2002. The Journal of Arachnology 30:211±218

THE TRIGONOTARBID ARACHNID ANTHRACOMARTUS VOELKELIANUS (ANTHRACOMARTIDAE)

Jason A. Dunlop: Institut fuÈr Systematische Zoologie, Museum fuÈr Naturkunde der Humboldt-UniversitaÈt zu Berlin, Invalidenstraûe 43, D-10115 Berlin, Germany. E-mail: [email protected] Ronny RoÈûler: Museum fuÈr Naturkunde, Theaterplatz 1, D-09111 Chemnitz, Germany

ABSTRACT. Anthracomartus voelkelianus Karsch 1882 from the Pennsylvanian (Langsettian) of Nowa Ruda, Poland was listed in a 1953 monograph by Petrunkevitch as an incertae sedis species with type material possibly in Dresden. Antharcomartus voelkelianus is the type species of the genus Anthracomartus Karsch 1882 and historically one of the ®rst described examples of the extinct order Trigonotarbida. It is a pivotal species for resolving the systematics of both Anthracomartus and a number of poorly de®ned, probably congeneric, taxa within Anthracomartidae. Karsch's ®gured types were overlooked by Petrunk- evitch, but have been traced to a repository in Berlin and are redescribed here. Additional type material from Dresden and Wrocøaw could not be traced. One of Karsch's ®gured Berlin specimens is regarded here as the holotype of A. voelkelianus, but his other ®gured fossil is evidently not conspeci®c and is tentatively referred here to Trigonotarbus sp. (Trigonotarbidae). Keywords: Trigonotarbida, Anthracomartidae, fossil, Pennsylvanian, Poland, systematics

Trigonotarbida is a group of diverse Pa- species, A. granulatus Fritsch 1904, based on laeozoic arachnids recorded from late Silurian some of Karsch's material? Petrunkevitch to early Permian strata, but occurring most (1953) overlooked repository data in the lit- frequently in the Coal Measures of Europe erature, missed the opportunity to study at and North America. Anthracomartus voelke- least some of the relevant fossils, and consid- lianus Karsch 1882 was described from Penn- ered both these species to be incertae sedis. sylvanian age rocks of Silesia (SW Poland) In this paper we aim to identify Karsch's orig- and is signi®cant as the type species of An- inal material and to redescribe the available thracomartus Karsch 1882, itself the type ge- type material of A. voelkelianus. nus of Anthracomartidae Haase 1890. The PREVIOUS WORK systematics of this family are poorly re- Original descriptions.ÐKarsch (1882) de- solvedÐsee e.g. Dunlop & Horrocks scribed new fossil arachnids from the Silesian (1996)Ðmostly due to Fritsch (1901, 1904) Coal Measures, above the 7th seam of the and Petrunkevitch (1945, 1949, 1953) erecting `Rubengrube, bei Neurode, Schlesien' (ϭ both genera and species based on what appear Ruben mine, near Nowa Ruda, Silesia). This to be super®cial and/or preservational differ- locality is in southern Poland, SE of Wrocøaw ences. Anthracomartus voelkelianus is poten- near the border with the Czech Republic. tially the senior synonym of some of the more Stratigraphically, the fossils come from the dubious anthracomartid species and restudy of `Schatzlarer Schichten' (ϭ the ZÏ aclerÏ forma- the type material is a necessary starting point tion). Karsch established Anthracomartus for a revision of the Anthracomartidae. There voelkelianus as a new genus and species re- are, however, discrepancies between Karsch's ferred to a new, extinct arachnid order, An- (1882) two ®gured specimens and between thracomarti. Karsch named the fossils after Karsch's and Haase's (1890) ®gures of what Mr. VoÈlkel, the pit foreman and collector of is ostensibly the same fossil. This raises the the specimens. He noted that the `small num- following questions: were Karsch's original ber' of specimens were in the possession of a specimens conspeci®c, and was Fritsch's new Mr. Schumann in Dresden, and that they were

