Rapid divergent evolution of internal female genitalia and the coevolution of male genital morphology revealed by micro-computed tomography.

Animal genitalia are thought to evolve rapidly and divergently in response to sexual selection. Studies of genital evolution have focused largely on male genitalia. The paucity of work on female genital morphology is probably due to problems faced in quantifying shape variation, due to their composition and accessibility. Here we use a combination of micro-computed tomography, landmark free shape quantification and phylogenetic analysis to quantify the rate of female genital shape evolution among 29 species of Antichiropus millipedes, and their coevolution with male genitalia. We found significant variation in female and male genital shape among species. Male genital shape showed a stronger phylogenetic signal than female genital shape, although the phylogenetic signal effect sizes did not differ significantly. Male genital shape was found to be evolving 1.2 times faster than female genital shape. Female and male genital shape exhibited strong correlated evolution, indicating that genital shape changes in one sex are associated with corresponding changes in the genital shape of the other sex. This study adds novel insight into our growing understanding of how female genitalia can evolve rapidly and divergently, and highlights the advantages of three-dimensional techniques and multivariate analyses in studies of female genital evolution.

A new pseudoscorpion genus (Garypinoidea: Garypinidae) from the Eocene supports extinction and range contraction in the European paleobiota.

During the Paleogene, the Holarctic experienced drastic climatic oscillations, including periods of extensive glaciation. These changes had a severe impact on both the flora and fauna causing widespread extinction and range shifts with some taxa retreating to refugia in the Mediterranean Basin. Here we provide evidence for this hypothesis using fossils from the pseudoscorpion family Garypinidae Daday, 1889 (Arachnida: Pseudoscorpiones). This family comprises 21 extant genera from all continents except Antarctica but is restricted to low mid-latitudes (<44°N) in the Northern Hemisphere. We provide the second record of garypinids from the European succinite ambers of the Eocene by describing the first extinct genus in Garypinidae, Baltamblyolpium gen. nov., which includes two species: Baltamblyolpium gizmotum sp. nov. from Baltic amber and Baltamblyolpium grabenhorsti sp. nov. from Bitterfeld amber. The new genus exhibits a morphology that closely resembles Neoamblyolpium Hoff, 1956 from western North America and the genus Amblyolpium Simon, 1898, which is widespread but includes taxa restricted to Mediterranean refugia in Europe. The discovery of a new fossil genus of Garypinidae from Europe confirms that the family was found at more northerly latitudes during the Eocene, however, extinction and range contraction resulted in their present-day relictual distribution in southern Europe like many other lineages that once thrived in the European "Baltic amber forest" of the Eocene.

A new genus and five new species of pseudoscorpions (Arachnida, Pseudoscorpiones, Withiidae) from Colombia.

The pseudoscorpion family Withiidae is widely distributed around the world, with most of its diversity in tropical areas. Five new species and a new genus from Colombia are described: Cystowithiusflorezisp. nov., Parawithiusbromelicolasp. nov., Oligowithiusachaguasp. nov., and the genus Paciwithiusgen. nov. with two species Paciwithiusvalduparensissp. nov. and Paciwithiuschimbilacussp. nov. A reassessment of the subgenus Dolichowithius (Oligowithius) Beier, 1936 allows the elevation to a full generic level, and the transfer of the only known species to Oligowithius, forming the new combination Oligowithiusabnormis (Beier, 1936), comb. nov.

Tightening the requirements for species diagnoses would help integrate DNA-based descriptions in taxonomic practice.

Modern advances in DNA sequencing hold the promise of facilitating descriptions of new organisms at ever finer precision but have come with challenges as the major Codes of bionomenclature contain poorly defined requirements for species and subspecies diagnoses (henceforth, species diagnoses), which is particularly problematic for DNA-based taxonomy. We, the commissioners of the International Commission on Zoological Nomenclature, advocate a tightening of the definition of "species diagnosis" in future editions of Codes of bionomenclature, for example, through the introduction of requirements for specific information on the character states of differentiating traits in comparison with similar species. Such new provisions would enhance taxonomic standards and ensure that all diagnoses, including DNA-based ones, contain adequate taxonomic context. Our recommendations are intended to spur discussion among biologists, as broad community consensus is critical ahead of the implementation of new editions of the International Code of Zoological Nomenclature and other Codes of bionomenclature.

