A new genus and species of arachnid (Chelicerata: Arachnida), Douglassarachne acanthopoda n. gen. n. sp., is described from the late Carboniferous (Moscovian) Coal Measures of the Mazon Creek Lagerstätte, Illinois, USA. This is a unique animal with distinctive large spines on the legs. It has a subovate body, a segmented opisthosoma, and a terminal anal tubercle. The legs are robust and appear to have been similar in construction throughout the limb series, with heavy spination of the preserved proximal podomeres. The mouthparts and coxo-sternal region are equivocal. The preserved character combination does not permit easy referral to any known arachnid order, living or extinct, thus the new fossil in placed as Arachnida/Pantetrapulmonata incertae sedis. It contributes to an emerging pattern of disparate body plans among late Carboniferous arachnids, ranging from anatomically modern members of living orders through to extinct taxa, such as the present fossil, whose phylogenetic position remains unresolved.

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Ticks are composed of 3 extant families (Argasidae, Ixodidae and Nuttalliellidae) and 2 extinct families (Deinocrotonidae and Khimairidae). The Nuttalliellidae possess one extant species (Nuttalliella namaqua) limited to the Afrotropic region. A basal relationship to the hard and soft tick families and its limited distribution suggested an origin for ticks in the Afrotropics. The Deinocrotonidae has been found in Burmese amber from Myanmar and Iberian amber from Spain, suggesting a wider distribution of the lineage composed of Deinocrotonidae and Nuttalliellidae. The current study describes 8 fossils from mid-Cretaceous (ca. 100 Ma) Burmese amber: 2 Deinocroton species (Deinocroton bicornis sp. nov.; Deinocroton lacrimus sp. nov.), 5 Nuttalliella species (Nuttalliella gratae sp. nov., Nuttalliella tuberculata sp. nov., Nuttalliella placaventrala sp. nov., Nuttalliella odyssea sp. nov., Nuttalliella tropicasylvae sp. nov.) and a new genus and species (Legionaris nov. gen., Legionaris robustus sp. nov.). The argument is advanced that Deinocroton do not warrant its own family, but forms part of the Nuttalliellidae comprising 3 genera, Deinocroton, Legionaris nov. gen. and Nuttalliella). Affinities of Burmese tick fossils to the Australasian region, specifically related to rifting of the Burma terrane from northern Australia ~150 million years ago, suggest that Nuttalliella had a much wider distribution than its current limited distribution. The distribution of Nuttalliella likely stretched from Africa over Antarctica and much of Australia, suggesting that extant members of this family may still be found in Australia. Considerations for the geographic origins of ticks conclude that an Afrotropic origin can as yet not be discarded.

Three examples of metastriate hard ticks (Ixodida: Ixodidae) with apparent affinities to modern Australasian genera are described from the mid-Cretaceous (ca. 100 Ma) Burmese amber of Myanmar. Two nymphs of Bothriocroton muelleri sp. nov. represent the oldest (and only) fossil record of this genus, living members of which are restricted to Australia and predominantly feed on monitor lizards, snakes and echidnas. A female of Archaeocroton kaufmani sp. nov. shares its basis capitulum shape with the tuatara tick Archaeocroton sphenodonti (Dumbleton, 1943), the only extant member of this genus and an endemic species for New Zealand. The presence of 2 Australasian genera in Burmese amber is consistent with a previous record of an Ixodes Latreille, 1795 tick from this deposit which resembles Australian members of this genus. They further support an emerging hypothesis that fauna of the amber forest, which may have been on an island at the time of deposition, was at least partly Gondwanan in origin. A revised evolutionary tree for Ixodida is presented compiling data from several new Burmese amber ticks described in the last few years.

Four fossil ticks (Arachnida: Parasitiformes: Ixodida) are described from mid-Cretaceous (ca. 100 Ma) Burmese amber of Myanmar. Ixodes antiquorum sp. nov. (Ixodidae) is the first Mesozoic record of Ixodes and the oldest representative of the most species-rich extant tick genus. Its affinities appear to lie with modern Australian forms, consistent with the hypothesis that Burmese amber hosted Gondwanan faunal elements. Even more remarkable is Khimaira fossus gen. et sp. nov. which combines a body resembling that of a soft tick (Argasidae) with a basis capitulum more like that of a hard tick (Ixodidae). We refer it to Khimairidae fam. nov. as a possible transitional form between the two main families of ticks alive today. Another member of the extinct Deinocrotonidae is described as Deinocroton copia sp. nov., while the first described adult female for Cornupalpatum burmanicum is associated with a dinosaur feather barb.

