Garwood et al.

BMC Evolutionary Biology (2017) 17:105
DOI 10.1186/s12862-017-0931-1


The phylogeny of fossil whip spiders
Russell J. Garwood1,2*, Jason A. Dunlop3, Brian J. Knecht4 and Thomas A. Hegna4

Background: Arachnids are a highly successful group of land-dwelling arthropods. They are major contributors to
modern terrestrial ecosystems, and have a deep evolutionary history. Whip spiders (Arachnida, Amblypygi), are one
of the smaller arachnid orders with ca. 190 living species. Here we restudy one of the oldest fossil representatives of
the group, Graeophonus anglicus Pocock, 1911 from the Late Carboniferous (Duckmantian, ca. 315 Ma) British Middle
Coal Measures of the West Midlands, UK. Using X-ray microtomography, our principal aim was to resolve details
of the limbs and mouthparts which would allow us to test whether this fossil belongs in the extant, relict family
Paracharontidae; represented today by a single, blind species Paracharon caecus Hansen, 1921.
Results: Tomography reveals several novel and significant character states for G. anglicus; most notably in the
chelicerae, pedipalps and walking legs. These allowed it to be scored into a phylogenetic analysis together with
the recently described Paracharonopsis cambayensis Engel & Grimaldi, 2014 from the Eocene (ca. 52 Ma) Cambay
amber, and Kronocharon prendinii Engel & Grimaldi, 2014 from Cretaceous (ca. 99 Ma) Burmese amber. We
recovered relationships of the form ((Graeophonus (Paracharonopsis + Paracharon)) + (Charinus (Stygophrynus
(Kronocharon (Charon (Musicodamon + Paraphrynus)))))). This tree largely reflects Peter Weygoldt’s 1996 classification
with its basic split into Paleoamblypygi and Euamblypygi lineages; we were able to score several of his characters for
the first time in fossils. Our analysis draws into question the monophyly of the family Charontidae.
Conclusions: Our data suggest that Graeophonus is a crown group amblypygid, and falls within a monophyletic
Paleoamblypgi clade, but outside the family Paracharontidae (= Paracharonopsis + Paracharon). Our results also suggest
a new placement for the Burmese amber genus Kronocharon, a node further down from its original position. Overall,
we offer a broad phylogenetic framework for both the fossil and Recent whip spiders against which future discoveries
can be tested.
Keywords: Amblypygi, Coal Measures, Amber, Fossil, Systematics, Pennsylvanian

Background defined by a ground pattern of two pairs of book lungs
Whip spiders (Arachnida: Amblypygi) are distinctive [1, 2]. The majority of recent analyses—molecular [3] and
creatures (Fig. 1) with a long, slender, antenniform first morphological [1]—suggest whip spiders are members of
pair of legs. These whip-like appendages give the group the Pedipalpi clade (Amblypygi, Uropygi and Schizomida),
its name, although they are occasionally referred to as although there has been historical discussion; see e.g.
tailless whip scorpions because they also resemble a re- Shultz [4]. Due to their highly modified sensory first pair
lated group of arachnids, the whip scorpions (Uropygi), of legs, whip spiders have to walk hexapodally using legs
albeit without the whip scorpion’s flagelliform telson. II–IV. They also possess a characteristic flattened body,
Both whip spiders and whip scorpions belong—together allowing the animals to crawl into narrow spaces under
with spiders (Araneae) and schizomids (Schizomida)—to rocks or tree bark, and they have spined, subchelate,
the arachnid clade Tetrapulmonata. This grouping is raptorial pedipalps to grasp and immobilise prey. Today,
the group has a tropical to subtropical distribution, with
around 190 extant species in five families. A detailed
* Correspondence:
School of Earth and Environmental Sciences, The University of Manchester,
overview of their biology and systematics can be found
Manchester M13 9PL, UK in Weygoldt [5]. A full species catalogue was offered by
Department of Earth Sciences, The Natural History Museum, Cromwell Road, Harvey ([6]; updated online as [7]) and further pub-
London SW7 5BD, UK
Full list of author information is available at the end of the article
lished species counts can be found in Prendini [8].

© The Author(s). 2017 Open Access This article is distributed under the terms of the Creative Commons Attribution 4.0
International License (, which permits unrestricted use, distribution, and
reproduction in any medium, provided you give appropriate credit to the original author(s) and the source, provide a link to
the Creative Commons license, and indicate if changes were made. The Creative Commons Public Domain Dedication waiver
( applies to the data made available in this article, unless otherwise stated.

scale bar equals 1 mm (redrawn from ref [34].5 mm (redrawn from ref [34]. g Damon variegatus ventral left palp trochanter. Figure 3). Ch—chelicerae. showing a prominent anterior ventral apophysis. BMC Evolutionary Biology (2017) 17:105 Page 2 of 14 Fig. drawn from an unnumbered specimen in the Museum für Naturkunde.Garwood et al. scale bar equals 2 mm (redrawn from ref [34]. scale bar equals 1 mm (redrawn from ref [34]. Dt—distitarsus. Figure 8). ventral anterior apophysis reduced to a spine. Figure 1). Abbreviations: 1–4—legs 1–4. scale bar equals 0. Ap—apotele. Tr—trochanter. b The chelicera of Paracharon caecus in lateral view. Ta—tarsus. Va—palpal ventral anterior apophysis. Pa—patella. Bti—basitibia. Figure 12).—a species typically 24–35 mm in length when fully grown—as seen in dorsal view. T1-4—cheliceral teeth 1–4 numbered from ventral to dorsal. Bt—basitarsus. and terminology used herein. f Ventral left trochanter of Charinus montanus palp. Vs—palpal ventral anterior spine . Pp—pedipalps.5 mm (redrawn from ref [34]. Fe—femur. the distalmost bicuspate. numbers proximally to distally. scale bar equals 0. Berlin. Figure 15). d Paracharon caecus palp in dorsal view. with only two patellar spines. Dti—distitibia. a Gross morphology of the extant amblypygid Damon sp. 1 Guide to the morphology of Amblypygids. c The chelicera of Charon grayi with three cheliceral teeth. Figure 7). e Pedipalp of Charon grayi which has three palpal spines forming a catching basket. S1-3—spines 1–3. Ti—tibia. scale bar equals 0.5 mm (redrawn from ref [34]. a species with four cheliceral teeth.

