Phylogenomic data reveal three new families of poorly studied Solifugae (camel spiders).

The systematics of the arachnid order Solifugae have been an enigma, owing to challenges in interpreting morphology, a paucity of molecular phylogenetic studies sampling across the group, and a dearth of taxonomic attention for many lineages. Recent work has suggested that solifuge families largely exhibit contiguous distributions and reflect patterns of vicariance, with the exception of three families: Melanoblossidae, Daesiidae and Gylippidae. Morphological studies have cast doubt on their existing circumscriptions and the present composition of these taxa renders their distributions as disjunct. We leveraged ultraconserved elements (UCEs) to test the phylogenetic placement of three key lineages of Solifugae that cause these anomalous distributions: Dinorhax rostrumpsittaci (putative melanoblossid), Namibesia (putative daesiid), and Trichotoma (putative gylippid). Phylogenetic placement of these three genera based on UCEs rendered the families that harbor them as para- or polyphyletic, recovering instead relationships that better accord with a biogeographic history driven by vicariance. Toward a stable and phylogenetically informed classification of Solifugae, we establish three new families, Dinorhaxidae new rank, Namibesiidae new rank and Lipophagidae new rank.

Invertebrate biodiversity continues to decline in cropland.

Modern agriculture has drastically changed global landscapes and introduced pressures on wildlife populations. Policy and management of agricultural systems has changed over the last 30 years, a period characterized not only by intensive agricultural practices but also by an increasing push towards sustainability. It is crucial that we understand the long-term consequences of agriculture on beneficial invertebrates and assess if policy and management approaches recently introduced are supporting their recovery. In this study, we use large citizen science datasets to derive trends in invertebrate occupancy in Great Britain between 1990 and 2019. We compare these trends between regions of no- (0%), low- (greater than 0-50%) and high-cropland (greater than 50%) cover, which includes arable and horticultural crops. Although we detect general declines, invertebrate groups are declining most strongly in high-cropland cover regions. This suggests that even in the light of improved policy and management over the last 30 years, the way we are managing cropland is failing to conserve and restore invertebrate communities. New policy-based drivers and incentives are required to support the resilience and sustainability of agricultural ecosystems. Post-Brexit changes in UK agricultural policy and reforms under the Environment Act offer opportunities to improve agricultural landscapes for the benefit of biodiversity and society.

An Opiliones-specific ultraconserved element probe set with a near-complete family-level phylogeny.

Sequence capture of ultraconserved elements (UCEs) has transformed molecular systematics across many taxa, with arachnids being no exception. The probe set available for Arachnida has been repeatedly used across multiple arachnid lineages and taxonomic levels, however more specific probe sets for spiders have demonstrated that more UCEs can be recovered with higher probe specificity. In this study, we develop an Opiliones-specific UCE probe set targeting 1915 UCEs using a combination of probes designed from genomes and transcriptomes, as well as the most useful probes from the Arachnida probe set. We demonstrate the effectiveness of this probe set across Opiliones with the most complete family-level phylogeny made to date, including representatives from 61 of 63 currently described families. We also test UCE recovery from historical specimens with degraded DNA, examine population-level data sets, and assess "backwards compatibility" with samples hybridized with the Arachnida probe set. The resulting phylogenies - which include specimens hybridized using both the Opiliones and Arachnida probe sets, historical specimens, and transcriptomes - are largely congruent with previous multi-locus and phylogenomic analyses. The probe set is also "backwards compatible", increasing the number of loci obtained in samples previously hybridized with the Arachnida probe set, and shows high utility down to shallow population-level divergences. This probe set has the potential to further transform Opiliones molecular systematics, resolving many long-standing taxonomic issues plaguing this lineage.

How many long branch orders occur in Chelicerata? Opposing effects of Palpigradi and Opilioacariformes on phylogenetic stability.

