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Alena Lukesova - One of the best experts on this subject based on the ideXlab platform.
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microwhip scorpions palpigradi feed on heterotrophic cyanobacteria in slovak caves a curiosity among Arachnida
PLOS ONE, 2013Co-Authors: Jaroslav Smrž, ĺubomir Kovac, Jaromir Mikes, Alena LukesovaAbstract:To date, only morphological and anatomical descriptions of microwhip scorpions (Arachnida: Palpigradi) have been published. This very rare group is enigmatic not only in its relationships to other arachnids, but especially due to the fact that these animals dwell only underground (in caves, soil, and interstitial spaces). We observed the curious feeding habit of the microwhip scorpion Eukoenenia spelaea over the course of one year in Ardovska Cave, located in Slovakia's Karst region. We chose histology as our methodology in studying 17 specimens and based it upon Masson's triple staining, fluorescent light and confocal microscopy. Single-celled cyanobacteria (blue-green algae) were conspicuously predominant in the gut of all studied palpigrades. Digestibility of the consumed cyanobacteria was supported by the presence of guanine crystals, glycogen deposits and haemocytes inside the palpigrade body. Cyanobacteria, the oldest cellular organisms on Earth, are very resistant to severe conditions in caves, including even darkness. Therefore, the cyanobacteria are able to survive in dark caves as nearly heterotrophic organisms and are consumed by cave palpigrades. Such feeding habit is extraordinary within the almost wholly predacious orders of the class Arachnida, and particularly so due to the type of food observed.
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Microwhip Scorpions (Palpigradi) Feed on Heterotrophic Cyanobacteria in Slovak Caves – A Curiosity among Arachnida
PloS one, 2013Co-Authors: Jaroslav Smrž, Jaromir Mikes, Ĺubomír Kováč, Alena LukesovaAbstract:To date, only morphological and anatomical descriptions of microwhip scorpions (Arachnida: Palpigradi) have been published. This very rare group is enigmatic not only in its relationships to other arachnids, but especially due to the fact that these animals dwell only underground (in caves, soil, and interstitial spaces). We observed the curious feeding habit of the microwhip scorpion Eukoenenia spelaea over the course of one year in Ardovska Cave, located in Slovakia's Karst region. We chose histology as our methodology in studying 17 specimens and based it upon Masson's triple staining, fluorescent light and confocal microscopy. Single-celled cyanobacteria (blue-green algae) were conspicuously predominant in the gut of all studied palpigrades. Digestibility of the consumed cyanobacteria was supported by the presence of guanine crystals, glycogen deposits and haemocytes inside the palpigrade body. Cyanobacteria, the oldest cellular organisms on Earth, are very resistant to severe conditions in caves, including even darkness. Therefore, the cyanobacteria are able to survive in dark caves as nearly heterotrophic organisms and are consumed by cave palpigrades. Such feeding habit is extraordinary within the almost wholly predacious orders of the class Arachnida, and particularly so due to the type of food observed.
Davide Pisani - One of the best experts on this subject based on the ideXlab platform.
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increasing species sampling in chelicerate genomic scale datasets provides support for monophyly of acari and Arachnida
Nature Communications, 2019Co-Authors: Jesus Lozanofernandez, Gregory D. Edgecombe, Alastair R Tanner, Mattia Giacomelli, Robert Carton, Jakob Vinther, Davide PisaniAbstract:Chelicerates are a diverse group of arthropods, represented by such forms as predatory spiders and scorpions, parasitic ticks, humic detritivores, and marine sea spiders (pycnogonids) and horseshoe crabs. Conflicting phylogenetic relationships have been proposed for chelicerates based on both morphological and molecular data, the latter usually not recovering arachnids as a clade and instead finding horseshoe crabs nested inside terrestrial Arachnida. Here, using genomic-scale datasets and analyses optimised for countering systematic error, we find strong support for monophyletic Acari (ticks and mites), which when considered as a single group represent the most biodiverse chelicerate lineage. In addition, our analysis recovers marine forms (sea spiders and horseshoe crabs) as the successive sister groups of a monophyletic lineage of terrestrial arachnids, suggesting a single colonisation of land within Chelicerata and the absence of wholly secondarily marine arachnid orders. Morphological and molecular data have led to conflicting phylogenetic hypotheses for the Chelicerata. Here, the authors reconstruct the phylogeny of the Chelicerata using genomic-scale datasets, finding evidence for a monophyletic Acari and a single terrestrialisation of Arachnida.