Approaches vary. In the same publication (Fig. The species is known from several well. Scudder later reinterpreted this find as a often surprisingly good preservation. focussing on the British Middle Coal Measures Pocock Both these harvestman studies were aided by the [13] described Graeophonus anglicus Pocock. [14]. stem groups and microtomography Whip spiders are rare as fossils. 2014. Petrunkevitch’s amber species was fossils. Tetrophthalmi [26. The next amblypygids found in the fossil record are For the present study. especially useful in revealing distal details of appendages. particularly in sider- whip spider and created a new genus Graeophonus ite nodules. 390 Ma) cuticle fragments named Ecchosis (i.Garwood et al. Coal Measures arachnids are it as a dragonfly larva. As previously noted. BMC Evolutionary Biology (2017) 17:105 Page 3 of 14 Fossil whip spiders Crown groups. Krono. 2a). 305 Ma) of Europe and North America. harvestman (Opiliones) has been successful in demon- narius. species descended from the most recent common pulchribothrium Selden & Shear. 1911 whi. 27]. These confirmed its commented on the similarity between G. 2014 from the ca. using the matrix of Weygoldt [34]. A full overview of historical work on Palaeozoic difficult to resolve using traditional methods of light taxa is provided by Dunlop et al. found in West African termite nests. Based on a re-examination of Pocock’s of Mexico [21. Unequivocal ies estimating dates of cladogenesis. their great antiquity—more than 300 million years—and see also [12]). Scudder [10] de. and Dunlop & Barov by Petrunkevitch [33] in the living (derived. 1890 to accommodate it as Graeophonus carbo. application of X-ray microtomography: a technique that ch—as the best preserved species—is the focus of the allows digital visualisation of the void left by the organism present study. and used as calibration points—so-called total- Figure 2. 1911. logical Nature Reserve known only from its ventral which are often buried deep in the host nodule. see below) [17] augmented this with further details of the sternal re. anglicus and the referral to the modern family Phrynidae. 315– created using node-dating priors for relaxed clocks [24]. it was first described by Pocock [13] and was later placed ing distinctive amblypygid pedipalps. anglicus to Paracharontidae. When studying fossils a key question is when the oldest dence for this group comprises some Middle Devonian crown-group representatives of a given clade first appear (Givetian: ca. ancestor of all extant members of that group). shape and the orientation and spination of its pedipalps. number of plesiomorphic traits. 2) for comparison with the fossils. which is thought to have a charon prendinii Engel & Grimaldi. microscopy [28–32]. recent molecular clock work on Scudder. & Martill [16] described a Coal Measures as a model organism. [18] documented a further Cretaceous species. The youngest records was described as blind. 115 Ma Crato whip spider Graeophonus anglicus from the British Middle Formation of Brazil—Dunlop. Weygoldt [5. includ. while at referred G. [14] went further and explicitly recently shown to be a nomen dubium [23]. 1 in the text) and interpreted evidence or tip-dating [25]. and recent developments have shown that fossils can be scribed an isolated opisthosoma from Cape Breton in coded directly into morphological matrices based on ex- Nova Scotia as Libellula carbonaria Scudder. species known from a ventral prosoma and limbs. Pocock [13] did strating that Carboniferous fossils which unequivocally re- not feel that Scudder’s two North American specimens solve as members of two modern suborders (Eupnoi and were conspecific and renamed the younger one Graeo. It larly high fidelity of preservation. Dunlop [15] described a Car. family Phrynichidae. present study was therefore to CT scan the fossils in the . This curious whip these authors described a whip spider from the Eocene spider. These chobothrium on the patella [9]—a character otherwise data provide calibration points for molecular clock stud- only seen in living Amblypygi today. as well as ornamentation such as spines or tubercles. 16 Ma) Dominican Republic amber are hints of lateral eye spots on the right side of the dorsal [19.e. which has a simi. Palaeozoic members of the clade include five species but time-calibrated phylogenies have traditionally been from the Late Carboniferous Coal Measures (ca. In a widely overlooked move. Engel & Grimaldi living whip spider species Paracharon caecus Hansen. 1876 (his tant taxa. Here. 20] and the probably contemporary Chiapas amber shield (Fig. It is a methodology that has proved preserved specimens [14]. More recently. 52 Ma) Cambay amber of India—Paracharonopsis similarities to the Coal Measures fossils in its carapace cambayensis Engel & Grimaldi. although in the photograph there are from Miocene (ca. The oldest potential evi. boniferous whip spider from the Writhlington Geo. solve sufficient features to test this placement cladistically guishable from a modern species of Phrynus Lamarck. Based on the discovery of a better particularly interesting for these purposes given both preserved specimen from Mazon Creek in Illinois ([11]. Dunlop et al. In the same monograph plans placed in a new suborder. 1921 (Paracharontidae). we selected the Carboniferous Cretaceous in age and come from the ca. Dyspnoi) lived alongside harvestmen with extinct body phonus scudderi Pocock. but could not re- least the Dominican amber material is barely distin. we offer the first 99 Ma Burmese amber from Myanmar. referred to as Fig. 22]. 1991 which have a tri. 34] gion and details of the walking limbs. and are surface. within the nodule. with high levels of photographs of type material belonging to this rare species morphological detail preserved. shows some (ca. For example. A principal aim of the 1801 (Phrynidae) found in the Caribbean.

The deposit is Late Carboniferous (Bashkirian/ important Carboniferous species—in particular from the Moscovian: Duckmantian) in age. During the scanning of this voids within siderite concretions.Garwood et al. which Pointon et al. f Original label.or a place at around 315 Ma. anglicus. Fossils were compared to Recent stem-group whip spider. using a Nikon HMX-ST 225 and tungsten reflec- In31257. 3) allowed us to productive site from which specimens were collected in the select one of the paratypes—NHM In31234—as the most late 1800s and early 1900s. London (NHMUK) We scanned all specimens at the Natural History Museum. Note the projecting anterior region of the carapace and the pedipalps which articulate up and down and have relatively weak spination. BMC Evolutionary Biology (2017) 17:105 Page 4 of 14 Fig. A primary objective was to Bashkirian/Moscovian boundary. a key criterion being well-preserved. Material Three specimens of Graeophonus anglicus. In carapace shape and details of the pedipalp this putatively plesiomorphic Recent species resembles the Carboniferous fossil Graeophonus anglicus Pocock. we also whip spiders in the collections of the Museum für Natur- chose to include all well-preserved fossil amblypygids in kunde Berlin. sometimes with partial specimen. Visualisations of the data (Fig. 2 Images of a syntype of the only living paleoamblypygid Paracharon caecus Hansen. Københavns Methods Universitet. or Westphalian B in appendages—and use this to explicitly test the phylogen. 1911 hope of yielding more details of the morphology of this kaolinite infill. c-d Close up of the pedipalp and its spination in dorsal and ventral view. were scanned Tomography from the Natural History Museum. 1921 (Paracharontidae) which lives in termites nests in Guinea Bissau (West Africa). As part of the study. traditional terminologies. scale bar equals 5 mm. 1921 (Fig. NHM In31248. a-b Entire animal in dorsal and ventral view. we collected 6284 projections at 195 kV/95 μA . and NHM London. The fossils—including these promising specimen. Comparative photographs of a syntype of the the same cladistic analysis to assess the phylogeny and rare living species Paracharon caecus Hansen. collections: NHM In31234. specimens—are typically preserved as three-dimensional three-dimensional pedipalps. e Enlarged lateral view of pedipalp in life position. a tion target. All originate from the Coseley Lagerstätte. Abbreviations as in Fig. 2) evolutionary history of the order. [35] demonstrate whether it is best considered a crown. were kindly provided by Jan Pedersen and Nikolaj Scharff from the Statens Naturhistoriske Museum. The Duckmantian straddles the etic position of G. 1.

