Fossil Record 9(1) (2006), 130–136 / DOI 10.1002/mmng.200600004

A new trigonotarbid from the Measures of Hagen-Vorhalle,

Jason A. Dunlop*,1 & Carsten Brauckmann**,2

1 Museum fu¨ r Naturkunde der Humboldt-Universita¨t zu Berlin, Invalidenstraße 43, D-10115 Berlin, Germany 2 Institut fu¨ r Geologie und Pala¨ontologie, Technische Universita¨t Clausthal, Leibnizstraße 10, D-38678 Clausthal-Zellerfeld, Germany

Received 7 August 2005, accepted 20 September 2005 Published online 01. 02. 2006

With 3 figures

Key words: , Namurian, , Trigonotarbidae, Archaeomartus, new .

Abstract

A new trigonotarbid (Arachnida: Trigonotarbida: Trigonotarbidae) is described as Archaeomartus roessleri n. sp. from the Upper Carboniferous (; Namurian B, higher Marsdenian) of Hagen-Vorhalle, Germany. Originally assigned to Trigonotarbus johnsoni Pocock, 1911, our new has a distinctly lobed carapace and thus resembles more closely the Early species Archaeomartus levis Størmer, 1970. In carapace morphology Archaeomartus approaches the condition seen in the larger and more heavily-armoured taxa Eophrynidae, Kreischeriidae and Aphantomartidae. Thus we provisionally re- solve Archaeomartus as sister-group to this probably monophyletic trio of families and discuss the possibility that Trigonotarbi- dae may be paraphyletic.

Schlu¨ sselwo¨ rter: Karbon, Namurium, Trigonotarbida, Trigonotarbidae, Archaeomartus, neue Art.

Zusammenfassung

Ein neuer Trigonotarbiden-Fund (Arachnida: Trigonotarbida: Trigonotarbidae) aus dem Ober-Karbon (Pennsylvanium; Na- murium B, ho¨ heres Marsdenium) von Hagen-Vorhalle, Deutschland, wird als Archaeomartus roessleri n. sp. beschrieben. Bei der Erst-Dokumentation wurde er noch zu Trigonotarbus johnsoni Pocock, 1911 gestellt; er unterscheidet sich hiervon aber durch die deutlichen Loben auf dem Carapax und a¨hnelt damit eher dem unterdevonischen Archaeomartus levis Størmer, 1970. In der Carapax-Morphologie na¨hert sich Archaeomartus mehr den gro¨ ßeren und kra¨ftiger skulptierten Eophrynidae, Kreischeriidae und Aphantomartidae. Daher fassen wir Archaeomartus zumindest vorla¨ufig als Schwestergruppe dieser wahrscheinlich monophyletischen Familien-Dreiergruppe auf; die Trigonotarbidae ko¨ nnten somit paraphyletisch sein.

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Introduction frequently from the Upper Carboniferous of and where they are often Trigonotarbids (Arachnida: Trigonotarbida) are one of the more abundant fossil arachnid groups an extinct order of which are cur- found in these Coal Measures ecosystems. They rently known from the late (Prˇı´dolı´: ca. are far more frequent than for example 414 Ma) to the early (Asselian: ca. in these assemblages. Around fifty trigonotarbid 290 Ma) (Dunlop 1996a; Ro¨ ssler 1998; Ro¨ ssler species have been described in total, although et al. 2003). They have been recovered most this is probably an over-estimate since revisions

* Corresponding author: e-mail: [email protected] ** E-mail: [email protected]

