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The tropical and subtropical rainforests of Australia’s eastern mesic zone have given rise to a complex and highly diverse biota. Numerous old endemic, niche-conserved groups persist in the montane rainforests south of Cooktown, where concepts of serial allopatric speciation resulting from the formation of xeric interzones have largely driven our biogeographic understanding of the region. Among invertebrate taxa, studies on less vagile arachnid lineages now complement extensive research on vertebrate taxa, and phylogenetic studies on mygalomorph spiders in particular are revealing significant insights about the biogeographic history of the Australian continent since the Eocene. One mygalomorph lineage entirely endemic to Australia’s tropical and subtropical eastern rainforests is the open-holed trapdoor spider genus Namea Raven, 1984 (family Anamidae). We explore, for the first time, the phylogenetic diversity and systematics of this group of spiders, with the aims of understanding patterns of rainforest diversity in Namea, of exploring the relative roles of lineage overlap versus in situ speciation in driving predicted high levels of congeneric sympatry, and of broadly reconciling morphology with evolutionary history. Original and legacy sequences were obtained for three mtDNA and four nuDNA markers from 151 specimens, including 82 specimens of Namea. We recovered a monophyletic genus Namea sister to the genus Teyl Main, 1975, and monophyletic species clades corresponding to 30 morphospecies OTUs, including 22 OTUs nested within three main species-complex lineages. Remarkable levels of sympatry for a single genus of mygalomorph spiders were revealed in rainforest habitats, with upland subtropical rainforests in south-eastern Queensland often home to multiple (up to six) congeners of usually disparate phylogenetic affinity living in direct sympatry or close parapatry, likely the result of simultaneous allopatric speciation in already co-occurring lineages, and more recent dispersal in a minority of taxa. In situ speciation, in contrast, appears to have played a relatively minor role in generating sympatric diversity within rainforest ‘islands’. At the population level, changes in the shape and spination of the male first leg relative to evolutionary history reveal subtle but consistent interspecific morphological shifts in the context of otherwise intraspecific variation, and understanding this morphological variance provides a useful framework for future taxonomic monography. Based on the phylogenetic results, we further provide a detailed taxonomic synopsis of the genus Namea, formally diagnosing three main species-complexes (the brisbanensis-complex, the dahmsi-complex and the jimna-complex), re-illustrating males of all 15 described species, and providing images of live spiders and burrows where available. In doing so, we reveal a huge undescribed diversity of Namea species from tropical and subtropical rainforest habitats, and an old endemic fauna that is beginning to shed light on more complex patterns of rainforest biogeography.

In Australia, climate change and continental drift have given rise to a complex biota comprising mesic specialists, arid-adapted lineages, and taxa that have arrived on the continent from Asia. We explore the phylogenetic diversity and biogeographic history of the Australian trapdoor spider genus Conothele Thorell, 1878 that is widespread in Australia’s monsoonal tropics and arid zone. We sequenced three mtDNA and five nuDNA markers from 224 specimens. We reconstructed the phylogenetic relationships among specimens and estimated the number of operational taxonomic units (OTUs) using species delimitation methods. The timing of divergences was estimated and ancestral area reconstructions were conducted. We recovered 61 OTUs, grouped into four major clades; a single clade represented by an arboreal ecomorph, and three fossorial clades. The Australian Conothele had a crown age of ~19 million years, and ancestral area reconstructions showed a complex history with multiple transitions among the monsoonal tropics, central arid zone, south-west and Pilbara bioregion. Conothele arrived on the continent during periods of biotic exchange with Asia. Since then, Conothele has colonised much of the Australian arid and monsoonal zones, during a period of climatic instability. The Pilbara bioregion harbours high lineage diversity, emphasising the role of climate refugia.

Compared with araneomorph spiders, relatively few mygalomorph spiders have evolved an obligate existence in subterranean habitats. The trapdoor spider genus Troglodiplura Main, 1969 and its sole named species T. lowryi Main, 1969 is endemic to caves on the Nullarbor Plain of southern Australia, and is one of the world’s most troglomorphic mygalomorph spiders. However, its systematic position has proved to be difficult to ascertain, largely due to a lack of preserved adults, with all museum specimens represented only by cuticular fragments, degraded specimens or preserved juveniles. The systematic placement of Troglodiplura has changed since it was first described as a member of the Dipluridae, with later attribution to Nemesiidae and then back to Dipluridae. The most recent hypothesis specifically allied Troglodiplura with the Neotropical subfamily Diplurinae, and therefore was assumed to have no close living relatives in Australia. We obtained mitochondrial sequence data from one specimen of Troglodiplura to test these two competing hypotheses, and found that Troglodiplura is a member of the family Anamidae (which was recently separated from the Nemesiidae). We also reassess the morphology of the cuticular fragments of specimens from several different caves, and hypothesise that along with T. lowryi there are four new troglobitic species, here named T. beirutpakbarai Harvey & Rix, T. challeni Harvey & Rix, T. harrisi Harvey & Rix, and T. samankunani Harvey & Rix, each of which is restricted to a single cave system and therefore severely threatened by changing environmental conditions within the caves. The first descriptions and illustrations of the female spermathecae of Troglodiplura are provided. The family Anamidae is further divided into two subfamilies, with the Anaminae Simon containing Aname L. Koch, 1873, Hesperonatalius Castalanelli, Huey, Hillyer & Harvey, 2017, Kwonkan Main, 1983, Swolnpes Main & Framenau, 2009 and Troglodiplura, and the Teylinae Main including Chenistonia Hogg, 1901, Namea Raven, 1984, Proshermacha Simon, 1909, Teyl Main, 1975 and Teyloides Main, 1985. ZooBank Registration: http://zoobank.org/References/2BE2B429-0998-4AFE-9381-B30BDC391E9C

