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Australian Assassins, Part III: A review of the Assassin Spiders (Araneae, Archaeidae) of tropical north-eastern Queensland

Abstract The assassin spiders of the family Archaeidae from tropical north-eastern Queensland are revised, with eight new species described from rainforest habitats of the Wet Tropics bioregion and Mackay-Whitsundays Hinterland: Austrarchaea griswoldi sp. n., Austrarchaea hoskini sp. n., Austrarchaea karenae sp. n., Austrarchaea tealei sp. n., Austrarchaea thompsoni sp. n., Austrarchaea wallacei sp. n., Austrarchaea westi sp. n. and Austrarchaea woodae sp. n. Specimens of the only previously described species, Austrarchaea daviesae Forster & Platnick, 1984, are redescribed from the southern Atherton Tableland. The rainforests of tropical eastern Queensland are found to be a potential hotspot of archaeid diversity and endemism, with the region likely to be home to numerous additional short-range endemic taxa. A key to species complements the taxonomy, with maps, natural history information and conservation assessments provided for all species.


Introduction
Few families of Australian spiders are as distinctive or as enigmatic as the 'assassin spiders' of the family Archaeidae, renowned for their unique cephalic morphology, strange araneophagic biology, great phylogenetic antiquity and relictual biogeography across the Southern Hemisphere. Although once considered to be one of the rarest of spiders families -and certainly one of the least understood in terms of taxonomy, phylogeny and biology -recent and ongoing research in the U.S.A., South Africa and Australia has shed increasing light on this remarkable lineage of spiders (see Platnick 1991a, 1991b, Lotz 1996, 2003, 2006, Harvey 2002a, Griswold et al. 2005, Wood et al. 2007, Wood 2008, 2012a, 2012b. Indeed, archaeids are now one of the better understood families of Araneae, with the southern African and Malagasy faunas continuing to be revised and phylogenetically tested, and the Australian fauna now almost completely revised taxonomically and phylogenetically. The last 20 years have seen large numbers of new species discovered and described in both the Afro-Malagasy and Australian regions (Platnick 2012), and archaeids (and their relatives) continue to feature heavily in studies exploring the classification and phylogeny of the basal Araneomorphae (e.g. Schütt 2000, Griswold et al. 2005, Rix et al. 2008. Australian Archaeidae have been progressively revised over the last two years, with 26 new species described since 2011, taking the total number of currently described Australian species to 30 (Platnick 2012). Rix and Harvey (2011) first documented the Archaeidae of 'mid-eastern' Australia, re-describing the only previously named species from the region, and describing 17 new species from south-eastern Queensland and eastern New South Wales. This revision first noted the possibility of two major taxonomic, phylogenetic and biogeographic disjunctions in eastern and southern Australia, highlighting the St Lawrence Gap (Webb and Tracey 1981) in central eastern Queensland, and the Australian Alps in south-eastern Australia as likely candidates (see Fig.  2). Subsequent molecular phylogenetic research by Rix and Harvey (2012b) tested and further confirmed these phylogenetic disjunctions, highlighting especially the allopatric separation of a deeply-divergent southern Australian (i.e. Victorian, South Australian and Western Australian) lineage from all other eastern Australian taxa. Rix and Harvey (2012b) also confirmed the importance of the St Lawrence Gap, between Gladstone and Mackay, as an important phylogenetic and biogeographic barrier between monophyletic clades of Austrarchaea Forster & Platnick, 1984 from tropical north-eastern Queensland and mid-eastern Australia (see Figs 2-3); a result congruent with morphological observations by Rix and Harvey (2011, figs 5E-G). A second taxonomic revision by Rix and Harvey (2012a) formally documented the distinctive southern Australian lineage, describing the new genus Zephyrarchaea Rix & Harvey, 2012a, along with nine new species from south-western Western Australia, Kangaroo Island (South Australia) and southern Victoria. As a result of these monographic and phylogenetic revisions, a surprisingly diverse Australian archaeid fauna has now been revealed, one dominated by large numbers of mostly allopatric, short-range endemic species, all of which are restricted to the 'mesic zone' (Byrne et al. 2011) of mainland Australia. Archaeids are ubiquitous in the tropical and subtropical upland rainforests of eastern Australia, and only those species from north of the St Lawrence Gap remain to be revised (Fig. 2).
