Address: Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity at the Humboldt University Berlin, Invalidenstrasse 43, D-10115 Berlin, Germany
Fossil arachnids date back more than 400 million years to the Silurian period, making them one of... more Fossil arachnids date back more than 400 million years to the Silurian period, making them one of the first animal groups to appear in terrestrial ecosystems. However, despite significant progress in the discovery and description of fossil arachnids in recent years there has been no basic overview of the arachnid fossil record published in a broadly accessible format. Previous studies were largely published in specialist academic journals or monograph series. There has never been a book dedicated to the fossils of this often maligned and neglected group of animals, suitable for students of biology or geology looking for basic palaeontological information; or for arachnologists interested in what their favourite animals were like millions of years ago. In this book we provide up-to-date background information on what the arachnids are, their relationships to one another, and how fossils may be relevant to these debates. For each individual order – spiders, scorpions, harvestmen, etc. – we have summarized their typical characteristics. This should help with the identification of newly discovered fossils from both amber and rock deposits. Related to this, we have provided generous high quality photographic illustrations which highlight the appearance and diversity of these animals, the different ways in which they can become fossilized, and their sometimes astonishing quality of preservation. Modern imaging techniques in particular are revealing specimens in new and exciting ways. We discuss the probable ecology of the extinct species based on the interpretation of their morphology and the behaviour of their living relatives. We also highlight important palaeontological studies, and offer an extensive repertoire of further reading. We hope this book will be of interest both to arachnologists and palaeontologists, and that it will serve as a reference work for biologists interested in the evolutionary history of spiders and their many relatives.
Three species of fossil smaridid mites (Parasitengona, Smarididae) are described based on postlar... more Three species of fossil smaridid mites (Parasitengona, Smarididae) are described based on postlarval forms. Fessonia wunderlichi sp. nov., Fessonia grabenhorsti sp. nov. and Fessonia groehni sp. nov. were discovered in Eocene (ca. 44–50 Ma) Baltic amber and the former two also in the probably younger Oligocene (ca. 23–25 Ma) Bitterfeld amber. Although Smarididae has been reported from Baltic (and other) Cenozoic amber localities, these are the first species from this group to be formally described. Fessonia wunderlichi sp. nov. and Fessonia grabenhorsti sp. nov. are also the first named mites from Bitterfeld amber, and at the same time the first examples of mite species common to both the Baltic and Bitterfeld deposits. Fessonia groehni sp. nov. is known from a single specimen originating from the Baltic deposit. Although the thickened hindlimbs seen in Fessonia wunderlichi sp. nov. resemble the modern smaridid genus Kraussiana Southcott, 1961, the structure of the crista metopica implies another extant genus – Fessonia von Heyden, 1826 – which we redefine here to accommodate the morphology of the amber species. An outline of previous findings of terrestrial Parasitengona in amber is also provided.
The purpose of this application, under Article 23.9.5 of the Code, is to conserve the specific na... more The purpose of this application, under Article 23.9.5 of the Code, is to conserve the specific name Phrynus mexicanus Poinar & Brown, 2004 which was proposed for a fossil whip spider (Arachnida, Amblypygi, PHRYNIDAE) preserved in Chiapas (Mexican) amber. This fossil species was initially described under the grammatically erroneous combination Phrynus mexicana. However, correcting the species epithet to mexicanus renders it a junior primary homonym of a Recent Mexican whip spider, Phrynus mexicanus Bilimek, 1867. The Recent species was subsequently transferred to the genus Paraphrynus Moreno, 1940, to give the current combination Paraphyrnus mexicanus (Bilimek, 1867), and neither species has ever been considered congeneric with the other.
The fossil whip spider Electrophrynus mirus Petrunkevitch 1971 (Arachnida: Amblypygi: Electrophry... more The fossil whip spider Electrophrynus mirus Petrunkevitch 1971 (Arachnida: Amblypygi: Electrophrynidae) from the Mexican Chiapas amber (Miocene: ca 15–20 Ma) is redescribed. Its original referral to an extinct family and genus based on an unusual configuration of the pedipalp is shown to be based on a misinterpretation. The Family Electrophrynidae has already been synonymized with Phrynidae. Although E. mirus closely resembles modern phrynids, a key diagnostic character relating to the dorsal pattern of spines on the pedipalp Patella is not preserved. Since this incomplete fossil cannot be meaningfully compared either to modern genera or to other whip spiders in Neotropical amber, we believe the pragmatic solution is to treat E. mirus as a nomen dubium.
The first camel spider (Arachnida, Solifugae) from the Upper Cretaceous (lowermost Cenomanian, ca... more The first camel spider (Arachnida, Solifugae) from the Upper Cretaceous (lowermost Cenomanian, ca. 99 Ma) Burmese amber from Myanmar is described as Cushingia ellenbergeri gen. et sp. nov. It represents one of only a handful of fossils belonging to this arachnid order, but ist precise systematic affinities are difficult to resolve. It shares characters with Karschiidae, the subfamily Gylippinae among the Gylippidae and the subfamily Dinorhaxinae containing a single, monotypic genus Dinorhax Simon, 1879 in the Family Melanoblossiidae; the latter genus occurring in South-East Asia today. Its general habitus is closest to Dinorhax, but differences between the fossil and this modern genus remain. On balance, the uncertain nature of some features precludes unequivocal referral to any one of the families above. We prefer to place this new genus as Solifugae incertae sedis and further discuss the wider biogeographical implications of this find. Our new fossil is also significant for coming from a presumed forest habitat, whereas most camel spiders today are associated with arid environments.