The pseudoscorpions of the genus Garypus L. Koch are restricted to seashore habitats where they occupy supralittoral and littoral zones primarily in tropical and subtropical areas. Few species have been recorded from the Indo-West Pacific region, and this project was devised to produce a review of the species found in museum collections and to test the relationships of the various garypid genera using a molecular analysis and an assessment of their morphology. A new subfamily classification is proposed with the subfamilies Garypinae, including Garypus and the new genus Anchigarypus Harvey (type species Garypus californicus Banks), and the Synsphyroninae for the other genera (Ammogarypus Beier, Anagarypus Chamberlin, Elattogarypus Beier, Eremogarypus Beier, Meiogarypus Beier, Neogarypus Vachon, Paragarypus Vachon, Neogarypus Vachon, Synsphyronus Chamberlin, and Thaumastogarypus Beier). The species-level revision of Garypus provides evidence for at least 14 species, most of which are known from only single localities. The following species are redescribed: G. insularis Tullgren from the Seychelles, G. krusadiensis Murthy & Ananthakrishnan from India and Sri Lanka, G. longidigitus Hoff from Queensland, Australia, G. maldivensis Pocock from the Maldives, G. nicobarensis Beier from the Nicobar Islands and G. ornatus Beier from the Marshall Islands. The holotype of G. insularis is a tritonymph, and not therefore readily identifiable. Nine new species are described: G. latens Harvey, sp. nov., G. malgaryungu Harvey, sp. nov., G. necopinus Harvey, sp. nov., G. postlei Harvey, sp. nov., G. ranalliorum Harvey, sp. nov. and G. weipa Harvey, sp. nov. from northern Australia, G. dissitus Harvey, sp. nov. from Cocos-Keeling Island, G. reong Harvey, sp. nov. and G. yeni Harvey, sp. nov. from Indonesia. A further possible new species from Queensland is described but not named, as it is represented by a single tritonymph. The subspecies of the Caribbean species G. bonairensis Beier are elevated to full species status: G. bonairensis, G. realini Hummelinck and G. withi Hoff. We supplement the descriptions with sequence data from five specimens from four species of Garypus and two species of Anchigarypus, and find COI divergence levels of 7–19% between Garypus species. http://zoobank.org/References/16463E29-6F13-4392-9E41-46A4312C852B

There is a yet uncovered multitude of species to be found among Western Australian Onychophora. Kumbadjena, one of the two genera that reside in this region, has been previously suggested to house an extensive species complex. Morphology alone has not been able to elucidate the diversity in this genus and has instead muddled species delineations. Topologies and species delimitation analyses resulting from the sequences of two mitochondrial ribosomal markers (12S rRNA and 16S rRNA), one nuclear ribosomal marker (18S rRNA), and one mitochondrial protein-coding gene (cytochrome c oxidase subunit I) are indicative of several undescribed species. Fixed diagnostic nucleotide changes in the highly conserved sequences of 18S rRNA warrant distinction of three new species of Kumbadjena: K. toolbrunupensis, sp. nov., K. karricola, sp. nov., and K. extrema, sp. nov. The geographic distributions of the proposed species suggest that Kumbadjena is another example of short-range endemism, a common occurrence in the flora and fauna of the region. The extensive biodiversity and endemism in the region necessitates conservation to preserve the species and processes that promote speciation harboured by Western Australia.

The Australasian spiny trapdoor spiders of the family Idiopidae (subfamily Arbanitinae) are revised at the generic level, using a multi-locus molecular phylogenetic foundation and comprehensive sampling of all known lineages. We propose a new family- and genus-group classification for the monophyletic Australasian fauna, and recognise 10 genera in four tribes. The Arbanitini Simon includes Arbanitis L. Koch, 1874 (61 species), Blakistonia Hogg, 1902 (one species) and Cantuaria Hogg, 1902 (43 species). The Aganippini Simon includes Bungulla Rix, Main, Raven & Harvey, gen. nov. (two species), Eucanippe Rix, Main, Raven & Harvey, gen. nov. (one species), Eucyrtops Pocock, 1897 (two species), Gaius Rainbow, 1914 (one species) and Idiosoma Ausserer, 1871 (14 species). The Cataxiini Rainbow and Euoplini Rainbow include just Cataxia Rainbow, 1914 (11 species) and Euoplos Rainbow, 1914 (12 species), respectively. Two distinctive new genera of Aganippini are described from Western Australia, and several previously valid genera are recognised as junior synonyms of existing genus-group names, including Misgolas Karsch, 1878 (= Arbanitis; new synonymy), Aganippe O. P.-Cambridge, 1877 (= Idiosoma; new synonymy) and Anidiops Pocock, 1897 (= Idiosoma; new synonymy). Gaius stat. rev. is further removed from synonymy of Anidiops. Other previously hypothesised generic synonyms are supported by both morphology and molecular phylogenetic data from 12 genes, including the synonymy of Neohomogona Main, 1985 and Homogona Rainbow, 1914 with Cataxia, and the synonymy of Albaniana Rainbow & Pulleine, 1918, Armadalia Rainbow & Pulleine, 1918, Bancroftiana Rainbow & Pulleine, 1918 and Tambouriniana Rainbow & Pulleine, 1918 with Euoplos. At the species level, the identifications of Eucy. latior (O. P.-Cambridge, 1877) and I. manstridgei (Pocock, 1897) are clarified, and three new species are described: Bungulla bertmaini Rix, Main, Raven & Harvey, sp. nov., Eucanippe bifida Rix, Main, Raven & Harvey, sp. nov. and Idiosoma galeosomoides Rix, Main, Raven & Harvey, sp. nov., the latter remarkable for its phragmotic abdominal morphology. The Tasmanian species Mygale annulipes C. L. Koch, 1842 is here transferred to the genus Stanwellia Rainbow & Pulleine, 1918 (family Nemesiidae), comb. nov., Arbanitis mestoni Hickman, 1928 is transferred to Cantuaria, comb. nov. and Idiosoma hirsutum Main, 1952 is synonymised with I. sigillatum (O. P.-Cambridge, 1870), new synonymy. In addition to the morphological synopses and an illustrated key to genera, molecular diagnoses are presented for all nominal taxa, along with live habitus and burrow images to assist in field identification. The Australasian idiopid fauna is highly diverse, with numerous new species known from all genera. As a result, this study provides a taxonomic and nomenclatural foundation for future species-level analyses, and a single reference point for the monographic documentation of a remarkable fauna. http://zoobank.org/?lsid=urn:lsid:zoobank.org:pub:BACE065D-1EF9-40C6-9134-AADC9235FAD8