The current paper -the third and final in a series revising the Archaeidae of Australia -presents a taxonomic revision of the assassin spiders from tropical north-eastern Queensland, including those species from the Mackay-Whitsundays Hinterland and the Wet Tropics bioregion, between Cooktown and Townsville (Figs 2,25). This revision takes the total number of described Australian Archaeidae to 38 species, with the genus Austrarchaea now including 27 described short-range endemic species.

Material and methods
All taxa were described and illustrated from specimens stored in 75% or 95% ethanol. Digital images were taken using a Leica MZ16A binocular microscope and a Leica DM2500 compound microscope, with auto-montage images captured using Leica DFC500 mounted cameras with Leica Application Suite Version 3.6.0 software. Male left pedipalps were dissected prior to imaging and bulbs were aligned for standardised comparison in the ventral and retrolateral positions illustrated. Female genitalia were dissected and cleared in a 10% lactic acid plus 90% glycerol solution, prior to mounting on temporary glass slides and imaging in a postero-ventral position ( Fig. 14G; see also Rix and Harvey 2011, fig . 5D), usually with genital plates removed (Fig. 7G). This postero-ventral alignment allowed for a much clearer presentation of the spermathecae, while still permitting inter-specific comparison at an equivalent (albeit opposite) plane to that of Harvey (2011, 2012a). Illustrations were made on Utoplex tracing paper, using printed template auto-montage images. Maps were generated using ArcMap version 9.3.1 (ESRI Inc.) with Virtual Earth (Microsoft Corp.).
Measurements are in millimetres (rounded to the nearest hundredth of a millimetre) and were taken using an ocular graticule on a Leica M80 binocular microscope. Left legs were removed from specimens prior to taking measurements and imaging lateral body profiles. Lateral profile images were standardised for inter-specific comparison by vertically aligning the centre of each left anterior median eye with the lower anterior margin of the carapace (above the labrum) Harvey 2011, 2012a). Carapace height was measured in lateral view, from the margin of the pars thoracica above coxa II to the highest point of the pars cephalica (Fig. 5). Carapace length was measured from the lower anterior margin of the carapace (above the labrum) to the posterior margin of the pars thoracica (above the pedicel) (Fig. 5). 'Neck' width was measured in lateral view, at the narrowest point of the carapace, with total length, carapace width, abdomen length and abdomen width all measured in dorsal view. To quantify inter-specific variation in the shape of the cephalothorax and 'head', three morphometric ratios (the carapace height to carapace length [CH/CL] ratio; the post-ocular ratio [P.O. ratio]; and the highest point of pars cephalica [HPC] to postocular length ratio) were derived from lateral profile images as defined and discussed by Harvey (2011, 2012a). For Material Examined sections, specimens not examined for the current revision, but currently housed at (or on loan to) the California Academy of Sci-ences (due to ongoing research) are listed separately, with identifications confirmed by H. Wood; one additional, unseen vial (QMB S50322) was identified as A. daviesae according to geographic proximity (Fig. 16). Species descriptions and numbering of the pedipalpal sclerites follows Rix and Harvey (2011) (but see also Remarks, below Diagnosis. Species of Austrarchaea can be distinguished from all southern Australian species of Zephyrarchaea by the significantly taller carapace (CH/CL ratio ≥ 2.0), by the presence of accessory setae on the distal bulge of the male cheliceral paturon, and by the fusion of the two conductor sclerites on the male pedipalp (Rix and Harvey 2012a, fig. 4). Australian Archaeidae are further distinguished from Old World taxa by the presence of numerous, clustered spermathecae in females (Fig. 7G), and by the presence of a long, wiry embolus on the pedipalp of males (Fig. 4).
Description. For a full generic description see Rix and Harvey (2011). For notes on genitalia and morphological differences among lineages of Austrarchaea, see Remarks (below).