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim Fossil Record 9(1) (2006) 131 have usually resulted in numerous synonyms Trigonotarbus johnsoni from the British Middle Coal Meas- (e.g. Ro¨ ssler 1998). Trigonotarbids resemble spi- ures (Westphalian B) of Coseley, Staffordshire, UK, was ex- amined, drawn and photographed some years ago by JAD in ders, but lack silk-producing . Trigono- the Natural History Museum, London (NHM). tarbids have a characteristic morphology of opisthosomal tergites divided into median and lateral plates. This they share with the rare Car- Geological setting boniferous–Recent , and the two orders have in some schemes been resolved as sister- The Hagen-Vorhalle locality in the German state taxa (Dunlop 1996b; Giribet et al. 2002), basal to of North Rhine-Westphalia is an internationally the so-called tetrapulmonate arachnids; i.e. a significant locality for Coal Measures fauna and phylogeny of the form: ((Trigonotarbida flora. The most productive horizon for are þ Ricinulei) þ (Araneae ( (Uropy- the so called “Vorhalle-Schichten” which have gi þ )))). The extinct trigonotarbids been dated as Upper Carboniferous, Namurian almost certainly fed on other and B (higher Marsdenian) based on goniatite strati- probably either ran down or ambushed their graphy. Hagen-Vorhalle is situated on the site of prey. Well-preserved Devonian examples have a former brickworks and the fossils were inten- mouthparts which strongly indicate that they sively collected between 1990 and 1997 by the were predatory, that they were not venomous “Pala¨ontologische Bodendenkmalpflege” of the and that they digested their food pre-orally like Westphalian Museum of Natural History. Part of many modern arachnids (Shear et al. 1987; Dun- the locality is now registered as a protected geo- lop 1994a). logical site for the town of Hagen. In addition to The majority of the described Carboniferous many plants, more than 300 fossils of terrestrial trigonotarbids were found at classic Westphalian arthropods are known from museum and private localities like the Saarland of Germany, the Brit- collections. The Hagen-Vorhalle locality has ish Middle Coal Measures, Mazon Creek in the yielded a diverse range of eighteen insect species, USA and Ny´rˇany in the . In re- six arachnids (including the new species described cent years a number of well-preserved fossil ara- here) and among the aquatic faunal component, chnids have been recovered from slightly older some incomplete eurypterids. A summary and (Namurian) sediments found in a former brick- further background literature can be found in yard quarry at Hagen-Vorhalle in the Ruhr area Brauckmann et al. (2003) and Hendricks (2005). of Germany (Brauckmann 1987a, 1991; Brauck- mann et al. 1985, 2003). These included a new trigonotarbid species, Eophrynus udus Brauck- Morphological interpretation mann in Brauckmann et al., 1985, an unnamed, incomplete trigonotarbid (Brauckmann 1987a) The new fossil (Figs 1, 2) is an almost complete and most recently (Brauckmann et al. 2003; specimen preserved essentially in dorsal view Brauckmann & Scho¨ llmann 2005) trigonotarbid material provisionally assigned to the British spe- cies Trigonotarbus johnsoni Pocock, 1911 (Trigo- notarbidae). Re-examination of the last specimen revealed that, unlike T. johnsoni, it has a cara- pace with lateral lobes – more like that of the Early Devonian species Archaeomartus levis Størmer, 1970 (see description by Brauckmann 1987b). The Hagen-Vorhalle fossil is thus reinter- preted as a new species of Archaeomartus Stør- mer, 1970 which we describe here.

Material

The holotype (P21 046) and only known specimen was ob- tained from its repository in the Westfa¨lisches Museum fu¨r Fig. 1. Holotype and only known specimen (WMfN P21 046) Naturkunde (WMfN), Mu¨ nster, Germany. Drawings were of Archaeomartus roessleri n. sp. from Upper Carboniferous prepared using a Leica MZ 12 stereomicroscope with a draw- (Namurian B, late Marsdenian) Coal Measures of Hagen- ing tube attachment. Pocock’s type and figured material of Vorhalle, Germany.

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1museum-fossilrecord.wiley-vch.de 132 Dunlop, J. A. & C. Brauckmann, Trigonotarbid arachnid from the Coal Measures of Hagen-Vorhalle

Fig. 2. Camera lucida drawing of the specimen shown in Fig. 1. Abbreviations: bt, basitarsus; cm, carapace margin; fe, femur; lb, carapace lobe; me, median eyes; pt, patella; py, superimposed ; st, superimposed sternite boundaries; ti, tibia; tr, trochanter; ts tarsus. Opisthosomal tergites numbered from 2–9, legs from 1–4; see text for details. Scale bar equals 1 mm. with a body length of about 6 mm. Under 70% gree of tectonic deformation, foreshortening the ethanol areas of original cuticle are revealed as body and legs for example. Such deformation darker patches and this area of preserved cuticle can occur at Hagen Vorhalle – see e.g. Brauck- is concentrated on the central region of the body, mann (1991, pl. 13: fig. 12) in which the wings of becoming more fragmentary at the margins of the insect figured here differ in length within a the and on the legs. Although com- single specimen – but we have only a single ex- pressed in a shale, some three-dimensionality ample of our new arachnid and unfortunately no (especially of the carapace and between adjacent reference point, such as a plant or ammonoid in tergites) is retained and surface relief is best re- the same slab, which would allow us to assess vealed under low-angle lighting. Two sets of the degree of any deformation which had oc- cracks run through the specimen, one close to curred. the midline, another at an oblique angle across the left side. They do not seriously obscure the C a r a p a c e – The carapace of the new fossil is morphology of the specimen. A reconstruction subtriangular, wider than long, and bluntly of its probable appearance in life is given in rounded at the front. The carapace bears a single Fig. 3 and suggests that this was a squat, robust pair of median eyes at its highest point. These little with short, rather heavily built legs. lack preserved cuticle in the fossil and are set We cannot, however, completely rule out the into slight recesses or sockets just in front of the possibility that the fossil has undergone some de- central point of the carapace. The whole median