211 212 THE JOURNAL OF ARACHNOLOGY made available to Karsch by Prof. Dames and also Silesian, and was described as being from Mr. Weiss. A repository for these fossils was `outside Bohemia'. Anthracomartus granula- not given. tus was differentiated from A. voelkelianus on Anthracomartus voelkelianus was brie¯y the grounds that it was shorter and wider with mentioned by Scudder (1884) and was sub- very clear granulation. Both species were list- sequently redescribed and reinterpreted by ed in the monographs of Pocock (1911) and Haase (1890) who, in contrast to Karsch, iden- Petrunkevitch (1913). ti®ed dorsal and ventral surfaces. Haase In a review of the Pennsylvanian arachnids (1890, p. 645) stated (correctly) that the two from Silesia, Schwarzbach (1935) mentioned specimens ®gured by Karsch were in the col- a Westphalian age for the Rubengrube type lection of the Geological Survey (formerly the locality. Schwarzbach noted that Karsch's KoÈniglich-Preuûische Geologische Landesan- type series of A. voelkelianus was not in stalt) of Berlin and were lent to him (i.e., Haa- Wrocøaw (formerly Breslau), except for the se) by Dames, who was director of the Ge- counterpart of the one ®gured by Karsch ology-Palaeontology Institute of the Museum (1882, ®g. 2). Schwarzbach cited a repository fuÈr Naturkunde (MfN) and who also worked number, No. 556, for this specimen in the freely for the Survey (W. Lindert, pers. Geological Institute of Wrocøaw, which im- comm.). Furthermore, Haase (p. 646) stated plies that Karsch's original material was di- that the `Gegendruck' (ϭ counterpart) of vided between Berlin, Dresden and Wrocøaw. Karsch's ®g. 1 was in the Mineralogical Mu- Schwarzbach also noted that it was uncertain seum of Dresden, which implies that the type whether the specimens in Karsch's two ®gures series was divided between at least two insti- belonged together. Presumably he was ques- tutions (see also below). Haase (1890, pl. 30, tioning whether they were conspeci®c, since ®g. 9) claimed to have ®gured the Berlin spec- the literature already implied that each of imen (the part) of Karsch's ®g. 1, but whereas Karsch's ®gured specimens consisted of a part Karsch's illustration shows a fossil with a and counterpart and that the specimen in quadrate carapace and a leg, Haase's shows Karsch's ®g. 1 ended up in Berlin and Dres- one with a more rounded carapace and no leg. den (Haase 1890) while Karsch's ®g. 2 ended In his monograph of Paleozoic arachnids, up in Berlin and Wrocøaw. Schwarzbach also Fritsch (1904) included an inverted copy of noted another Wrocøaw specimen (no. 555) as Karsch's (1882, ®g. 1) illustration of A. voelk- having been collected by VoÈlkel, further sup- elianusÐin Fritsch's version the leg is on the porting the idea that the original Rubengrube left sideÐand noted that the cuticle of this material ended up in more than one institu- species is ®nely granulated. Note that the tion. Although originally labelled as A. voelk- Czech author Anton FricÏ is sometimes cited elianus, based on its wide body Schwarzbach under this Czech spelling of his name but, like referred no. 555 to A. granulatus. many non-Germans in the Austro-Hungarian Petrunkevitch's monographs.ÐPetrunk- empire, published the papers mentioned here evitch (1949) recognized the signi®cance of A. under the Germanized spelling `Fritsch'. voelkelianus as the type species of Anthraco- Fritsch (1904, p. 40) also created a new spe- martus, and discussed the differences between cies, the somewhat broader A. granulatus, Karsch's and Haase's interpretations (see also based on material in Dresden which he im- above). He concluded that the specimen plied was described as A. voelkelianus, i.e., matching Karsch's (1882, ®g. 1) must be re- `. . . ein Exemplar das A. VoÈlkelianus Fig. 2. garded as the holotype and questioned wheth- bezeichnet war...'