Distribution. Species of Austrarchaea occur in mesic habitats throughout eastern Queensland and New South Wales (Fig. 3), usually in montane rainforests (Figs 1E-F), but also in lowland rainforests or wet eucalypt forests on or adjacent to the Great Dividing Range . In north-eastern Queensland, archaeids occur throughout the Wet Tropics bioregion, from the Mount Finnigan Uplands (near Cooktown) south to Mount Elliot (near Townsville) 25). In the Mackay and Whitsundays Hinterland region, archaeids can be found in the Eungella National Park (near Mackay), north to Mount Dryander (south of Bowen) (Figs 24-25). The genus is not known to occur south or west of the Australian Alps (Fig. 2), which may be a vicariant biogeographic barrier between populations of Austrarchaea and Zephyrarchaea Harvey 2012a, 2012b).
Remarks. The genus Austrarchaea includes three major lineages in eastern Australia (Figs 3-4), each readily distinguished by the morphology of the abdomen and the structure of the male pedipalp (Fig. 4). The most widespread lineage (the A. nodosa species-group) occurs south of the St Lawrence Gap, from Kroombit Tops National Park in central Queensland, south to the Badja State Forest in southern New South Wales (Fig. 3); species in this lineage possess six dorsal hump-like tubercles on the abdomen and an exposed tegular cavity with a variably scutiform conductor (Fig. 4). The second, most restricted lineage (the A. monteithi lineage) is known only from the Gibraltar Range National Park in northern New South Wales (Fig. 3); the single known species, A. monteithi, possesses five dorsal hump-like tubercles on the abdomen and an exposed tegular cavity with a hooked conductor (Fig. 4). The third lineage (the A. daviesae species-group; revised in this paper) occurs north of the St Lawrence Gap, from Eungella National Park north to Cooktown (Figs 3,25); species in this lineage possess only four dorsal hump-like tubercles on the abdomen (recumbent in A. woodae sp. n.) and a more enclosed tegular cavity with a very large, arched conductor (Figs 4,(6)(7)(8)(9)(10)(11)(12)(13)(14)(15).
Although the derived pedipalpal morphology of A. daviesae and its relatives is strikingly different to that of congeners further south, the distal tegular sclerites can nonetheless be broadly homologised with those of A. nodosa and A. monteithi on the basis of their shape and relative position in the unexpanded tegular cavity. The embolus in all nine known north-eastern Queensland species is a long, sinuous, strongly sclerotized process emerging from the distal bulb pro-ventrally, in some species bearing an additional accessory spur. Tegular sclerite 3 (TS 3) is always a prominent, pro-ventrally directed process, which is fused to the retro-ventral margin of the tegular bulb (the latter of which is usually also concomitantly modified). Tegular sclerite 2 (plus 2a, i.e. TS 2-2a) is usually inserted just behind TS 3 in the unexpanded tegular cavity, forming a distinctive, mesally-looped and distally whip-like structure common to all taxa in the A. daviesae species-group; the extent of this very long, whip-like TS 2a is usually proximate to the distal extension of the embolus in the unexpanded state. This TS 2-2a morphology is in stark contrast to that of A. monteithi, A. nodosa and related species, in which TS 2a is usually covered and largely obscured by a more spur-like TS 2 process. Tegular sclerite 1 (TS 1) -generally the most prominent sclerite in species of Zephyrarchaea and other species of Austrarchaea -is reduced and often obscured in most archaeid species from north-eastern Queensland, although a few taxa possess a larger, more distinctive TS 1 posterior to the TS 2-2a complex (e.g. Fig. 9D). Inter-specific variation among taxa in the A. daviesae species-group is pronounced, with male pedipalp morphologies usually highly autapomorphic for each species. Five broad pedipalp types (Types A-E) can be distinguished among north-eastern Queensland taxa, with Type A being the most common form, shared between five of the nine known species, and Types B-E each currently unique to single species. Figure 6 highlights differences between these different pedipalp morphologies, which are further diagnosed in the Key to species (see below).
Distribution and habitat. Austrarchaea daviesae is known from the 'Misty Mountains' region of the southern Atherton Tableland, in the vicinity of Ravenshoe and Millaa Millaa, with additional specimens also known from Mount Bartle Frere in the adjacent Wooroonooran National Park (see Remarks, below) (Figs 16,25). Specimens have been collected in pitfall and flight intercept traps, by beating vegetation, or by beating and sifting elevated leaf litter at the bases of lawyer vine palms (Calamus spp.) in dense tropical rainforest (Fig. 1F).
Conservation status. This species has a relatively widespread distribution in several National Parks protected under World Heritage legislation, and is not considered to be of conservation concern.