A Case for Below-Ground Dispersal? Insights into the Biology, Ecology and Conservation of Blind Cave Spiders in the Genus Troglodiplura (Mygalomorphae: Anamidae).

Previously described from only fragments of exoskeleton and juvenile specimens, the cave spider genus Troglodiplura (Araneae: Anamidae), endemic to the Nullarbor Plain, is the only troglomorphic member of the infraorder Mygalomorphae recorded from Australia. We investigated the distribution of Troglodiplura in South Australia, collecting and observing the first (intact) mature specimens, widening the number of caves it has been recorded in, and documenting threats to conservation. Phylogenetic analyses support the placement of Troglodiplura as an independent lineage within the subfamily Anaminae (the 'Troglodiplura group') and provide unequivocal evidence that populations from apparently isolated cave systems are conspecifics of T. beirutpakbarai Harvey & Rix, 2020, with extremely low or negligible inter-population mitochondrial divergences. This is intriguing evidence for recent or contemporary subterranean dispersal of these large, troglomorphic spiders. Observations of adults and juvenile spiders taken in the natural cave environment, and supported by observations in captivity, revealed the use of crevices within caves as shelters, but no evidence of silk use for burrow construction, contrasting with the typical burrowing behaviours seen in other Anamidae. We identify a range of threats posed to the species and to the fragile cave ecosystem, and provide recommendations for further research to better define the distribution of vulnerable taxa within caves and identify actions needed to protect them.

Correlation with a limited set of behavioral niches explains the convergence of somatic morphology in mygalomorph spiders.

Understanding the drivers of morphological convergence requires investigation into its relationship with behavior and niche space, and such investigations in turn provide insights into evolutionary dynamics, functional morphology, and life history. Mygalomorph spiders (trapdoor spiders and their kin) have long been associated with high levels of morphological homoplasy, and many convergent features can be intuitively associated with different behavioral niches. Using genus-level phylogenies based on recent genomic studies and a newly assembled matrix of discrete behavioral and somatic morphological characters, we reconstruct the evolution of burrowing behavior in the Mygalomorphae, compare the influence of behavior and evolutionary history on somatic morphology, and test hypotheses of correlated evolution between specific morphological features and behavior. Our results reveal the simplicity of the mygalomorph adaptive landscape, with opportunistic, web-building taxa at one end, and burrowing/nesting taxa with structurally modified burrow entrances (e.g., a trapdoor) at the other. Shifts in behavioral niche, in both directions, are common across the evolutionary history of the Mygalomorphae, and several major clades include taxa inhabiting both behavioral extremes. Somatic morphology is heavily influenced by behavior, with taxa inhabiting the same behavioral niche often more similar morphologically than more closely related but behaviorally divergent taxa, and we were able to identify a suite of 11 somatic features that show significant correlation with particular behaviors. We discuss these findings in light of the function of particular morphological features, niche dynamics within the Mygalomorphae, and constraints on the mygalomorph adaptive landscape relative to other spiders.

The systematics of the pseudoscorpion genus Indohya (Pseudoscorpiones: Hyidae) in Australia.