. Fritsch based his new er Karsch's two specimens were conspeci®c species and the reconstruction (his ®g. 48) on and if Karsch (®g. 1) and Haase (pl. 30, ®g. a number of specimens, but this reference to 9) had actually ®gured the same fossil. Pe- `Fig. 2' is confusing. If could mean the ®g. 2 trunkevitch did not study the original material, of Karsch's plate, but this particular fossil is and remarked that he was unable to obtain in Berlin (see below). It could refer to the permission to visit Dresden during a post-war counterpart of Karsch's ®g. 2 specimen, but tour of European museums. Correspondence this specimen has been reported from in the MfN, Berlin reveals that in 1951 Pe- Wrocøaw (see below). No locality for A. gran- trunkevitch wrote to Alfred KaÈstner (then at ulatus is stated, but the material is probably the MfN) to ask if someone from Berlin could DUNLOP & ROÈ ûLERÐTRIGONOTARBID ANTHRACOMARTUS VOELKELIANUS 213 visit Dresden and establish the identity of the pository numbers 09446 (Karsch's ®g. 1) and type from the material there seen by Karsch; 09447 (Karsch's ®g. 2). The only difference himself a former curator in the MfN, Berlin. is that in the original plates the specimens Curiously, Petrunkevitch did not ask KaÈst- were drawn on larger, squarer slabs of matrix ner about the Karsch types cited as being in with associated plant material. The actual Berlin and Wrocøaw. It appears that Petrunk- slabs (Figs. 1, 2) are smaller and irregular and evitch simply overlooked the repository de- do not look to have been trimmed, thus it ap- tails in Haase (1890) and Schwarzbach (1935) pears that a certain amount of artistic licence and assumed that all the types of both A. was used in the illustrations (W. Lindert, pers. voelkelianus and A. granulatus were in Dres- comm.). This is not to say that the drawings den (see also Petrunkevitch 1953), as implied of the animals themselves are inaccurate (they by a cursory reading of both Karsch (1882) are actually very good) only that there is a and Fritsch (1904). The unfortunate irony is discrepancy concerning the matrix. that the specimens ®gured by Karsch were all Specimens 09446 and 09447 are the only the time in the Geological Survey of Berlin examples of A. voelkelianus in the BGR, Ber- (see below), a building located at Invaliden- lin collection. The building of the `Geologis- straûe 44, adjacent to KaÈstner in the MfN (In- che Landesanstalt' was damaged in the war, validenstraûe 43). KaÈstner contacted the material was lost (W. Lindert, pers. comm.) `Staatliches Museum fuÈr Tierkunde', Dresden, and thus it is possible that additional speci- but the paleontology department there was un- mens were originally present. Neither speci- able to locate types of either species. Conse- men exactly matches Haase's (1890. pl. 30, quently, Petrunkevitch (1953, p. 68) regarded ®g. 9) illustration, supposedly also of 09446, A. voelkelianus as an incertae sedis species, and it remains unclear what Haase actually citing it as `Carboniferous of Silesia. (In Dres- drew. Although Petrunkevitch (1949) tended den?)'