Excepting a handful of nodes, phylogenetic relationships between chelicerate orders remain poorly resolved, due to both the incidence of long branch attraction artifacts and the limited sampling of key lineages. It has recently been shown that increasing representation of basal nodes plays an outsized role in resolving the higher-level placement of long-branch chelicerate orders. Two lineages have been consistently undersampled in chelicerate phylogeny. First, sampling of the miniaturized order Palpigradi has been restricted to a fragmentary transcriptome of a single species. Second, sampling of Opilioacariformes, a rarely encountered and key group of Parasitiformes, has been restricted to a single exemplar. These two lineages exhibit dissimilar properties with respect to branch length; Opilioacariformes shows relatively low evolutionary rate compared to other Parasitiformes, whereas Palpigradi possibly acts as another long-branch order (an effect that may be conflated with the degree of missing data). To assess these properties and their effects on tree stability, we constructed a phylogenomic dataset of Chelicerata wherein both lineages were sampled with three terminals, increasing the representation of these taxa per locus. We examined the effect of subsampling phylogenomic matrices using (1) taxon occupancy, (2) evolutionary rate, and (3) a principal components-based approach. We further explored the impact of taxon deletion experiments that mitigate the effect of long branches. Here, we show that Palpigradi constitutes a fourth long-branch chelicerate order (together with Acariformes, Parasitiformes, and Pseudoscorpiones), which further destabilizes the chelicerate backbone topology. By contrast, the slow-evolving Opilioacariformes were consistently recovered within Parasitiformes, with certain subsampling practices recovering their placement as the sister group to the remaining Parasitiformes. Whereas the inclusion of Opilioacariformes always resulted in the non-monophyly of Acari with support, deletion of Opilioacariformes from datasets consistently incurred the monophyly of Acari except in matrices constructed on the basis of evolutionary rate. Our results strongly suggest that Acari is an artifact of long- branch attraction.

Locomotion in the pseudoscorpion Chelifer cancroides: forward, backward and upside-down walking in an eight-legged arthropod.

While insect locomotion has been intensively studied, there are comparably few studies investigating octopedal walking behaviour, and very little is known about pseudoscorpions in particular. Therefore, we performed an extensive locomotion analysis during forward, backward and upside-down walking in the cosmopolitan pseudoscorpion Chelifer cancroides. During forward locomotion, we observed C. cancroides to freeze locomotion frequently for short time periods. These microstops were barely visible to the naked eye with a duration of 100-200 ms. Our locomotion analysis revealed that C. cancroides performs a statically stable and highly coordinated alternating tetrapod gait during forward and backward walking, with almost complete inversion of the tetrapod schemes, but no rigidly fixed leg coordination during upside-down walks with low walking speeds up to 4 body lengths per second. Highest speeds (up to 17 body lengths per second), mainly achieved by consistent leg coordination and strong phase shifts, were observed during backward locomotion (escape behaviour), whereas forward walking was characterised by lower speeds and phase shifts of ∼10% between two loosely coupled leg groups within one tetrapod. That is, during the movement of one tetrapod group, the last and the third leg are almost synchronous in their swing phases, as are the second and the first leg. A special role of the second leg pair was demonstrated, probably mainly for stability reasons and related to the large pedipalps.

The genome of a daddy-long-legs (Opiliones) illuminates the evolution of arachnid appendages.

Chelicerate arthropods exhibit dynamic genome evolution, with ancient whole-genome duplication (WGD) events affecting several orders. Yet, genomes remain unavailable for a number of poorly studied orders, such as Opiliones (daddy-long-legs), which has hindered comparative study. We assembled the first harvestman draft genome for the species Phalangium opilio, which bears elongate, prehensile appendages, made possible by numerous distal articles called tarsomeres. Here, we show that the genome of P. opilio exhibits a single Hox cluster and no evidence of WGD. To investigate the developmental genetic basis for the quintessential trait of this group-the elongate legs-we interrogated the function of the Hox genes Deformed (Dfd) and Sex combs reduced (Scr), and a homologue of Epidermal growth factor receptor (Egfr). Knockdown of Dfd incurred homeotic transformation of two pairs of legs into pedipalps, with dramatic shortening of leg segments in the longest leg pair, whereas homeosis in L3 is only achieved upon double Dfd + Scr knockdown. Knockdown of Egfr incurred shortened appendages and the loss of tarsomeres. The similarity of Egfr loss-of-function phenotypic spectra in insects and this arachnid suggest that repeated cooption of EGFR signalling underlies the independent gains of supernumerary tarsomeres across the arthropod tree of life.

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.

Genomic Determination of Reproductive Mode in Facultatively Parthenogenetic Opiliones.