Six species of spider (Arachnida: Araneae) are newly recorded from a series of localities in the ... more Six species of spider (Arachnida: Araneae) are newly recorded from a series of localities in the eastern part of the Republic of the Sudan. Crossopriza pristina (Simon, 1890) (Pholcidae) was found at Tokar and New Halfa. These represent the first African records of this species. Hippasa cinerea Simon, 1898 (Lycosidae) was found at New Halfa, Pardosa oncka Lawrence, 1927 (Lycosidae) and Wadicosa fidelis (O. Pickard-Cambridge, 1872) (Lycosidae) were both found in the Dinder National Park. Plexippoides flavescens (O. Pickard-Cambridge, 1872) (Salticidae) was found at New Halfa, and Pseudicius spiniger (O. Pickard-Cambridge, 1872) (Salticidae) – already known from South Sudan – is here reported from the Dinder National Park in the Republic. Additionally, we offer several new locality records for five species previously documented from the Republic of the Sudan. Afrofilistata fradei (Berland & Millot, 1940) (Filistatidae) was found at New Halfa, Ocyale pilosa (Roewer, 1960) (Lycosidae) was found in the Dinder National Park, Pardosa injucunda (O. Pickard-Cambridge, 1876) (Lycosidae) was found at New Halfa, Kassala, and in the Dinder National Park, Thomisus daradioides Simon, 1890 (Thomisidae) at Khashm El-Gerba and at New Halfa, and Plexippus paykulli (Audouin, 1825) (Salticidae) at New Halfa, Kassala, and in the Dinder National Park.
One of the oldest known fossil scorpions, a new species from the mid-Silurian Eramosa Formation (... more One of the oldest known fossil scorpions, a new species from the mid-Silurian Eramosa Formation (430 myr) of Ontario, Canada, exhibits several surprising features. The depositional environment and associated biota indicate a marine habitat; however, the leg morphology of this scorpion, which has a short tarsus in common with all Recent scorpions, suggests that a key adaptation for terrestrial locomotion, the ability to support its weight on a subterminal ‘foot’, appeared remarkably early in the scorpion fossil record. Specimens are preserved intact and undisturbed in a splayed posture typical of moults rather than carcasses. We postulate that these animals were aquatic, but occasionally ventured into extremely shallow water, or onto a transient subaerially exposed surface while moulting, before returning to deeperwater. Shed exuviae were preserved in situ by rapid overgrowth of bacterial biofilm.
A new specimen of Lacinius Thorell, 1876; (Opiliones: Phalangiidae) from Eocene Baltic amber is d... more A new specimen of Lacinius Thorell, 1876; (Opiliones: Phalangiidae) from Eocene Baltic amber is described. We interpret it as conspecific with a slightly younger record from the German Bitterfeld amber, originally referred to as the extant species L. erinaceus Starega, 1966. Our new specimen reveals pedipalpal apophyses on both the patella and the tibia, features which we can now confirm in the Bitterfeld fossil too. This unique character combination for the genus justifies a new, extinct species: Lacinius bizleyi sp. nov. The Baltic amber inclusion dates to ca. 44–49 Ma, and is thus the oldest putative example of Lacinius in the fossil record. It is a further example of an arachnid species shared between Baltic and Bitterfeld amber.
Arachnids are an important group of arthropods. They are diverse and abundant; a major constituen... more Arachnids are an important group of arthropods. They are diverse and abundant; a major constituent of many terrestrial ecosystems; and possess a deep and extensive fossil record. In recent years a number of exceptionally preserved arachnid fossils have been investigated using tomography and associated techniques, providing valuable insights into their morphology. Here we use X-ray microtomography to reconstruct members of two extinct arachnid orders. In the Haptopoda, we demonstrate the presence of ‘clasp-knife’ chelicerae, and our novel redescription of a member of the Phalangiotarbida highlights leg details, but fails to resolve chelicerae in the Group due to their small size. As a result of these reconstructions, tomographic studies of three-dimensionally preserved fossils now exist for three of the four extinct orders, and for fossil representatives of several extant ones. Such studies constitute a valuable source of high fidelity data for constructing phylogenies. To illustrate this, here we present a cladistic analysis of the chelicerates to accompany these reconstructions. This is based on a previously published matrix, expanded to include fossil taxa and relevant characters, and allows us to: cladistically place the extinct arachnid orders; explicitly test some earlier hypotheses from the literature; and demonstrate that the addition of fossils to phylogenetic analyses can have broad implications. Phylogenies based on chelicerate morphology—in contrast to molecular studies—have achieved elements of consensus in recent years. Our work suggests that these results are not robust to the addition of novel characters or fossil taxa. Hypotheses surrounding chelicerate phylogeny remain in a state of flux.