. In fact the entire genus Anthracomar- towards the idea that Haase drew a different tus became an incertae sedis taxon as a result fossil (perhaps now lost), we suspect that Haa- of Petrunkevitch's (1953) revision, although se's ®gure could also be based on 09446, but Petrunkevitch (1955a) appeared to revalidate with the leg omitted and a different emphasis the taxon in the Treatise on Invertebrate Pa- to Karsch's, more accurate, version. leontology, listing it among the other anthra- Unfortunately, the material cited by comartid genera but without any detailed dis- Schwarzbach in Wrocøaw could not be traced. cussions. An outline drawing based on Martin Schwarzbach was associated with the Karsch's (1882, ®g. 1) was included by Pe- geological institute of the University of trunkevitch (1955a), but since then there has Wrocøaw before the war and later material been no further mention of A. voelkelianus in from this institute passed to the Muzeum Geo- the literature. logiczne (Geological Institute, Wrocøaw Uni- versity: Cybulskiego 30, 50±205 Wrocøaw: A. METHODS Pacholska, pers. comm.). However, a lot of The KoÈniglich-Preuûische Geologische material is known to have been lost during the Landesanstalt is now included in the Bunde- war, no inventory books were preserved and sanstalt fuÈr Geowissenschaften und Rohstoffe no fossil arachnids could be traced in the Mu- (BGR), Hannover (branch Berlin) and in 1996 zeum Geologiczne collections (A. Pacholska, the collections were moved from Invaliden- pers. comm.). The Wrocøaw material, includ- straûe to a new repository at Wilhelmstraûe ing the possible counterpart of Karsch's ®g- 25±30, D-13593 Berlin. Both the type cata- ured specimen, therefore appears to be lost. logues of Dienst (1928, p. 125), and more re- Furthermore, the material cited by Haase cently Daniels et al. (1998, p. 39), con®rm (1890) as being in Dresden could not be that Karsch's material is present in this col- traced either. In the State Museum of Miner- lection. alogy and Geology of Dresden there are sev- The BGR, Berlin fossils consist of the two eral slabs with plant fossils from the Nowa specimens actually ®gured by Karsch (1882) Ruda site collected at the end of the 19th cen- in his original description. Both match tury. No arachnid remains could be found in Karsch's original illustrations, i.e. the append- them (L. Kunzmann, pers. comm.). During ages are on the correct side, and have the re- World War II the complete Paleozoic collec- 214 THE JOURNAL OF ARACHNOLOGY tion was moved to Pillnitz for safe keeping, SYSTEMATIC PALEONTOLOGY thus loss of specimens during that time ap- pears unlikely. Order Trigonotarbida Petrunkevitch 1949 Anthracomartus voelkelianus was com- Remarks.ÐPetrunkevitch (1949) divided pared to other anthracomartid material, prin- Anthracomarti into two orders: Anthracomar- cipally in the Natural History Museum, Lon- tida and Trigonotarbida. The features he used don (BMNH) and the National Museum to separate these taxa were rejected by Dunlop Prague (NMP). The types of A. buchi (Gol- (1996) as either misinterpretations of the fos- denberg 1873) and A. hageni (Goldenberg sils or as insuf®cient grounds for maintaining 1873)Ðboth poorly preserved, isolated tri- distinct orders. They were reunited under the gonotarbid opisthosomasÐwere examined in more clearly de®ned Trigonotarbida, with An- the palaeontological institute of the University thracomartidae representing one family, diag- of Bonn (repository numbers IPB GuthoÈrl 5 nosed on tergites divided laterally into 5 plates & 6), but the type of A. granulatus could not as opposed to 3 plates in all other trigonotar- be found in Dresden (L. Kunzmann, pers. bids (see Dunlop 1996 for further discus- comm.). Specimens were drawn under alcohol sions). using a camera lucida. All measurements are in mm. Family Anthracomartidae Haase 1890 Remarks.