3. BMC Evolutionary Biology (2017) 17:105 Page 5 of 14 Fig. the latter used in the current study. [36]. recovering instead a for whether the cheliceral serrula is more rounded—as in . Cladistic Taxa and Characters Added character 2). e A single chelicera. and polytomy with no reliable placement for fossil taxa. (Additional file 1). Instead we created using the SPIERS software suite. f-g Close up of the left pedipalp and its spination in dorsal and ventral view. scale bar equals 0. on taxon (after Weygoldt [34].5 mm copper filter.5 mm. tibial spination numbered. morphological terminology Character 38 reflects the number of cheliceral teeth within follows Weygoldt [5. [40]. We modelled and based on the matrix of Pepato et al. Character 54 further codes unable to reproduce these results.8 μm. redescription of the species. Abbreviations as in Fig. modified after Garwood & Dunlop [1]. 34]. apart from the pedipalps where the amblypygids. and character 39 whether the distal-most tooth is bicuspate ([34]. Digital visualisations were TNT. We created volumes using CT Pro.4 s. scale bar equals 1 mm. Models are provided as SI to the numerous characters. Character 53 reflects the presence of a In order to assess the phylogeny of the fossil whip spiders serrula. c-d Close up of the right pedipalp and its spination in dorsal and ventral view. dered in the open source ray tracer Blender. but were amblypygids and Schizomida. and likely to result from differences in the search strategies the 4MP (2000 × 2000) Perkin Elmer detector panel and consensus tree calculations between Hennig86 and provided a voxel size of 15. The are fully described in the current paper in the VAXML interchange format [38] supplementary character statements (Additional file 2). we added tially transparent in Fig. 1 over 360 degrees rotation. which is either four or three depending we use the scheme of Shultz [4]. character 1). patellar spination numbered. scale bar equals 5 mm. Garwood et al [39]. which in missing elements of the anatomy in this program following turn built on that of Giribet et al. [41]. showing four teeth. 1911. This is exposure of 1.Garwood et al. We used this model as the basis of our and we summarise relevant character additions here. inferred/reconstructed anatomy partially transparent. and the model was ren. following the chose to code fossil whip spiders into the matrix of methods of Garwood et al. 3 Tomographic reconstruction of Graeophonus anglicus Pocock. In order to obtain the methods of Garwood and Dunlop [37] through com. with a 0. and to fully parison with living species. modified setae found on the cheliceral free finger in we first re-ran the matrix of Weygoldt [34]. scale bar equals 1 mm. a-b Whole animal in dorsal and ventral view. and then rendered them par. encompass their morphological disparity. internal resolution within the amblypygids.

The modification in plane of motion is probably associated The former were coded from Weygoldt ([5. For all analyses. but palp. with the distal-most largest.000 replicates). Our searches comprised tree proximally) is coded as absent or present in character 73 bisection-reconnection [TBR] with 1000 replicates. 16 segments. frequencies. Stygophrynus [44] was coded on the suggestion spines ([34]. 20] which is essentially modern. TNT was used to create a ones is highly reduced ([34]. added the genera Charon.Garwood et al. Whilst this is largely only replicates). We further explored the matrix by likely to be preserved in fossils. and for the amblypygids. Inkscape. character 18). in contrast with other whip spiders. tarsus is subdivided or fused.0. and Paracharonopsis and Kronocharon from pedipalp tarsus is equivalent to Weygoldt’s distitarsus (the Engel & Grimaldi [18]. Hence we have coded lateral eyes as present herein. supports are included through symmetric resampling tion of Engel & Grimaldi [18]. 34]. for the amber fossils Paracharonopsis from the descrip. and in others (e.1. and distal part of this we term the descriptions unreliable. We analysed this matrix using TNT v. We also added the three Character 69 has been modified to reflect whether palpal known and reasonably complete fossil genera: Graeophonus. 25 segments. saving ([34]. For the implied weights analysis. change probability 33%. We omitted the three species two parts of which are fused within the Neoamblypygi described by Petrunkevitch [12] as we consider the species clade. character 14). is a more representative of every extant family—and to test the pos- vertical plane of motion (Figs. 2a).000 replicates) and Bremer informative for the living species. For our implied weights analyses. as it differs in some important aspects of the chanter has a distinctive ventral apophysis. character 4). it can also be coded support [48].1. dorsal row of patellar spines on the pedipalp which. Character 71 reflects the absence or presence of a Schawaller [19. character 11). three principal spines. TNT-file ready format. This segment was chosen as it is more from 0. and we provide jackknife ([46]. character 83 is 3. Paracharon and Stygophrynus. unmodified. BMC Evolutionary Biology (2017) 17:105 Page 6 of 14 the whip spiders and the two Protoschizomidae genera of The previously published arachnid phylogeny of Garwood the schizomids—or toothlike as seen in the schizomid et al [39] included three whip spider genera. which using a traditional search and unordered multistate charac- is coded in character 72 ([34]. Character 76 strict consensus tree which was exported as an SVG into codes for the presence of spines on the palpal tarsus. In order to include a Graeophonus. ([45] . Morphological interpretation These are listed in the character statements supplementary Much of the basic somatic morphology in Graeophonus file (Additional file 2). on the arrangement and nature of the spines of the pedi. We recognise five states: was conducted within TNT. This character is modified coded from the results herein and redescription of Dunlop after Weygoldt ([34]. in some groups. anglicus was covered by Pocock [13]. The data matrix is available as Additional file 3 in a sequentially decrease in length towards the base (i. being more proximal running the analyses with differing taxa and characters than the (also subdivded) tarsus. 10. These were carried out under equal and records if the most proximal spine of the three principal implied weights. Where a Kronocharon species figured by Wunderlich [43] is applicable. Resampling of all analyses Weygoldt ([34]. We highlight that as part of ongoing development of the Results matrix we have added a number of further characters. For our equal weight analysis.001 to 122. we have coded this as unknown for this spe. associated with a ition of the extinct Kronocharon in more detail—we have presumably plesiomorphic pediform state for these limbs. suggest lateral eyespots are present (Fig. 10. Cladistic Analysis Phrynichidae) is more of a prehensile structure (a distinc. Whether these ters. 33% removal probability. character 10). Musicodamon. 74. character with prey capture. [14]. 10. Petrunkevitch [50] . k =3. character 8). up to 23. 2 and 3). cies due to difficulty in differentiating the podomeres. see below). we highlight that our et al. and character pedipalp from Charon even though both genera are cur- 66 on whether this is present as a spine ([34]. This row often consists of available with the sponsorship of the Willi Hennig Society). character 7). and the species described by apotele. and also on the basis of excluded to explore their impact. We note that the new photos of the type specimen of Much of the internal resolution within Amblypygi is based Paracharon. Charinus. Whilst a rough figure for ([49] . nodal support values are shown as absolute obtainable. we present the We have additionally added a limb character to reflect strict consensus of an arachnid-wide analysis at a k value of the whip-like first limbs of whip spiders. an additional strict consensus coded based on the number of subdivisions of the tibia of all tree topologies recovered from 88 k values ranging of the first leg. of a reviewer. associated with the limbs’ raptorial nature. bootstrap ([47] .g. and the subsequent character. rently placed in the same family. and has been added as character 62. family Hubbardiidae (after Cokendolpher & Reddell [42]). made tion coded in character 75). Character the number of lenses and the nature of the rhabdomes as 64 reflects whether the pedipalps have a row of femoral unknown.e. 100 trees per cycle. as well as a generic A key difference seen in the pedipalps of Paracharon and coding for Amblypygi as a whole. form a catching basket.000 more than 25 segments. character 65 on whether the tro. and Paraphrynus. 18).

