Prashant P Sharma

University of Wisconsin–Madison | UW · Department of Integrative Biology

Despite significant advances in invertebrate phylogenomics over the past decade, the higher-level phylogeny of Pycnogonida (sea spiders) remains elusive. Due to the inaccessibility of some small-bodied lineages, few phylogenetic studies have sampled all sea spider families. Previous efforts based on a handful of genes have yielded unstable tree top...

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 Phalang...

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. While conflicting hypotheses prevail in morphological and molecular datasets alike, the monophyly of Arac...

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 ha...

Neofunctionalization of duplicated gene copies is thought to be an important process underlying the origin of evolutionary novelty and provides an elegant mechanism for the origin of new phenotypic traits. One putative case where a new gene copy has been linked to a novel morphological trait is the origin of the arachnid patella, a taxonomically re...

Asymmetrical rates of cladogenesis and extinction abound in the Tree of Life, resulting in numerous minute clades that are dwarfed by larger sister groups. Such taxa are commonly regarded as phylogenetic relicts or “living fossils” when they exhibit an ancient first appearance in the fossil record and prolonged external morphological stasis, partic...

Asymmetrical rates of cladogenesis and extinction abound in the Tree of Life, resulting in numerous minute clades that are dwarfed by larger sister groups. Such taxa are commonly regarded as phylogenetic relicts or "living fossils" when they exhibit an ancient first appearance in the fossil record and prolonged external morphological stasis, partic...

Vestigial organs provide a link between ancient and modern traits and therefore have great potential to resolve the phylogeny of contentious fossils that bear features not seen in extant species. Here we show that extant daddy-longlegs (Arachnida, Opiliones), a group once thought to possess only one pair of eyes, in fact additionally retain a pair...

The biota of cave habitats faces heightened conservation risks, due to geographic isolation and high levels of endemism. Molecular datasets, in tandem with ecological surveys, have the potential to delimit precisely the nature of cave endemism and identify conservation priorities for microendemic species. Here, we sequenced ultraconserved elements...

Recent advances in higher-level invertebrate phylogeny have leveraged shared features of genomic architecture to resolve contentious nodes across the tree of life. Yet, the interordinal relationships within Chelicerata have remained recalcitrant given competing topologies in recent molecular analyses. As such, relationships between topologically un...

The armored harvestman family Assamiidae (Arachnida: Opiliones: Laniatores) is widely distributed throughout the Old World tropics, specifically throughout tropical Asia and Central Africa. However, the systematics and intrafamilial relationships of the group remain poorly understood. This can be largely attributed to the complicated taxonomic hist...

The location of Israel at the junction of three continents leads to a unique fauna of both Palearctic and Afrotropical zoogeographic origins. Following systematic revisions over the past sixty years and the discovery of new species, the only available key to the pseudoscorpions of Israel has become outdated. We provide here an up-to-date checklist...

The chelicerate body plan is distinguished from other arthropod groups by its division of segments into two tagmata: the anterior prosoma (cephalothorax) and the posterior opisthosoma (abdomen). Little is understood about the genetic mechanisms that establish the prosomal-opisthosomal (PO) boundary. To discover these mechanisms, we created high-qua...

Advanced sequencing technologies have expedited resolving higher-level arthropod relationships. Yet, dark branches persist, principally among groups occurring in cryptic habitats. Among chelicerates, Solifugae (“camel spiders”) is the last order lacking a higher-level phylogeny and thus, historically characterized as “neglected [arachnid] cousins”....

Scorpions are ancient and historically renowned for their potent venom. Traditionally, the systematics of this group of arthro-pods was supported by morphological characters, until recent phylogenomic analyses (using RNAseq data) revealed most of the higher-level taxa to be non-monophyletic. While these phylogenomic hypotheses are stable for almost...

The proliferation of genomic resources for Chelicerata in the past ten years has revealed that the evolution of chelicerate genomes is more dynamic than previously thought, with multiple waves of ancient whole genome duplications affecting separate lineages. Such duplication events are fascinating from the perspective of evolutionary history becaus...

The location of Israel at the junction of three continents leads to a unique faunal combination of Palearctic and Afrotropic zoogeographic origins. Following systematic revisions over the past sixty years and the discovery of new species, the only available key to the pseudoscorpions (Arachnida: Pseudoscorpiones) of Israel (Beier 1963) has become o...