Herein, Amblyomma tenellum Koch, 1844 (Ixodidae) is reestablished as a valid tick name and remove... more Herein, Amblyomma tenellum Koch, 1844 (Ixodidae) is reestablished as a valid tick name and removed from the synonymy list of Amblyomma cajennense (Fabricius, 1787), while Amblyomma imitator Kohls, 1958, is relegated to a junior synonym of A. tenellum. Amblyomma tenellum is redescribed based on the examination of male type specimens collected by Deppe at the beginning of the 19th century in Mexico and described by Koch in 1844.
Fossil mesostigmatid mites (Acari: Parasitiformes:Mesostigmata) are extremely rare, and specimens... more Fossil mesostigmatid mites (Acari: Parasitiformes:Mesostigmata) are extremely rare, and specimens from only nine families, including four named species, have been described so far. A new record of Myrmozercon sp. described here from Eocene (ca 44–49 Myr) Baltic amber represents the first—and so far only—fossil example of the derived, extant family Laelapidae. Significantly, modern species of this genus are habitually myrmecophilous and the fossil mite described here is preserved attached to the head of the dolichoderine ant Ctenobethylus goepperti (Mayr, 1868). It thus offers the oldest unequivocal evidence for an ecological association between mesostigmatid mites and social insects in the order Hymenoptera.
Trigonotarbids are a group of Palaeozoic arachnids, best known from Coal Measures sites in Europe... more Trigonotarbids are a group of Palaeozoic arachnids, best known from Coal Measures sites in Europe and North America. Here, we review previous work on their higher systematics and redescribe Trigonotarbus johnsoni Pocock, 1911, from the Coseley Lagerstätte, UK, on the basis of both hand specimens and X-ray microtomography reconstruction. This is a historically important species – being the eponymous taxon for the entire order – but lacks recent or comprehensive description. We also present the first cladistic analysis of Trigonotarbida, based on a novel matrix of 49 characters, 20 trigonotarbids, and four outgroups. Two families, Archaeomartidae and Lissomartidae, were only represented by a single exemplar. At least two exemplars were scored to test monophyly for other families. Anthracomartidae, Anthracosironidae, and Eophrynidae were all recovered as monophyletic, but Palaeocharinidae, Trigonotarbidae, Aphantomartidae, and Kreischeriidae were not. In a wider context, our data support a (Trigonotarbida + Ricinulei) clade; albeit based on a limited set of outgroups. Relationships within Trigonotarbida were not so well resolved, although two previously anticipated clades were consistently recovered in the Agreement subtrees with strong support: (1) (Palaeocharinus (Archaeomartidae + Anthracomartidae)) and (2) Lissomartus as sister group to a so called ‘eophrynid assemblage’ (Aphantomartus (Alkenia (Pseudokreischeria (Kreischeria (Eophrynus + Pleophrynus))))).
New finds of Late Palaeozoic arachnids, based on three well-preserved carapaces from the Carbonif... more New finds of Late Palaeozoic arachnids, based on three well-preserved carapaces from the Carboniferous of Russia and Ukraine and one complete, albeit poorly preserved, specimen from the Permian of Kazakhstan, are described. The spider genus Arthrolycosa is reported from the Late Carboniferous (Late Pennsylvanian: Kasimovian–Gzhelian) of Chunya in the Tunguska Basin of Siberia; it is the first find of a spider outside the Carboniferous tropics. Another fossil assigned to the same genus comes from the Late Carboniferous (Early Pennsylvanian: Bashkirian) of Kamensk–Shakhtinsky in the Donets Basin of Russia; it is probably the oldest fossil spider known. A thelyphonid(whip scorpion) carapace is described from the Late Carboniferous (Late Pennsylvanian: Kasimovian) of the adjacent Lugansk Province of the Donets Basin of Ukraine.
A fossil opilioacarid mite (Parasitiformes: Opilioacarida) in Burmese amber is described as ?Opil... more A fossil opilioacarid mite (Parasitiformes: Opilioacarida) in Burmese amber is described as ?Opilioacarus groehni sp. nov. This ca. 99 Ma record (Upper Cretaceous: Cenomanian) represents only the third fossil example of this putatively basal mite lineage, the others originating from Eocene Baltic amber (ca. 44–49 Ma). Our new record is not only the oldest record of Opilioacarida, but it is also one of the oldest examples of the entire Parasitiformes clade. The presence of Opilioacarida—potentially Opiloacarus—in the Cretaceous of SE Asia suggests that some modern genus groups were formerly more widely distributed across the northern hemisphere, raising questions about previously suggested Gondwanan origins for these mites.
This paper serves two roles. First, it acts as an introduction to Blender, an open-source compute... more This paper serves two roles. First, it acts as an introduction to Blender, an open-source computer graphics program, which can be of utility to paleontologists. To lessen the software’s otherwise steep learning curve, a step-by-step guide to create an idealized reconstruction of a fossil in the form of a three-dimensional model in Blender, or to use the software to render results from ‘virtual paleontology’ techniques, is provided as an online supplemental data file. Second, here we demonstrate the use of Blender with a case study on the extinct trigonotarbid arachnids. We report the limb articulations of members of the Devonian genus Palaeocharinus on the basis of exceptionally preserved fossils from the Rhynie Cherts of Scotland. We use these newly reported articulations to create a Blender model, and draw comparisons with the gait of extant arachnids to produce as accurate a representation of the trigonotarbid flexing its limbs and walking as possible, presented in additional online supplemental data files. Knowledge of the limb articulations of trigonotarbid arachnids also allows us to discuss their functional morphology: trigonotarbids’ limbs and gait were likely comparable to extant cursorial spiders, but lacked some innovations seen in more derived arachnids.