The trapdoor spider family Nemesiidae comprises 14 genera in Australia, the majority of which are included in the subfamily Anaminae. Here we provide evidence from a multigene molecular analysis of most Australian genera of Anaminae for a previously unrecognised clade that also differs from its sister-genus, Aname L. Koch, by the lack of a prominent asetose ventral depression on the pedipalpal tibia and the medium-sized mating spur on tibia I of males. This depression is a characteristic of all species of Aname examined to date, and represents a newly recognised character system in the subfamily. The new genus, named Hesperonatalius, is represented by three new species – H. maxwelli, sp. nov., H. harrietae, sp. nov. and H. langlandsi, sp. nov. – all from arid Western Australia. http://zoobank.org/References/D5352390-5D21-49DD-A123-A074422EB860

Fully troglobitic pseudoscorpions are rare in the Afrotropical Region, and we explored the identity and phylogenetic relationships of specimens of a highly modified troglobite of the family Gymnobisiidae in the dark zone of the Wynberg Cave system, on Table Mountain, South Africa. This large pseudoscorpion – described as Gymnobisium inukshuk Harvey & Giribet, sp. nov. – lacks eyes and has extremely long appendages, and has been found together with other troglobitic fauna endemic only to this cave system. Phylogenetic analyses using the nuclear ribosomal genes 18S rRNA and 28S rRNA and the mitochondrial protein-encoding gene cytochrome c oxidase subunit I unambiguously place the new species with other surface Gymnobisium from South Africa. This placement receives strong support and is stable to analytical treatments, including static and dynamic homology, parsimony and maximum likelihood, and data removal for ambiguously aligned sites. This species is the first troglobitic species of the family and one of the most highly modified pseudoscorpions from the Afrotropical Region. http://zoobank.org/urn:lsid:zoobank.org:pub:5227092B-A64B-4DB3-AD90-F474F0BA6AED

The phylogenetic relationships of the Australian species of Feaellidae are assessed with a molecular analysis using mitochondrial (CO1) and nuclear (ITS2) data. These results confirm the morphological analysis that three previously undescribed species occur in the Pilbara bioregion, which are named and described: Feaella (Tetrafeaella) callani, sp. nov., F. (T.) linetteae, sp. nov. and F. (T.) tealei, sp. nov. The males of these three species, as well as males of F. anderseni Harvey and other unnamed species from the Kimberley region of north-western Australia, have a pair of enlarged, thick-walled bursa that are not found in other feaellids. Despite numerous environmental impact surveys for short-range endemic invertebrates in the Pilbara, very few specimens have been collected, presumably due to their relictual distributions, restricted habitat preferences and low densities. http://zoobank.org/urn:lsid:zoobank.org:pub:131F0587-F2EE-405F-BE5A-772F072D9915

The trapdoor spider family Migidae has a classical Gondwanan distribution and is found on all southern continents except the Indian region. The Australian fauna consists of three genera including Moggridgea O. P. Cambridge from south-western Australia and Kangaroo Island, South Australia; Moggridgea is otherwise widespread throughout Africa. The sole named species of Moggridgea from Western Australia, M. tingle Main, and its unnamed relatives are the subject of the present paper, which was stimulated by concern for the long-term persistence of populations, and the discovery of deep genetic divergences between populations. A phylogeny of the Western Australian species relative to African and South Australian Moggridgea was generated using molecular COI and ITS rDNA data, and based on both molecular and morphological criteria we conclude that the Western Australian taxa should be removed from Moggridgea and transferred to a new genus, Bertmainius. The seven species are delimited using both morphological and molecular criteria: B. tingle (Main) (the type species), and six new species, B. colonus, B. monachus, B. mysticus, B. opimus, B. pandus and B. tumidus. All seven species are considered to be threatened using IUCN criteria, with the major threatening processes being inappropriate fire regimes and climate change.