Despite an abundance of gene expression surveys, comparatively little is known about Hox gene function in Chelicerata. Previous investigations of paralogs of labial (lab) and Deformed (Dfd) in a spider have shown that these play a role in tissue maintenance of the pedipalpal segment (lab-1) and in patterning the first walking leg identity (Dfd-1),...

Caves have long been recognized as a window into the mechanisms of diversification and convergent evolution, due to the unique conditions of isolation and life in the dark. These lead to adaptations and reduce dispersal and gene flow, resulting in high levels of speciation and endemism. The Israeli cave arachnofauna remains poorly known, but likely...

Opiliones (harvestmen) have come to be regarded as an abundant source of model groups for study of historical biogeography, due to their ancient age, poor dispersal capability, and high fidelity to biogeographic terranes. One of the least understood harvestman groups is the Paleotropical Assamiidae, one of the more diverse families of Opiliones. Du...

Despite an abundance of gene expression surveys, comparatively little is known about Hox gene function in Chelicerata, with emphasis on the Hox logic of the anterior prosomal segments, which bear the mouthparts. Previous investigations of individual paralogs of labial ( lab ) and Deformed ( Dfd ) in the spider Parasteatoda tepidariorum have shown t...

Here, we report the draft genome and annotation of "Candidatus Nardonella dryophthoridicola" strain NARMHE1, obtained via Oxford Nanopore sequencing of the ovaries of its host, the weevil Metamasius hemipterus, from a population from southeast Brazil. M etamasius (family, Curculionidae; subfamily, Dryophthorinae) harbors obligate endo-symbiont Nard...

Considerable progress has been achieved in resolving higher-level relationships of Arthropoda in the past two decades, largely precipitated by advances in sequencing technology. Yet, dark branches persist in the arthropod tree of life, principally among groups that are difficult to collect, occur in cryptic habitats, or are characterized by minute...

Four new laniatorean harvestmen specimens (Arachnida: Opiliones: Laniatores) are described from the mid-Cretaceous (upper Albian–lower Cenomanian) Burmese amber of Northern Myanmar. One is placed as Insidiatores indet., but is not formally named as it is probably immature. Burmalomanius circularis n. gen. n. sp. and Petroburma tarsomeria n. gen. n....

Metamasius is a large genus of dryophthorine weevils, with nearly 85 species. Among the economically important pests in the genus, M. hemipterus is currently separated in three subspecies, based largely on color patterns of the elytra, pronotum, and sternum. The tenuous limits of M. hemipterus subspecies were created over fifty years ago and never...

Caves constitute ideal study systems for investigating adaptation and speciation, as the abiotic conditions shared by aphotic habitats exert a set of environmental filters on their communities. Arachnids constitute an important component of many cave ecosystems worldwide. We investigated the population genomics of two whip spider species: Sarax ioa...

Asymmetrical rates of cladogenesis and extinction abound in the Tree of Life, resulting in numerous minute clades that are dwarfed by larger sister groups. Such taxa are commonly regarded as phylogenetic relicts or "living fossils" when they exhibit an ancient first appearance in the fossil record and prolonged external morphological stasis, partic...

Background The comparative embryology of Chelicerata has greatly advanced in recent years with the integration of classical studies and genetics, prominently spearheaded by developmental genetic works in spiders. Nonetheless, the understanding of the evolution of development and polarization of embryological characters in Chelicerata is presently l...

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...

The basal phylogeny of Chelicerata is one of the opaquest parts of the animal Tree of Life, defying resolution despite application of thousands of loci and millions of sites. At the forefront of the debate over chelicerate relationships is the monophyly of Arachnida, which has been refuted by most analyses of molecular sequence data. A number of ph...

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. While conflicting hypotheses prevail in morphological and molecular datasets alike, the monophyly of Arac...

The Sox family of transcription factors regulate many processes during metazoan development, including stem cell maintenance and nervous system specification. Characterising the repertoires and roles of these genes can therefore provide important insights into animal evolution and development. We further characterised the Sox repertoires of several...

The homology of the arachnid chelicera with respect to other head appendages in Panarthropoda has long been debated. Gene expression data and the re-interpretation of early transitional fossils have supported the homology of the deutocerebrum and its associated appendages, implying a homology between primary antennae (mandibulates), chelicerae (euc...