The first fossil potentially assignable to the extant hard tick genus Haemaphysalis CL Koch (1844) (Ixodida: Ixodidae) is described from the Late Cretaceous (ca. 99 Ma) Burmese amber of Myanmar. Haemaphysalis (Alloceraea) cretacea sp. nov. is the oldest and only fossil representative of this genus; living members of which predominantly feed on mammals. Their typical hosts are known since at least the Jurassic and the discovery of a mid-Cretaceous parasite, which might have fed on mammals raises again the question of to what extent ticks are coupled to their (modern) host groups. An inferred Triassic split of Argasidae (soft ticks) into the bird-preferring Argasinae and mammal-preferring Ornithodorinae dates to about the time when dinosaurs (later including birds) and mammaliaforms as potential hosts were emerging. Ixodidae may have split into Prostriata and Metastriata shortly after the end-Permian mass extinction, an event which fundamentally altered the terrestrial vertebrate fauna. Prostriata (the genus Ixodes) prefer birds and mammals today, and some may have used groups like cynodonts in the Triassic. Basal metastriate ticks (e.g. Amblyomma) prefer reptiles, but derived metastriates (including Haemaphysalis) again prefer mammals. Here, we may be looking at a younger (Cretaceous?) shift associated with more recent mammalian radiations.

Amblyomma birmitum sp. nov. is formally described as a new record from 99 Ma old Burmese amber from Myanmar. This confirms the presence of the extant hard tick genus Amblyomma C.L. Koch, 1844 (Ixodida: Ixodidae) in the Late Cretaceous. This discovery is placed in its wider context and some reports of fossil hard ticks, such as a Hyalomma C.L. Koch, 1844 in Eocene Baltic amber, are misidentifications. The genus Amblyomma belongs to the clade Metastriata, a group which probably also accommodates two extinct genera, Cornupalpatum Poinar and Brown, 2003 and Compluriscutata Poinar and Buckley, 2008, also found in Burmese amber. All three fossils are thus only a little younger than published molecular divergence time estimates (ca. 124 ± 17 Ma) for the Metastriata lineage. Amblyomma has a largely Gondwanan distribution today. However, in some biogeographical scenarios, e.g. the Samafrica model, its predicted radiation time postdates the dissolution of the original Gondwana supercontinent raising questions about how its current distribution pattern was achieved.

A relatively diverse chelicerate fauna has been detected in Early Devonian, Siegenian strata of the Westerwald area, Rhineland-Palatinate, Germany. The arachnids, comprising trigonotarbids and the oldest phalangiotarbids, are described and figured here along with the chasmataspidids. To accomodate the phalangiotarbid a new genus and species in the family Architarbidae, Devonotarbus hombachensis, is raised. Devonotarbus n. gen. is characterized by an approximately straight posterior carapace margin, abbreviated and undivided anterior tergites, a large sixth tergite, and fused posterior tergites. The chasmataspidid closely resembles Diploaspis casteri from the Emsian assemblage of Alken an der Mosel, but is readily discernible as a new species, D. muelleri, by a strong tuberculation of its dorsal integument. With only fragmentary opisthosomal remains available, the trigonotarbids cannot be placed in known taxa with any certainty at this time.

The whip scorpions (Arachnida: Uropygi) from the Pennsylvanian Coal Measures of Mazon Creek, Illinois, USA are restudied. Four Mazon Creek uropygid specimens have previously been described and a fifth is added here. Contrary to earlier reports, none of the specimens lack eyes and the sternum is similarly shaped in all the specimens. Consequently, the two younger names, Geralinura similis Petrunkevitch, 1913 and G. gigantea Petrunkevitch, 1913 are here treated as junior synonyms of G. carbonaria Scudder, 1884. The morphology of the pedipalps is more like those of some modern amblypygids, being spiniferous and non-chelate, suggesting the subchelate palps found in all Recent whip scorpions is an apomorphy not yet incorporated into the bodyplan of G. carbonaria. Indeed, subchelate palps also seem to be absent in other species of this age. For completeness, we summarise here the six Pennsylvanian whip scorpions currently recognised—including sketch reconstructions—with comments on further differences between the Paleozoic and living taxa. Geralinura Scudder, 1884 is reserved for G. carbonaria and the British species G. britannica Pocock, 1911. Prothelyphonus Friĉ, 1904 is reinstated for the Czech species P. bohemicus (Kuŝta, 1884). Parageralinura gen. nov. is proposed for the widely overlooked Dutch species P. neerlandica (Laurentiaux-Vieria and Laurentiaux, 1961) and the German species P. naufraga (Brauckmann and Koch, 1983). All the Coal Measures whip scorpion genera are treated here as plesion taxa, basal to the Thelyphonidae, which can be defined as whip scorpions with a fully subchelate pedipalp, the chelae formed from large and distinct patellar and tibial apophyses.