Birds harbour a wide array of other taxa in their nests or in their plumage, which either have an ectoparasitic or commensal relationship with the host. We report on the discovery of a cheliferid pseudoscorpion found in the plumage and nests of the sociable weaver bird (Philetairus socius) in southern Africa. The nests of these communal birds are the largest of any bird, and may contain up to 500 individuals. The pseudoscorpion is likely to have a mutualistic relationship with the birds, most likely preying on other small invertebrates in the nests. Molecular data derived from two populations of the pseudoscorpion found divergence levels of 1.1% in cytochrome oxidase 1 (CO1), and an analysis of CO1 and two rRNA genes (18S and 28S) found a close relationship with Chelifer and Parachelifer in the tribe Cheliferini, which is supported by the morphology of the male genitalia. The molecular analysis also suggests that Beierius may not belong to the Cheliferini. The pseudoscorpion found in association with the sociable weaver represents a new genus and species, Sociochelifer metoecus Harvey, sp. nov.

Palpigradi are a poorly understood group of delicate arachnids, often found in caves or other subterranean habitats. Concomitantly, they have been neglected from a phylogenetic point of view. Here we present the first molecular phylogeny of palpigrades based on specimens collected in different subterranean habitats, both endogean (soil) and hypogean (caves), from Australia, Africa, Europe, South America and North America. Analyses of two nuclear ribosomal genes and COI under an array of methods and homology schemes found monophyly of Palpigradi, Eukoeneniidae and a division of Eukoeneniidae into four main clades, three of which include samples from multiple continents. This supports either ancient vicariance or long-range dispersal, two alternatives we cannot distinguish with the data at hand. In addition, we show that our results are robust to homology scheme and analytical method, encouraging further use of the markers employed in this study to continue drawing a broader picture of palpigrade relationships.

The Yilgarn and Pilbara regions of Western Australia are considered biodiversity hotspots for subterranean invertebrates. While the relatively well studied (aquatic) stygofauna are typically constrained to geographically isolated habitats (‘subterranean islands’) and have likely originated from multiple independent epigean ancestors, the troglofauna found in cavernicolous calcretes and fractured rock remains largely unstudied. Here we focus on the pseudoscorpion genera Tyrannochthonius Chamberlin, 1929 and Lagynochthonius Beier, 1951, as common components of the troglofauna, to determine whether they also display highly restricted distributional patterns, and have independent origins. Bayesian and maximum likelihood analyses of sequence data from the mtDNA cytochrome c oxidase I (COI) and the small subunit 18S nuclear genes for subterranean and epigean species from both genera reveal divergent mtDNA lineages that are restricted to single aquifers and/or geographic locations. This strong geographic structuring of troglobitic pseudoscorpions is indicative of short-range endemism and supports the ‘subterranean island’ hypothesis. Further, independent sister relationships between subterranean and epigean taxa indicate multiple invasions into subterranean habitats, likely driven by post-Miocene aridification, consistent with that predicted for the stygofauna. The phylogeny also reveals that Tyrannochthonius + Lagynochthonius is monophyletic but that Lagynochthonius is polyphyletic and nested inside Tyrannochthonius. The results of this study point to common processes that have shaped the diversity and uniqueness of both stygofaunal and troglofaunal communities in Western Australia.

The Pilbara bioregion of Western Australia is an area that contains vast mineral deposits and unique ecosystems. To ensure that mineral deposits are mined with minimal impact on the natural environment, impact assessment surveys are required to determine what fauna and flora species are located within proposed development areas, in particular, by determining the distributions of short-range endemic species (SREs). One infraorder of Arachnida, the Mygalomorphae (trapdoor spiders and their kin), are frequently identified as SREs. These identifications are traditionally performed using morphological techniques; however, only males can be reliably identified to species. Furthermore, the majority of species have not been formally described and males comprise only ~5% of specimens collected. To assess mygalomorph diversity and the distribution of species in the Pilbara, we employed a molecular barcoding approach. Sequence data from the mitochondrial DNA cytochrome c oxidase subunit I (COI) gene were obtained from 1134 specimens, and analysed using Bayesian methods. Only a fraction of the total mygalomorph fauna of the Pilbara has been documented, and using a species boundary cut-off of 9.5% sequence divergence, we report an increase in species richness of 191%. Barcoding provides a rapid, objective method to help quantify mygalomorph species identifications and their distributions, and these data, in turn, provide crucial information that regulatory authorities can use to assess the environmental impacts of large-scale developments.

A major challenge confronting many contemporary systematists is how to integrate standard taxonomic research with conservation outcomes. With a biodiversity crisis looming and ongoing impediments to taxonomy, how can systematic research continue to document species and infer the ‘Tree of Life’, and still maintain its significance to conservation science and to protecting the very species it strives to understand? Here we advocate a systematic research program dedicated to documenting short-range endemic taxa, which are species with naturally small distributions and, by their very nature, most likely to be threatened by habitat loss, habitat degradation and climate change. This research can dovetail with the needs of industry and government to obtain high-quality data to inform the assessment of impacts of major development projects that affect landscapes and their biological heritage. We highlight how these projects are assessed using criteria mandated by Western Australian legislation and informed by guidance statements issued by the Environmental Protection Authority (Western Australia). To illustrate slightly different biological scenarios, we also provide three case studies from the Pilbara region of Western Australia, which include examples demonstrating a rapid rise in the collection and documentation of diverse and previously unknown subterranean and surface faunas, as well as how biological surveys can clarify the status of species thought to be rare or potentially threatened. We argue that ‘whole of biota’ surveys (that include all invertebrates) are rarely fundable and are logistically impossible, and that concentrated research on some of the most vulnerable elements in the landscape – short-range endemics, including troglofauna and stygofauna – can help to enhance conservation and research outcomes.