ÐFollowing Petrunkevitch GEOLOGICAL SETTING (1953, 1955a), the family Anthracomartidae The type material of A. voelkelianus comes includes nine valid genera. Two of these, Bra- from the Lower Silesian coal basin. This chypyge Woodward 1878 (known only from makes up a portion of the Intra-Sudetic basin, an opisthosoma) and Maiocercus Pocock the largest geological unit in the Middle Su- 1911 are both distinct in having a scalloped detes. The Pennsylvanian sedimentary succes- opisthosomal margin (see also Dunlop & Hor- sion of the Intra-Sudetic basin is distinctly dif- rocks 1996). Most of the remaining genera: ferentiated into several complexes of different Promygale Fritsch 1901, Brachylycosa Fritsch clastic material composition, color and pale- 1904, Coryphomartus Petrunkevitch 1913, ontological inventory (Bossowski et al. 1995). Pleomartus Petrunkevitch 1945, Cleptomartus The ®rst lithostratigraphic subdivision was Petrunkevitch 1949, Cryptomartus Petrunk- proposed in the 19th century and was based on evitch 1949 and Oomartus Petrunkevitch informal mining terminology. The sediments 1953 were based on specimens which were of the ZÏ aclerÏ formation, from which the A. either originally, or at some stage (Pocock voelkelianus fossils were obtained, were ac- 1910), referred to Anthracomartus;anincer- cumulated under a ¯uviolacustrine regime and tae sedis taxon in Petrunkevitch's (1953) represent sub-environments of river channels scheme. In his key in this paper, Petrunkevitch diagnosed the anthracomartid genera based of low and high sinuosity. In the vicinity of mostly on carapace morphology, but provi- Nowa Ruda the lower (Langsettian age) por- sional work suggests that many of the sup- tion of the ZÏ aclerÏ formation is dominated by posed differences between the carapaces are ®ne-grained clastics that contain as many as preservational artifacts, typically based on 20 coal seams. The origin of the coal-bearing missing features and strongly in¯uenced by sequence, whose thickness reaches 160 m, whether the fossils were compressed to a was peat swamp development in extended al- greater or lesser extent in shales or preserved luvial plains. The upper (Duckmantian age) more three-dimensionally in ironstone concre- Ï portion of the ZaclerÏ formation is as much as tions. The taxonomy of the ¯attened NyÂrÏany 230 m thick and is composed primarily of material in Prague in especially suspect and coarse-grained clastics. Today it is dif®cult to includes specimens, probably identi®ed by Pe- say exactly where the 7th seam of the 1882 trunkevitch, (NMP A/165b & A/22b) where terminology should be placed within the se- the part and counterpart have been assigned quence. All indications point to the lower ZÏ a- to different genera. This study revalidates An- clerÏ formation (upper Langsettian age, [ϭ thracomartus, rediagnosed below based on the Westphalian A]) in recent terminology. redescription of the genotype. Anthracomartus DUNLOP & ROÈ ûLERÐTRIGONOTARBID ANTHRACOMARTUS VOELKELIANUS 215

Figures 1±2.ÐPhotographs of Karsch's fossils housed in the BGR, Berlin. Both from the Pensylvanian (Langsettian) of the Ruben mine, Nowa Ruda, Intra-Sudetic basin, Poland. 1, No. 09446, holotype of A. voelkelianus; 2, No. 09447, a specimen which is not the counterpart of, nor even conspeci®c with, 09446 and which is referred here to? Trigonotarbus sp. Scale ϭ 5 mm.

is potentially the senior synonym of some of Anthracomartus voelkelianus Karsch 1882 these poorly diagnosed anthracomartid genera. Figs. 1, 3 Anthracomartus VoÈlkelianus Karsch 1882: 560± Anthracomartus Karsch 1882 561, pl. 21, ®g. 1; Scudder 1884:14, 17; Haase 1890: 645±646. Pl. 30, ®gs. 8, 9; Fritsch 1904: Type species.ÐAnthracomartus voelkeli- 40, ®g. 47. anus Karsch 1882 Anthracomartus voÈlkelianus Karsch: Pocock 1911: Included species.ÐAnthracomartus gran- 3±4, 63; Schwarzbach 1935: 5; Petrunkevitch ulatus Fritsch 1904, A. buchi (Goldenberg 1949: 195Ð198, ®gs. 192, 193; Petrunkevitch 1873) and A. hageni (Goldenberg 1873), the 1953: 58, 68; Petrunkevitch 1955a: 107, ®g. 67 latter two species both nomina dubia. (1). Emended diagnosis.