A new eophrynid trigonotarbid (Arachnida: Trigonotarbida: Eophrynidae) from the Pennsylvanian (Ka... more A new eophrynid trigonotarbid (Arachnida: Trigonotarbida: Eophrynidae) from the Pennsylvanian (Kasimovian) Astrasado Formation of the Kinney Brick Quarry, New Mexico is described. This fossil – the first arachnid to be recorded from the Astrasado Formation – is preserved primarily as a dorsal opisthosoma. Pleophrynus hawesi sp. nov. diagnostically preserves evidence for three pairs of posterior opisthosomal spines rather than the two usually seen in other eophrynids. A comparison of opisthosomal tuberculation patterns among the best known eophrynid species is included. At ca. 304.0–306.5 Ma, our new taxon represents one of the youngest records of Eophrynidae, while the Kinney Brick Quarry is only the twelfth site in North America to yield trigonotarbid arachnids; compared to more than 60 such localities in Europe.
A provisional checklist of spiders (Arachnida: Araneae) documented from Sudan – here differentiat... more A provisional checklist of spiders (Arachnida: Araneae) documented from Sudan – here differentiated into the Republic of Sudan and South Sudan – is compiled based on published records in the literature. Collectors of historical note include Carl Jickeli, Antonín Stecker, Louis Vossion, Maurice Potter, Stanley Flower, Harold King, Eberhard Fraas, Franz Werner, Leonard Jägerskiöld, Willoughby Lowe, Hubert Lynes, Mary Steele, James Macdonald, Pierre Benoit, John Cloudsley-Thompson and John Lewis. 113 species from 28 families have been documented from the Republic of Sudan, and 59 species in 18 families from South Sudan. Nineteen species are common to both countries, yielding a total of 153 species in 30 families for Sudan in its traditional sense. The most diverse families (with more than three species recorded) are Salticidae, Araneidae, Thomisidae, Lycosidae, Sparassidae, Philodromidae, Oxyopidae, Theridiidae and Gnaphosidae. Published records largely derive from Khartoum State, Red Sea State, South Kurdufan State and North Kurdufan State (all with 10 or more species records) in the Republic of Sudan, and from Bahr El Jabal [Central Equatoria] and Upper Nile State (with 38 and 26 species respectively) from South Sudan.
An exceptionally preserved specimen of the extinct trigonotarbid arachnid Eophrynus prestvicii (A... more An exceptionally preserved specimen of the extinct trigonotarbid arachnid Eophrynus prestvicii (Arachnida: Trigonotarbida) from the Late Carboniferous (Duckmantian) British Middle Coal Measures of the UK is redescribed with the help of X-ray micro-tomography (XMT). Previous work is summarised and the correct spelling of the species Name confirmed. Reconstruction of the void left by the original specimen within a siderite (ironstone) nodule reveals ist three-dimensional structure as well as novel details which are difficult to resolve using traditional methods of study. The pedipalps and legs can now be reconstructed almost in their entirety and we can confirm that the chelicerae hung beneath the prosoma with a backwards-directed fang. Opisthosomal segmentation is clarified: a narrow tergite 1 is followed by eight further tergites (2–9), clearly demarked by their ornamentation. In general, a much more accurate picture of the spines and tuberculation ornamenting the dorsal surface of the body was resolved. Ventrally the opisthosoma bears so-called ventral sacs and terminates in a two-segmented pygidium. Based on the XMT model, a new reconstruction of E. prestvicii is presented. Although phylogenetically unrelated, the sometimes similar-looking, spiny and tuberculate lanitorid harvestmen (Opiliones: Lanitores) offer a possible ecological model for these highly ornamented trigonotarbids.
An annotated catalogue of the rare mesothele spiders (Araneae: Mesothelae: Liphistiidae) held in ... more An annotated catalogue of the rare mesothele spiders (Araneae: Mesothelae: Liphistiidae) held in the Museum für Naturkunde Berlin is presented. The museum hosts non-type specimens of nine species representing all three currently recognised genera, namely: Liphistius desultor Schiödte, 1849, L. malayanus cameroni Haupt, 1983, L. cf. thaleban Schwendinger, 1990, Heptathela kikuyai Ono, 1998, H. kimurai (Kishida, 1920), H. yanbaruensis Haupt, 1983, Ryuthela ishigakiensis Haupt, 1983, R. nishihirai (Haupt, 1979) and R. tanikawai Ono, 1997 (spec. reval.). The geographical focus of this collection is Malaysia and Japan, and most of the material was collected by the Berlin-based zoologist Joachim Haupt.