The orb-weaving spider subfamily Arkyinae L. Koch, 1872 is exclusively found in the Australasian region and its taxonomy and the systematic relationships within and between genera of this subfamily are poorly understood. We here revise the arkyine genus Demadiana Strand, 1929 to include six Australian species, four of which are described as new: Demadiana simplex (Karsch, 1878) (type species), D. carrai, sp. nov., D. cerula (Simon, 1908), comb. nov., D. complicata, sp. nov., D. diabolus, sp. nov., and D. milledgei, sp. nov. A phylogenetic analysis based on an updated araneid morphological data matrix including 57 genera of orb-weaving spiders identified Demadiana as a member of the araneid subfamily Arkyinae. A separate phylogenetic analysis for the genus at the species level showed little resolution within Demadiana, but did identify a monophyletic Demadiana supported by three putative synapomorphies: small unique setal pits with spherical sockets covering the carapace, sternum and the bases of the paturon (chelicerae), an extreme elongation of the trumpet-like aggregate spigots of the posterior lateral spinnerets and a distinct curvature of the embolus. We detail several new generic and species synonymies within Arkyinae. Aerea Urquhart, 1891 (type species Aerea alticephala Urquhart, 1891) and Neoarchemorus Mascord, 1968 (type species N. speechleyi Mascord, 1968) are regarded as junior synonyms of Arkys Walckenaer, 1837 (type species A. lancearius Walckenaer, 1837), resulting in Arkys speechleyi (Mascord, 1968), comb. nov. Aerea magnifica Urquhart, 1893 and Archemorus simsoni Simon, 1893 are regarded as junior synonyms of Aerea alticephala Urquhart, 1891, and Arkys nitidiceps Simon, 1908 is proposed as a junior synonym of Arkys walckenaeri Simon, 1879.

South-western Western Australia is a biodiversity hotspot, with high levels of local endemism and a rich but largely undescribed terrestrial invertebrate fauna. Very few phylogeographic studies have been undertaken on south-western Australian invertebrate taxa, and almost nothing is known about historical biogeographic or cladogenic processes, particularly on the relatively young, speciose Quaternary sand dune habitats of the Swan Coastal Plain. Phylogeographic and taxonomic patterns were studied in textricellin micropholcommatid spiders belonging to the genus Raveniella Rix & Harvey. The Micropholcommatidae is a family of small spiders with a widespread distribution in southern Western Australia, and most species are spatially restricted to refugial microhabitats. In total, 340 specimens of Raveniella were collected from 36 surveyed localities on the Swan Coastal Plain and 17 non-Swan Coastal Plain reference localities in south-western Western Australia. Fragments from three nuclear rRNA genes (5.8S, 18S and ITS2), and one mitochondrial protein-coding gene (COI) were used to infer the phylogeny of the genus Raveniella, and to examine phylogeographic patterns on the Swan Coastal Plain. Five new species of Raveniella are described from Western Australia (R. arenacea, sp. nov., R. cirrata, sp. nov., R. janineae, sp. nov., R. mucronata, sp. nov. and R. subcirrata, sp. nov.), along with a single new species from south-eastern Australia (R. apopsis, sp. nov.). Four species of Raveniella were found on the Swan Coastal Plain: two with broader distributions in the High Rainfall and Transitional Rainfall Zones (R. peckorum Rix & Harvey, R. cirrata); and two endemic to the Swan Coastal Plain, found only on the western-most Quindalup dunes (R. arenacea, R. subcirrata). Two coastally restricted species (R. subcirrata, R. janineae) were found to be morphologically cryptic but genetically highly distinct, with female specimens morphologically indistinguishable from their respective sister-taxa (R. cirrata and R. peckorum). The greater Perth region is an important biogeographic overlap zone for all four Swan Coastal Plain species, where the ranges of two endemic coastal species join the northern and south-western limits of the ranges of R. peckorum and R. cirrata, respectively. Most species of Raveniella were found to occupy long, highly autapomorphic molecular branches exhibiting little intraspecific variation, and an analysis of ITS2 rRNA secondary structures among different species of Raveniella revealed the presence of an extraordinary hypervariable helix, ranging from 31 to over 400 nucleotides in length.

Australia was historically considered a poor prospect for subterranean fauna but, in reality, the continent holds a great variety of subterranean habitats, with associated faunas, found both in karst and non-karst environments. This paper critically examines the diversity of subterranean fauna in several key regions for the mostly arid western half of Australia. We aimed to document levels of species richness for major taxon groups and examine the degree of uniqueness of the fauna. We also wanted to compare the composition of these ecosystems, and their origins, with other regions of subterranean diversity world-wide. Using information on the number of ‘described’ and ‘known’ invertebrate species (recognised based on morphological and/or molecular data), we predict that the total subterranean fauna for the western half of the continent is 4140 species, of which ~10% is described and 9% is ‘known’ but not yet described. The stygofauna, water beetles, ostracods and copepods have the largest number of described species, while arachnids dominate the described troglofauna. Conversely, copepods, water beetles and isopods are the poorest known groups with less than 20% described species, while hexapods (comprising mostly Collembola, Coleoptera, Blattodea and Hemiptera) are the least known of the troglofauna. Compared with other regions of the world, we consider the Australian subterranean fauna to be unique in its diversity compared with the northern hemisphere for three key reasons: the range and diversity of subterranean habitats is both extensive and novel; direct faunal links to ancient Pangaea and Gondwana are evident, emphasising their early biogeographic history; and Miocene aridification, rather than Pleistocene post-ice age driven diversification events (as is predicted in the northern hemisphere), are likely to have dominated Australia’s subterranean speciation explosion. Finally, we predict that the geologically younger, although more poorly studied, eastern half of the Australian continent is unlikely to be as diverse as the western half, except for stygofauna in porous media. Furthermore, based on similar geology, palaeogeography and tectonic history to that seen in the western parts of Australia, southern Africa, parts of South America and India may also yield similar subterranean biodiversity to that described here.