Sexual reproduction may pose myriad short-term costs to females. Despite these costs, sexual reproduction is near ubiquitous. Facultative parthenogenesis is theorized to mitigate some of the costs of sex, as individuals can participate in occasional sex to limit costs while obtaining many benefits. However, most theoretical models assume sexual reproduction is fixed following mating, with no possibility of clutches of mixed reproductive ontogeny. Therefore, we asked: if coercive males are present at high frequency in a population of facultative parthenogens, will their clutches be solely sexually produced, or will there be evidence of sexually and asexually-produced offspring? How will their offspring production compare to conspecifics in low-frequency male populations? We addressed our questions by collecting females and egg clutches of the facultatively parthenogenetic Opiliones species Leiobunum manubriatum and L. globosum. In L. manubriatum, females from populations with few males were not significantly more fecund than females from populations with higher male relative frequency, despite the potential release of the former from sexual conflict. We used 3 genotyping methods along with a custom set of DNA capture probes to reveal that offspring of L. manubriatum from these high male populations were primarily produced via asexual reproduction. This is surprising because sex ratios in these southern populations approach equality, increasing the probability for females to encounter mates and produce offspring sexually. We additionally found evidence for reproductive polymorphisms within populations. Rapid and accurate SNP genotyping data will continue to allow us to address broader evolutionary questions regarding the role of facultative reproductive modes in the maintenance of sex.

Developmental gene expression as a phylogenetic data class: support for the monophyly of Arachnopulmonata.

Despite application of genome-scale datasets, the phylogenetic placement of scorpions within arachnids remains contentious between two different phylogenetic data classes. Paleontologists continue to recover scorpions in a basally branching position, partly owing to their morphological similarity to extinct marine orders like Eurypterida (sea scorpions). Phylogenomic datasets consistently recover scorpions in a derived position, as the sister group of Tetrapulmonata (a clade of arachnids that includes spiders). To adjudicate between these hypotheses using a rare genomic change (RGC), we leveraged the recent discovery of ancient paralogy in spiders and scorpions to assess phylogenetic placement. We identified homologs of four transcription factors required for appendage patterning (dachshund, homothorax, extradenticle, and optomotor blind) in arthropods that are known to be duplicated in spiders. Using genomic resources for a spider, a scorpion, and a harvestman, we conducted gene tree analyses and assayed expression patterns of scorpion gene duplicates. Here we show that scorpions, like spiders, retain two copies of all four transcription factors, whereas arachnid orders like mites and harvestmen bear a single copy. A survey of embryonic expression patterns of the scorpion paralogs closely matches those of their spider counterparts, with one paralog consistently retaining the putatively ancestral pattern found in the harvestman, as well as the mite, and/or other outgroups. These data comprise a rare genomic change in chelicerate phylogeny supporting the inference of a distal placement of scorpions. Beyond demonstrating the diagnostic power of developmental genetic data as a phylogenetic data class, a derived placement of scorpions within the arachnids, together with an array of stem-group Paleozoic scorpions that occupied marine habitats, effectively rules out a scenario of a single colonization of terrestrial habitat within Chelicerata, even in tree topologies contrived to recover the monophyly of Arachnida.

Morphology and performance of the 'trap-jaw' cheliceral strikes in spiders (Araneae, Mecysmaucheniidae).

Mecysmaucheniidae spiders have evolved ultra-fast cheliceral strikes 4 times independently. The mechanism for producing these high-speed strikes is likely due to a latch/spring system that allows for stored energy to be rapidly released. This study examined two different sister lineages: Zearchaea has ultra-fast cheliceral strikes and Aotearoa, based on external morphology of the clypeus, is hypothesized to have slower strikes. Using high-speed videography, I first gathered kinematic data on each taxon. Then, using histology and data from micro-computed tomography scanning, I examined internal cheliceral muscle morphology to test whether shifts in muscle anatomy correspond to performance differences in cheliceral strike. Results from high-speed video analysis revealed that Zearchaea achieves peak angular velocities of 25.0×103±4.8×103 rad s-1 (mean±s.d.) in durations of 0.0843±0.017 ms. The fastest recorded strike had a peak angular and linear velocity of 30.8×103 rad s-1 and 18.2 m s-1, respectively. The slower striking sister species, Aotearoa magna, was three orders of magnitude slower in velocity and longer in duration. Histology revealed sarcomere length differences, with some muscles optimized for force, and other muscles for speed. 3D printed models revealed structural differences that explain how the chelicerae hinge open and close. Combining all of this evidence, I put forth a hypothesis for the ultra-fast trap-jaw mechanism. This research documents the morphological shifts that accompany ultra-fast movements that result in increased rotation in joints and increased muscle specialization.

RNA virome screening in diverse but ecologically related citrus pests reveals potential virus-host interactions.