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.

The Late Carboniferous whip spider Graeophonus anglicus Pocock, 1911 (Arachnida: Amblypygi), is redescribed on the basis of the holotype and nine other specimens all preserved in sideritic nodules from the British Middle Coal Measures of Coseley, Staffordshire, UK. This species is clearly basal with respect to most living whip spiders, expressing numerous plesiomorphic character states and can be referred to both the suborder Paleoamblypygi and the ‘living fossil’ family Paracharontidae (with one Recent species), the latter based on an explict character of dorsal spination on the pedipalp femur. This suggests that crown-group Amblypygi originated by at least the mid-Palaeozoic.

A harvestman (Arachnida: Opiliones) is described from the Early Devonian (Pragian) Rhynie cherts, Aberdeenshire, Scotland. Eophalangium sheari gen. et sp. nov. is the oldest known harvestman. The material includes both males and a female preserving, respectively, a cuticle-lined penis and ovipositor within the opisthosoma. Both these structures are of essentially modern appearance. The Rhynie fossils also show tracheae which are, again, very similar to those of living harvestmen. This is the oldest unequivocal record of arachnid tracheal respiration and indicates that E. sheari was terrestrial. An annulate, setose ovipositor in the female suggests that it can be excluded from the clades Dyspnoi and Laniatores, in which the ovipositor lacks such annulations. However, the penis shows evidence of two muscles, a feature of uncertain polarity seen in modern Troguloidea (Dyspnoi). The presence of median eyes and long legs excludes Cyphophthalmi, and thus, E. sheari is tentatively referred to the suborder Eupnoi. Therefore, this remarkable material is implicitly a crown-group harvestman and is one of the oldest known crown-group chelicerates. It also suggests an extraordinary degree of morphological stasis within the eupnoid line, with the Devonian forms differing little in gross morphology – and perhaps in reproductive behaviour – from their modern counterp

The type material of Chasmataspis laurencii Caster & Brooks, 1956 (Chelicerata: Chasmataspidida) from the Middle Ordovician (Tremadoc to Caradoc) of Sevier County, Tennessee, USA, is redescribed, and comparisons are drawn with recently discovered Devonian chasmataspids from Scotland, Germany and Russia. The depositional setting of the C. laurencii fossils is reinterpreted as an ash fall into shallow marine/tidal sediments. Chasmataspis laurencii confirms the presence of 13 opisthosomal segments in Chasmataspidida, a character of unresolved polarity. The phylogenetic position ofC. laurencii is difficult to resolve. A monophyletic Chasmataspidida has one convincing autapomorphy in the nine-segmented postabdomen, but C. laurencii shares a number of characters with xiphosurans (i.e. cardiac lobe, pre-abdomen with axial region, biramous and chelate limbs), while the Devonian taxa more closely resemble eurypterids (i.e. through pediform limbs and a genital appendage). Earlier interpretations of the respiratory system in C. laurencii appear unconvincing in the light of new evidence from Devonian forms with opisthosomal opercula. Resting impressions of a Chasmataspis-like animal from the Upper Cambrian Hickory Sandstone of Texas, USA, also appear to show evidence of opercula. These Texan fossils could represent the oldest record of Euchelicerata. Two chasmataspid families are recognised, and C. laurencii is placed in the monotypic Chasmataspididae Caster & Brooks, 1956, redefined here on the fused pre-abdominal buckler with an axial region and the narrow, elongate post-abdomen ending in a long, lanceolate telson.

The first fossil opilioacariform mite (Acari: Opilioacariformes), Paracarus pristinus sp. n., is described from Baltic amber (Tertiary: Eocene). This well-preserved inclusion has six eyes and, thus, can be excluded from the derived, four-eyed genera. Based on characters such as the extremely long leg 4, it is assigned to the extant central Asian genus Paracarus Chamberlin & Mulaik, 1942. As with many Baltic amber inclusions, we lack an exact provenance for the type specimen, but the Baltic region is further north than the present day range of Recent opilioacariform species and suggests that the range of this group was once more extensive within Eurasia. The first solifuge (Solifugae) from Baltic amber is described as Palaeoblossia groehni gen. et sp. n. Based on its size and general appearance, it can be assigned to Daesiidae, and again represents an arachnid group which previously had a much wider geographical range. If adult, it may represent the smallest solifuge known.