Remarks. The identification and distribution of Austrarchaea daviesae has, until recently, been difficult to ascertain, as the holotype male (QMB S1091; Fig. 7B) is without pedipalps (these presumably having been mounted on SEM stubs as per Forster and Platnick 1984, figs 70-74). Similarly, no adult male specimens had been collected from the Atherton Tableland since the original holotype collection in 1978. Fortunately, an adult male and female were collected in early 2012, from the paratype locality (Malaan National Park), near the type locality of Majors Mountain. These specimens (WAM T125183), described above, closely conform to original descriptions, and the male pedipalp appears indistinguishable from that illustrated in Forster and Platnick (1984, figs 70-74). Interestingly, the distribution of A. daviesae appears to extend beyond the Atherton Tableland, with eastern populations apparently sympatric or at least partly sympatric with A. woodae sp. n. on Mount Bartle Frere, in the Wooroonooran National Park. Adult Mount Bartle Frere specimens collected by the California Academy of Sciences in 2009 are conspecific with specimens from Malaan National Park, as confirmed by pedipalp images supplied by H. Wood (pers. comm.). Another juvenile specimen from Mount Bartle Frere (CASENT 9023672), collected in 2006, is also conspecific with these adult Mount Bartle Frere specimens, as determined by almost identical COI sequences (H. Wood, pers. comm.). Interestingly, female specimens collected by the QM from Boulder Caves, near the type locality of A. woodae sp. n., also appear to be A. daviesae (rather than A. woodae sp. n.), due to the presence of fully developed (rather than recumbent) abdominal tubercles, and a similar carapace morphology and similar genitalia to specimens from the Atherton Tableland. Austrarchaea woodae sp. n. thus appears to be much rarer than A. daviesae at altitudes ≤ ~1000 m, and may actually be restricted to higher altitude montane rainforest on the summit of Mount Bartle Frere. Etymology. The specific epithet is a patronym in honour of the late Doug Wallace OAM , for his passion and enthusiasm for arachnology, for his contributions to the study of Australian (and especially Queensland) spiders, for his efforts in founding and fostering the Rockhampton Arachnological Society, and for his encouragement of MGR over many years.

Austrarchaea wallacei
Diagnosis. Austrarchaea wallacei can be distinguished from all other Archaeidae from north-eastern Queensland except A. karenae sp. n., A. tealei sp. n. and A. thomp-soni sp. n. by the presence of a triangular spur on the embolus (Fig. 8D); from A. thompsoni sp. n. by the presence of a prominent, triangular tegular sclerite 1 (TS 1) (Fig. 8D); and from A. karenae sp. n. and A. tealei sp. n. by the shape of tegular sclerite 3 (TS 3), which has a bluntly pointed, triangular apex (Figs 8C-D).
Female: Unknown. Distribution and habitat. Austrarchaea wallacei is known only from the summit of Mount Misery, 34 km north-west of Cape Tribulation (Figs 17,25). The single known specimen was collected in a pitfall trap in tropical rainforest at 850 m elevation.
Conservation status. Unknown (data deficient). Etymology. The specific epithet is a patronym in honour of Dr Karen Edward, for her contributions to our understanding of Wet Tropics biogeography, and for her great friendship to MGR and MSH over many years.

Austrarchaea karenae
Diagnosis. Austrarchaea karenae can be distinguished from all other Archaeidae from north-eastern Queensland except A. tealei sp. n., A. thompsoni sp. n. and A. wallacei by the presence of a triangular spur on the embolus (Figs 9D-E); from A. thompsoni sp. n. by the presence of a prominent, triangular tegular sclerite 1 (TS 1), which is vis-ible in ventral view (Fig. 9D); and from A. tealei sp. n. and A. wallacei by the shape of tegular sclerite 3 (TS 3), which has a single, sharply pointed process distally (Fig. 9D).
Female: Unknown. Distribution and habitat. Austrarchaea karenae is known only from the Windsor Tableland, 44 km north-west of Mossman (Figs 18, 25). The single known specimen was collected in high altitude tropical rainforest.
Conservation status. Unknown (data deficient). Etymology. The specific epithet is a patronym in honour of Geoff Thompson, for his ongoing efforts in collecting and documenting the invertebrate rainforest fauna of the Wet Tropics, and for collecting the only known specimen of this species.