ÐAnthracomartids Anthracomartus voelkelianus Karsch: Petrunkevitch with a smooth opisthosomal margin, lacking 1913, pp. 94, 99. the marginal scalloping seen in Brachypyge Material.ÐBGR, Berlin No. 09446 (Ho- and Maiocercus. The status of the remaining lotype). From the `Rubengrube', Nowa Ruda, anthracomartid genera (see above) is question- Intra-Sudetic basin, Poland. Pennsylvanian able and merits revision. (Langsettian). Not BGR, Berlin No. 09447 Remarks.ÐThe two Goldenberg species (see below). were referred to Anthracomartus by GuthoÈrl Emended diagnosis.ÐCarapace with (1934), but they are based on poor specimens slightly bilobed anterior region divided by (IPB GuthoÈrl 5 & 6) which are effectively un- median sulcus. Opisthosoma broadly oval, al- identi®able (Petrunkevitch 1953). Both are re- most circular in outline, but slightly longer garded here as nomina dubia. To date, we than wide and widest midway along its length. have been unable to trace the type of A. gran- Based on published descriptions, the opistho- ulatus, supposedly in Dresden (see above), soma of A. granulatus is wider than long. thus we have been unable to con®rm its ge- Description.ÐOnly carapace, opisthosoma neric af®nities. and one leg preserved in dorsal view with 216 THE JOURNAL OF ARACHNOLOGY

Figures 3±4.ÐCamera lucida drawings of the specimens shown in Figs. 1±2. Abbreviations: cx ϭ coxa, et ϭ probable median eye tubercle, fe ϭ femur, lb ϭ anterior lobe on carapace, li ϭ limb, pp ϭ pedipalp, py ϭ pygidium, sm ϭ sternum, st ϭ sternites (superimposed through onto dorsal surface in Fig. 3), tr ϭ trochanter. Scale ϭ 5 mm. some ventral features superimposed posteri- sion of most tergites into 5 plates with median orly (Figs. 1, 3). Total length 18.0. Whole plate wider than lateral plates, clearly visible, specimen with a somewhat granular appear- at least anteriorly. Posterior segmentation less ance, in places resolved into a distinct pattern distinct because ventral elements (sternites of tiny tubercles. Carapace subquadrate, 6.1 which are distinctly angled on the midline) are long, 6.6 wide, slightly rounded at anterolat- superimposed. Outline of circular pygidium eral corners. Carapace with slight relief, but (ventral, diameter 1.0) impressed through onto margins on all sides irregular and full depth dorsal surface. of carapace obscured within matrix. Carapace Remarks.ÐKarsch (1882) did not desig- with somewhat bilobed appearance in anterior nate a type from among his `small number' of half; lobes approximately symmetrical. Pos- specimens which implies a series of syntypes. terior half of carapace somewhat depressed. However, since only two of these fossils have Carapace lacks lateral eye tubercles and pro- been positively identi®ed, and since they ap- jecting anterior clypeus seen in more three- pear not to be conspeci®c (see below), we re- dimensionally preserved anthracomartids gard the Berlin specimen corresponding to (Dunlop 1996; Dunlop & Horrocks 1996), but Karsch's ®g. 1 as the holotype; see also com- small, raised area near anterior margin consis- ments by Petrunkevitch (1949, p. 198). If tent with median eye tubercle. Single, almost more of the original Rubengrube material is complete, but rather poorly preserved, leg oc- subsequently identi®ed and con®rmed to be A. curs on right side, total length c. 9. Individual voelkelianus then BGR, Berlin No. 09446 podomeres indistinguishable, probably leg IV may have to be redesignated as a lectotype. judging from its position. An adequate diagnosis of both the genus Opisthosoma broadly oval, slightly longer and species is dif®cult without a revision of (11.7) than wide (11.2). Cuticle preserved as the anthracomartids and an assessment of the dark regions (best