in more considered equivocal in previous studies. 2c). caecus (Fig. the femur of the pedipalp in G.Garwood et al. This is again similar to the condition ([34]. like P. Some legs show evidence for a pair of tarsal claws. The centre of the dorsal shield in in Paracharon caecus (Fig. but the CT data reveals them In detail. but the presence or absence of a pad-like pulvillus Fig. Legs II-IV of G. 3f ). we could resolve that the trochanter of the as two-segmented. 3a. anglicus has a deep depression (the fovea) which prob. only two prominent tibial spines (Fig. 3d. tarsus and apotele. caecus (Fig.b. anglicus is equivocal. In this scheme. character 18) remains equivocal. In modern whip spiders the fang for all Amblypygi—and that in Graeophonus anglicus it opposes a series of internal teeth (Fig. In all other pedipalp article in whip spiders as the tibia. 3e). Additional file 1). the prosomal dorsal shield coxae of legs II-IV are very clearly preserved and coxa II in (or carapace) is a single plate. In more derived whip spiders (namely the Unidistitarsata. clasp-knife structures similar to those pedipalp has a ventral apophysis (Fig. In brief. and structure can be phylogenetically informative ([34]. BMC Evolutionary Biology (2017) 17:105 Page 7 of 14 and Dunlop et al. outline. whose number is more like a flange and not reduced to a spine ([34]. as elucidated by Weygoldt Graeophonus anglicus. Three prominent tibial spines are seen in more derived living species ([34]. character 28). most proximal one may not be preserved at its full length. like many authors. referred to the next uppermost or distal tooth has only one cusp. This implies that leg I was antenniform as in Recent taxa. 3a and 4. The CT scan can trace leg I down to the first few articles of what is clearly a subdivided tibia (Fig. g). derived whip spiders the pedipalps primarily articulate from The chelicerae of Graeophonus anglicus were also largely side to side. somewhat reniform in particular is quite tuberculate (Fig. caecus (Fig. distal parts between these claws in G. equivocal in previous studies. 3b). 2c). We could confirm the general depression. the first leg of Graeophonus anglicus appears to be long and slender. 3b). This is the presence of lateral eye tubercles (Fig. 2c) and was used as evidence for G. Additional file 1. [14]. We could confirm that. see below) the tarsus and apotele are fused. Also of significance is the fact that. f. It also has. at least in legs II and III. patella. referring the Carboniferous fossil to Paracharontidae. 3a). We prefer to inter- pret and score it as a tarsus which is (still) separate from its apotele.f )—a character of living species (Fig. and III and IV can now be traced down to near their tips (Fig. anglicus are more complete. 1). Previous studies [14] recognised included as additional file 4) similar to the condition in two dorsal spines on the femur of the pedipalp in Paracharon caecus (Fig. 2e). anglicus primarily eye tubercle in G. As in living whip spiders. however we living whip spiders the upper tooth has two cusps. The distal tip of the pedi- palp in G. d. it is preserved . like Paracharon character 13). anglicus. 2c) there is a small (proximal) tibial spine which was not shown in Weygoldt’s drawings. we can argue that G. as in Paracharon. 3g). 3a) which were more like the condition in P. 3c). but based on comparisons with extant material is quite robust and. movie whip spider relationships. the Weygoldt [34]. in the photo- graph of P. anglicus appears to be offset against the preced- ing element forming a pretarsus or apotele (Fig. 3c. 4 Graeophonus anglicus as it may have appeared in life. character 8). Previous interpretations picked up the median observation that the pedipalps in G. Several grooves radiate out from this more detail (Fig. caecus (Fig. 3a-b). but with a distinct projection of the anterior Pedipalp morphology is a key character for resolving median region (Figs. tibia. anglicus ence of four internal teeth in G. 2a). The CT scan was able to resolve the pres. The follow Shultz [4] and recognise a more conventional coxo-sternal region is well-preserved and for the first time series of articles for an arachnid pedipalp: namely a the CT scan picks up the small coxae of leg I (Fig. However. and the CT data confirmed articulate up and down in a vertical plane (Fig. anglicus. character 7) and has only two short spines here (Fig. The femur of leg I hypothetical. The ably acted as an attachment site for the muscles of the CT scans allowed us to investigate pedipalp morphology in sucking stomach. The femur. In Weygoldt’s [34] character 14 terminology this would be referred to as a “divided distitarsus”. 3g). but the exact number of tibial segments ([34]. character 4). although the lacks a prominent row of dorsal spines (Weygoldt [34]. anglicus has two patellar spines (Fig.

Characters relating to this spination + Paracharon)) + (Charinus (Stygophrynus ((Kronocharon explicitly group Paracharon closer to Paracharonopsis (Charon (Musicodamon + Paraphrynus))))). 7). and parsimony in anterior and posterior ends. [14]. (5) presence of a pulvillus on legs II-IV. Weygoldt [34] tested this These are: (1) the presence of serrula on the chelicerae. The opisthosoma is oval with a series are unchanged. We note that discussions regarding the of tergites (Figs. we must entertain the Unidistitarsata. albeit with the modification of the infraorder dorsal shield. 53]. Within the below). recognising this is potentially a more accurate form of parsimony analysis than implied weights. Equal weights analysis recovers Amblypygi in a polytomy Our justification for this is that Graeophonus anglicus has with Uropygi and Haptopoda. whip spiders were broadly divided into so. [51]) based on the detail. pulvillus at the ends of legs II-IV. in which the anterior margin is straight or only neoamblypygids. and equivalent to the ‘apullvillates’ in Paracharon and Paracharonopsis (Fig. consisting of Graeophonus. of the same name) and Neoamblypygi. These define the (Cretaceous-Recent) whip formations and the pedipalp structure of our terminal taxa scorpion family Thelyphonidae as a crown group which (Fig. (4) a dorsal row of patellar spines rendering the ‘pulvillate’ taxa paraphyletic.and distitibia as of equal weights at high k values. Looking at the character distribution on this tree in called pulvillate and apulvillate taxa (e. Outgroup comparison does suggest that other paleoamblypygid features like the vertically Amblypygi articulating pedipalps are probably plesiomorphic (as it is Cladistic analysis of our matrix using traditional search for other Pedipalpi). Weygoldt’s model was largely apomorphy supports Paleoamblypygi in our analysis. Palaeontology) and treat it as a stem-paleoamblypygid. (3) presence (Charinidae (Charontidae (Phrynichidae + Phrynidae)))). which encompassed their extinct genus possibility that the paleoamblypygids are a grade rather Kronocharon Engel & Grimaldi. (2) hypothesis cladistically and recovered (Paracharontidae a ventral apophysis on the pedipalp trochanter. but this was not the most parsimonious result see below). The basic (see below). weighted analysis.Garwood et al. We present these trees in Fig. but shows some internal instabil- have a similar leg configuration to help them to crawl into ity within the amblypygids. This may well Implied weights analyses recovers ((Amblypygi) (Haptopoda reflect its orientation in life since modern whip spiders (Uropygi + Schizomida)). 5. The patella is small and bell shaped. the pedipalp. Note that the phrynoids are the most derived blypygi sensu Weygoldt [34]. Cladistic results and discussion but could be overly-precise. which has a placement matching that tibia of leg IV is divided into a basi. Other relationships ([34]. neoamblypy- gids were divided into the superfamilies Charontoidea Paleoamblypygi (for Charontidae) and Phrynoidea (for Phrynichidae and The analysis supports a monophyletic suborder Paleoam- Phrynidae). The taxon Kronocharon. We also present our results earliest fossil whip scorpions lack projecting apophyses on from this analysis mapped onto the main character trans. This is not seen in the Euamblypygi (see name Charinidae to Charinina in Prendini [8]. of the opisthosoma are equivocal. group in this scheme.g. This results from the fossil narrow spaces. BMC Evolutionary Biology (2017) 17:105 Page 8 of 14 with the prolateral surface uppermost. he recognised two suborders: albeit with a reversal in the most derived genera. of a palpal cleaning organ. Were this carapace projection to be inter- Engel & Grimaldi [18] introduced another clade name. 4. 2014 + Phrynoidea (but than a clade. and ingroup whip spider a much simpler pattern of pedipalp spines compared to relationships of the form (Graeophonus (Paracharonopsis other whip spiders. a pulvillus the other four families. we now formally ex- of 520 steps. adopted in subsequent classifications. As an alternative on the pedipalp. such as the catalogue namely the anterior projection at the front of the prosomal of Harvey [6]. preted as a plesiomorphic character.38 steps. and the distitibia itself appears to be undivided solved as sister group to Musicodamon. and plotted against geological time (Fig. character 23). 5). Ventral sacs on the underside comparison to probabilistic methods are ongoing [52. A single putative Quintero’s earlier scheme. 6). Traditionally. and (6) Paleoamblypygi (for Paracharontidae) and Euamblypygi for the presence of ventral sacs. Implied weights analysis (k = 3). A reconstruction of the In this work we opt to focus our discussion on the equally likely appearance in life is presented in Fig. 3a-b and 4) which are shorter towards the merits of different weighting schemes. Finally. can be defined by more raptorial—and presumably . This result is similar to Tetlie & Dunlop’s [54] phylogenetic structure is thus largely in accordance with conclusions about the Coal Measures whip scorpions. although not all of them can be seen in all the fossils. The the results of Weygoldt [34]. in the present analysis. but at lower ones is re- expected. the recent study of amber fossils by slightly rounded. options (TBR) and equal weights (EW) resulted in 96 trees In contrast to Dunlop et al. The euamblypygids were further and ventral sacs are not unique for Amblypygi and can also subdivided into the infraorders Charinidae (for the family occur in some other arachnid orders. Note that serrula. resulted in clude Graeophonus from Paracharontidae (see Systematic 24 trees of 41. Amblypygi is defined in our analysis by six apomor- presence or absence of this small fleshy pad called the phies. way of expressing this.