Study of abundant phalangiotarbid (Arachnida: Phalangiotarbida) material – provisionally assigned here to Bornatarbus mayasii (Haupt in Nindel 1955) – from the Upper Carboniferous of Writhlington, UK, has revealed new information about some previously equivocal characters. The present authors report a trifurcate apotele, possible spiracles on sternite 5, and confirm the presence of 10 opisthosomal tergites plus a dorsal anal operculum. The affinities of phalangiotarbids are obscure, with most authors favouring affinities with Opiliones (harvestmen) and/or Acari (mites and ticks). Phalangiotarbida is scored for characters used in previous studies of arachnid relationships. A cladistic analysis based on 63 characters using 13 terminal arachnid taxa (plus a hypothetical outgroup), resolves Phalangiotarbida as sister group to (Palpigradi + Tetrapulmonata): the taxon Megoperculata sensu Shultz (1990). Even under cladistic analysis, the position of the Phalangiotarbida remains hard to resolve, but a prosomal sternite with distinct sclerites potentially groups them with the Megoperculata.

The first whipspider (Arachnida: Amblypygi) and three new examples of whipscorpions (Arachnida: Uropygi: Thelyphonida) are described from Lower Cretaceous (Aptian) strata of the Crato Formation, Ceará State, Brazil. The whipspider is described as Britopygus weygoldti gen. et sp. nov. It resembles members of the extant family Charinidae, but cannot be unequivocally placed. Two of the whipscorpions are referred to Mesoproctus rowlandi Dunlop, 1998 and one could be a subadult, since, like juveniles of extant species, it has a relatively broad anterior sternum. There are differences between these two specimens in pedipalp and flagellum morphology, but both characters are variable within extant species. The third and largest whipscorpion is incomplete and referred to Mesoproctus sp. Its carapace length is equal to or exceeds that of the largest known extant genus Mastigoproctus Pocock, 1894.

Plesiosiro madeleyi Pocock, 1911, the monotypic representative of the fossil order Haptopoda Pocock, 1911 from the British Middle Coal Measures (Upper Carboniferous, Westphalian B) of Coseley, Staffordshire, is redescribed. A specimen from Sparth Bottoms, Lancashire is not a Plesiosiro and is referred to Arthropoda incertae sedis. Plesiosiro was probably a predator, having tooth-like projections on the femora for prey capture, and shows adaptations of the limbs for crawling in narrow spaces. The uncertain nature of the chelicerae, eyes and respiratory organs makes resolving the phylogenetic position of Plesiosiro difficult, though Haptopoda deserves its status as a separate order. Plesiosiro resembles both cythophthalmid and trogulid opilionids. Its broad prosoma–opisthosoma junction could be interpreted as a synapomorphy for Plesiosiro + Opiliones, but this character is not unique to these orders. Alternatively, Plesiosiro has a large, ventral opisthosomal sclerite. This is interpreted here as a genital operculum, a synapomorphy for Tetrapulmonata, and Plesiosiro is tentatively referred to this clade. Based on the synapomorphies of a divided sternum and subdivided telotarsi a relationship of the form: (Plesiosiro (Amblypygi (Thelyphonida + Schizomida))) is proposed.

A new chasmataspid (Chelicerata, Chasmataspida) is described as Forfarella mitchelli gen. et sp. nov. from the Lower Devonian of Scotland. This species is problematic in that it was catalogued under the name Forfarella mitchelli Waterston (no date), but was never formally described. Furthermore, its type locality is uncertain. Chasmataspids are a poorly known taxon within Chelicerata which appear to be closely related to eurypterids. F. mitchelli is only the fourth chasmataspid species to be described. Though poorly preserved, it shows affinities with other Devonian chasmataspids and supports the hypothesis that this group may have been more geographically widespread in the Devonian than has previously been recognised.

Four new specimens of phalangiotarbid (Arachnida: Phalangiotarbida) from the Upper Carboniferous (upper Westphalian A) of Westhoughton, Lancashire, UK, are referred to Mesotarbus peteri sp. nov. An additional Lancashire phalangiotarbid, Phalangiotarbus subovalis (Woodward, 1872), from the Upper Carboniferous (lower/middle Westphalian A) of Burnley, is redescribed and designated the neotype of this species. This material allows new interpretations of the opisthosomal segmentation and respiratory organs of phalangiotarbids, and a reconstruction of Mesotarbus peteri is presented.