The pseudoscorpion genus Indohya Beier, 1974 is known to occur in three Gondwanan fragments around the Indian Oceansouthern India, Madagascar and north-western Australiasuggesting that the genus had evolved prior to the breakup of Gondwana and was present on each landmass as they rifted apart during the Mesozoic. The Australian fauna is the most diverse, with nine species previously described from Cape Range and the Kimberley region of north-western Australia. The present study documents the genus Indohya in Australia using a combination of morphology and DNA sequence data. We found a total of 36 species, including 27 new species. The majority of the Pilbara fauna consist of blind troglobites collected from subterranean ecosystems, with an additional three eyed species from epigean habitats. The new species consist of one from Cape Range (I. anastomosa Harvey & Burger, n. sp.), 21 from the Pilbara (I. adlardi Harvey & Burger, n. sp., I. alexanderi Harvey & Burger, n. sp., I. aphana Harvey & Burger, n. sp., I. aquila Harvey & Burger, n. sp., I. arcana Harvey & Burger, n. sp., I. arnoldstrongi Harvey & Burger, n. sp., I. boltoni Harvey & Burger, n. sp., I. cardo Harvey & Burger, n. sp., I. catherineae Harvey & Burger, n. sp., I. cockingi Harvey & Burger, n. sp., I. cribbi Harvey & Burger, n. sp., I. draconis Harvey & Burger, n. sp., I. furtiva Harvey & Burger, n. sp., I. incomperta Harvey & Burger, n. sp., I. jessicae Harvey & Burger, n. sp., I. lynbeazlyeae Harvey & Burger, n. sp., I. morganstrongi Harvey & Burger, n. sp., I. rixi Harvey & Burger, n. sp., I. sagmata Harvey & Burger, n. sp., I. scanloni Harvey & Burger, n. sp. and I. silenda Harvey & Burger, n. sp.) and five from the Kimberley (I. currani Harvey & Burger, n. sp., I. finitima Harvey & Burger, n. sp., I. julianneae Harvey & Burger, n. sp., I. karenae Harvey & Burger, n. sp. and I. sachsei Harvey & Burger, n. sp.). The study is augmented with sequence data from 29 species of Indohya, including all of the 24 species recorded from the Pilbara and Cape Range, and five of the 12 known Kimberley species. Seven clades recovered during the molecular analysis are only represented by nymphs, but we used COI sequence data to diagnose these species in the absence of adult morphological data.

Two new species of Rhopalochernes Beier, 1932 (Pseudoscorpiones: Chernetidae) from Colombia.

The pseudoscorpion genus Rhopalochernes Beier, 1932 is restricted to the Neotropical region and currently includes eight species. Recent field collections in the Caribbean region of Colombia have revealed the presence of two previously undescribed species, which are here described as Rhopalochernes luiscarlosi sp. nov. and Rhopalochernes catalinae sp. nov. They differ from one another by the shape of the carapace, the position of the trichobothria and the length of the nodus ramosus. They differ from other species of the genus by the absence of eye spots, and the number of the cheliceral setae. One of these features, the presence of only four setae on the cheliceral hand, is shared with R. panamensis Heurtault, 1998 and R. chamberlini Heurtault, 1998, with other species possessing a higher number. Additionally, the key for the genus proposed by Beier (1932) is updated for included all the species.

One-way ticket to the blue: A large-scale, dated phylogeny revealed asymmetric land-to-water transitions in acariform mites (Acari: Acariformes).

Acariform mites are an ancient and megadiverse lineage that may have experienced a complex pattern of invasions into terrestrial and aquatic habitats. These among-realm transitions may relate to periods of turmoil in Earth's history or be simply results of uneven biodiversity patterns across habitats. Here, we inferred a dated, representative acariform phylogeny (five genes, 9,200 bp aligned, 367 terminals belonging to 150 ingroup plus 15 outgroup families, 23 fossil calibration points) which was used to infer transitions between marine/freshwater/terrestrial habitats. We detected four unambiguous transitions from terrestrial to freshwater habitats (Hydrozetes, Naiadacarus, Fusohericia, Afronothrus, Homocaligus); one from freshwater to marine (Pontarachnidae), and four from marine to brackish or freshwater transitions (all among Halacaridae: Acarothrix; Halacarellus petiti; Copidognathus sp.; clade Limnohalacarus + Soldanellonyx + Porohalacarus + Porolohmannella). One transition to the sea was inferred ambiguously with respect to the ancestor being either terrestrial or freshwater (Hyadesiidae), and another must be most carefully examined by adding potential related taxa (Selenoribatidae + Fortuyniidae). Finally, we inferred a single, remarkable transition from aquatic to terrestrial habitats involving early evolution of the large and ecologically diverse lineage: the ancestor of the Halacaridae + Parasitengona clade was probably freshwater given our dataset, thus making terrestrial Parasitengona secondarily terrestrial. Overall, our results suggested a strong asymmetry in environmental transitions: the majority occurred from terrestrial to aquatic habitats. This asymmetry is probably linked to mites' biological properties and uneven biodiversity patterns across habitats rather than Earth's geological history. Since the land holds more acariform diversity than water habitats, a shift from the former is more likely than from the latter. We inferred the following relationships: alicid endeostigmatid + eriophyoid (Alycidae, (Nanorchestidae, (Nematalycidae, Eriophyoidea))) being sister group to the remaining Acariformes: (proteonematalycid Endeostigmata, alicorhagiid Endeostigmata, Trombidiformes, Oribatida (including Astigmata)). Trombidiform relationships had several novel rearrangements: (i) traditional Eupodina lacked support for the inclusion of Bdelloidea; (ii) Teneriffidae, traditionally placed among Anystina, was consistently recovered in a clade including Heterostigmata in Eleutherengona; (iii) several lineages, such as Adamystidae, Paratydeidae, Caeculidae and Erythracaridae, were recovered in a large clade along other Anystina and Eleutherengona, suggesting single origins of several fundamental character states, such as the reduction of the cheliceral fixed digit and development of the palpal thumb-claw complex.