Austrarchaea thompsoni
Diagnosis. Austrarchaea thompsoni can be distinguished from all other Archaeidae from north-eastern Queensland except A. karenae, A. tealei sp. n. and A. wallacei by the presence of a triangular spur on the embolus (Fig. 10D); and from A. karenae, A. tealei sp. n. and A. wallacei by the very small tegular sclerite 1 (TS 1), which is not visible in ventral view (Fig. 10D), and by the more proximally positioned embolic spur, which is situated near the base of the exposed embolic portion (Fig. 10D).
Female: Unknown. Distribution and habitat. Austrarchaea thompsoni is known only from Devils Thumb, on the Carbine Tableland 10 km west-north-west of Mossman (Figs 19,25). The single known specimen was collected in high altitude tropical rainforest.
Conservation status. Unknown (data deficient). Diagnosis. Austrarchaea tealei can be distinguished from all other Archaeidae from north-eastern Queensland except A. karenae, A. thompsoni and A. wallacei by the presence of a triangular spur on the embolus (Figs 11E-F); from A. thompsoni by the presence of a prominent, triangular tegular sclerite 1 (TS 1), which is visible in ventral view (Fig. 11E); and from A. karenae and A. wallacei by the shape of tegular sclerite 3 (TS 3), which has a second short, pointed process distally (Fig. 11E).
Distribution and habitat. Austrarchaea tealei is known only from Mossman Gorge, 4.5 km west-south-west of Mossman (Figs 20, 25). Specimens have been collected under logs (as newly-hatched juveniles; H. Wood, pers. comm.), or by beating and sifting elevated leaf litter at the bases of lawyer vine palms (Calamus spp.) in lowland tropical rainforest.
Remarks. The female specimen described above (from the ANIC) is tentatively identified as conspecific with the holotype of A. tealei, despite a somewhat different carapace morphology and a fairly imprecise collection locality. Austrarchaea thompsoni does occur on nearby mountains above the Mossman River, and thus it possible (albeit unlikely) that the female specimen from "Mossman Gorge" (collected in 1967) may actually belong to another species. We have described it here in the absence of evidence suggesting any sympatry in the Mossman Gorge region, given the fact that all other recently collected Mossman Gorge material appears to be conspecific with the holotype (including CAS material; H. Wood, pers. comm.), and given the similarly small body size of this female specimen and the holotype male (Fig. 5). Other material examined. AUSTRALIA: Queensland: Dinden National Park: same data as holotype, 1 juvenile (QMB S59537).

Austrarchaea westi
Etymology. The specific epithet is a patronym in honour of Paul West, for his friendship to MSH over many years, and for helping fund the Western Australian Museum's 'archaeid project' from 2009-2012.
Diagnosis. Austrarchaea westi can be distinguished from all other Archaeidae from north-eastern Queensland by the presence of a unique Type B pedipalp (Fig. 6), with very small bulb (width << 0.30 mm) (Figs 6, 12D), and by the relatively short embolus, which is distally enclosed within the conductor (Figs 6, 12D). This species can be further distinguished by the very short, barely differentiated accessory setae on the male chelicerae (Fig. 12B).
Female: Unknown. Distribution and habitat. Austrarchaea westi is known only from Mount Williams, on the Lamb Range 11 km west of Cairns (Figs 21, 25). The two known specimens were collected in high altitude tropical rainforest.
Conservation status. Unknown (data deficient). Diagnosis. Austrarchaea woodae can be distinguished from all other Archaeidae from north-eastern Queensland by the presence of a unique Type C pedipalp (Fig. 6), with a proximally constricted conductor (Figs 6, 13D), large, flattened, distally folded tegular sclerite 3 (TS 3) (Figs 6, 13D-E), and apple-shaped bulb profile in ventral view (Figs 6, 13C-D). This species can be further distinguished by the dense, pick-like tuft of accessory setae on the male chelicerae ( Fig. 13B; similar only to A. harmsi among Australian Archaeidae), and by the almost spherical abdomen with recumbent hump-like tubercles ( Fig. 13A; similar only to species of Zephyrarchaea among Australian Archaeidae).
