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Arachnida: Opiliones) from the Upper Carboniferous of Missouri, USA EUROPEAN ARACHNOLOGY 2003 LOGUNOV (D.V. & PENNEY D. eds.), pp. 6774. © ARTHROPODA SELECTA (Special Issue No.1, 2004). ISSN 0136-006X (Proceedings of the 21st European Colloquium of Arachnology, St.-Petersburg, 49 August 2003) A spiny harvestman (Arachnida: Opiliones) from the Upper Carboniferous of Missouri, USA Øèïàñòûé ñåíîêîñåö (Arachnida: Opiliones) èç ïîçäíåãî êàðáîíà Ìèññóðè, ÑØÀ J.A. DUNLOP Institut für Systematische Zoologie, Museum für Naturkunde der Humboldt-Universität zu Berlin, Invaliden- strasse 43, D-10115 Berlin, Germany. email: [email protected] ABSTRACT. A new fossil harvestman (Arachnida: Opiliones) is described from the Upper Carboniferous Coal Measures of western Missouri, USA. Echinopustulus samuelnelsoni gen. et sp.n. is tentatively assigned to the suborder Dyspnoi and shares features with both trogulid and, less convincingly, ceratolasmatid opilionids. This fairly long-legged fossil is only known from the dorsal surface which exhibits a pustulate ornament and a remarkable, autapomorphic, morphology of two pairs of large opisthosomal spines: an anterior, paramedian pair emerging from a single, central tubercle and a posterior pair positioned more laterally. Like the French fossil Eotrogulus fayoli Thevenin, 1901, this new material may provide further evidence that the Dyspnoi clade extends back to at least the late Carboniferous (c. 300 Ma). ÐÅÇÞÌÅ. Îïèñàí íîâûé èñêîïàåìûé ñåíîêîñåö (Arachnida: Opiliones) èç âåðõíåêàðáîíî- âûõ óãëåé çàïàäíîãî Ìèññóðè, ÑØÀ. Echinopustulus samuelnelsoni gen. et sp.n., êîòîðûé ïðåäâàðèòåëüíî îòíåñåí ê ïîäîòðÿäó Dyspnoi, íåñåò ïðèçíàêè êàê òðîãóëèä, òàê è â ìåíüøåé ñòåïåíè öåðàòîëÿñìèä. Ýòî îò÷åòëèâî äëèííîíîãîå èñêîïàåìîå èçâåñòíî òîëüêî ñ äîðçàëü- íîé ïîâåðõíîñòè, íà êîòîðîé èìååòñÿ ïóïûð÷àòûé îðíàìåíò è çàìåòíàÿ, àóòàïîðìîðôíàÿ, ñòðóêòóðà, ñîñòîÿùàÿ èç äâóõ ïàð êðóïíûõ îïèñòîìàëüíûõ øèïîâ: ïåðåäíåé, ïàðàìåäèàëü- íîé ïàðû, âûõîäÿùåé èç åäèíñòâåííîãî öåíòðàëüíîãî áóãîðêà, è çàäíåé ïàðû, ðàñïîëîæåí- íîé áîëåå ëàòåðàëüíî. Êàê è èñêîïàåìîå Eotrogulus fayoli Thevenin, 1901 èç Ôðàíöèè, íîâûé ìàòåðèàë îòîäâèãàåò íàõîäêè êëàäû Dyspnoi íàçàä, êàê ìèíèìóì äî ïîçäíåãî êàðáîíà (ïðèìåðíî 300 ìëí.). KEY WORDS: fossil, Carboniferous, taxonomy, harvestman, Dyspnoi, new species, USA. ÊËÞ×ÅÂÛÅ ÑËÎÂÀ: èñêîïàåìîå, êàðáîí, òàêñîíîìèÿ, ñåíîêîñåö, Dyspnoi, íîâûé âèä, ÑØÀ. Introduction kevitch, 1913], and from the slightly older east Kirkton site in Scotland [Wood et al., 1985]. Fossil harvestmen (Arachnida: Opiliones) Restudy of the extinct arachnid order Kusta- are rare. The oldest examples date back to the rachnida from Mazon Creek has indicated that Early Devonian [Dunlop et al., 2003] and a these fossils are misidentified harvestmen [Beall, small number of Carboniferous harvestmen have 1986; Dunlop, 2004]. By contrast, the entire also been described. These come from the Coal Mesozoic has yielded only two incompletely Measures of Commentry in France [Thevenin, described specimens, while the Tertiary has a 1901] and Mazon Creek in the USA [Petrun- much more diverse and better known fauna 68 EUROPEAN ARACHNOLOGY 2003 including specimens in shales [Cokendolpher Germany. This incomplete and not very well & Cokendolpher, 1982] and inclusions in am- illustrated fossil is somewhat consistent with a ber [e.g., Cokendolpher & Poinar, 1998; Starê- living trogulid in overall shape, but this assign- ga, 2002; Dunlop & Giribet, 2003]. All four ment is difficult to confirm from the description suborders sensu Giribet et al. [2002] are repre- and again the status of the holotype is uncertain. sented in Tertiary amber and most of these In this paper a new fossil arachnid is de- fossils can be