Phylogenomics of Scorpions Reveal Contemporaneous Diversification of Scorpion Mammalian Predators and Mammal-Active Sodium Channel Toxins.

Scorpions constitute a charismatic lineage of arthropods and comprise more than 2500 described species. Found throughout various tropical and temperate habitats, these predatory arachnids have a long evolutionary history, with a fossil record that began in the Silurian. While all scorpions are venomous, the asymmetrically diverse family Buthidae harbors nearly half the diversity of extant scorpions, and all but one of the 58 species that are medically significant to humans. However, the lack of a densely sampled scorpion phylogeny has hindered broader inferences of the diversification dynamics of scorpion toxins. To redress this gap, we assembled a phylogenomic data set of 100 scorpion venom gland transcriptomes and genomes, emphasizing the sampling of highly toxic buthid genera. To infer divergence times of venom gene families, we applied a phylogenomic node dating approach for the species tree in tandem with phylostratigraphic bracketing to estimate the minimum ages of mammal-specific toxins. Our analyses establish a robustly supported phylogeny of scorpions, particularly with regard to relationships between medically significant taxa. Analysis of venom gene families shows that mammal-active sodium channel toxins (NaTx) have independently evolved in five lineages within Buthidae. Temporal windows of mammal-targeting toxin origins are correlated with the basal diversification of major scorpion mammal predators such as shrews, bats, and rodents. These results suggest an evolutionary model of relatively recent diversification of buthid NaTx homologs in response to the diversification of scorpion predators. [Adaptation; arachnids; phylogenomic dating; phylostratigraphy; venom.].

Comprehensive Species Sampling and Sophisticated Algorithmic Approaches Refute the Monophyly of Arachnida.

Deciphering the evolutionary relationships of Chelicerata (arachnids, horseshoe crabs, and allied taxa) has proven notoriously difficult, due to their ancient rapid radiation and the incidence of elevated evolutionary rates in several lineages. Although conflicting hypotheses prevail in morphological and molecular data sets alike, the monophyly of Arachnida is nearly universally accepted, despite historical lack of support in molecular data sets. Some phylotranscriptomic analyses have recovered arachnid monophyly, but these did not sample all living orders, whereas analyses including all orders have failed to recover Arachnida. To understand this conflict, we assembled a data set of 506 high-quality genomes and transcriptomes, sampling all living orders of Chelicerata with high occupancy and rigorous approaches to orthology inference. Our analyses consistently recovered the nested placement of horseshoe crabs within a paraphyletic Arachnida. This result was insensitive to variation in evolutionary rates of genes, complexity of the substitution models, and alternative algorithmic approaches to species tree inference. Investigation of sources of systematic bias showed that genes and sites that recover arachnid monophyly are enriched in noise and exhibit low information content. To test the impact of morphological data, we generated a 514-taxon morphological data matrix of extant and fossil Chelicerata, analyzed in tandem with the molecular matrix. Combined analyses recovered the clade Merostomata (the marine orders Xiphosura, Eurypterida, and Chasmataspidida), but merostomates appeared nested within Arachnida. Our results suggest that morphological convergence resulting from adaptations to life in terrestrial habitats has driven the historical perception of arachnid monophyly, paralleling the history of numerous other invertebrate terrestrial groups.

The first true millipede-1306 legs long.