Female: Unknown. Distribution and habitat. Austrarchaea woodae is known only from near the summit of Mount Bartle Frere, 12 km south-west of Babinda (Figs 22, 25). The only known specimen was collected in high altitude tropical rainforest.
Etymology. The specific epithet is a patronym in honour of Dr Conrad Hoskin, for his contributions to our understanding of Wet Tropics biogeography, and for his efforts in documenting the remarkable endemic biota of Mount Elliot.
Distribution and habitat. Austrarchaea hoskini is known only from Mount Elliot, 30 km south-east of Townsville (Figs 23, 25). The few known specimens were collected in high altitude rainforest along North Creek.

Austrarchaea spp. (unidentified specimens)
Note. In the absence of adult male specimens or molecular data, the following female and juvenile specimens (see Figs 16-23, 25) could not be confidently identified as known species. Species of Austrarchaea are difficult to identify (and diagnose) by females alone, and in the Wet Tropics these difficulties were compounded by the absence of representative adults from across the region. Material is thus here listed according to upland subregional zones of faunal endemism, as proposed for the Wet Tropics bioregion (see Discussion, below; Table 1 Paratypes: Allotype female, same data as holotype (QMB S92213); 2 males, 1 female and 2 juveniles, same data as holotype (WAM T125630); 1 female, same data as holotype except Broken River Rainforest Discovery Circuit, hand collecting at night, 24.III.2012 (QMB S92214).
Other Etymology. The specific epithet is a patronym in honour of Dr Charles Griswold, for his outstanding contributions to arachnology, and for his contributions to the study of Archaeidae and other basal Araneomorphae.
Diagnosis. Austrarchaea griswoldi can be distinguished from all other Archaeidae from north-eastern Queensland by the presence of a unique Type E pedipalp (Fig. 6), with a very large bulb (width >> 0.30 mm) (Figs 6, 15E), modified ventro-distal rim of the tegulum forming rectangular opercular plate (Figs 6, 15E), and very large, flattened tegular sclerite 3 (TS 3), the latter extending along the entire retrolateral edge of the conductor (Fig. 15F). This species can be further distinguished by the very short, barely differentiated comb of accessory setae on the male chelicerae (Fig. 15C), and by the presence of only two pairs of female spermathecae (Fig. 15G).
Distribution and habitat. Austrarchaea griswoldi is known only from Eungella National Park, 70 km west of Mackay (Figs 24-25). Specimens have been collected by beating and sifting elevated leaf litter in tropical rainforest (Fig. 1E), especially under the dead fronds of Eungella Fan Palms (Livistona sp.).
Natural history. A single female specimen was collected by MGR during night collecting in March 2012, suspended with her egg-sac in a tangled maternal web decorated with hanging debris, at the base of a large standing rainforest tree trunk. This egg-sac (Fig. 1D) was carried with both legs IV, positioned behind and against the posterior face of the abdomen, and was composed of soft brown silk. The shape of the egg-sac was irregular, with two large projections, and 18 spiderlings hatched out of the egg-sac on 3-4 April 2012.