Fossil arachnids date back more than 400 million years to the Silurian period, making them one of... more Fossil arachnids date back more than 400 million years to the Silurian period, making them one of the first animal groups to appear in terrestrial ecosystems. However, despite significant progress in the discovery and description of fossil arachnids in recent years there has been no basic overview of the arachnid fossil record published in a broadly accessible format. Previous studies were largely published in specialist academic journals or monograph series. There has never been a book dedicated to the fossils of this often maligned and neglected group of animals, suitable for students of biology or geology looking for basic palaeontological information; or for arachnologists interested in what their favourite animals were like millions of years ago. In this book we provide up-to-date background information on what the arachnids are, their relationships to one another, and how fossils may be relevant to these debates. For each individual order – spiders, scorpions, harvestmen, etc. – we have summarized their typical characteristics. This should help with the identification of newly discovered fossils from both amber and rock deposits. Related to this, we have provided generous high quality photographic illustrations which highlight the appearance and diversity of these animals, the different ways in which they can become fossilized, and their sometimes astonishing quality of preservation. Modern imaging techniques in particular are revealing specimens in new and exciting ways. We discuss the probable ecology of the extinct species based on the interpretation of their morphology and the behaviour of their living relatives. We also highlight important palaeontological studies, and offer an extensive repertoire of further reading. We hope this book will be of interest both to arachnologists and palaeontologists, and that it will serve as a reference work for biologists interested in the evolutionary history of spiders and their many relatives.
Three species of fossil smaridid mites (Parasitengona, Smarididae) are described based on postlar... more Three species of fossil smaridid mites (Parasitengona, Smarididae) are described based on postlarval forms. Fessonia wunderlichi sp. nov., Fessonia grabenhorsti sp. nov. and Fessonia groehni sp. nov. were discovered in Eocene (ca. 44–50 Ma) Baltic amber and the former two also in the probably younger Oligocene (ca. 23–25 Ma) Bitterfeld amber. Although Smarididae has been reported from Baltic (and other) Cenozoic amber localities, these are the first species from this group to be formally described. Fessonia wunderlichi sp. nov. and Fessonia grabenhorsti sp. nov. are also the first named mites from Bitterfeld amber, and at the same time the first examples of mite species common to both the Baltic and Bitterfeld deposits. Fessonia groehni sp. nov. is known from a single specimen originating from the Baltic deposit. Although the thickened hindlimbs seen in Fessonia wunderlichi sp. nov. resemble the modern smaridid genus Kraussiana Southcott, 1961, the structure of the crista metopica implies another extant genus – Fessonia von Heyden, 1826 – which we redefine here to accommodate the morphology of the amber species. An outline of previous findings of terrestrial Parasitengona in amber is also provided.
The purpose of this application, under Article 23.9.5 of the Code, is to conserve the specific na... more The purpose of this application, under Article 23.9.5 of the Code, is to conserve the specific name Phrynus mexicanus Poinar & Brown, 2004 which was proposed for a fossil whip spider (Arachnida, Amblypygi, PHRYNIDAE) preserved in Chiapas (Mexican) amber. This fossil species was initially described under the grammatically erroneous combination Phrynus mexicana. However, correcting the species epithet to mexicanus renders it a junior primary homonym of a Recent Mexican whip spider, Phrynus mexicanus Bilimek, 1867. The Recent species was subsequently transferred to the genus Paraphrynus Moreno, 1940, to give the current combination Paraphyrnus mexicanus (Bilimek, 1867), and neither species has ever been considered congeneric with the other.
The fossil whip spider Electrophrynus mirus Petrunkevitch 1971 (Arachnida: Amblypygi: Electrophry... more The fossil whip spider Electrophrynus mirus Petrunkevitch 1971 (Arachnida: Amblypygi: Electrophrynidae) from the Mexican Chiapas amber (Miocene: ca 15–20 Ma) is redescribed. Its original referral to an extinct family and genus based on an unusual configuration of the pedipalp is shown to be based on a misinterpretation. The Family Electrophrynidae has already been synonymized with Phrynidae. Although E. mirus closely resembles modern phrynids, a key diagnostic character relating to the dorsal pattern of spines on the pedipalp Patella is not preserved. Since this incomplete fossil cannot be meaningfully compared either to modern genera or to other whip spiders in Neotropical amber, we believe the pragmatic solution is to treat E. mirus as a nomen dubium.
The first camel spider (Arachnida, Solifugae) from the Upper Cretaceous (lowermost Cenomanian, ca... more The first camel spider (Arachnida, Solifugae) from the Upper Cretaceous (lowermost Cenomanian, ca. 99 Ma) Burmese amber from Myanmar is described as Cushingia ellenbergeri gen. et sp. nov. It represents one of only a handful of fossils belonging to this arachnid order, but ist precise systematic affinities are difficult to resolve. It shares characters with Karschiidae, the subfamily Gylippinae among the Gylippidae and the subfamily Dinorhaxinae containing a single, monotypic genus Dinorhax Simon, 1879 in the Family Melanoblossiidae; the latter genus occurring in South-East Asia today. Its general habitus is closest to Dinorhax, but differences between the fossil and this modern genus remain. On balance, the uncertain nature of some features precludes unequivocal referral to any one of the families above. We prefer to place this new genus as Solifugae incertae sedis and further discuss the wider biogeographical implications of this find. Our new fossil is also significant for coming from a presumed forest habitat, whereas most camel spiders today are associated with arid environments.