One new family, six new genera and six new species of laniatorean harvestmen (Arachnida: Opiliones: Laniatores) from the mid-Cretaceous (Cenomanian, ca. 99 Ma) Burmese amber of Northern Myanmar are described. Ellenbergellus tuberculatus gen. et sp. nov. is the first fossil assignable to the extant family Tithaeidae. Mesokanus oehmkuehnlei gen. et s...

Segmentation is a key characteristic of Arthropoda that is linked to the evolutionary success of this lineage. It has previously been shown in both vertebrates and short germ insects that posterior segmentation requires canonical Wnt (cWnt) signaling, which maintains the expression of Caudal and the posterior growth zone; disruption of cWnt signali...

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 LBA 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 despit...

Chelicerates exhibit dynamic evolution of genome architecture, with multiple whole genome duplication events affecting groups like spiders, scorpions, and horseshoe crabs. Yet, genomes remain unavailable for several chelicerate orders, such as Opiliones (harvestmen), which has hindered comparative genomics and developmental genetics across arachnid...

The Cyphophthalmi genus Troglosiro (the only genus of the family Troglosironidae) is endemic to New Caledonia, representing one of the oldest lineages of this emerged part of Zealandia. Its species are short-range endemics, many known from single localities. Here we examined the phylogenetic relationships of Troglosironidae using standard Sanger-se...

Background Arachnids are important components of cave ecosystems and display many examples of troglomorphisms, such as blindness, depigmentation, and elongate appendages. Little is known about how the eyes of arachnids are specified genetically, let alone the mechanisms for eye reduction and loss in troglomorphic arachnids. Additionally, duplicatio...

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 LBA 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 despit...

Sexually dimorphic traits are widespread in animals, and include sex-specific weapons, ornamentation and, although less noticed, glands and associated structures. In arachnids, certain lineages of the order Opiliones exhibit diverse forms of dimorphism in the armature and length of appendages (common in Laniatores), as well as in the presence of se...

Southern Australian waters feature remarkably diverse assemblages of the sea spider family Callipallenidae Hilton, 1942. The most speciose of the three Australian-endemic genera currently recognised has been known as Meridionale Staples, 2014, but is here reinstated under the name Pallenella Schimkewitsch, 1909 based on its type species Pallenella...

Scorpions constitute a charismatic lineage of arthropods and comprise more than 2,500 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 h...

Background: The resurgence of interest in the comparative developmental study of chelicerates has led to important insights, such as the discovery of a genome duplication shared by spiders and scorpions, inferred to have occurred in the most recent common ancestor of Arachnopulmonata (a clade comprising the five arachnid orders that bear book lungs...

Invasive, nuisance aquatic species such as Eurasian watermilfoil (Myriophyllum spicatum) are rapidly spreading across the United States. One common active ingredient used to control this invasive species is 2,4-Dichlorophenoxyacetic acid (2,4-D). Application of 2,4-D to aquatic environments typically occurs while many freshwater fish are spawning a...

Background The resurgence of interest in the comparative developmental study of chelicerates has led to important insights, such as the discovery of a genome duplication shared by spiders and scorpions, inferred to have occurred at the most recent common ancestor of Arachnopulmonata (a clade comprised of the five book lung-bearing arachnid orders)....

The Sox family of transcription factors regulate many different processes during metazoan development, including stem cell maintenance, nervous system specification and germline development. In addition, it has recently become apparent that SoxB genes are involved in embryonic segmentation in several arthropod species. Segmentation in arthropods oc...

A new species of Cyphophthalmi belonging to the Southeast Asian genus Miopsalis Thorell, 1890 is described and illustrated using SEM and confocal microscopy. The species is known from two localities in the northern part of Mindanao and constitutes only the second described Cyphophthalmi species endemic to the Philippines. Sequence data obtained fro...

Arachnids are important components of cave ecosystems and display many examples of troglomorphisms, such as blindness, depigmentation, and elongate appendages. Little is known about how the eyes of arachnids are specified genetically, let alone the mechanisms for eye reduction and loss in troglomorphic arachnids. Additionally, paralogy of Retinal D...

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 scorp...

Developmental modes in pycnogonids are variable, even between congeneric species. Most of our previous knowledge of pycnogonid life history stages is based on morphological descriptions of the larval and those postlarval instars still retained on male ovigers, whereas the free-living instars are largely unknown. In this paper, a combination of morp...