Conservation status. This species appears to be a short-range endemic taxon (Harvey 2002b, Harvey et al. 2011), which although potentially restricted in distribution, is abundant within the Eungella National Park (MGR, pers. obs.). It is not considered to be of conservation concern. Austrarchaea spp. (unidentified specimens) Note. In the absence of adult male specimens or molecular data, the following female specimens (see

Discussion
The Wet Tropics World Heritage Area. The Australian Wet Tropics bioregion, situated in north-eastern Queensland between Cooktown and Townsville (Figs 16-23, 25), is a World Heritage area renowned for its rich rainforest biota and very high levels of local endemism (e.g. see Williams et al. 1996, Crisp et al. 2001, Yeates et al. 2002, Slatyer et al. 2007, and references therein). Much has been written about the biogeography of the region, and numerous seminal contributions over several decades have resulted in the Wet Tropics becoming a model landscape for understanding processes of rainforest biogeography, speciation and diversification, in both plant and animal taxa (e.g. Williams et al. 1996, Schneider et al. 1998, Moritz et al. 2000, Crisp et al. 2001, Yeates et al. 2002. Much of this research has focussed on Pleistocene climatic fluctuations, and the concomitant effects these fluctuations have had on the vicariant biogeography, phylogeography and/or speciation of different taxa, especially vertebrates (e.g. Schneider et al. 1998, Schneider and Moritz 1999, Hoskin et al. 2011, Bell et al. 2012). However, as highlighted by Hoskin et al. (2011), few vertebrate lineages have undergone in situ radiation within the Wet Tropics, and most show little phenotypic divergence despite often strong phylogeographic signal; evidence for a deeper and more complex history of speciation. Different taxa also highlight a variety of responses and wildly different patterns of distribution and endemism at different spatial scales (Hoskin et al. 2011), and this is especially true of invertebrates, which often show "extraordinarily high" levels of diversity and endemism compared to vertebrates (Bell et al. 2007: 4995; see also Yeates et al. 2002). Indeed, for flightless or low vagility arthropods, the Wet Tropics have aptly been described as an "epicentre of evolution" (Bouchard 2002: 449), and much remains to be tested in order to understand historical mechanisms of speciation (and subsequent extinction, range contraction or dispersal) in both space and time. Allopatric speciation within Pliocene or Pleistocene refugia has been suggested for at least several endemic insect and vertebrate lineages (e.g. Bell et al. 2007, Hoskin et al. 2011, although deeper, Miocene-age divergences are increasingly being implicated in the major diversification of the Australian Wet Tropics fauna (see Moritz et al. 2000).
Patterns of distribution within the Wet Tropics have historically been assessed in terms of 'regional endemism' (i.e. those species confined to the Wet Tropics) versus 'subregional endemism' (i.e. those species confined to a single subregion within the Wet Tropics) (Yeates et al. 2002), and an extensive subregional classification has been developed and modified for the entire Wet Tropics over nearly 30 years (e.g. see Winter et al. 1984, Monteith 1995, Williams et al. 1996, Schneider et al. 1998, Moritz et al. 2001, Edward 2011 (Table 1; Figs 16B-23B). This subregional classification, separating adjacent upland and lowland forest blocks, has provided a useful foundation for assessing patterns of distribution, diversity and endemism throughout the Wet Tropics, and has been widely tested or applied in studies of vertebrates (Williams et al. 1996, Schneider et al. 1998, Schneider and Moritz 1999, Dolman and Moritz 2006, Hoskin et al. 2011, Bell et al. 2012) and invertebrates (e.g. Bouchard 2002, Yeates et al. 2002, Bell et al. 2007, Edward 2011, Boyer and Reuter 2012. These studies include comparative analyses of endemicity, biogeographical concordance and conservation significance within and between subregions (e.g. Moritz et al. 2001, Yeates et al. 2002, as well as more traditional estimates of phylogeny, biogeography and phylogeography across a suite of co-occurring taxa. For all such analyses, the historical biogeographic significance of the major upland subregions has been consistently demonstrated, with often strong concordance between palaeoclimatic modelling and phylogeographic structure (Yeates et al. 2002, Hoskin et al. 2011. Archaeidae in the Wet Tropics. Assassin spiders appear to be largely ubiquitous in upland rainforests throughout most of the Wet Tropics, extending from the Finnigan Uplands near Cooktown south to the Elliot Uplands near Townsville (Fig. 25). Ar- Table 1. List of upland subregional zones of faunal endemism identified for the Wet Tropics bioregion (by Winter et al. 1984, Williams et al. 1996 and other authors) (see Discussion; Figs 16B-23B), noting current collection records of Archaeidae, including the presence of any described species. Subregional zones are listed from the northern-most Mt Finnigan Uplands (Fig. 16B) to the southern-most Elliot Uplands (Fig. 23B). Note the addition of Hinchinbrook Island (as per Edward 2011), and the current absence of archaeid collections from four of the southern subregions. F = female specimen/s; J = juvenile specimen/s; M = male specimen/s; N.R. = no recorded specimens. chaeidae seem to be less common in lowland tropical rainforests (true of most species of Austrarchaea throughout their range), however populations from Mossman Gorge and near Cape Tribulation in the northern Daintree National Park suggest that they may be more widespread in lowland forest systems than current collection records suggest. Indeed, given the relatively high proportion of sites represented only by juvenile specimens or unidentified females (Fig. 25), the tendency for short-range endemism in Austrarchaea generally, and the very small number of adult male specimens available for taxonomic research, the Wet Tropics may actually be home to a significantly larger number of archaeid species than documented in this revision. For example, sites like Mount Bartle Frere support two sympatric or at least partially sympatric species (Figs 16,22), and in the Mossman River region of the southern Daintree National Park, different taxa appear to occupy different lowland (A. tealei) versus upland (A. thompsoni) habitats in relatively close proximity (Figs 19-20). Apart from A. daviesae, which is known from two adjacent subregions (Fig. 16B) Table 1).