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Exploring the evolution and terrestrialization of scorpions (Arachnida: Scorpiones) with rocks and clocks
Organisms Diversity & Evolution, 2019Co-Authors: Richard J. Howard, Gregory D. Edgecombe, David A. Legg, Davide Pisani, Jesus Lozano-fernandezAbstract:Scorpions (Arachnida: Scorpiones Koch, 1837) are an ancient chelicerate arthropod lineage characterised by distinctive subdivision of the opisthosoma and venomous toxicity. The crown group is represented by over 2400 extant species, and unambiguous fossil representatives are known at least from the Cretaceous Period. However, a number of extinct scorpion lineages existed in the Palaeozoic Era, many of which are of a contentious marine (or at least semi-aquatic) lifestyle, and have long caused confusion regarding the nature of arachnid terrestrialization and arachnid phylogeny more broadly. To clarify the process of terrestrialization, there is a need to marry fossil and extant scorpions in a common evolutionary framework utilising modern advances in phylogenetics. Here, we review phylogenetic hypotheses of arachnid and scorpion interrelationships, relevant advances in phylogenetic divergence time estimation and the scorpion fossil record—especially with reference to terrestrialization. In addition, we provide a list of scorpion fossil calibrations for use in molecular dating and demonstrate their utility in deriving a novel scorpion time tree using Bayesian relaxed-clock methods. Our results reveal a window of divergence from 335 to 266 Mya for the scorpion crown group, consistent with a Pangean origin of crown scorpions inferred from the biogeographical distribution of the extant fauna.
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Homeobox gene duplication and divergence in arachnids.
Molecular Biology and Evolution, 2018Co-Authors: Daniel J. Leite, Luis Baudouin-gonzalez, Sawa Iwasaki-yokozawa, Natascha Turetzek, Nikola-michael Prpic, Yasuko Akiyama, Davide Pisani, Jesus Lozano-fernandez, Prashant P SharmaAbstract:: 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 genes, including two Hox clusters, appear to have been retained in arachnopulmonates. To better understand the consequences of this ancient WGD and the evolution of arachnid homeobox genes, we have characterised and compared the homeobox repertoires in a range of arachnids. We found that many families and clusters of these genes are duplicated in all studied arachnopulmonates (Parasteatoda tepidariorum, Pholcus phalangioides, Centruroides sculpturatus and Mesobuthus martensii) compared with non-arachnopumonate arachnids (Phalangium opilio, Neobisium carcinoides, Hesperochernes sp. and Ixodes scapularis). To assess divergence in the roles of homeobox ohnologs, we analysed the expression of P. tepidariorum homeobox genes during embryogenesis and found pervasive changes in the level and timing of their expression. Furthermore, we compared the spatial expression of a subset of P. tepidariorum ohnologs with their single copy orthologs in P. opilio embryos. We found evidence for likely subfunctionlisation and neofunctionalisation of these genes in the spider. Overall our results show a high level of retention of homeobox genes in spiders and scorpions post WGD, which is likely to have made a major contribution to their developmental evolution and diversification through pervasive subfunctionlisation and neofunctionalisation, and paralleling the outcome of WGD in vertebrates.
Prashant P Sharma - One of the best experts on this subject based on the ideXlab platform.
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Developmental gene expression as a phylogenetic data class: support for the monophyly of Arachnopulmonata
Development Genes and Evolution, 2020Co-Authors: Erik D. Nolan, Carlos E. Santibáñez-lópez, Prashant P SharmaAbstract: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.
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a critical appraisal of the placement of xiphosura chelicerata with account of known sources of phylogenetic error
Systematic Biology, 2019Co-Authors: Jesus A Ballesteros, Prashant P SharmaAbstract: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 re-evaluated the placement of Xiphosura among chelicerates using the most complete phylogenetic data set to date, expanding outgroup sampling, and including data from whole genome sequencing projects. In spite of uncertainty in the placement of some arachnid clades, all analyses show Xiphosura consistently nested within Arachnida as the sister group to Ricinulei (hooded tick spiders). It is apparent that the radiation of arachnids is an old one and occurred over a brief period of time, resulting in several consecutive short internodes, and thus is a potential case for the confounding effects of incomplete lineage sorting (ILS). We simulated coalescent gene trees to explore the effects of increasing levels of ILS on the placement of horseshoe crabs. In addition, common sources of systematic error were evaluated, as well as the effects of fast-evolving partitions and the dynamics of problematic long branch orders. Our results indicated that the placement of horseshoe crabs cannot be explained by missing data, compositional biases, saturation, or ILS. Interrogation of the phylogenetic signal showed that the majority of loci favor the derived placement of Xiphosura over a monophyletic Arachnida. Our analyses support the inference that horseshoe crabs represent a group of aquatic arachnids, comparable to aquatic mites, breaking a long-standing paradigm in chelicerate evolution and altering previous interpretations of the ancestral transition to the terrestrial habitat. Future studies testing chelicerate relationships should approach the task with a sampling strategy where the monophyly of Arachnida is not held as the premise.