The name "millipede" translates to a thousand feet (from mille "thousand" and pes "foot"). However, no millipede has ever been described with more than 750 legs. We discovered a new record-setting species of millipede with 1,306 legs, Eumillipes persephone, from Western Australia. This diminutive animal (0.95 mm wide, 95.7 mm long) has 330 segments, a cone-shaped head with enormous antennae, and a beak for feeding. A distant relative of the previous record holder, Illacme plenipes from California, it belongs to a different order, the Polyzoniida. Discovered 60 m below ground in a drill hole created for mineral exploration, E. persephone possesses troglomorphic features; it lacks eyes and pigmentation, and it has a greatly elongated body-features that stand in stark contrast to its closest surface-dwelling relatives in Australia and all other members of its order. Using phylogenomics, we found that super-elongation (> 180 segments) evolved repeatedly in the millipede class Diplopoda. The striking morphological similarity between E. persephone and I. plenipes is a result of convergent evolution, probably for locomotion in similar soil habitats. Discovered in the resource-rich Goldfields-Esperance region and threatened by encroaching surface mining, documentation of this species and conservation of its habitat are of critical importance.

A national-scale dataset for threats impacting Australia's imperiled flora and fauna.

Australia is in the midst of an extinction crisis, having already lost 10% of terrestrial mammal fauna since European settlement and with hundreds of other species at high risk of extinction. The decline of the nation's biota is a result of an array of threatening processes; however, a comprehensive taxon-specific understanding of threats and their relative impacts remains undocumented nationally. Using expert consultation, we compile the first complete, validated, and consistent taxon-specific threat and impact dataset for all nationally listed threatened taxa in Australia. We confined our analysis to 1,795 terrestrial and aquatic taxa listed as threatened (Vulnerable, Endangered, or Critically Endangered) under Australian Commonwealth law. We engaged taxonomic experts to generate taxon-specific threat and threat impact information to consistently apply the IUCN Threat Classification Scheme and Threat Impact Scoring System, as well as eight broad-level threats and 51 subcategory threats, for all 1,795 threatened terrestrial and aquatic threatened taxa. This compilation produced 4,877 unique taxon-threat-impact combinations with the most frequently listed threats being Habitat loss, fragmentation, and degradation (n = 1,210 taxa), and Invasive species and disease (n = 966 taxa). Yet when only high-impact threats or medium-impact threats are considered, Invasive species and disease become the most prevalent threats. This dataset provides critical information for conservation action planning, national legislation and policy, and prioritizing investments in threatened species management and recovery.

Diversification of the mygalomorph spider genus Aname (Araneae: Anamidae) across the Australian arid zone: Tracing the evolution and biogeography of a continent-wide radiation.

The assembly of the Australian arid zone biota has long fascinated biogeographers. Covering over two-thirds of the continent, Australia's vast arid zone biome is home to a distinctive fauna and flora, including numerous lineages which have diversified since the Eocene. Tracing the origins and speciation history of these arid zone taxa has been an ongoing endeavour since the advent of molecular phylogenetics, and an increasing number of studies on invertebrate animals are beginning to complement a rich history of research on vertebrate and plant taxa. In this study, we apply continent-wide genetic sampling and one of the largest phylogenetic data matrices yet assembled for a genus of Australian spiders, to reconstruct the phylogeny and biogeographic history of the open-holed trapdoor spider genus Aname L. Koch, 1873. This highly diverse lineage of Australian mygalomorph spiders has a distribution covering the majority of Australia west of the Great Dividing Range, but apparently excluding the high rainfall zones of eastern Australia and Tasmania. Original and legacy sequences were obtained for three mtDNA and four nuDNA markers from 174 taxa in seven genera, including 150 Aname specimen terminals belonging to 102 species-level operational taxonomic units, sampled from 32 bioregions across Australia. Reconstruction of the phylogeny and biogeographic history of Aname revealed three radiations (Tropical, Temperate-Eastern and Continental), which could be further broken into eight major inclusive clades. Ancestral area reconstruction revealed the Pilbara, Monsoon Tropics and Mid-West to be important ancestral areas for the genus Aname and its closest relatives, with the origin of Aname itself inferred in the Pilbara bioregion. From these origins in the arid north-west of Australia, our study found evidence for a series of subsequent biome transitions in separate lineages, with at least eight tertiary incursions back into the arid zone from more mesic tropical, temperate or eastern biomes, and only two major clades which experienced widespread (primary) in situ diversification within the arid zone. Based on our phylogenetic results, and results from independent legacy divergence dating studies, we further reveal the importance of climate-driven biotic change in the Miocene and Pliocene in shaping the distribution and composition of the Australian arid zone biota, and the value of continent-wide studies in revealing potentially complex patterns of arid zone diversification in dispersal-limited invertebrate taxa.