As an evergreen ecosystem, citrus orchards have specialized pest species and stable ecological homeostasis; thus, they provide an ideal model for investigating RNA viromes in diverse but ecologically related species. For this purpose, we collected specialized citrus pests from three classes of invertebrates, Insecta, Arachnida, and Gastropoda and we constructed two kinds of libraries (RNA and small RNA) for the pests by deep sequencing. In total, six virus-derived sequences were identified, including four Picornavirales, one Jingchuvirales and one Nidovirales. The picornavirus-derived small RNAs showed significant small RNA peaks and symmetric distribution patterns along the genome, which suggests these viruses infected the hosts and triggered host antiviral immunity RNA interference. Screening of virus-derived sequences in multiple species of citrus pests (n = 10 per species) showed that Eotetranychus kankitus picorna-like virus and Tetranychus urticae mivirus may be present in multiple pests. Our investigation in citrus pests confirmed that RNA viruses revealed by metagenomics could impact host immunity (e.g. RNAi). An approach with parallel deep sequencing of RNAs and small RNAs is useful not only for viral discoveries but also for understanding virus-host interactions of ecologically related but divergent pest species.

Ordered phylogenomic subsampling enables diagnosis of systematic errors in the placement of the enigmatic arachnid order Palpigradi.

The miniaturized arachnid order Palpigradi has ambiguous phylogenetic affinities owing to its odd combination of plesiomorphic and derived morphological traits. This lineage has never been sampled in phylogenomic datasets because of the small body size and fragility of most species, a sampling gap of immediate concern to recent disputes over arachnid monophyly. To redress this gap, we sampled a population of the cave-inhabiting species Eukoenenia spelaea from Slovakia and inferred its placement in the phylogeny of Chelicerata using dense phylogenomic matrices of up to 1450 loci, drawn from high-quality transcriptomic libraries and complete genomes. The complete matrix included exemplars of all extant orders of Chelicerata. Analyses of the complete matrix recovered palpigrades as the sister group of the long-branch order Parasitiformes (ticks) with high support. However, sequential deletion of long-branch taxa revealed that the position of palpigrades is prone to topological instability. Phylogenomic subsampling approaches that maximized taxon or dataset completeness recovered palpigrades as the sister group of camel spiders (Solifugae), with modest support. While this relationship is congruent with the location and architecture of the coxal glands, a long-forgotten character system that opens in the pedipalpal segments only in palpigrades and solifuges, we show that nodal support values in concatenated supermatrices can mask high levels of underlying topological conflict in the placement of the enigmatic Palpigradi.

Phylogenomic interrogation resolves the backbone of the Pseudoscorpiones tree of life.

Pseudoscorpiones, with nearly 3700 described species, are an ancient and globally distributed group of arachnids with a fossil record dating back to the Middle Devonian. Previous attempts to reconstruct their phylogenetic history have used morphology or a few amplicons, mostly of rRNAs and mitochondrial genes, which have not been able to completely resolve family-level relationships nor the earliest nodes in the pseudoscorpion tree-those which are most informative about the origins of key characters like venoms and silk. Here we undertake a phylogenetic approach using 41 pseudoscorpion transcriptomes and a series of analyses that account for many of the common pitfalls faced in large phylogenomic analyses. All analyses, using concatenation methods and coalescent approaches, supported monophyly of Iocheirata (the venomous pseudoscorpions), which diversified mostly during the Mesozoic, but paraphyly of Epiocheirata, with a sister group relationship of Feaelloidea to Iocheirata, with Chthonioidea as their sister group. These three main lineages were established during the mid-to-late Paleozoic. Our phylogenetic scheme is consistent with the prior hypothesis that the lack of venom in Pseudoscorpiones is plesiomorphic and not a synapomorphy of Epiocheirata. Based on the results of this study, a new classification is proposed for Pseudoscorpiones including the following new nomenclatural and taxonomic acts: the new suborders Palaeosphyronida Harvey and Atoposphyronida Harvey for Dracochelidae and Feaelloidea, respectively; the newly recognized superfamily Garypinoidea for Garypinidae and Larcidae; the revised rank for Lechytiidae and Tridenchthoniidae, which are regarded as subfamilies of Chthoniidae; the revised rank for Tridenchthoniini and Verrucadithini which are regarded as tribes of Tridenchthoniinae; and the elevation of Hesperolpiinae as a distinct family, Hesperolpiidae.

Discovering the silk road: Nuclear and mitochondrial sequence data resolve the phylogenetic relationships among theraphosid spider subfamilies.