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Homeobox gene duplication and divergence in arachnids.
Molecular Biology and Evolution, 2018Co-Authors: Daniel J. Leite, Luis Baudouin-gonzalez, Sawa Iwasaki-yokozawa, Natascha Turetzek, Nikola-michael Prpic, Yasuko Akiyama, Davide Pisani, Jesus Lozano-fernandez, Prashant P SharmaAbstract:: 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 genes, including two Hox clusters, appear to have been retained in arachnopulmonates. To better understand the consequences of this ancient WGD and the evolution of arachnid homeobox genes, we have characterised and compared the homeobox repertoires in a range of arachnids. We found that many families and clusters of these genes are duplicated in all studied arachnopulmonates (Parasteatoda tepidariorum, Pholcus phalangioides, Centruroides sculpturatus and Mesobuthus martensii) compared with non-arachnopumonate arachnids (Phalangium opilio, Neobisium carcinoides, Hesperochernes sp. and Ixodes scapularis). To assess divergence in the roles of homeobox ohnologs, we analysed the expression of P. tepidariorum homeobox genes during embryogenesis and found pervasive changes in the level and timing of their expression. Furthermore, we compared the spatial expression of a subset of P. tepidariorum ohnologs with their single copy orthologs in P. opilio embryos. We found evidence for likely subfunctionlisation and neofunctionalisation of these genes in the spider. Overall our results show a high level of retention of homeobox genes in spiders and scorpions post WGD, which is likely to have made a major contribution to their developmental evolution and diversification through pervasive subfunctionlisation and neofunctionalisation, and paralleling the outcome of WGD in vertebrates.
Jaroslav Smrž - One of the best experts on this subject based on the ideXlab platform.
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microwhip scorpions palpigradi feed on heterotrophic cyanobacteria in slovak caves a curiosity among Arachnida
PLOS ONE, 2013Co-Authors: Jaroslav Smrž, ĺubomir Kovac, Jaromir Mikes, Alena LukesovaAbstract:To date, only morphological and anatomical descriptions of microwhip scorpions (Arachnida: Palpigradi) have been published. This very rare group is enigmatic not only in its relationships to other arachnids, but especially due to the fact that these animals dwell only underground (in caves, soil, and interstitial spaces). We observed the curious feeding habit of the microwhip scorpion Eukoenenia spelaea over the course of one year in Ardovska Cave, located in Slovakia's Karst region. We chose histology as our methodology in studying 17 specimens and based it upon Masson's triple staining, fluorescent light and confocal microscopy. Single-celled cyanobacteria (blue-green algae) were conspicuously predominant in the gut of all studied palpigrades. Digestibility of the consumed cyanobacteria was supported by the presence of guanine crystals, glycogen deposits and haemocytes inside the palpigrade body. Cyanobacteria, the oldest cellular organisms on Earth, are very resistant to severe conditions in caves, including even darkness. Therefore, the cyanobacteria are able to survive in dark caves as nearly heterotrophic organisms and are consumed by cave palpigrades. Such feeding habit is extraordinary within the almost wholly predacious orders of the class Arachnida, and particularly so due to the type of food observed.