Taxonomic Sampling and Rare Genomic Changes Overcome Long-Branch Attraction in the Phylogenetic Placement of Pseudoscorpions.

Long-branch attraction is a systematic artifact that results in erroneous groupings of fast-evolving taxa. The combination of short, deep internodes in tandem with long-branch attraction artifacts has produced empirically intractable parts of the Tree of Life. One such group is the arthropod subphylum Chelicerata, whose backbone phylogeny has remained unstable despite improvements in phylogenetic methods and genome-scale data sets. Pseudoscorpion placement is particularly variable across data sets and analytical frameworks, with this group either clustering with other long-branch orders or with Arachnopulmonata (scorpions and tetrapulmonates). To surmount long-branch attraction, we investigated the effect of taxonomic sampling via sequential deletion of basally branching pseudoscorpion superfamilies, as well as varying gene occupancy thresholds in supermatrices. We show that concatenated supermatrices and coalescent-based summary species tree approaches support a sister group relationship of pseudoscorpions and scorpions, when more of the basally branching taxa are sampled. Matrix completeness had demonstrably less influence on tree topology. As an external arbiter of phylogenetic placement, we leveraged the recent discovery of an ancient genome duplication in the common ancestor of Arachnopulmonata as a litmus test for competing hypotheses of pseudoscorpion relationships. We generated a high-quality developmental transcriptome and the first genome for pseudoscorpions to assess the incidence of arachnopulmonate-specific duplications (e.g., homeobox genes and miRNAs). Our results support the inclusion of pseudoscorpions in Arachnopulmonata (new definition), as the sister group of scorpions. Panscorpiones (new name) is proposed for the clade uniting Scorpiones and Pseudoscorpiones.

A systematic revision of Draculoides (Schizomida: Hubbardiidae) of the Pilbara, Western Australia, Part I: the Western Pilbara.

The schizomid fauna of mainland Australia currently comprises 60 species within seven named genera, of which five are endemic to the continent: Attenuizomus Harvey, 2000, Brignolizomus Harvey, 2000, Draculoides Harvey, 1992, Julattenius Harvey, 1992, Notozomus Harvey, 2000. Most Australian schizomids have been described from eastern and northern Australia, but there is also a significant subterranean fauna that has been found in hypogean habitats in the semi-arid Pilbara region of Western Australia. The vast majority of these species can be assigned to the genus Draculoides and this study is the first in a proposed series to revise this highly diverse genus. We treat the species found in the western Pilbara region, which includes 13 new species and 13 previously named species, using morphological characters and multi-locus sequence data. We also incorporate a molecular "mini-barcode" approach for COI, 12S and ITS2 to diagnose the new species. The new species are named: Draculoides akashae Abrams and Harvey, n. sp., D. belalugosii Abrams and Harvey, n. sp., D. carmillae Abrams and Harvey, n. sp., D. christopherleei Abrams and Harvey, n. sp., D. claudiae Abrams and Harvey, n. sp., D. immortalis Abrams and Harvey, n. sp., D. karenbassettae Abrams and Harvey, n. sp., D. mckechnieorum Abrams and Harvey, n. sp., D. minae Abrams and Harvey, n. sp., D. noctigrassator Abrams and Harvey, n. sp., D. nosferatu Abrams and Harvey, n. sp., D. piscivultus Abrams and Harvey, n. sp. and D. warramboo Abrams and Harvey, n. sp. We also provide the first descriptions of males of D. anachoretus (Harvey, Berry, Edward and Humphreys, 2008) and D. gnophicola (Harvey, Berry, Edward and Humphreys, 2008). All of the new species are subterranean-dwelling, short-range endemic species that occur in regions subject to mining activities, rendering them of high conservation significance.

The open-holed trapdoor spiders (Mygalomorphae: Anamidae: Namea) of Australia's D'Aguilar Range: revealing an unexpected subtropical hotspot of rainforest diversity.