The mygalomorph spiders in the family Theraphosidae, also known as "tarantulas", are one of the most popular and diverse groups of arachnids, but their evolutionary history remains poorly understood because morphological analyses have only provided mostly controversial results, and a broad molecular perspective has been lacking until now. In this study we provide a preliminary molecular phylogenetic hypothesis of relationships among theraphosid subfamilies, based on 3.5 kbp of three nuclear and three mitochondrial markers, for 52 taxa representing 10 of the 11 commonly accepted subfamilies. Our analysis confirms the monophyly of the Theraphosidae and of most recognized theraphosid subfamilies, supports the validity of the Stromatopelminae and Poecilotheriinae, and indicates paraphyly of the Schismatothelinae. The placement of representatives of Schismatothelinae also indicates possible non-monophyly of Aviculariinae and supports the distinction of the previously contentious subfamily Psalmopoeinae. Major clades typically corresponded to taxa occurring in the same biogeographic region, with two of them each occurring in Africa, South America and Asia. Because relationships among these major clades were poorly supported, more extensive molecular data sets are required to test the hypothesis of independent colonization and multiple dispersal events among these continents.

A revised dated phylogeny of scorpions: Phylogenomic support for ancient divergence of the temperate Gondwanan family Bothriuridae.

The scorpion family Bothriuridae occupies a subset of landmasses formerly constituting East and West temperate Gondwana, but its relationship to other scorpion families is in question. Whereas morphological data have strongly supported a sister group relationship of Bothriuridae and the superfamily Scorpionoidea, a recent phylogenomic analysis recovered a basal placement of bothriurids within Iurida, albeit sampling only a single exemplar. Here we reexamined the phylogenetic placement of the family Bothriuridae, sampling six bothriurid exemplars representing both East and West Gondwana, using transcriptomic data. Our results demonstrate that the sister group relationship of Bothriuridae to the clade ("Chactoidea" + Scorpionoidea) is supported by the inclusion of additional bothriurid taxa, and that this placement is insensitive to matrix completeness or partitioning by evolutionary rate. We also estimated divergence times within the order Scorpiones using multiple fossil calibrations, to infer whether the family Bothriuridae is sufficiently old to be characterized as a true Gondwanan lineage. We show that scorpions underwent ancient diversification between the Devonian and early Carboniferous. The age interval of the bothriurids sampled (a derived group that excludes exemplars from South Africa) spans the timing of breakup of temperate Gondwana.

Also looking like Limulus? - retinula axons and visual neuropils of Amblypygi (whip spiders).

BACKGROUND: Only a few studies have examined the visual systems of Amblypygi (whip spiders) until now. To get new insights suitable for phylogenetic analysis we studied the axonal trajectories and neuropil architecture of the visual systems of several whip spider species (Heterophrynus elaphus, Damon medius, Phrynus pseudoparvulus, and P. marginemaculatus) with different neuroanatomical techniques. The R-cell axon terminals were identified with Cobalt fills. To describe the morphology of the visual neuropils and of the protocerebrum generally we used Wigglesworth stains and µCT. RESULTS: The visual system of whip spiders comprises one pair of median and three pairs of lateral eyes. The R-cells of both eye types terminate each in a first and a second visual neuropil. Furthermore, a few R-cell fibres from the median eyes leave the second median eye visual neuropil and terminate in the second lateral eye neuropil. This means R-cell terminals from the lateral eyes and the median eyes overlap. Additionally, the arcuate body and the mushroom bodies are described. CONCLUSIONS: A detailed comparison of our findings with previously studied chelicerate visual systems (i.e., Xiphosura, Scorpiones, Pseudoscorpiones, Opiliones, and Araneae) seem to support the idea of close evolutionary relationships between Xiphosura, Scorpiones, and Amblypygi.

HEMOLYMPH CHEMISTRY REFERENCE RANGES OF THE CHILEAN ROSE TARANTULA GRAMMOSTOLA ROSEA (WALKENAER, 1837) USING THE VETSCAN BIOCHEMISTRY ANALYZER BASED ON IFCC-CLSI C28-A3.