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-fossilrecord.wiley-vch.de Fossil Record 9(1) (2006) 133 region of the carapace is raised – the ‘glabella’ widening of both the patella and tibia can be de- sensu Brauckmann 1987b – and forms three dis- tected. In legs 3 and 4 the femur, tibia and tarsus tinct lobes, the anteriormost of which bears the are the longest articles, the patella is shorter, eyes. The two lobes behind this are divided from while the very short basitarsus (or metatarsus) is the anterior lobe, and each other, by strong su- reminiscent of early derivative trigonotarbids like ture lines. Either side of this raised, central re- palaeocharinids; see e.g. illustrations in Dunlop gion the lateral flanks of the carapace exhibit a (1996b) and Fayers et al. (2005). The tarsus in series of procurved folds or suture lines defining our fossil is bluntly rounded distally. A pair of three subrectangular to subtriangular lobes on tarsal claws would be predicted (see Fig. 3) from each side. This fossil bears smooth and strongly better preserved Devonian material (e.g. Shear demarcated lateral carapace margins which were et al. 1987), but are not seen here. probably slightly raised in life. This closely re- sembles the condition seen in Lissomartus schu- O p i s t h o s o m a – The opisthosoma in the new cherti (Petrunkevitch, 1913); see illustrations in fossil is rounded and, unusually for trigonotar- Dunlop (1995a). The posterior margin of the car- bids, slightly wider than long. Tergites are apace in our new fossil is straight, not raised, strongly differentiated from each other along the and further ornament in the form of spines or length of the opisthosoma and have straight mar- tubercles on the carapace is lacking. gins in the median region. However, the charac- teristic trigonotarbid division into median and Appendages – The coxo-sternal region, lateral plates is weakly expressed and best de- mouthparts and in our new fossil are fined by cuticle distribution which seems to be equivocal. Although cuticle preservation is pat- more or less complete on the median plates only chy, boundaries of leg articles can be resolved (Fig. 2). The inner region of the lateral plates is under low-angle lighting. Leg 1 is known only thus outlined by wide gaps in the preserved cuti- from the proximal articles which seem to indi- cle on both sides of the opisthosoma, with cate a femur and patella of about the same further (tergal) cuticle fragments reappearing at length. Leg 2 is missing, but legs 3 and 4 are es- the margins. Unlike the median plates, divisions sentially complete. In general the legs are notice- between the lateral plates are poorly defined ably short and stubby with rather wide articles and interpretations are complicated by the super- compared to other known Carboniferous trigo- imposition of ventral elements onto the dorsal notarbids (e.g. Dunlop 1995a). Articles appear to surface. This is a typical feature of trigonotarbids have been basically cylindrical, but a slight distal compressed in shales. Here, the outline of the ventral sternites can be seen towards the back and sides of the opisthosoma – presumably be- cause the animal was thinner here in life. Traces of up to five sternites are visible in our fossil. Unlike the tergites, these are all somewhat pro- curved, more rounded elements, with the degree of curvature increasing posteriorly. The poster- iormost sternite surrounds another typical trigo- notarbid feature, a ventral pygidium (or postab- domen) formed from two small, circular segments which are also impressed here through onto the dorsal surface. This pygidium represents either opisthosomal segments 10–11 (Shear et al. 1987) or 11–12 (Dunlop 1996b) and probably bore the anal opening in life. Eight clear tergal elements are preserved in the fossil; however, assigning tergites unequivo- cally to segments in trigonotarbids is challenging. Petrunkevitch (1949), for instance, claimed (probably incorrectly) that segment number was Fig. 3. Sketch reconstruction of the probable appearance of variable within Trigonotarbida. Studies of well- Archaeomartus roessleri n. sp. in life. Setae and other details based on better preserved Devonian trigonotarbids and com- preserved Devonian material (Shear et al. 1987; parisons with extant arachnids. Dunlop 1996b; Fayers et al. 2005) revealed that