Six species of spider (Arachnida: Araneae) are newly recorded from a series of localities in the ... more Six species of spider (Arachnida: Araneae) are newly recorded from a series of localities in the eastern part of the Republic of the Sudan. Crossopriza pristina (Simon, 1890) (Pholcidae) was found at Tokar and New Halfa. These represent the first African records of this species. Hippasa cinerea Simon, 1898 (Lycosidae) was found at New Halfa, Pardosa oncka Lawrence, 1927 (Lycosidae) and Wadicosa fidelis (O. Pickard-Cambridge, 1872) (Lycosidae) were both found in the Dinder National Park. Plexippoides flavescens (O. Pickard-Cambridge, 1872) (Salticidae) was found at New Halfa, and Pseudicius spiniger (O. Pickard-Cambridge, 1872) (Salticidae) – already known from South Sudan – is here reported from the Dinder National Park in the Republic. Additionally, we offer several new locality records for five species previously documented from the Republic of the Sudan. Afrofilistata fradei (Berland & Millot, 1940) (Filistatidae) was found at New Halfa, Ocyale pilosa (Roewer, 1960) (Lycosidae) was found in the Dinder National Park, Pardosa injucunda (O. Pickard-Cambridge, 1876) (Lycosidae) was found at New Halfa, Kassala, and in the Dinder National Park, Thomisus daradioides Simon, 1890 (Thomisidae) at Khashm El-Gerba and at New Halfa, and Plexippus paykulli (Audouin, 1825) (Salticidae) at New Halfa, Kassala, and in the Dinder National Park.
One of the oldest known fossil scorpions, a new species from the mid-Silurian Eramosa Formation (... more One of the oldest known fossil scorpions, a new species from the mid-Silurian Eramosa Formation (430 myr) of Ontario, Canada, exhibits several surprising features. The depositional environment and associated biota indicate a marine habitat; however, the leg morphology of this scorpion, which has a short tarsus in common with all Recent scorpions, suggests that a key adaptation for terrestrial locomotion, the ability to support its weight on a subterminal ‘foot’, appeared remarkably early in the scorpion fossil record. Specimens are preserved intact and undisturbed in a splayed posture typical of moults rather than carcasses. We postulate that these animals were aquatic, but occasionally ventured into extremely shallow water, or onto a transient subaerially exposed surface while moulting, before returning to deeperwater. Shed exuviae were preserved in situ by rapid overgrowth of bacterial biofilm.
A new specimen of Lacinius Thorell, 1876; (Opiliones: Phalangiidae) from Eocene Baltic amber is d... more A new specimen of Lacinius Thorell, 1876; (Opiliones: Phalangiidae) from Eocene Baltic amber is described. We interpret it as conspecific with a slightly younger record from the German Bitterfeld amber, originally referred to as the extant species L. erinaceus Starega, 1966. Our new specimen reveals pedipalpal apophyses on both the patella and the tibia, features which we can now confirm in the Bitterfeld fossil too. This unique character combination for the genus justifies a new, extinct species: Lacinius bizleyi sp. nov. The Baltic amber inclusion dates to ca. 44–49 Ma, and is thus the oldest putative example of Lacinius in the fossil record. It is a further example of an arachnid species shared between Baltic and Bitterfeld amber.
Arachnids are an important group of arthropods. They are diverse and abundant; a major constituen... more Arachnids are an important group of arthropods. They are diverse and abundant; a major constituent of many terrestrial ecosystems; and possess a deep and extensive fossil record. In recent years a number of exceptionally preserved arachnid fossils have been investigated using tomography and associated techniques, providing valuable insights into their morphology. Here we use X-ray microtomography to reconstruct members of two extinct arachnid orders. In the Haptopoda, we demonstrate the presence of ‘clasp-knife’ chelicerae, and our novel redescription of a member of the Phalangiotarbida highlights leg details, but fails to resolve chelicerae in the Group due to their small size. As a result of these reconstructions, tomographic studies of three-dimensionally preserved fossils now exist for three of the four extinct orders, and for fossil representatives of several extant ones. Such studies constitute a valuable source of high fidelity data for constructing phylogenies. To illustrate this, here we present a cladistic analysis of the chelicerates to accompany these reconstructions. This is based on a previously published matrix, expanded to include fossil taxa and relevant characters, and allows us to: cladistically place the extinct arachnid orders; explicitly test some earlier hypotheses from the literature; and demonstrate that the addition of fossils to phylogenetic analyses can have broad implications. Phylogenies based on chelicerate morphology—in contrast to molecular studies—have achieved elements of consensus in recent years. Our work suggests that these results are not robust to the addition of novel characters or fossil taxa. Hypotheses surrounding chelicerate phylogeny remain in a state of flux.
Herein, Amblyomma tenellum Koch, 1844 (Ixodidae) is reestablished as a valid tick name and remove... more Herein, Amblyomma tenellum Koch, 1844 (Ixodidae) is reestablished as a valid tick name and removed from the synonymy list of Amblyomma cajennense (Fabricius, 1787), while Amblyomma imitator Kohls, 1958, is relegated to a junior synonym of A. tenellum. Amblyomma tenellum is redescribed based on the examination of male type specimens collected by Deppe at the beginning of the 19th century in Mexico and described by Koch in 1844.