We present a description of the first indigenous member of the arachnid order Palpigradi from Australia. Eukoenenia guzikae, sp. nov. was collected from subterranean environments in the Yilgarn region of Western Australia. The sole male specimen differs from all other members of the genus in several small but significant ways, including by the combined presence of six blades in the prosomal lateral organs, nine pairs of setae on the propeltidium, the presence of a spur on coxa IV, the chaetotaxy of sternites IV–VIII, and the shape of the male genital lobes. It shows some similarities in the male genital region to a group of species found in Madagascar.

We studied a variety of chthoniid pseudoscorpions collected from the semiarid zone of Western Australia. Five new species of Tyrannochthonius Chamberlin, 1929 and three new species of Lagynochthonius Beier, 1951 are named and described from the Pilbara and Gasgoyne regions, and surrounding areas such as Barrow Island. Tyrannochthonius basme, sp. nov. and Lagynochthonius asema, sp. nov. from pisolitic mesas near Pannawonica, T. garthhumphreysi, sp. nov. from limestone karst on Barrow Island, T. souchomalus, sp. nov. from calcrete deposits near Cue, T. billhumphreysi, sp. nov. and L. polydentatus, sp. nov. from a calcrete deposit on Sturt Meadows Station and L. leemouldi, sp. nov. from calcrete near Marble Bar are all considered to represent hypogean species as all exhibit typical troglomorphic adaptations including total loss of eyes and attenuated appendages. New records are provided for T. brooksi Harvey and T. butleri Harvey from Cape Range peninsula. A new epigean species, T. aridus, sp. nov., was found on Barrow Island and the Pilbara mainland. Two further putative new species based upon nymphal specimens from subterranean environments are described but not named owing to the lack of adult specimens. Although the epigean species T. aridus, sp. nov. is relatively widespread, all of the subterranean species are thought to represent short-range endemic species as they have been found at very few locations, all of which occur in localised habitats such as limestone or within mesa formations. Tyrannochthonius chamorro Chamberlin, 1947 from Guam is transferred to the genus Lagynochthonius, creating the new combination Lagynochthonius chamorro (Chamberlin 1947).

We used molecular and morphological techniques to study troglobitic schizomids inhabiting a variety of subterranean landforms in semiarid Western Australia. The study was designed to explore the taxonomic and phylogenetic status of newly discovered populations of subterranean schizomids. Molecular sequences of the mitochondrial DNA (mtDNA) cytochrome oxidase subunit 1 (COI) and small subunit rRNA (12S) were obtained from a total of 73 schizomid specimens. Populations sampled from boreholes within mesa landforms in the Robe Valley were highly genetically distinct from species of Draculoides Harvey, 1992 found elsewhere in the Pilbara (Cape Range and Barrow Island). Pronounced genetic structuring was also evident at a fine spatial scale within the Robe Valley, with populations from each of the mesas examined exhibiting unique and highly divergent mtDNA lineages. These molecular data were generally supported by small but significant morphological features, usually in the secondary male structures, but some species were represented only by female specimens that possessed more conservative morphologies. The molecular data defined two major in-group clades, which were supported by morphological differences. One clade was widespread and included the type species of Draculoides, D. vinei (Harvey), along with D. bramstokeri Harvey & Humphreys, D. brooksi Harvey, D. julianneae Harvey, D. mesozeirus, sp. nov. and D. neoanthropus, sp. nov. The second clade was restricted to the Robe Valley and deemed to represent a new genus, Paradraculoides, which included four new species P. anachoretus, sp. nov., P. bythius, sp. nov., P. gnophicola, sp. nov. and P. kryptus, sp. nov. (type species).

Recent surveys of groundwater invertebrates (stygofauna) worldwide are yielding rich troves of biodiversity, with significant implications for invertebrate systematists and phylogeneticists as well as ecologists and groundwater managers. What is the ecological significance of this high biodiversity of invertebrates in some aquifers? How might it influence groundwater ecosystem services such as water purification or bioremediation? In terrestrial ecosystems, biodiversity is typically positively correlated with rates of ecosystem functions beneficial to humans (e.g. crop pollination). However, the links between biodiversity, ecosystem function, and ecosystem services in groundwater are unknown. In some aquifers, feeding, movement and excretion by diverse assemblages of stygofauna potentially enhance groundwater ecosystem services such as water purification, bioremediation and water infiltration. Further, as specific taxa apparently play ‘keystone’ roles in facilitating ecosystem services, declines in abundance or even their extinction have serious repercussions. One way to assess the functional significance of biodiversity is to identify ‘ecosystem service providers’, characterise their functional relationships, determine how service provision is affected by community structure and environmental variables, and measure the spatio-temporal scales over which these operate. Examples from Australian and New Zealand alluvial aquifers reveal knowledge gaps in understanding the functional importance of most stygofauna, hampering effective protection of currently undervalued groundwater ecosystem services.