The carapace is figured as more rounded tarsus and apotele also fuse into a single (and spineless) anteriorly. More generally. and Bremer support values. The status of Thelyphrynus elongatus analysis—present in Charinus too. 6). Shown is the strict consensus tree run under an equal weighting scheme. BMC Evolutionary Biology (2017) 17:105 Page 9 of 14 Fig. It is defined in our dataset by (Petrunkevitch. In lieu of a formal amber and living palaeoamblypyid genera are more heavily . The family Paracharontidae is restricted here to Paracharon the original figures of Sorellophrynus carbonarius and Paracharonopsis (Fig. bootstrap. 6). 2c). a strict consensus tree recovered using implied weighting (k = 3). 1913) from Mazon Creek suggest that the presence of several small spines on the pedipalp tarsus this genus also has the paleoamblypygid character of a (Fig. a character which is—convergently in our carapace projection. redescription we provisionally place this genus as a the palp of Graeophonus suggests that whip spiders also paleoamblypygid too. but the original figures imply that the carapace tip. 5 The results of the cladistic analysis of chelicerates and arachnids used to place fossil amblypygids. is harder reversed in the more derived whip spiders in which the to assess. seem to show a trend in which the pedipalps become increasingly better adapted for restraining their prey (Fig. we suggest that the pedipalps of the and pedipalp are not well-preserved. also from Mazon Creek. This character is Petrunkevitch. jackknife. Paracharontidae With respect to the other Coal Measures whip spiders.Garwood et al. Likewise. 1913. and a strict consensus of the trees recovered under 88 implied weights analyses at varying k values more efficient—pedipalp used for prey capture.

as shown in Paracharon caecus is unknown but we suspect that both Figure 15 of Weygoldt [34]. 3 and 6). The biology of living has. Traditionally Stygophrynus and Charon were placed together as the family Charontidae. 6. but this would still leave the family paraphyletic and containing genera having quite different pedipalp architectures. However. our dataset by (1) pedipalps which articulate primarily in a forming the Unidistitarsata based on the diagnostic apo- horizontal plane and (2) a row of dorsal spines on the morphy of an undivided pedipalp tip (Fig. by contrast. see also Fig. anglicus (Figs. an undivided distitarsus. To accommodate this result we could assign Kronocharon to Charontidae too. and reflected in the scoring of this species. 6 Results of the cladistic analysis highlighting key character Charontidae sensu stricto. we pedipalp femur. As noted above on prey items immediately in front of them. BMC Evolutionary Biology (2017) 17:105 Page 10 of 14 broad sweeping movements to capture prey which is further away. side-to-side. action of their pedipalps greatest number of elements in the leg I tibia.Garwood et al. (Charinus differs from the original interpretation of Engel & Grimaldi (Stygophrynus ((Kronocharon (Charon (Musicodamon + [18]. 6). The elongation of the pedipalps in derived euamblypygids supports this general hypothesis. enabling possible to score the number of tibial elements in leg I for . but only two in G. Almost all Stygophrynus. It may be conferred a considerable evolutionary advantage. Musicodamon + Paraphrynus). who proposed that the fossil genus was the sister Paraphrynus))))) among our terminal taxa—is defined in group of the Phrynoidea (i. Unnamed clade The next clade recovered in our analysis is (Stygophrynus (Kronocharon (Charon (Musicodamon + Paraphrynus)))). like phrynoids.e. and the fossil paleoamblypygids. together with sketch reconstructions of the pedipalps of our selected terminal taxa (not to scale). It is also supported by a longer number of tibial elements in the first pair of legs and the loss of the coxal gland opening on the segment bearing the third pair of legs.e. For This clade is defined in our dataset by the key character of example the patella bears three spines in Paracharontidae. structure and spination going up but we are reluctant to add a monogeneric family to the through the phylogenetic tree nomenclature at this stage. Our results challenge the monophyly of this family (see also below) especially given the fact that Stygophrynus retains a separate tarsus and apotele in the pedipalp while Charon has the more derived character of a fusion of these elements. 6. The principal apomorphy which supports this group is the tendency for the distal ventral patellar spines on the pedi- palp to form a ‘catching basket’ (subsequently modified into the phrynichid ‘hand’). In our analysis the Burmese amber genus Kronocharon resolved as sister group to the Neoamblypygi (but see also Euamblypygi implied weights analysis). Since we were not comprehensive in covering extant genera we prefer for now to place Stygophrynus as the sister group to the Unidistitarsata (see below) and restrict Charontidae to the genus Charon as Fig. In our analysis the two extant genera are also split apart from each other by the extinct Kronocharon which also has a fused pedipalp tip (Fig. Note the necessary to recognise a separate family for Stygophrynus. grab down his character 14 (see also [55] for example). changes in pedipalp orientation. We concede that it may prove transformations. the fusion of the tarsus and apotele into a single element. Unidistitarsata spined than the pedipalps of Graeophonus anglicus. The most derived whip spiders also have the the horizontal. rather than just a couple of isolated spines contend that—contra Engel & Grimaldi [18]—Charon also as in Paleoamblypygi. This position for Kronocharon Weygoldt’s [34] suborder Euamblypygi—i. retains a pedipalp tip which is living whip spiders are euamblypygids. We presume that divided. although this does move up-tree in some implied weights analyses).