Fossil mesostigmatid mites (Acari: Parasitiformes:Mesostigmata) are extremely rare, and specimens... more Fossil mesostigmatid mites (Acari: Parasitiformes:Mesostigmata) are extremely rare, and specimens from only nine families, including four named species, have been described so far. A new record of Myrmozercon sp. described here from Eocene (ca 44–49 Myr) Baltic amber represents the first—and so far only—fossil example of the derived, extant family Laelapidae. Significantly, modern species of this genus are habitually myrmecophilous and the fossil mite described here is preserved attached to the head of the dolichoderine ant Ctenobethylus goepperti (Mayr, 1868). It thus offers the oldest unequivocal evidence for an ecological association between mesostigmatid mites and social insects in the order Hymenoptera.
Trigonotarbids are a group of Palaeozoic arachnids, best known from Coal Measures sites in Europe... more Trigonotarbids are a group of Palaeozoic arachnids, best known from Coal Measures sites in Europe and North America. Here, we review previous work on their higher systematics and redescribe Trigonotarbus johnsoni Pocock, 1911, from the Coseley Lagerstätte, UK, on the basis of both hand specimens and X-ray microtomography reconstruction. This is a historically important species – being the eponymous taxon for the entire order – but lacks recent or comprehensive description. We also present the first cladistic analysis of Trigonotarbida, based on a novel matrix of 49 characters, 20 trigonotarbids, and four outgroups. Two families, Archaeomartidae and Lissomartidae, were only represented by a single exemplar. At least two exemplars were scored to test monophyly for other families. Anthracomartidae, Anthracosironidae, and Eophrynidae were all recovered as monophyletic, but Palaeocharinidae, Trigonotarbidae, Aphantomartidae, and Kreischeriidae were not. In a wider context, our data support a (Trigonotarbida + Ricinulei) clade; albeit based on a limited set of outgroups. Relationships within Trigonotarbida were not so well resolved, although two previously anticipated clades were consistently recovered in the Agreement subtrees with strong support: (1) (Palaeocharinus (Archaeomartidae + Anthracomartidae)) and (2) Lissomartus as sister group to a so called ‘eophrynid assemblage’ (Aphantomartus (Alkenia (Pseudokreischeria (Kreischeria (Eophrynus + Pleophrynus))))).
New finds of Late Palaeozoic arachnids, based on three well-preserved carapaces from the Carbonif... more New finds of Late Palaeozoic arachnids, based on three well-preserved carapaces from the Carboniferous of Russia and Ukraine and one complete, albeit poorly preserved, specimen from the Permian of Kazakhstan, are described. The spider genus Arthrolycosa is reported from the Late Carboniferous (Late Pennsylvanian: Kasimovian–Gzhelian) of Chunya in the Tunguska Basin of Siberia; it is the first find of a spider outside the Carboniferous tropics. Another fossil assigned to the same genus comes from the Late Carboniferous (Early Pennsylvanian: Bashkirian) of Kamensk–Shakhtinsky in the Donets Basin of Russia; it is probably the oldest fossil spider known. A thelyphonid(whip scorpion) carapace is described from the Late Carboniferous (Late Pennsylvanian: Kasimovian) of the adjacent Lugansk Province of the Donets Basin of Ukraine.
A fossil opilioacarid mite (Parasitiformes: Opilioacarida) in Burmese amber is described as ?Opil... more A fossil opilioacarid mite (Parasitiformes: Opilioacarida) in Burmese amber is described as ?Opilioacarus groehni sp. nov. This ca. 99 Ma record (Upper Cretaceous: Cenomanian) represents only the third fossil example of this putatively basal mite lineage, the others originating from Eocene Baltic amber (ca. 44–49 Ma). Our new record is not only the oldest record of Opilioacarida, but it is also one of the oldest examples of the entire Parasitiformes clade. The presence of Opilioacarida—potentially Opiloacarus—in the Cretaceous of SE Asia suggests that some modern genus groups were formerly more widely distributed across the northern hemisphere, raising questions about previously suggested Gondwanan origins for these mites.
This paper serves two roles. First, it acts as an introduction to Blender, an open-source compute... more This paper serves two roles. First, it acts as an introduction to Blender, an open-source computer graphics program, which can be of utility to paleontologists. To lessen the software’s otherwise steep learning curve, a step-by-step guide to create an idealized reconstruction of a fossil in the form of a three-dimensional model in Blender, or to use the software to render results from ‘virtual paleontology’ techniques, is provided as an online supplemental data file. Second, here we demonstrate the use of Blender with a case study on the extinct trigonotarbid arachnids. We report the limb articulations of members of the Devonian genus Palaeocharinus on the basis of exceptionally preserved fossils from the Rhynie Cherts of Scotland. We use these newly reported articulations to create a Blender model, and draw comparisons with the gait of extant arachnids to produce as accurate a representation of the trigonotarbid flexing its limbs and walking as possible, presented in additional online supplemental data files. Knowledge of the limb articulations of trigonotarbid arachnids also allows us to discuss their functional morphology: trigonotarbids’ limbs and gait were likely comparable to extant cursorial spiders, but lacked some innovations seen in more derived arachnids.
