( ): 9- resemblance to tropical rainforest, Memoirs of the Queensland Museum 32 1 reminders of an earlier, wetter scenario in the 30. scattered presence of the kurrajong trees, Bra- Ewing, T.L., 1983. Beetles and other insects of tropical forest canopies at Manaus, Brazil, sampled by chychiton gregorii . Cycads and the Livistona insecticidal fogging. Pp. 59-75 in: Sutton, ST.. palms persist in gorges of the Central Australian Whitmore, T.C. and Chadwick, A.C. (eds). Black- Ranges and the latter in some isolated gorges in well: Edinburgh. et al., northwestern Australia (Humphreys Gray, M.R., 1987. Distribution of the funnel web these spiders demonstrate 1990). Nor do spiders. Pp. 3 1 3-32 1 in: Covacevich, .1., Davie. P. specific behavioural specialisations comparable and Pearn, P. (eds). Twde plants and — a to those of the endemic, autochthonous genera. guide for Australia. Queensland Museum: Bris- bane. Thus it seems that the apparent rainforest 346 B. Y. MAIN

Gray, M.R.. 1992. Funnel-webs: separating fact Main. B.Y., 1986. Further studies on the systematics from fiction. Australian Natural History 24(3): of Australian Diplurinae (Araneae: Mygalomor- 32-39. phae: Dipluridae): a new genus from south-west- Hofer. H., 1990. The spider community (Araneae) of a ern Australia. Records of the Western Australian Central Amazonian blackwater inundation forest Museum 12 (4): 395-402. (igapo). Acta Zoologica Fennica 190: 173-179. Main, BY., 1990. Dwarf males in mygalomorph Humphreys. W.F.. Brooks, R.D. and Bine. B., 1990. spiders: adaptation to environmental hazards. Rediscovery of the palm Livistona alfredii on Acta Zoologica Fennica 190: 273-278. North West cape Peninsula. Records of the West- Main. B.Y., 1991a. Kimberley spiders: rainforest ern Australian Museum 14: 647-650. strongholds. Pp. 271-294 in: McKenzie. N.L.,

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