as noted above. pulvillus. Paracharontidae from the Eocene to Recent. are defined in this same clade in the literature is Apullvillata. Finally the two most derived genera in our dataset (Musicodamon and Paraphrynus) belong to the Phrynoidea (or Phrynida). As noted above. BMC Evolutionary Biology (2017) 17:105 Page 11 of 14 Fig. an older name for the nus)) from our selected terminal taxa. which is defined here by (1) the reduction Neoamblypygi of four cheliceral teeth to three and (2) the loss of the The neoamblypgids. However. Euamblypygi can be dated from the Cretaceous to Recent. By contrast Systematic palaeontology in Kronocharon this apophysis is described as being a Order AMBLYPYGI Thorell. analysis by the apomorphy of a spine-like ventral apophysis on the trochanter of the pedipalp. Circles indicate fossil occurrences with their approximate dates in millions of years. in our scheme Charontidae is no longer Suborder PALEOAMBLYPYGI Weygoldt. 1996 monophyletic. and we recognise Charontidae sensu Stem-PALEOAMBLYPYGI . Phrynoidea) were also present during the late Cretaceous Kronocharon based on Wunderlich [43]. number of tibial as opposed to tarsal elements is not given here we have chosen to treat this character in the amber Phyrinoidea genus as equivocal. (Charon (Musicodamon + Paraphry. 1883 large and carina-like [18]. while the phrynid from the ca. 115 Ma Crato Formation of Brazil implies that the remaining lineages (Unidistitarsata. Amblypygi and Palaeoamblypgyi can be dated from the Carboniferous to Recent. 7 The recovered phylogeny expressed as the current higher taxon/family structure and superimposed on the the known fossil record. since the exact Phrynoidea. rather than explicitly a spine. Neoamblypygi.Garwood et al. The enigmatic Ecchosis is tentatively treated as being close to whip spider origins. Indeed. who reported 65 stricto (consisting of Charon only) as the sister-group to elements beyond the patella.

with terminal claws visible in leg III. but tarsus divided into †Graeophonus Scudder. Femur broadens distally and bears only two small dorsal spines. 1996 curved on the mesal margin and here bearing two spines. trochanters bell- shaped.4 mm). †?Thelyphrynus Petrunkevitch. total length ~7 mm. microtomography (Fig. Coxae subtriangular.6 mm. Sarax tear-drop shaped tubercle. Included genera Patellae short and bell shaped.5 mm. From Coseley. Tarsus separate from terminal apotele. small lateral eye tubercles in Simon. 2002. but with wide anterior projection. more distal elements equivocal but Martill.d. 1996 Description Included genera Description as Dunlop et al. three tarsomeres. Charon Karsch. †Kronocharon Engel & Grimaldi. projecting forwards beyond the anterior margin prosomal shield Included genus and composed of two articles (Fig. Dorsal Family CHARINIDAE Quintero. Femur IV with hook-like ventral projection at its distal margin. proximal shorter (0. Patella broad. maximum width 5. Weygoldt . 1892. number of individual lenses equivocal. Phrynichodamon. Infraorder NEOAMBLYPGI Weygoldt. tri. both without spines. Pair Included genera of median eyes on the anterior projection.L. Legs Moreno. length 7 mm. 1921. 2002. and a prominent outer distal spine. 2014. Tibia with two mesal Included genera spines.and distitibia. 1852 angular. Family PARACHARONTIDAE Weygoldt. 3e): a basal element Stygophrynus Kraepelin. borne on a Catageus Thorell. Leg I trochanter more than twice as wide as long. min. Leg I coxa small. 1996 shield length 4. Tibia slender and divided Damon C. maximum †Graeophonus anglicus Pocock. 1839. Carboniferous (Duckmantian) (Figs. 1892. Femora somewhat flattened. BMC Evolutionary Biology (2017) 17:105 Page 12 of 14 Included genera subdivided.Garwood et al. Basitarsus undivided. Leg 1 tibia incomplete but was evi. II-IV more robust. Additional file 1). 1892 the proximal one slightly shorter (0. apparently with thin Family PHRYNICHIDAE Simon. Included genera tella small.3 mm) than distal (0. bell shaped. Superfamily CHARONTOIDEA Simon. Pulvillus equivocal. Trochanter with flange. the most distal of which has only one cusp. Basal element Clade UNIDISTITARSATA Engel & Grimaldi. Ventral sternite pattern NHM 31234 (paratype). (or paturon. Unnamed clade Chelicerae small. 1913. 1879. Prosomal dorsal shield reniform. posteriormost four visible Material tergites increasingly shorter. near Dudley. 3 and 4. Total length 11. Pedipalps robust and bearing numerous spines Included genus (Fig. 1895. Leg 1 femur slender. 1801. First tergite short. Leg 1 pa. 1911 width 4 mm. Paraphrynus overall habitus implies an antenniform appendage. 1894. next six longer and approximately of equal length. Late sacs on underside of the opisthosoma equivocal. 1940. of the ‘clasp-knife type. flattened.3 mm. into a basi. 1850. 2014 bears four teeth. 1852 Legs gracile.6 mm). 8 mm long) opposed by a gently curving and tapering distal fang (1 mm in length). The same article has a further distal spine on the outer surface (0. British Middle Coal Measures. 3c. Genus †Graeophonus Scudder. an anterolateral position can also be resolved. †Sorellophrynus Harvey. largely matches that of the corresponding tergites. like ventral apophysis. Family PHRYNIDAE Blanchard. slightly pro. 1890 Opisthosoma oval. 1986 shield with median longitudinal depression (the fovea) and several depressions radiating out from this structure. 1892 margins and becoming slightly narrower distally. 3). 2014. Phrynus Lamarck. narrowing slightly distally. 1892 sensu stricto one (0. Euphrynichus Weygoldt 1995. 1890. Tritosternum or additional expected sternal elements between the leg coxae equivocal. Charinus Simon. Heterophrynus Pocock. [14]: here we primarily Paracharon Hansen.3 mm) than the distal Family CHARONTIDAE Simon. Suborder EUAMBLYPYGI Weygoldt. leg I antenniform. Superfamily PHRYNOIDEA Blanchard.6 mm). †Britopygus Dunlop & dently subdivided.g). these elements are not further Musicodamon Fage. Ventral Staffordshire UK.f. †Paracharonopsis Engel & focus on novel morphological features revealed through Grimaldi. Acanthophrynus Kraepelin 1899. Koch. 1899.