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1museum-fossilrecord.wiley-vch.de 134 Dunlop, J. A. & C. Brauckmann, Trigonotarbid arachnid from the Coal Measures of Hagen-Vorhalle the first tergite in trigonotarbids is usually a morphology can be found in the redescriptions modified into a so-called locking ridge which of Dunlop (1995a, b) and Ro¨ ssler & Dunlop tucked into a fold at the back of the carapace (1997) respectively. and thus ‘locked’ the prosoma and opisthosoma E v o l u t i o n a r y t r e n d s – Within this suite of together. Something similar is present in extant taxa we can detect an evolutionary trend to- ricinuleids. This first tergite seems to become re- wards increased structural complexity in the car- duced in more derived trigonotarbids (Dunlop apace. This is reflected in the appearance of a 1994b), thus it is rarely detected in Carbonifer- raised, central carapace region, present in Trigo- ous material – the present fossil included – be- notarbus Pocock, 1911 (Trigonotarbidae) and cause it is either: (a) hidden beneath the cara- Lissomartus Petrunkevitch, 1949 (Lissomartidae). pace and/or (b) reduced in size. Thus the first The next stage seems to have been the appear- tergite visible in our fossil is recognised as tergite ance of subtriangular lateral lobes flanking this 2. central carapace region. The earliest stages of Furthermore, some trigonotarbids have the this were documented by Dunlop (1995a, figs 3, second and third tergites fused together into a 7) in Lissomartus, where there are faint lateral diplotergite (2 þ 3) (e.g. Shear et al. 1987; Dun- divisions on the flanks of the carapace. At a lop 1996b; Fayers et al. 2005). This condition is more derived grade, these carapace lobes on the usually easily recognisable as a noticeably larger flanks appear to have become more strongly de- tergite at the front of the opisthosoma. This is fined. Exactly this stage of organisation is re- not the case in our fossil, in which the anterior- flected in the Alken species, Archaeomartus levis most tergites are all approximately the same (Trigonotarbidae), as figured by Brauckmann length. For this reason we suspect that tergal fu- (1987b, figs 3–4), and our new Hagen-Vorhalle sion – which is probably homoplastic for Trigo- fossil (Figs 1–3), which we consequently assign notarbida (see e.g. Dunlop 1995b for further ex- to the same genus (see Systematic Palaeontol- amples of unfused tergites) – is not present in ogy). Provisionally, the next level of organisation our fossil and we can simply number the eight – perhaps deriving from the Archaeomartus con- tergites present as 2–9. This fits studies of other dition – would be the appearance of larger, trigonotarbids in which the posteriormost tergite much more heavily ornamented with is invariably the ninth. This tergite 9 is divided in dense pustules or tubercles covering the cara- some taxa, undivided in others. Shear (2000) dis- pace and dorsal opisthosoma. Such ornamenta- cussed the polarity and significance of this char- tion is characteristic for the eophrynid-kreischer- acter. In our fossil the ninth tergite appears to iid-aphantomartid group (which we here term be undivided. the ‘eophrynid assemblage’), and could well be synapomorphic for them.