The neobisioid pseudoscorpion family Hyidae has a distribution spanning the margins of the Indian Ocean and its phylogenetic position has been suggested to lie midway between a clade containing the Bochidae and Ideoroncidae, and a clade containing the remaining neobisioid families, Gymnobisiidae, Neobisiidae, Parahyidae and Syarinidae. Their central position within the Neobisioidea, and their biogeographic patterns, have prompted the present study, which is designed to test the monophyly of the family, deduce the phylogeny of all known hyids and to describe several newly discovered species. The phylogenetic analysis of the 14 species of the pseudoscorpion family Hyidae along with 20 other species of Neobisioidea demonstrates the monophyly of the family and its distinctness from all other neobisioid families. Two genera, Hya Chamberlin and Indohya Beier, are recognised, even though the phylogenetic analysis did not consistently recover the monophyly of Indohya. The effect of outgroup selection was tested on the Hyidae and revealed markedly different tree topologies. Hyella Harvey, with the type species H. humphreysi Harvey, is newly synonymised with Indohya owing to similarities between the previously described species and some new troglobitic species. The Indohyinae Harvey is synonymised with Hyidae. Two new epigean species and three new troglobitic species of Indohya are described: I. damocles, sp. nov., I. gollum, sp. nov., I. haroldi, sp. nov. and I. napierensis, sp. nov. from northern Western Australia, and I. jacquelinae, sp. nov. from Madagascar. All species of Indohya are short-range endemics, having very small distributional ranges. New data are presented for Indohya humphreysi (Harvey), including the first description of the male. The first protonymphs of the family Hyidae are described, including those of Hya minuta (Tullgren) and Indohya gollum, sp. nov.

Five species of the nephilid genus Nephila Leach are found in the Australasian region, which for the purposes of this study was defined as Australia and its dependencies (including Lord Howe I., Norfolk I., Christmas I., Cocos (Keeling) Is), New Guinea (including Papua New Guinea and the Indonesian province of West Papua), Solomon Is, Vanuatu, New Caledonia, Fiji, Tonga, Niue, New Zealand and other parts of the south-west Pacific region. All species are redescribed and illustrated. Nephila pilipes (Fabricius) occurs in the closed forests of eastern and northern Australia, New Guinea, Solomon Is and Vanuatu (through to South-East Asia); N. plumipes (Latreille) is found in Australia (including Lord Howe I. and Norfolk I.), New Guinea, Vanuatu, Solomon Is and New Caledonia; N. tetragnathoides (Walckenaer) inhabits Fiji, Tonga and Niue; N. antipodiana (Walckenaer) occurs in northern Australia (as well as Christmas I.), New Guinea and Solomon Is (through to South-East Asia); and N. edulis (Labillardière) is found in Australia (including Cocos (Keeling) Is), New Guinea, New Zealand and New Caledonia. Epeira (Nephila) walckenaeri Doleschall, E. (N.) hasseltii Doleschall, N. maculata var. annulipes Thorell, N. maculata jalorensis Simon, N. maculata var. novae-guineae Strand, N. pictithorax Kulczyński, N. maculata var. flavornata Merian, N. pictithorax Kulczyński, N. maculata var. flavornata Merian, N. maculata piscatorum de Vis, and N. (N.) maculata var. lauterbachi Dahl are proposed as new synonyms of N. pilipes. Nephila imperialis var. novaemecklenburgiae Strand, N. ambigua Kulczyński, N. sarasinorum Merian and N. celebesiana Strand are proposed as new synonyms of N. antipodiana. Meta aerea Hogg, N. meridionalis Hogg, N. adelaidensis Hogg and N. meridionalis hermitis Hogg are proposed as new synonyms of N. edulis. Nephila picta Rainbow is removed from the synonymy of N. plumipes and treated as a synonym of N. edulis, and N. nigritarsis insulicola Pocock is removed from the synonymy of N. plumipes and treated as a synonym of N. antipodiana. Allozyme data demonstrate that N. pilipes is distinct at the 80% FD level from N. edulis, N. plumipes and N. tetragnathoides. Nephila plumipes and N. tetragnathoides, deemed to represent sister-taxa owing to the shared presence of a triangular protrusion of the male pedipalpal conductor, were found to differ at 15% FD in the genetic study. No genetic differentiation was found between 10 populations of N. edulis sampled across mainland Australia. Species of the genus Nephila have been extensively used in ecological and behavioural studies, and the biology of Nephila species in the Australasian region is extensively reviewed and compared with studies on Nephila species from other regions of the world.

The Australian fauna is assessed for short-range endemism at the species level, i.e. the prevalence of species with naturally small ranges of less than 10,000 km2. The phenomenon is found to be widespread and several groups are found to consist principally of short-range endemics: Gastropoda (snails and slugs, both freshwater and terrestrial), Oligochaeta (earthworms), Onychophora (velvet worms), Araneae (mygalomorph spiders), Schizomida (schizomids), Diplopoda (millipedes), Phreatoicidea (phreatoicidean crustaceans), and Decapoda (freshwater crayfish). The majority of taxa with high numbers of short-range endemics possess similar ecological and life-history characteristics, such as poor powers of dispersal and confinement to discontinuous habitats. The conservation of such groups is often hampered by poor taxonomic knowledge, but modern, comprehensive biotic surveys will be helpful in identifying short-range endemics.