The D'Aguilar Range of subtropical south-eastern Queensland (Australia), harbours an upland rainforest biota characterised by high levels of endemic diversity. Following recent phylogenetic and biogeographic research into the open-holed trapdoor spiders of the genus Namea Raven, 1984 (family Anamidae), remarkable levels of sympatry for a single genus of mygalomorph spiders were recorded from the D'Aguilar Range. It is now known that eight different species in the genus can be found in the D'Aguilar uplands, with five apparently endemic to rainforest habitats. In this paper we present a phylogenetic and taxonomic synopsis of the remarkable anamid fauna of the D'Aguilar Range: a key to the eight species is provided, and four new species of Namea are described (N. gloriosa sp. nov., N. gowardae sp. nov., N. nebo sp. nov. and N. nigritarsus sp. nov.). In shining a spotlight on the mygalomorph spiders of this region, we highlight the D'Aguilar Range as a hotspot of subtropical rainforest diversity, and an area of considerable conservation value.

The pseudoscorpion genus Verrucachernes (Pseudoscorpiones: Chernetidae) in the Indian region.

The pseudoscorpion genus Verrucachernes Chamberlin, 1947 is widely distributed in the Old World tropics, with three named Australasian and West Pacific species and two from Africa. A review of some pseudoscorpions described from India has revealed that Withius parvus Beier, 1930 (currently in the genus Metawithius) and Pselaphochernes indicus Beier, 1974 are misplaced and actually belong to the genus Verrucachernes Chamberlin, 1947, forming the new combinations V. parvus (Beier, 1930) comb. nov. and V. indicus (Beier, 1974) comb. nov., respectively. Both species possess the single, large, rounded spermatheca and other features typical of Verrucachernes.

Open-holed trapdoor spiders of the genus Teyl (Mygalomorphae: Nemesiidae: Anamini) from Western Australia's Pilbara bioregion: a new species and expanded phylogenetic assessment.

The open-holed trapdoor spiders of the genus Teyl Main, 1975 from the Pilbara bioregion of Western Australia are investigated. A single endemic species from the southern Pilbara, T. heuretes sp. nov., is newly described, representing the northern-most occurrence of the genus in Australia. Legacy molecular data for Australian Nemesiidae, along with newly generated sequences for all described species of Teyl known from Western Australia, are analysed using Maximum Likelihood methods, providing molecular data for T. heuretes and an expanded phylogenetic assessment of the genus.

The millipede genus Antichiropus (Diplopoda: Polydesmida: Paradoxosomatidae), part 3: species of the Pilbara bioregion of Western Australia.

The species of the millipede genus Antichiropus Attems, 1911 found in the Pilbara region of Western Australia are reviewed, and 33 new species are described. The new species are: A. anguinus Car, n. sp., A. antius Car, n. sp., A. apricus Car, n. sp., A. cirratus Car, n. sp., A. confragus Car, n. sp., A. cristatus Car, n. sp., A. cucumeraceous Car, n. sp., A. cunicularis Car, n. sp, A. echinus Car, n. sp., A. filiolus Car, n. sp., A. forcipatus Car, n. sp., A. georginae Car, n. sp., A. gibbus Car, n. sp., A. hystricosus Car, n. sp., A. julianneae Car, n. sp., A. literulus Car, n. sp., A. lucyae Car, n. sp., A. nicholasi Car, n. sp., A. nimbus Car, n. sp., A. patriciae Car, n. sp., A. pendiculus Car, n. sp., A. picus Car, n. sp., A. procerus Car, n. sp., A. quaestionis Car, n. sp., A. rupinus Car, n. sp., A. salutus Car, n. sp., A. servulus Car, n. sp., A. simmonsi Car, n. sp., A. sloanae Car, n. sp., A. spathion Car, n. sp., A. uvulus Car, n. sp., A. verutus Car, n. sp. and A. vindicatus Car, n. sp.. The number of described Antichiropus species now stands at 72. Two species (A. julianneae Car, n. sp. and A. pendiculus Car, n. sp.) lack one diagnostic feature of the genus, namely a solenomere process, but are included here because they conform to the genus definition in all other characters. We also obtained sequence data from four mitochondrial genes (cytochrome c oxidase subunit 1 [COI], cytochrome c oxidase subunit 3 [COIII], cytochrome B [CytB], and 12S rRNA [12S]), and one nuclear gene (28S rRNA [28S]) for 19 species. Three main clades were recovered: one in the northern Pilbara, one in the southern Pilbara, and one just outside the south-western margin of the Pilbara.