2:e641. RJG is a member of the Interdisciplinary Centre for Ancient Life relatively few spines to facilitate prey capture: namely two (UMRI). USA. the fossil species can be excluded from the WIU Undergraduate Research Grant and the WIU Department of Geology. (2) appearance of a spine-like ventral apophysis on the pedi. 5) largely support the 1996 phylogeny of Weygoldt. BJK and TAH. for providing photographs of Paracharon. 1999. Kaluziak ST. 1996. 1911 presented herein. D-10115 Berlin. Tillman Hall 113. Garwood RJ. London) for providing access to fossils in common with typical arachnid walking legs. Three-dimensional reconstruction and the phylogeny of extinct chelicerate orders. inferred/ Conclusions reconstructed anatomy partially transparent. Title: Tomographic 2. UK. living or extinct. Germany. The Natural History Museum. evolution and modification of the group’s pedipalps Ethics approval (Fig. 1 University Circle. González VL. IL 61455. Sharma PP. which was funded in part by the EPSRC (grants EP/ the projecting anterior part of the carapace (Fig. although the recently described Burmese amber genus Competing interests The authors declare that they have no competing interests. appendages still primarily articulate up and down. in Københavns Universitet). i. Wheeler WC. Description: phylogenetic signal. Phrynichus Karsch.pdf. 7). Kronocharon is placed a node deeper in the tree compared to its original interpretation. and its evolutionary significance.Garwood et al. Phrynoidea. Hormiga G. .3 mb) et al. of the tarsus and apotele to define the Unidistitarsata. Shultz JW. (ZIP 45. The University of Manchester. 1935. specifically three patella spines and The dataset supporting the conclusions of this article are available in a additional small tarsal spines. Xerophrynus Weygoldt. and a tendency to concentrate the patella Springer Nature remains neutral with regard to jurisdictional claims in spines distally into a ‘catching basket’ which define the published maps and institutional affiliations.5281/zenodo. 1911. 7). The relationships we recover All authors read and approved the final manuscript. UK. testing the position of future fossil discoveries. patella and three (two mesal. and (3) loss of the pulvillus on the legs and Author details 1 reduction in the number of cheliceral teeth defining the School of Earth and Environmental Sciences. one lateral) on the tibia. Title: Character statements. A phylogenetic analysis of the arachnid orders based on reconstruction of Graeophonus anglicus Pocock. Description: A 3D morphological characters.2 mb) Tomographic investigation of the Carboniferous amblypy- gid Graeophonus anglicus reveals the least modified Acknowledgements pedipalps of any whip spider. 7). Eocene–Recent group Paracharontidae which has more Availability of data and materials pedipalp spines. Trends which can be recognised include (1) fusion Not applicable. (PDF 507 kb) (Arachnida: Amblypygi). Pérez-Porro AR. London SW7 5BD.150:221–65. Shultz JW. which the pedipalps primarily articulate from side to side. They have her care. (TNT 44 kb) Additional file 4: A video showing the tomographic reconstruction of Graeophonus anglicus Pocock. Cromwell Road.vaxml (see [38]). Muscular anatomy of a whipspider. 2Department of Earth Sciences. Several of the most derived phrynoid genera Manchester M13 9PL. Description: Cladistic matrix used in the current analysis. Most whip spiders belong to Zenodo repository (DOI: 10. Title: Cladistic matrix. with a horizontal plane of motion. Neoamblypygi. F007906/1. Dunlop J. Zool J Linn Soc. The Cretaceous Authors’ contributions phrynid from the Crato Formation of Brazil implies that RJG and JAD wrote most of the text and descriptions and carried out the all three euamblypyid families should have been present in cladistic analysis. 3Museum für Naturkunde. Invalidenstraße highly concentrated patellar spines form the so-called 43. Zool J Linn Soc. G. small dorsal spines on the femur. EP/F001452/1 and EP/I02249X/1).tnt (see [45]). 3. 2014. Trichodamon Mello-Leitao.31:2963–84. These results provide a framework for University. 2007. 2014. Tomographic images and the reconstruction were produced by RJG. the Cretaceous (Fig.437900). BMC Evolutionary Biology (2017) 17:105 Page 13 of 14 1996. 6). in which the distally Leibniz Institute for Research on Evolution and Biodiversity. 4. Western Illinois phrynichid ‘hand’.e. mesh model of Graeophonus anglicus in the VAXML interchange format. Received: 14 September 2016 Accepted: 9 March 2017 References Additional files 1. and Claire Mellish (Natural History Museum. Additional file 3: File format:. All authors contributed intellectually to the paper. Macomb. These We thank Jan Pedersen and Nikolaj Scharff (Statens Naturhistoriske Museum. We are grateful for the comments of two anonymous reviewers. 6). Paleoamblypygi (Fig. Additional file 1: File format:. Publisher’s Note palp trochanter. Funding anglicus is placed in the Carboniferous–Recent suborder RJG’s visit to Berlin was supported by a European Union SYNTHESYS grant. This analysis demonstrates Consent for publication that whip spider phylogeny is effectively reflected in the All authors have provided consent to publish. 4Department of Geology. BJK’s work was supported by a However. PeerJ. (Fig. in a TNT ready format. Phrynus longipes (Pocock) Morphological characters statements for the characters used in the current analysis. Mol Biol Evol. also have extremely long pedipalps. (AVI 87. in The datasets supporting the conclusions of this article are included within the article (and its additional files). 1879. Phylogenomic interrogation of Arachnida reveals systemic conflicts in Additional file 2: File format:. the Cretaceous–Recent suborder Euamblypygi (Fig. two larger spines on the which improved the manuscript.126:81–116. which can be defined based on We would like to acknowledge the assistance provided by the Manchester X-ray Imaging Facility.