seen under alcohol) but characters used to de®ne taxa. Despite their preservation patchy in posterior region and to- apparent diversity in the literature, anthraco- wards margins of opisthosoma. Characteristic martid fossils are morphologically rather ho- anthracomartid tergite pattern, including a mogeneous. Furthermore, we are cautious large diplotergite (segments 2 ϩ 3) and divi- about relying too heavily on reduction (e.g., DUNLOP & ROÈ ûLERÐTRIGONOTARBID ANTHRACOMARTUS VOELKELIANUS 217 absence of eyes or ornament) and/or propor- ved (Figs. 3, 4) and overall this fossil, with an tion-based characters (e.g., length±width ra- apparently triangular prosoma ending in a tios) in material which has potentially been bluntly rounded `snout', is much more like ex- distorted during preservation. That said, pro- amples of the family Trigonotarbidae; com- visional observations of other anthracomartid pare with ®gures in Pocock (1911) and Pe- fossils suggest that gross morphological car- trunkevitch (1949, 1955b). BGR 09447 is apace ornamentation and the shape of the op- tentatively referred to the genus Trigonotar- isthosoma (i.e., circular, oval or pear-shaped) bus. In size and general shape BGR 09447 may be useful characters. This information resembles the French Stephanian species Tri- has therefore been used in the diagnosis gonotarbus arnoldi Petrunkevitch 1955b, but above. since the Rubengrube fossil is only incom- pletely known from a rather poorly preserved Trigonotarbidae Petrunkevitch 1949 ventral surface we are reluctant to assign it to ?Trigonotarbus sp. Pocock 1911 a species. Figs. 2, 4 Anthracomartus VoÈlkelianus Karsch 1882: 560± ACKNOWLEDGMENTS 561, pl. 21, ®g. 2. We thank Wolfgang Lindert (BGR, Berlin) Material.ÐBGR, Berlin No. 09447. From for providing specimens in his care and valu- the Rubengrube, Nowa Ruda, Poland. Penn- able information on the history of the BGR sylvanian (Langsettian). Reported counterpart material, Lutz Kunzmann (State Museum of (Geologisches Institut in Breslau [ϭ Mineralogy and Geology of Dresden) for in- Wrocøaw], No. 556) missing, presumed lost. formation on their collections and Antonina Description.ÐIncomplete specimen pre- Pacholska (Muzeum Geologiczne, Wrocøaw) served in ventral view. Total preserved length for information on the Breslau University ma- 16.6. Prosoma subtriangular, converging to terial. Stephan Shultka (MfN, Berlin) and An- blunt point anteriorly. Mouthparts not pre- drzej Wiktor (Wrocøaw) helped track down re- served, but a series of subtriangular coxae, in- positories. We thank the reviewers for creasing in size from anterior to posterior, sur- comments on the manuscript and Martin rounds relatively large, heart-shaped sternal Sander (IPB), Andrew Ross (BMNH) and Vo- element. Proximal limb elements present, best jtech Turek (NMP) for access to material in preserved on left side, but incomplete. Pedi- their collections. palps poorly preserved. All legs robust with somewhat rounded trochanters and short prox- LITERATURE CITED imal podomeres; probably femora and patel- Bossowski, A., A. Ihnatowicz, K. Mastalerz, L. Ku- lae, although podomere boundaries mostly not rowski & G.J. Nowak. 1995. Intra-Sudetic De- clearly de®ned (Fig. 4). Opisthosoma incom- pression. Pp. 142±147, In The Carboniferous plete, but preserved outline suggests it was al- System of Poland. (A. Zdanowski & H. Zakova, most circular (diameter c. 9.5.) in life; poste- eds.). Polish Geological Institute, Warszawa. rior, including pygidium, missing. At least ®ve Daniels, C.H.v., A. Heinke, C. Heunisch, W. Lin- dert & T. Wiese. 1998. Wissenschaftliche Origin- opisthosomal sternites visible, all lacking or- ale in den Sammlungen BGR/NLfb, Hannover namentation and gently procurved with angle und BGR, Berlin. Bundesanstalt fuÈr Geowissen- of procurvature increasing posteriorly. schaften und Rohstoffe, Berlin. Remarks.