Discussion

There is a lineage of trigonotarbids with sub- Systematic palaeontology triangular carapaces, beginning with the oldest known (Silurian) species (Dunlop 1996a). By Order Trigonotarbida Petrunkevitch, 1949 contrast, other trigonotarbids have more quad- Family ? Trigonotarbidae Petrunkevitch, 1949 rate or rounded carapaces and, in some species, these retain an additional pair of lateral eye tu- R e m a r k s. Trigonotarbidae currently comprises bercles (Shear et al. 1987; Fayers et al. 2005); al- two genera: Trigonotarbus and Archaeomartus. most certainly the plesiomorphic character state. However, since a strongly lobed carapace occurs The ‘triangular-carapace’ trigonotarbids, which only in Archaeomartus and the ‘eophrynid as- have lost their lateral eyes, may represent a de- semblage’ (see above), we must entertain the rived (monophyletic) lineage. Provisionally, this possibility that Trigonotarbidae is paraphyletic. group comprises the Silurian plesion genus and Archaeomartus could be sister-group to the ‘eo- the families Trigonotarbidae (including our new phrynid assemblage’ based on the synapomorphy species), Lissomartidae, Eophrynidae, Kreischer- of a strongly lobed carapace. Yet the first stages iidae and Aphantomartidae; although in the last of lobation occur in Lissomartus, a genus which the carapace seems to have reversed to a less tri- also shares some features with our new Archaeo- angular outline (see e.g. figures in Ro¨ ssler 1998). martus species, like the strongly defined carapace Details of lissomartid, eophrynid and kreischeriid margin. In terms of the degree of lobation, Lis-

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim museum-fossilrecord.wiley-vch.de Fossil Record 9(1) (2006) 135 somartus also seems to be intermediate between (1911) and Petrunkevitch (1949). Based on simi- Trigonotarbus and Archaeomartus. Clearly, more larities with A. levis, we refer our new fossil to work is required to resolve relationships among Archaeomartus as redefined above. these genera, but a phylogeny of the form (Trigonotarbus (Lissomartus (Archaeomartus þ ‘eophrynids’))) is conceivable based on car- Archaeomartus roessleri n. sp. apace characters; which would render Trigono- Figs 1–3. tarbidae paraphyletic. Indeed the unpublished thesis of Dunlop (1994b) proposed raising Ar- 2003 Trigonotarbus johnsoni – Brauckmann et al.: 48, text- chaeomartus to a new family; albeit for slightly fig. 19, pl. 10., fig. 1. (misidentification) 2005 Trigonotarbus johnsoni – Brauckmann & Scho¨ llmann: different reasons to those outlined here. We pre- 71, fig. 2. (misidentification) fer not to create monogeneric families at this D e r v a t i o n o f n a m e. For our colleague Dr Ronny Ro¨ ss- stage pending careful restudy of the three species ler (Chemnitz), in recognition of his work on trigonotarbids currently assigned to Trigonotarbus (cf. Pocock and late Palaeozoic terrestrial ecosystems in general. 1911; Petrunkevitch 1955; Schultka 1991) and a H o l o t y p e a n d o n l y s p e c i m e n . WMfN P21 046; almost more thorough cladistic analysis of Trigonotarbi- complete specimen in dorsal view. da. Diagnosis. Archaeomartus with three lobes forming the central carapace region, divided from each other by a Y- shaped suture; legs relatively short, leg length about equal to or less than the width of the opisthosoma. Archaeomartus Størmer, 1970. Ty p e l o c a l i t y. Hagen-Vorhalle, Germany. Ty p e - h o r i z o n . Vorhalle beds, goniatite subzone R2c, Type species. Archaeomartus levis Størmer, 1970 (by ori- Upper Carboniferous (Pennsylvanian), Namurian B, late ginal designation). Marsdenian (ca. 319 Ma). Additional species. Archaeomartus tuberculatus Stør- D e s c r i p t i o n. Body squat and compact, lacking mer, 1970, Archaeomartus roessleri n. sp. spines, tubercles or other ornament. Total length D i a g n o s i s. Trigonotarbidae with a lobed, sub- ca. 6 mm. Carapace subtriangular, wider than triangular carapace and rounded opisthosoma, long, bluntly rounded anteriorly and with broad, wider than long; body lacking spines, dense tu- strongly demarcated lateral margins. Length berculation or other strong ornament (emended 2.3 mm, maximum width ca. 4 mm. Median area from Brauckmann 1987b). of carapace raised and divided into three princi- Remarks. Archaeomartus was originally raised pal lobes. Anteriormost lobe bears a single pair by Størmer (1970) for two species from the Early of median eyes, each located in a slightly de- Devonian of Alken an der Mosel, both described pressed socket. Behind this front lobe, a further from isolated, rounded opisthosomas and differ- pair of lobes occur, strongly demarcated from entiated from each other by the presence or ab- the anterior lobe and from each other by a Y- sence of marginal tuberculation. Brauckmann shaped depression. Lateral carapace regions be- (1987b) described a more complete specimen of tween the median area and the broad margin di- A. levis which included for the first time the car- vided on each side into three rectangular to sub- apace, which was found to have a distinctly triangular lobes, outlined by slightly procurving lobed morphology. This material also confirmed sutures. Posterior margin of carapace straight. that the opisthosoma is slightly wider than long All legs short and robust with thick, stubby arti- in A. levis which matches our new fossil, cles. Patella and tibia widening slightly distally. although as noted above further specimens may Basitarsus short, cylindrical. Tarsus bluntly be needed to exclude the possibility that this is rounded distally. Individual article lengths as fol- due to deformation in the Hagen Vorhalle mate- lows. Leg 3: femur 0.7, patella 0.6, tibia 0.8, basi- rial. In its carapace lobation and (perhaps) tarsus 0.3, tarsus 0.8 mm. Leg 4: femur 1.2, patel- opisthosomal proportions our new species la 0.7, tibia 0.9, basitarsus 0.3, tarsus at least matches A. levis, and can be excluded from the 0.7 mm. Opisthosoma rounded, slightly wider type species of Trigonotarbidae, Trigonotarbus than long. Length 3.7 mm, maximum width johnsoni, to which it was originally referred 4.5 mm. Median tergites decreasing successively (Brauckmann et al. 2003; Brauckmann & Scho¨ ll- in width posteriorly; lateral tergites preserved as mann 2005). T. johnsoni has smooth marginal fragments only. Posterior margins of all median carapace regions, a large diamond-shaped de- tergites straight. Eight tergites, probably repre- pression in the back of the carapace and a more senting segments 2–9, visible. Approximate circular opisthosoma; see e.g. figures in Pocock lengths: 2: 0.4, 3: 0.5, 4: 0.4, 5: 0.6, 6: 0.5, 7: 0.4, 8:

# 2006 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim 1museum-fossilrecord.wiley-vch.de 136 Dunlop, J. A. & C. Brauckmann, Trigonotarbid arachnid from the Coal Measures of Hagen-Vorhalle

0.5 and 9: 0.4 mm. Margins of five ventral sternites – 1996a. A trigonotarbid arachnid from the Upper Silurian and two segments constituting the small, circular of Shropshire. – Palaeontology 39: 605–614. – 1996b. Evidence for a relationship between pygidium, diameter ca. 0.3 mm. Sternites pro- Ricinulei and Trigonotarbida. – Bulletin of the British curved, especially posteriorly around pygidium. arachnological Society 10: 193–204. Fayers, S. R., Dunlop, J. A. & Trewin, N. H. 2005. A new early Devonian trigonotarbid arachnid from the Windy- field chert, Rhynie, Scotland. – Journal of Systematic Pa- Acknowledgements laeontology 2: 269–284. Giribet, G., Edgecombe, G. D., Wheeler, W. C. & Babbitt, C. 2002. Phylogeny and systematic position of : a We thank Lothar Scho¨llmann (WMfN) and Andrew Ross combined analysis of chelicerate relationships using mor- (NHM) for access to material in their care, Mrs G. Thomas phological and molecular data. – 18: 5–70. (WMfN) for preparing Fig. 1 and Paul Selden (Manchester) and Hendricks, A. (ed.) 2005. Als Hagen am quator lag. Die Ronny Ro¨ßler (Chemnitz) for helpful comments during review. Fossilien der Ziegeleigrube Hagen-Vorhalle. 222 pp., Landschaftsverband Westfalen-Lippe, Westfa¨lisches Mu- seum fu¨ r Naturkunde, Mu¨ nster. References Petrunkevitch, A. I. 1913. A monograph of the terrestrial Pa- laeozoic Arachnida of North America. – Transactions of the Connecticut Academy of Arts and Science 18: 1–137. Brauckmann, C. 1987a. Neue Arachniden (Ricinuleida, Tri- – 1949. A study of Palaeozoic Arachnida. – Transactions of gonotarbida) aus dem Namurium B von Hagen-Vorhalle the Connecticut Academy of Arts and Science 37:69– (Ober-Karbon, West-Deutschland). – Dortmunder Bei- 315. tra¨ge zur Landeskunde, Naturwissenschaftliche Mitteilun- – 1955. Trigonotarbus arnoldi, a new species of fossil ara- gen 20: 97–109. chnid from Southern France. – Journal of Paleontology – 1987b. 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