Wet Tropics Upland Subregion
Estimating the actual number of Archaeidae in north-eastern Queensland is a difficult task, given the surprisingly small number of collection records for the region, the very small number of adult male specimens available, and the related absence in all but two instances of anything other than single-point distributions for most known species (Fig. 25). However, available records do provide some tantalising clues, and hint at the likelihood of a possible hotspot of archaeid diversity in the Wet Tropics. Indeed, with (i) at least four other upland subregional zones with known archaeid records but for which adult male specimens are unavailable, (ii) four additional upland subregions which may harbour archaeid populations but are currently without collections (Table  1), (iii) the likelihood that at least a minority of Wet Tropical subregions may harbour multiple endemic species, either sympatrically or in upland versus lowland habitats, and (iv) the likely presence of at least one additional species in a separate upland zone of the Whitsundays region (Fig. 24), the actual number of taxa in the A. daviesae species-group is almost certainly > 50% larger than currently recognised. Thus, at a conservative estimate, there may be 15 or more short-range endemic species in tropical Queensland, a number almost equivalent to the total archaeid diversity of mid-eastern Australia. These figures are perhaps not surprising, given the remarkable levels of diversity and endemism seen in other invertebrate groups (see Yeates et al. 2002), but raise the question of how (and when) this diversity was generated. Rix and Harvey (2012b) inferred an Eocene divergence date for taxa in the A. daviesae species-group (relative to mid-eastern Australian Austrarchaea), suggesting that the monophyletic archaeid fauna of north-eastern Queensland has evolved in isolation for 35-50 million years -a result at least consistent with the high levels of interspecific genitalic variation seen across this lineage relative to other Australian clades (Fig. 6). However, geographic sampling for both molecules and morphology is currently inadequate across the Wet Tropics, and a more detailed, thoroughly-sampled molecular study is required to properly assess patterns of speciation in the A. daviesae species-group, and address the significant gaps in our understanding of divergence dates, distributional boundaries and inter-specific relationships within this lineage. The group's diversity, strict reliance on rainforest habitats and relative ubiquity in the Wet Tropics certainly makes them an ideal candidate for testing patterns of speciation and biogeography throughout the region, and it is our hope that this revision will provide a solid taxonomic foundation for future research. at the Queensland Museum who have, over three decades, carefully and systematically documented the biodiversity and conservation values of the Queensland Wet Tropics World Heritage Area. We acknowledge the enormous funding and logistical investments that have been made to this end, and recognise the importance of the Queensland Museum's collections as repositories of this remarkable natural heritage.     Fig. 3). Note the variation in the shape of the male pedipalp and the marked differences in the shape and orientation of the conductor (C), embolus (E) and the distal tegular sclerites. Note also the number of abdominal hump-like tubercles (1-6): four in the A. daviesae species-group; five in A. monteithi; and six in the A. nodosa species-group. Figure 5. Graphs depicting the relationship between carapace length (CL) and carapace height (CH) for species of Austrarchaea from north-eastern Queensland. Smaller grey dots denote species of Austrarchaea from mid-eastern Australia (see Rix and Harvey 2011); smaller grey squares denote species of Zephyrarchaea from southern Australia (see Rix and Harvey 2012a). Note the relatively large body sizes of A. hoskini sp. n. and A. woodae sp. n., and the body size variation between populations of A. daviesae from the Atherton Tableland (AT) and Mount Bartle Frere (Mt BF), respectively.                    . Summary distribution of the Austrarchaea daviesae species-group in tropical north-eastern Queensland, showing collections records for described species (labelled, with black circles) and unidentified juveniles or females (yellow circles) (see Table 1). Note the high proportion of unidentified specimens, especially within the Wet Tropics bioregion between Cooktown and Ingham.