Despite significant advances in invertebrate phylogenomics over the past decade, the higher-level phylogeny of Pycnogonida (sea spiders) remains elusive. This group of arthropods appeared early in the fossil record, with the oldest unambiguous fossils dating to the Silurian. Due to the inaccessibility of some small-bodied lineages, few phylogenetic...

Sexually dimorphic traits are widespread in animals, and include sex-specific weapons, ornamentation and, although less noticed, glands and associated structures. In arachnids, certain lineages of the order Opiliones exhibit diverse forms of dimorphism in the armature and length of appendages (common in Laniatores), as well as in the presence of se...

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 arachn...

Extended materials and methods

A detailed account of the errors and anomalies in the construction of Matrix A and Matrix B in the work of Lozano Fernández et al. (2019).

Morphological characters historically have supported the monophyly, composition and phylogenetic relationships among scorpion families. While recent phylogenomic analyses have recovered most of these traditional higher-level relationships as non-monophyletic, certain key taxa have yet to be sampled using a phylogenomic approach. Salient among these...

Scorpion toxins are thought to have originated from ancestral housekeeping genes that underwent diversification and neofunctionalization, as a result of positive selection. Our understanding of the evolutionary origin of these peptides is hindered by the patchiness of existing taxonomic sampling. While recent studies have shown phylogenetic inertia...

Horseshoe crabs (Xiphosura) are traditionally regarded as sister group to the clade of terrestrial chelicerates (Arachnida). This hypothesis has been challenged by recent phylogenomic analyses, but the non-monophyly of Arachnida has consistently been disregarded as artifactual. We reevaluated the placement of Xiphosura among chelicerates using the...

After tumultuous revisions to the family-level systematics of Laniatores (the armored harvestmen), the basally branching family Phalangodidae presently bears a disjunct and irregular distribution, attributed to the fragmentation of Pangea. One of the curious lineages assigned to Phalangodidae is the monotypic Israeli genus Haasus, the only Laniator...

Phylogenetic relationships in Opiliones (Arachnida) at the suborder level have greatly stabilized in recent years, largely due to advances in molecular systematics. Nonetheless, identifying morphological characters in the context of well-resolved phylogenies is essential for testing new systematic hypotheses and establishing diagnostic markers. Her...

The Neartic family Vaejovidae (Scorpiones: Chactoidea) has long been treated as a diverse and systematically cohesive group of scorpions, but its monophyly and relationship to other scorpion families have historically been questioned. Morphological data have supported its monophyly and a variety of phylogenetic placements within the superfamily Cha...

The mygalomorph spider Aphonopelma hentzi (Girard, 1852) (Texas brown tarantula) is a geographically widespread and accessible system for the study of comparative development in spiders. However, remarkably little information is available regarding the timing of egg sac deposition and duration of embryogenesis in this species, a gap that can direct...

The Philippine archipelago harbors a remarkable diversity of harvestmen, with respect to both taxonomy and complexity of biogeographic origins. Among the armored harvestmen (suborder Laniatores), six families of distantly related groups occur in this archipelago. Here we describe a new species of the family Tithaeidae, Tithaeus odysseus sp. nov., d...

Two new species of Petrobunus Sharma & Giribet, 2011, (Petrobunidae) are described: P. chongqing sp. nov. (♀♂) and P. hebei sp. nov. (♀♂). Petrobunus chongqing sp. nov. is similar to P. spinifer Sharma & Giribet, 2011, in having two large spines on the male sternite 7. However, the new species can be distinguished from P. spinifer by the absence of...

Scorpions have evolved a variety of toxins with a plethora of biological targets, but characterizing their evolution has been limited by the lack of a comprehensive phylogenetic hypothesis of scorpion relationships grounded in modern, genomescale datasets. Disagreements over scorpion higher-level systematics have also incurred challenges to previou...

The transcription factors spineless (ss) and tiptop/teashirt (tio/tsh) have been shown to be selectors of distal appendage identity in an insect, but it is unknown how they regulate one another. Here, we examined the regulatory relationships between these two determinants in the milkweed bug Oncopeltus faciatus, using maternal RNA interference (RNA...