ÐAs previous authors have hint- Dienst, P. 1928. Zusammenstellung der im Geolo- ed, this fossil cannot be the counterpart of gischen Landesmuseum zu Berlin aufbewahrten 09446 (described above), nor is it conspeci®c Originale. I. PalaÈozoologischer Teil. Preuûische with it. The proportions of the prosoma and Geologische Landesanstalt, Berlin. opisthosoma, and of the appendages, are sig- Dunlop, J.A. 1996. Systematics of the fossil arach- ni®cantly different (compare Figs. 1 & 3 with nids. Revue suisse de Zoologie, hors seÂrie:173± 2 & 4). BGR 09447 cannot even be included 184. Dunlop, J.A. & C.A. Horrocks. 1996. A new spec- in Anthracomartidae, a family which charac- imen of the Carboniferous trigonotarbid arachnid teristically preserves a number of sharply an- Maiocercus celticus (Pocock 1902) from Lanca- gled ventral sternites in front of the pygidium shire, UK. Proceedings of the Yorkshire Geolog- (see also above; Fig. 2). The corresponding ical Society 51:23±31. sternites in BGR 09447 are smoothly procur- Fritsch, A. 1901. Fauna der Gaskohle und der Kalk- 218 THE JOURNAL OF ARACHNOLOGY

stein der Permformation BoÈhmens, vol. 4, fasc. Petrunkevitch, A.I. 1955a. Arachnida. Pp. 44±175, 2, pp. 56±63 (sometimes cited as 1899). In Treatise on Invertebrate Paleontology, Pt. P, Fritsch, A. 1904. Palaeozoische Arachniden. Pri- Arthropoda 2 (R. C. Moore, ed.). Geological So- vately published with support from the Imperial ciety of America and University of Kansas Press, Academy, Vienna. Dr. Eduard GreÂgr, Prague. Lawrence. Karsch, F. 1882. Ueber ein neues Spinnenthier aus Petrunkevitch, A.I. 1955b. Trigonotarbus arnoldi,a der schlesischen Steinkohle und die Arachniden new species of fossil arachnid from Southern der Steinkohlenformation uÈberhaupt. Zeitschrift France. Journal of Paleontology 29:475±477. der Deutschen geologischen Gesellschaft 34: Pocock, R.I. 1910. Notes on the morphology and 556±561. generic nomenclature of some Carboniferous Goldenberg, F. 1873. Fauna saraepontana fossilis. Arachnida. Geological Magazine, Decade 5, 7: Die fossilen Thiere aus der Steinkohlenformation 505±512. von SaarbruÈcken, SaarbruÈcken 1:12. Pocock, R.I. 1911. A monograph of the terrestrial GuthoÈrl, P. 1934. Die Arthropoden aus dem Karbon Carboniferous Arachnida of Great Britain. und Perm des Saar-Nahe-Pfalz-Gebietes (Arach- Monographs of the Palaeontographical Society, noidea, pp. 34±44). Abhandlungen der London, 84 pp. preuûische geologischen Landesanstalt, N.F. 164: Schwarzbach, M. 1935. Spinnentiere (Arachnoidea) 1±219. aus dem schlesischen Oberkarbon. Jahresberi- Petrunkevitch, A.I. 1913. A monograph of the ter- chten der Geologischen Vereinigung Oberschle- restrial Palaeozoic Arachnida of North America. siens 1935:1±8. Transactions of the Connecticut Academy of Arts Scudder, S.H. 1884. A contribution to our knowl- and Science 18:1±137. edge of Paleozoic Arachnida. Proceedings of the Petrunkevitch, A.I. 1945. Palaeozoic Arachnida of American Academy of Arts and Sciences 20:13± Illinois. An inquiry into their evolutionary 22. trends. Illinois State Museum, Scienti®c Papers 3(2):1±72. Woodward, H. 1878. Discovery of the remains of a Petrunkevitch, A.I. 1949. A study of Palaeozoic fossil crab (Decapoda Brachyura) in the Coal- Arachnida. Transactions of the Connecticut Measures of the environs of Mons, Belgium. Academy of Arts and Science 37:69±315. Geological Magazine, Decade 2, 5:433±436. Petrunkevitch, A.I. 1953. Paleozoic and Mesozoic Arachnida of Europe. Memoirs of the Geological Manuscript received 10 May 2001, revised 28 No- Society of America 53:1±122. vember 2001.