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Microwhip Scorpions (Palpigradi) Feed on Heterotrophic Cyanobacteria in Slovak Caves – A Curiosity among Arachnida
PloS one, 2013Co-Authors: Jaroslav Smrž, Jaromir Mikes, Ĺubomír Kováč, Alena LukesovaAbstract:To date, only morphological and anatomical descriptions of microwhip scorpions (Arachnida: Palpigradi) have been published. This very rare group is enigmatic not only in its relationships to other arachnids, but especially due to the fact that these animals dwell only underground (in caves, soil, and interstitial spaces). We observed the curious feeding habit of the microwhip scorpion Eukoenenia spelaea over the course of one year in Ardovska Cave, located in Slovakia's Karst region. We chose histology as our methodology in studying 17 specimens and based it upon Masson's triple staining, fluorescent light and confocal microscopy. Single-celled cyanobacteria (blue-green algae) were conspicuously predominant in the gut of all studied palpigrades. Digestibility of the consumed cyanobacteria was supported by the presence of guanine crystals, glycogen deposits and haemocytes inside the palpigrade body. Cyanobacteria, the oldest cellular organisms on Earth, are very resistant to severe conditions in caves, including even darkness. Therefore, the cyanobacteria are able to survive in dark caves as nearly heterotrophic organisms and are consumed by cave palpigrades. Such feeding habit is extraordinary within the almost wholly predacious orders of the class Arachnida, and particularly so due to the type of food observed.
Jason A. Dunlop - One of the best experts on this subject based on the ideXlab platform.
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Arachnids in Bitterfeld amber: A unique fauna of fossils from the heart of Europe or simply old friends?
Pensoft Publishers, 2018Co-Authors: Jason A. Dunlop, Ulrich Kotthoff, Jörg U. Hammel, Jennifer Ahrens, Danilo HarmsAbstract:Bitterfeld amber, sometimes referred to as Saxon or Saxonian amber, is a potentially significant but poorly known source of arthropod data for the Palaeogene of northern Europe. An important aspect is a long-standing controversy about the age of this amber: namely whether it is equivalent to, and perhaps merely a southerly extension of, the better-known Baltic amber, or whether it is a unique and geological younger deposit sampling a different fauna. Here, we briefly review the Bitterfeld arachnids with particular emphasis on how these data could be used to elucidate the age of this deposit. Five arachnid orders have been recorded from Bitterfeld amber: spiders (Araneae), acariform mites (Acariformes), parasitiform mites (Parasitiformes), harvestmen (Opiliones) and pseudoscorpions (Pseudoscorpiones). This is a lower diversity than Baltic amber, where scorpions (Scorpiones) and camel spiders (Solifugae) have also been recorded. Spiders are the most comprehensively studied group, with more than 75 described species. Other groups such as pseudoscorpions and mites appear to be very diverse, but are virtually undescribed. Morphological overlap is apparent in the arachnid fauna and 40 species are currently shared between Baltic and Bitterfeld amber whilst 50 species are unique to the Bitterfeld deposit. At the family level overlap is even higher, but in all groups Baltic amber appears more diverse than Bitterfeld. This overlap may be interpreted as evidence for temporal conspecifity of the Baltic and Bitterfeld ambers, albeit with the Bitterfeld and Baltic ambers possibly representing independent localities within a larger Eocene European amber area which also included the Rovno amber from the Ukraine. However, caution should be exercised because the taxonomic foundation for such assumptions is far from comprehensive, most of the material remains to be studied in detail using modern techniques of morphological reconstruction. There are further issues with date estimates because some arachnid groups show extraordinary morphological stasis over time, even at species level, which may bias the analyses available. Here, we review the available knowledge on Bitterfeld arachnids and discuss how a detailed assessment of this fauna, and other arthropod taxa, could be generated. Several natural history museums – including Hamburg and Berlin – as well as private collectors host major assemblages of Bitterfeld fossils which may help to clarify the debate about the age and provenance of the material, and the extent to which (morpho)-species were maintained both over geographical distances and potentially geological time
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A fossil whip-scorpion (Arachnida: Thelyphonida) from the Upper Carboniferous of the Carnic Alps (Friuli, NE Italy)
Rivista Italiana Di Paleontologia E Stratigrafia, 2016Co-Authors: Paul A. Selden, Jason A. Dunlop, Luca SimonettoAbstract:A new and well-preserved fossil whip scorpion (Arachnida: Uropygi: Thelyphonida) is described from the Late Carboniferous of the Carnic Alps, Friuli, Italy. It is referred to Parageralinura marsiglioi n. sp. The new specimen is the first Carboniferous arachnid to be described from mainland Italy and is possibly the youngest Palaeozoic thelyphonid.