Compared to other arthropods, such as crustaceans or insects, the term 'chelicerate' often does not evoke a similar sense of recognition or familiarity. Yet the subphylum Chelicerata has been encountered by every living person today, frequently to the effect of fear, awe, or outright revulsion. Chelicerates include such familiar groups as spiders,...

Homeobox genes are key toolkit genes that regulate the development of metazoans and changes in their regulation and copy number have contributed to the evolution of phenotypic diversity. We recently identified a whole genome duplication (WGD) event that occurred in an ancestor of spiders and scorpions (Arachnopulmonata), and that many homeobox gene...

The study of sensory structures has the potential to provide insights into the natural history and evolution of animals. The sensory structures of arachnids are usually concentrated on the pedipalps (the tritocerebral appendages) or on the distal podomere (tarsus) of the anterior walking legs, the latter being the case for armored harvestmen (Opili...

Austropurcellia, a genus of dispersal-limited arachnids endemic to isolated patches of coastal rainforest in Queensland, Australia, has a remarkable biogeographic history. The genus is a member of the family Pettalidae, which has a classical temperate Gondwanan distribution; previous work has suggested that Austropurcellia is an ancient lineage, wi...

Transcriptomic and genomic analyses have illuminated the diversity of venoms in three of the four venomous arachnid orders (scorpions, spiders, and ticks). To date, no venom gland transcriptome analysis has been available for pseudoscorpions, the fourth venomous arachnid lineage. To redress this gap, we sequenced an mRNA library generated from the...

Background: The morphological and functional evolution of appendages has played a key role in the diversification of arthropods. While the ancestral arthropod appendage is held to be polyramous, terrestriality is associated with the reduction or loss of appendage rami, which may obscure the homology of different appendage derivatives. Proxies for...

Significance In the fruit fly, ventral appendage (leg) identity is specified by a gene network including Wnt-1/wg , Sp6-9 , and Dll , but little is known about the conservation of this network beyond insects. We disrupted Wnt signaling and Sp6-9 in a spider, a member of Chelicerata, the sister group to all remaining arthropods. Our results provide...

Supplementary Appendix accompanying Setton and Sharma (2018).

We provide a first scanning electron microscopy examination of the Paleotropical harvestman family Podoctidae (Opiliones: Laniatores), focusing on the distitarsus of the legs of Metibalonius sp. Distitarsi I and II are mostly equipped with olfactory sensilla chaetica with wall pores, while those of legs III and IV have gustatory sensilla chaetica w...

The scorpion family Bothriuridae occupies a subset of landmasses formerly constituting East and West 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 ba...

Paragraph In animals, small RNA molecules termed PIWI-interacting RNAs (piRNAs) silence transposable elements (TEs), protecting the germline from genomic instability and mutation. piRNAs have been detected in the soma in a few animals, but these are believed to be specific adaptations of individual species. Here, we report that somatic piRNAs were...

During the present study, we examined the phylogeography and systematics of two species of velvet worm (Peripatopsis Pocock, 1894) in the forested region of the southern Cape of South Africa. A total of 89 P. moseleyi (Wood-Mason, 1879) and 65 P. sedgwicki (Purcell, 1899) specimens were collected and sequenced for the cytochrome c oxidase subunit I...

Arachnids exhibit tremendous species richness and adaptations of biomedical, industrial, and agricultural importance. Yet genomic resources for arachnids are limited, with the first few spider and scorpion genomes becoming accessible in the last four years. We review key insights from these genome projects, and recommend additional genomes for sequ...

In animals, small RNA molecules termed PIWI-interacting RNAs (piRNAs) silence transposable elements (TEs), protecting the germline from genomic instability and mutation. piRNAs have been detected in the soma in a few animals, but these are believed to be specific adaptations of individual species. Here, we report that somatic piRNAs were likely pre...

Synopsis The internal phylogeny of Chelicerata and the attendant evolutionary scenario of arachnid terrestrialization have a long and contentious history. Previous studies of developmental gene expression data have suggested that respiratory systems of spiders, crustaceans, and insects are all serially homologous structures derived from the epipods...

Background: The duplication of genes can occur through various mechanisms and is thought to make a major contribution to the evolutionary diversification of organisms. There is increasing evidence for a large-scale duplication of genes in some chelicerate lineages including two rounds of whole genome duplication (WGD) in horseshoe crabs. To investi...