A new eophrynid trigonotarbid (Arachnida: Trigonotarbida: Eophrynidae) from the Pennsylvanian (Ka... more A new eophrynid trigonotarbid (Arachnida: Trigonotarbida: Eophrynidae) from the Pennsylvanian (Kasimovian) Astrasado Formation of the Kinney Brick Quarry, New Mexico is described. This fossil – the first arachnid to be recorded from the Astrasado Formation – is preserved primarily as a dorsal opisthosoma. Pleophrynus hawesi sp. nov. diagnostically preserves evidence for three pairs of posterior opisthosomal spines rather than the two usually seen in other eophrynids. A comparison of opisthosomal tuberculation patterns among the best known eophrynid species is included. At ca. 304.0–306.5 Ma, our new taxon represents one of the youngest records of Eophrynidae, while the Kinney Brick Quarry is only the twelfth site in North America to yield trigonotarbid arachnids; compared to more than 60 such localities in Europe.
A provisional checklist of spiders (Arachnida: Araneae) documented from Sudan – here differentiat... more A provisional checklist of spiders (Arachnida: Araneae) documented from Sudan – here differentiated into the Republic of Sudan and South Sudan – is compiled based on published records in the literature. Collectors of historical note include Carl Jickeli, Antonín Stecker, Louis Vossion, Maurice Potter, Stanley Flower, Harold King, Eberhard Fraas, Franz Werner, Leonard Jägerskiöld, Willoughby Lowe, Hubert Lynes, Mary Steele, James Macdonald, Pierre Benoit, John Cloudsley-Thompson and John Lewis. 113 species from 28 families have been documented from the Republic of Sudan, and 59 species in 18 families from South Sudan. Nineteen species are common to both countries, yielding a total of 153 species in 30 families for Sudan in its traditional sense. The most diverse families (with more than three species recorded) are Salticidae, Araneidae, Thomisidae, Lycosidae, Sparassidae, Philodromidae, Oxyopidae, Theridiidae and Gnaphosidae. Published records largely derive from Khartoum State, Red Sea State, South Kurdufan State and North Kurdufan State (all with 10 or more species records) in the Republic of Sudan, and from Bahr El Jabal [Central Equatoria] and Upper Nile State (with 38 and 26 species respectively) from South Sudan.
An exceptionally preserved specimen of the extinct trigonotarbid arachnid Eophrynus prestvicii (A... more An exceptionally preserved specimen of the extinct trigonotarbid arachnid Eophrynus prestvicii (Arachnida: Trigonotarbida) from the Late Carboniferous (Duckmantian) British Middle Coal Measures of the UK is redescribed with the help of X-ray micro-tomography (XMT). Previous work is summarised and the correct spelling of the species Name confirmed. Reconstruction of the void left by the original specimen within a siderite (ironstone) nodule reveals ist three-dimensional structure as well as novel details which are difficult to resolve using traditional methods of study. The pedipalps and legs can now be reconstructed almost in their entirety and we can confirm that the chelicerae hung beneath the prosoma with a backwards-directed fang. Opisthosomal segmentation is clarified: a narrow tergite 1 is followed by eight further tergites (2–9), clearly demarked by their ornamentation. In general, a much more accurate picture of the spines and tuberculation ornamenting the dorsal surface of the body was resolved. Ventrally the opisthosoma bears so-called ventral sacs and terminates in a two-segmented pygidium. Based on the XMT model, a new reconstruction of E. prestvicii is presented. Although phylogenetically unrelated, the sometimes similar-looking, spiny and tuberculate lanitorid harvestmen (Opiliones: Lanitores) offer a possible ecological model for these highly ornamented trigonotarbids.
An annotated catalogue of the rare mesothele spiders (Araneae: Mesothelae: Liphistiidae) held in ... more An annotated catalogue of the rare mesothele spiders (Araneae: Mesothelae: Liphistiidae) held in the Museum für Naturkunde Berlin is presented. The museum hosts non-type specimens of nine species representing all three currently recognised genera, namely: Liphistius desultor Schiödte, 1849, L. malayanus cameroni Haupt, 1983, L. cf. thaleban Schwendinger, 1990, Heptathela kikuyai Ono, 1998, H. kimurai (Kishida, 1920), H. yanbaruensis Haupt, 1983, Ryuthela ishigakiensis Haupt, 1983, R. nishihirai (Haupt, 1979) and R. tanikawai Ono, 1997 (spec. reval.). The geographical focus of this collection is Malaysia and Japan, and most of the material was collected by the Berlin-based zoologist Joachim Haupt.
Raubmilben sind häufige Vertreter der Spinnentiere, die im Boden oder Höhlensystemen leben. Als F... more Raubmilben sind häufige Vertreter der Spinnentiere, die im Boden oder Höhlensystemen leben. Als Fossilien sind sie mit weltweit nur vier beschriebenen Arten extrem selten. Ein überraschender Fund aus dem Siegener Bergland, der in einem Nachlass aufgetaucht ist, gehört zu einer Raubmilbe vermutlich aus der Familie Parasitidae und ist in Aragonit konserviert. Leider ist eine genauere Altersbestimmung kaum möglich. Wahrschienlich ist die Raubmilbe subfossil und wurde vom nassen Kalksinter eingeschlossen.
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