The use of invertebrate hemolymph chemistry analysis has the potential to become a major diagnostic tool. The goal of this study was to generate statistically sound hemolymph reference ranges from healthy tarantulas. Hemolymph was drawn from wild caught, acclimatized, and apparently healthy female Chilean rose tarantulas Grammostola rosea (Walkenaer, 1837) ( n = 43) using a modified technique. Hemolymph samples were separately analyzed using the Avian-Reptilian Profile Plus reagent rotor for VetScan® for the following chemistries: aspartate aminotransferase, bile acids, creatine kinase, uric acid, glucose, total calcium, phosphorus, total protein, albumin, potassium, and sodium. With this method the authors were able to establish statistically sound reference ranges for aspartate aminotransferase, creatine kinase, glucose, phosphorus, and total protein. Further in situ studies will determine the practical usability of these values in the evaluation of tarantula health.

The Opiliones tree of life: shedding light on harvestmen relationships through transcriptomics.

Opiliones are iconic arachnids with a Palaeozoic origin and a diversity that reflects ancient biogeographic patterns dating back at least to the times of Pangea. Owing to interest in harvestman diversity, evolution and biogeography, their relationships have been thoroughly studied using morphology and PCR-based Sanger approaches to infer their systematic relationships. More recently, two studies utilized transcriptomics-based phylogenomics to explore their basal relationships and diversification, but sampling was limiting for understanding deep evolutionary patterns, as they lacked good taxon representation at the family level. Here, we analysed a set of the 14 existing transcriptomes with 40 additional ones generated for this study, representing approximately 80% of the extant familial diversity in Opiliones. Our phylogenetic analyses, including a set of data matrices with different gene occupancy and evolutionary rates, and using a multitude of methods correcting for a diversity of factors affecting phylogenomic data matrices, provide a robust and stable Opiliones tree of life, where most families and higher taxa are precisely placed. Our dating analyses using alternative calibration points, methods and analytical parameters provide well-resolved old divergences, consistent with ancient regionalization in Pangea in some groups, and Pangean vicariance in others. The integration of state-of-the-art molecular techniques and analyses, together with the broadest taxonomic sampling to date presented in a phylogenomic study of harvestmen, provide new insights into harvestmen interrelationships, as well as an overview of the general biogeographic patterns of this ancient arthropod group.

Numerical simulation of colloidal self-assembly of super-hydrophobic arachnid cerotegument structures.

Certain arachnids exhibit complex coatings of their exoskeleton, consisting of globular structures with complex surface features. This, so-called, cerotegument is formed by a multi-component colloidal secretion that self-assembles and cures on the body surface, and leads to high water repellency. Previous ultrastructural studies revealed the involvement of different glandular cells that contribute different components to the secretion mixture, but the overall process of self-assembly into the complex regular structures observed remained highly unclear. Here we study this process from a theoretical point of view, starting from the so-called Tammes-problem. We show that slight changes of simple parameters lead to a variety of morphologies that are highly similar to the ones observed in the species specific cerotegument structures of whip-spiders. These results are not only important for our understanding of the formation of globular hierarchical structures in nature, but also for the fabrication of novel surface coatings by colloidal lithography.

Permian scorpions from the Petrified Forest of Chemnitz, Germany.

BACKGROUND: Paleozoic scorpions (Arachnida: Scorpiones) have been widely documented from the Carboniferous Period; which hosts a remarkable assemblage of more than sixty species including both putative stem- and crown-group fossils. By contrast the succeeding Permian Period is almost completely devoid of records, which are currently restricted to a trace fossil from the early Permian of New Mexico, USA and some limb fragments from the late Permian of the Vologda Region, Russia. RESULTS: ?Opsieobuthus tungeri sp. nov. from the Petrified Forest of Chemnitz, Germany represents the first complete body fossils of scorpions from the Permian. Explosive volcanism preserved these remarkable specimens in situ as part of the palaeosol horizon and bedrock of the Petrified Forest, immediately beneath the Zeisigwald tuff horizon. This dates to the early Permian (Sakmarian) or ca. 291 Ma. Intriguingly, the specimens were obtained from a palaeosol horizon with a compacted network of different-sized woody roots and thus have been preserved in situ in their likely life position, even within their original burrows. Differences in the structure of the comb-like pectines in the two fossils offer evidence for sexual dimorphism, and permit further inferences about the ecology and perhaps even the reproductive biology of these animals. CONCLUSIONS: As putative members of a Coal Measures genus, these fossils suggest that at least some Carboniferous scorpion lineages extended their range further into the Permian. This contributes towards a picture of scorpion evolution in which both basal and derived (orthostern) forms coexisted for quite some time; probably from the end of the Carboniferous through to at least the mid Triassic.