Goloboff PA. 2015. 2002. Rev Ibérica Arachnol. Garwood RJ. Anim Biodivers Outl High-Level Classif implications for time scale calibration. Edgecombe GD. Revision of the Protoschizomidae (Arachnida: Crato Formation of Brazil. 2002.135:179–90. Wheeler WC. The affinities of the Carboniferous whip 35. mexicana. Garwood RJ. Smithsonian Institution Press.9:409–36.6:699–702. Cladistics. Illustrations of the Carboniferous Arachnida of North America 2014.24:774–86. Charbonnier S. USA. Cladistics. Grimaldi DA. The first whipspider (Arachnida: Amblypygi) and morphological and molecular data. creacion de subordenes. Braddy SJ. Proc R Soc B. 33. et al.105:245–50. Giribet G. 47. Raffles Bull Zool. a free program for phylogenetic Beitr Zur Naturkunde Ser B Geol Paläontol. Sharma PP. 1955. Quintero D. 2011. development. Farris JS. 2008.24:1017–23. Cladistics. TNT. high-precision U–Pb dates from western European Carboniferous tuffs. Catalogue of the smaller arachnid orders of the world. Petrunkevitch AI. (Amblypygi: Charontidae) from Java and adjacent islands. Clark J. Dunlop JA. Part P Arthropoda 2. Dunlop JA. A total-evidence approach to dating with fossils. Warnock RCM.18:5–70. Schawaller W. Szumik CA. North America. Giribet G. Beitr Zur Araneol. Palaeontology. 2012. Dunlop JA. Palaeontol Electron. and a format for virtual 12. Tomographic reconstruction of neopterous Carboniferous insect nymphs. 2004. New and interesting insects from the Carboniferous of Cape 37. Garwood RJ. Dunlop JA. Panama 1983 of the Hymenoptera. 1st ed. is described from Mexican amber. Giribet G. Rocha CEF. Treatise Invertebr. Zhou GRS. Garwood RJ. Garwood RJ.50:48–66. . Engel MS. 2014. Kallersjo M. Nat Commun.0 [Internet]. Sutton MD. Brown AE. 18. 28. The enigmatic arthropod Camptophyllia. Harvey MS. Stuttg 45. Ninth Int. 2008. 2010. Barov V. 2014. Arachnol. 10. Lamsdell JC. Schulmeister S. Scudder SH.39:783–91. Curr Biol. 20. Ronquist F. nuevas especies (Arachnida: Amblypygi). Available from: http://museum. Källersjö M. 27. 2008. Felsenstein J. Combined methodologies for three- dimensional reconstruction of fossil plants preserved in siderite nodules: Stephanospermum braidwoodensis nov. Giribet G. 31. Albert VA. Electrophrynus mirus (Amblypygi). Revision de la clasificacion de amblypygidos pulvinados: calibration of the molecular clock. Palaeontology. PLoS One. J Arachnol.63:1–44. Goloboff PA. Proc Geol Assoc. Reddell JR. Oxelman B. Petrunkevitch AI. Perth. Mrugalla B. Cokendolpher JC. Chabard D. 1911. Pointon MA. Congr. analysis. 43.188:1–17. system (Chelicerata: Arachnida: Amblypygi).56:281–8. 34.5:841–4. 42–162.26:273–84. Aust. dammermani Roewer. Rahmadi C. Tanner AR. Exploring uncertainty in the 51. Bonamo PMP. Earth mites: sensitivity to homology assessment under total evidence. Chiapas amber spiders. Lubin YD. Beitr.8:88–90. Evolution. et al. 1994. 8. Congreve CR. Charbonnier S. Whip Spiders. Invertebr Biol. Improvements to resampling measures of group support. West. phylogenetic analysis of phenotype data. Chew DM. Amblypygi). 38. bootstrap. Phrynus 49. Sotty D. Dunlop JA. 1971. Scudder SH. p. 13. In: Ebehard WG. una nueva familia y un nuevo genero con tres 25. the periodicity of late Carboniferous Surv Taxon Richness Zootaxa. Indonesia with 19. Prendini L. Redescription of the Chiapas amber whip spider 50. Proc. Puttick MN. New whip-spiders. A Study of Palaeozoic Arachnida.3:31–74. Uropygi) from reconstruction of siderite-hosted Carboniferous arachnids. Weygoldt P. Sutton MD. In: Moore RC. Phylogeny and systematic Geological Nature Reserve. Phylogenetic position of the acariform spider Graeophonus anglicus Pocock. 1992. 2016.54:145–61.wa. Dunlop JA.4:443–56. 32. Ross A. Carboniferous harvestmen demonstrate early cladogenesis and stasis in Uncertain-tree: discriminating among competing approaches to the Opiliones. 1991. position of Opiliones: A combined analysis of chelicerate relationships using 16. 1986 [cited 2016 Sep 5]. Paleontol. Neue Befunde an Geißelspinne in Dominikanischem 46. An Upper Carboniferous amblypygid from the Writhlington 41. Dunlop JA. Nixon KC. 9. Sutton MD. J Zool Syst Evol Res.15:12p. Erstnachweis der Ordnung Geißelspinne in Dominikanischem notes on S.12:99–124. 2010. Cladistics. Parsimony Bernstein (Stuttgarter Bernsteinsammlung: Arachnida. Biol Lett. Donoghue PCJ. Biol Lett. Collingwood: CSIRO Publishing. Cladistics. Arachnida.9:1–17. 2017. Garwood RJ. software toolkit for tomographic visualisation. Dunlop JA. Lawrence: University of Kansas Press. Ramirez MJ. A new whip spider (Arachnida: Amblypygi). Stenstrup: Apollo 2000. p. Harvey MS. p. Edgecombe GD. 2003. New species of Charon (Amblypygi. Hilton J. 22. 39. Babbitt C. 1949. 2012.92:325–34. West PLJ. 29. 6.34:185–202. Mem Boston Soc Nat Hist. [Internet]. Univ Calif Publ Entomol. Mus. France. New and rare fossil Arachnida in Cretaceous Burmese amber (Amblypygi. Confidence limits on phylogenies: An approach using the 21. Garwood RJ. Poinar Jr GO. Martill DM. 2005. Selden PA. Branch support and tree stability. Environ Sci Trans R Soc Edinb. Schizomida) with notes on the phylogeny of the order. Accessed 5 Sept 2016. BMC Evolutionary Biology (2017) 17:105 Page 14 of 14 5. Spencer ART. Farris JS. Dunlop JA.15:14p. 1878. Harvey MS. paleontologists with a case study on extinct arachnids.88:735–46. sp. The walking dead: Blender as a tool for Breton. Siveter DJ.82:299–312. Trans Conn Acad Arts Sci. Monogr Palaeontogr Soc. Petrunkevitch A. Novit 44. A monograph of the terrestrial Palaeozoic Arachnida of specimen interchange. Chabard D. Anatomically modern 52.18:1–137. Whip Spiders of the World. Charontidae) from (Arachnida: Trigonotarbida). 2012. In: Wunderlich J. J Geol Soc. Klopfstein S. 2016. 1996. Schawaller W. Rev Palaeobot Palynol. Siveter DJ. editor.59:447–62. Tex Meml Mus 17.Garwood et al.8:156–9. Stuttg jackknifing outperforms neighbor-joining. 1982.37:69–315. II. BMC Evol Biol. Northern Australia and Christmas Island. Beitr Zur Naturkunde Ser B Geol Paläontol.86:1–12. including maternal care. Sutton MD. Pocock RI. Dunlop JA.7:e45779. Syst Biol. A first epigean species of Stygophrynus Kraepelin Paleoentomologicae.2:444.19:324–32. and the origin of subchelate pedipalps in whip 2009. and reinterpretations of Devonian Araneae. Bernstein:(Stuttgarter Bernsteinsammlung: Arachnida. Devonian of New York. Lipscomb D. 2007. 1881–5. Sutton MD. applied to the early radiation Robinson BD. High-fidelity X-ray micro-tomography 54. 2012. Sevastopulo GD. Dunlop JA. Trans R Soc Edinb Earth Sci.61:973–99.12:53–62. Amblypygi Thorell. Bremer K. Holloway L. Garwood RJ. A new fossil whip spider (Arachnida: Amblypygi) from Speleol Monogr. 2012. editors. A spider and other arachnids from the 36. 1911 (Arachnida: Amblypygi). (Medullosales) from the Mazon Creek lagerstätte. Zur Araneol. 48. Pepato AR. Geralinura carbonaria (Arachnida. and Britain.10:1–23. Trans Conn Acad Arts Sci. Fleming JF. Amblypygi). Ricinulei amd Uropygi: Thelyphonida). 15.3148:154. Weygoldt P. Crowley QG. 2012. 1928. Palaeontology. Tetlie OE. onychophoran terrestrialization. Farris JS. 11. Dunlop JA. Evolutionary morphology of whip spiders: towards a phylogenetic 7. 2003. 1998. O’Reilly JE. 1913.10:295–304. 23. 1883. A monograph of the terrestrial Carboniferous Arachnida of Great Carboniferous Onychophora from Montceau-les-Mines. 30. Mazon Creek. Sutton MD. Can Nat Q J Sci. 24. 1994. Garwood RJ. editor. Rasnitsyn AP. A Paleozoic stem group to 53.284:20162290. 2012.169:713–21. 1996. J Paleontol. 1979. Illinois. Yang Z. Garwood RJ. 14. Palaeontol Electron. 2011.98:165–78. Shear WA. cycles and stratigraphical correlation.43:220–3. Kluge AG. X-ray micro-tomography of Carboniferous stem- Dictyoptera: new insights into early insects. 1985. 2014. Implied weighting and its utility in palaeontological mite harvestmen revealed through integration of phylogenetics and datasets: a study using modelled phylogenetic matrices. 203–212 26. Vilhelmsen L. Morphology and systematics of Anthracomartidae 55. Wunderlich J. version 1. Murray DL. Sutton MD.64:1–84. J Arachnol. Biol Lett. of the Orders Anthracomarti and Pedipalpi. Sotty D. the Early Eocene and mid-Cretaceous. 2011. scorpions. Whipspiders (Arachnida: Amblypygi) in amber from 2015. Spiers and VAXML: A 1890. Ovtcharova M. three new whipscorpions (Arachnida: Thelyphonida) from the Lower Cretaceous 42. Harvey MS. J Paleontol. Petrunkevitch AI. the Crato Formation of Brazil.34:241–81.