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Sperm carriers in Silurian sea scorpions
Naturwissenschaften, 2011Co-Authors: Carsten Kamenz, Andreas Staude, Jason A. DunlopAbstract:Invasion of the land by arachnids required adaptations of numerous organs, such as gills evolving into lungs, as well as mechanisms facilitating sperm transfer in a terrestrial environment. Many modern arachnids use spermatophores for this purpose, i.e. sperm transmitters detached from the body. Exceptionally preserved Silurian (423 Ma) fossils of Eurypterus tetragonophthalmus Fischer, 1839 (Chelicerata: Eurypterida) preserve so-called ‘horn organs’ which we here demonstrate as being equivalent to the spermatophore-producing parts of the genital tract in certain modern arachnids. This clarifies a long-running debate about sexing eurypterids based on the shape of the median abdominal (or genital) appendage. To our knowledge this is also the oldest direct evidence for spermatophore-mediated sperm transfer in the fossil record and suggests that eurypterids had evolved mating techniques using spermatophores as early as the Silurian, a valuable prerequisite for life on land. Spermatophores are absent in sea spiders (Pycnogonida) and horseshoe crabs (Xiphosura); thus the shared presence of sclerotized sperm-transfer devices in eurypterids and arachnids is a novel character, newly elucidated here, which offers explicit support for (Eurypterida + Arachnida). For this clade the name Sclerophorata n. nov. is proposed. Arachnida can be further defined by fusion of the originally paired genital opening.
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Geological history and phylogeny of Chelicerata.
Arthropod structure & development, 2010Co-Authors: Jason A. DunlopAbstract:Chelicerata probably appeared during the Cambrian period. Their precise origins remain unclear, but may lie among the so-called great appendage arthropods. By the late Cambrian there is evidence for both Pycnogonida and Euchelicerata. Relationships between the principal euchelicerate lineages are unresolved, but Xiphosura, Eurypterida and Chasmataspidida (the last two extinct), are all known as body fossils from the Ordovician. The fourth group, Arachnida, was found monophyletic in most recent studies. Arachnids are known unequivocally from the Silurian (a putative Ordovician mite remains controversial), and the balance of evidence favours a common, terrestrial ancestor. Recent work recognises four principal arachnid clades: Stethostomata, Haplocnemata, Acaromorpha and Pantetrapulmonata, of which the pantetrapulmonates (spiders and their relatives) are probably the most robust grouping. Stethostomata includes Scorpiones (Silurian-Recent) and Opiliones (Devonian-Recent), while Haplocnemata includes Pseudoscorpiones (Devonian-Recent) and Solifugae (Carboniferous-Recent). Recent works increasingly favour diphyletic mite origins, whereby Acaromorpha comprises Actinotrichida (Devonian-Recent), Anactinotrichida (Cretaceous-Recent) and Ricinulei (Carboniferous-Recent). The positions of the Phalangiotarbida (Devonian-Permian) and Palpigradi (Neogene-Recent) are poorly resolved. Finally, Pantetrapulmonata includes the following groups (listed here in their most widely recovered phylogenetic sequence): Trigonotarbida (Silurian-Permian), Uraraneida (Devonian-Permian), Araneae (Carboniferous-Recent), Haptopoda (Carboniferous), Amblypygi (?Devonian-Recent), Thelyphonida (Carboniferous-Recent) and Schizomida (Paleogene-Recent).
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A fossil trigonotarbid arachnid with a ricinuleid-like pedipalpal claw
Zoomorphology, 2009Co-Authors: Jason A. Dunlop, Carsten Kamenz, Giovanni TalaricoAbstract:A fossil trigonotarbid (Arachnida: Trigonotarbida) assigned to Palaeocharinus sp. from the Early Devonian ( c . 410 Ma) Rhynie cherts of Aberdeenshire, Scotland, UK is described, specifically for a previously unrecognised feature of the distal end of the pedipalp. This exhibits a small chela formed from a movable, unpaired apotele articulating against a slightly shorter, fixed projection from the inferior surface of the tarsus. Among other arachnids, this morphology has only previously been observed in the rare and enigmatic Carboniferous—Recent taxon Ricinulei. This character offers explicit support for a monophylum (Trigonotarbida + Ricinulei); a hypothesis previously based on opisthosomal characters which (if correct) would draw Ricinulei within the so-called Pantetrapulmonata clade (i.e. spiders and their closest relatives). Our data is not consistent with Ricinulei’s traditional position as sister-group of mites (Acari) and is an example of how considering extinct taxa may help to resolve the position of a ‘difficult’ living group.