The schizomid fauna of the Seychelles is reviewed and the following taxa are recognised: Apozomus gerlachi, sp. nov., Bamazomus aviculus, sp. nov., Mahezomus apicoporus, sp. nov., Anepsiozomus sobrinus, sp. nov., Ovozomus similis (Hirst), Secozomus latipes (Hansen) and Zomus bagnallii (Jackson). Four new genera are recognised, each represented by a single species: Mahezomus, gen. nov. (type species M. apicoporus, sp. nov.); Anepsiozomus, gen. nov. (type species A. sobrinus, sp. nov.); Ovozomus, gen. nov. (type species Schizomus similis Hirst); and Secozomus, gen. nov. (type species Schizomus latipes Hansen). Three species of Schizomus described from Madagascar are transferred to Bamazomus: B. madagassus (Lawrence), comb. nov., B. milloti (Lawrence), comb. nov. and B. vadoni (Lawrence), comb. nov. Problems in the systematics of the subfamily Hubbardiinae are explored, especially with regard to the high number of genera currently recognised. Biogeographic patterns of the arachnid fauna of the Seychelles are briefly explored, and a list of the Scorpiones, Pseudoscorpiones, Amblypygi, Schizomida and Palpigradi is included.

Systematists have frequently interpreted lateral compression of the idiosoma in post-larval instars of water mites as a synapomorphy indicating common ancestry. This paper re-examines the evidence to support this assumption and concludes that lateral flattening has occurred independently several times during water mite evolution, and especially often in various basal clades of the superfamily Hygrobatoidea. Reassessment of the phylogenetic relationships of the genera attributed to these clades results in improved understanding of the early evolution and biogeography of hygrobatoid water mites and necessitates redefinition of several clades and significant revision of the family classification. The South African genus Karlvietsia K.O. Viets, 1962 is transferred from the subfamily Frontipodopsinae of the family Aturidae to the Hygrobatidae. The Australian genera Tasmanaxona Cook, 1986, Wheenyella Cook, 1986 and Wheenyoides Harvey, 1990 are also removed from Frontipodopsinae and placed with the genus Wettina Piersig, 1892, here removed from the family Pionidae, in the family Wettinidae Cook, stat. nov. Based on apparent relationship with members of Wettinidae, the South African genus Stormaxonella K.O. Viets, 1962 is transferred from the Aturidae and provisionally placed in the Wettinidae. The clade including the predominately Holarctic genus Lethaxona K.H. Viets, 1932 and the Western Hemisphere genus LethaxonellaCook, 1963 is recognised as the probable sister group of Wettinidae, and these genera are consequently removed from the Aturidae and placed in Lethaxonidae, fam. nov. The genus Frontipodopsis Walter is reassigned from Aturidae to the now monobasic family Frontipodopsidae K.H. Viets, stat. nov. Finally, five new species are described, including Frontipodopsis (Frontipodopsella) sudafricanus, sp. nov., Wettina occidentalis, sp. nov., Karlvietsia simplex, sp. nov., Hygrobates (Hygrobates) gereckei, sp. nov. and Hygrobates (Hygrobatides)frontipodoides, sp. nov., and both Karlvietsia brevipalpis K.O. Viets and K. angustipalpis K.O. Viets are redescribed.

The Hyidae are redefined and three new synapomorphies are recognised which define the family: basi-dorsal mound on femora I and 11; minute setae on female sternite 11; and 2–3 stout setae on the pedipalpal femur. Three genera are included: Hya Chamberlin, with H. rninuta (Tullgren) and H. chamberlini, sp, nov.; Indohya Beier, with I. besucheti Beier, I. caecata Beier, I. panops, sp. nov., I. beieri, sp, nov., I. pusilla, sp. nov. and I. typhlops, sp. nov.; and Hyella, gen. nov., with H. humphreysi, sp. nov. (type species), the first known cavernicolous species of the family. Hya heterodonta Chamberlin is considered a junior synonym of Ideobisium minutum Tullgren. The Hyidae are divided into two subfamilies, Hyinae for Hya, and Indohyinae, subfam. nov., for Indohya and Hyella. A new distribution record is given for Tyrannochthonius krakatau Harvey (Chthoniidae), based on part of the type series of I. minutum.

A new pseudoscorpion classification is proposed with two new suborders, Epiocheirata and Iocheirata, based upon a cladistic analysis of relationships within the order. The Epiocheirata contains two superfamilies: Chthonioidea for Chthoniidae, Tridenchthoniidae and Lechytiidae, stat. nov. (for Lechytia), and Feaelloidea for Feaellidae and Pseudogarypidae. The Iocheirata is divided into two infraorders: Hemictenata Balzan and Panctenata Balzan. The Hemictenata contains a single superfamily, Neobisioidea for Bochicidae, Gymnobisiidae, Hyidae, Ideoroncidae, Neobisiidae, Parahyidae, fam. nov. (for Parahya) and Syarinidae. The Panctenata contains two microorders: Mestommatina, nov, with Garypoidea for Cheiridiidae, Garypidae, Geogarypidae, Larcidae, fam. nov. (for Archeolarca and Larca) and Pseudochiridiidae, and Olpioidea for Menthidae and Olpiidae; and Elassommatina, nov. with Sternophoroidea, stat. nov, for Sternophoridae, and Cheliferoidea for Atemnidae, Cheliferidae, Chernetidae and Withiidae. The Vachoniidae is synonymised with the Bochicidae, and the Cheiridioidea is treated as a synonym of Garypoidea. Philomaoria Chamberlin and Philomaoriini are transferred from the Withiidae to the Cheliferidae. The chthoniid tribe Pseudotyrannochthoniini is elevated to subfamily rank, and the systematic position of the Devonian family Dracochelidae is discussed.