assigned to Recent genera. scribed from the Coal Measures of western The Upper Carboniferous harvestman fau- Missouri, USA. In overall appearance this spiny na currently consists of the rare Commentry and specimen is quite different from previously de- Mazon Creek forms which can now be divided scribed Carboniferous arachnids and clearly among three genera: Eotrogulus Thevenin, merits a new taxon. It is interpreted as the 1901, Nemastomoides Thevenin, 1901 and Kust- second example of an Upper Carboniferous arachne Scudder, 1890. The position of these dyspnoid harvestman; albeit of uncertain affin- fossils within Opiliones remains problematic. ities within the Dyspnoi clade. Both Eotrogulus and Nemastomoides have been included in the superfamily Troguloidea of the Material and methods suborder Dyspnoi. These two fossil genera were raised to new families by Petrunkevitch [1955]; The fossil was acquired from a private deal- albeit on questionable characters relating to the er (Charles Isbon) by Dr Elliot Nelson (St Lou- coxo-sternal region which should not be part of is, USA), who recognized its significance and the harvestman ground pattern. Restudy is re- made it available for study to the author. The quired to confirm their familial status and Dun- fossil was reportedly collected from private lop [in press] compared both Kustarachne and land in western Missouri, USA, but the exact Nemastomoides to living Eupnoi; the group locality could not be traced. It may come from which includes the most common daddy-long- near the town of Windsor. However, a number legs type of harvestmen. of sites in western Missouri yield Coal Measure Dyspnoi are fairly abundant and widespread fossils similar to the famous Braidwood section today across the northern hemisphere, although of the Mazon Creek locality [B. Stinchcomb, they do tend to be predominantly soil animals pers. comm.] and in general these Missouri coal and are thus more cryptic than the more familiar deposits are not as extensive or as well re- Eupnoi. Modern-looking Dyspnoi assignable to searched as those of Illinois, which include the living genus Nemastoma have long been Mazon Creek. The Missouri fossils are pre- known from Tertiary amber [see e.g., Petrun- served in a similar way to Mazon Creek materi- kevitch, 1955], but the only reliable Palaeozoic al as impressions in clay-ironstone concretions. dyspnoid harvestman is the French fossil The sediments yielding the Missouri concre- Eotrogulus from the Coal Measures of Com- tions belong to the Pleasanton Group, which is mentry. From its description in the literature dated to upper Middle Pennsylvanian; equiva- Eotrogulus looks superficially like a member of lent to the Upper Carboniferous in European the trogulid clade, with a somewhat elongate, stratigraphy. A summary of the geological set- lozenge-shaped body in which the opisthosoma ting of this region can be found in Howe [1982]. is broadly joined to a subtriangular carapace. The fossils were studied under a stereomicro- The legs are quite long and slender, but nowhere scope with a camera lucida attachment used to near as extreme as in some phalangiid harvest- prepare the drawings. A fine needle was used to men. Eotrogulus was raised to a new, monotyp- prepare the anterior part of the carapace and also to ic family and potentially represents the oldest excavate the soft sediment infill from the spines on the back of the opisthosoma. Other Upper Carbonif- record of both Troguloidea and Dyspnoi. Un- erous harvestmen from Mazon Creek were obtained fortunately the status of the holotype is uncer- from the United States National Museum (USNM) tain. The only other fossil trogulid described in and the Peabody Museum, Yale (YPM). Of the non- the literature is Trogulus longipes Haupt, 1956 amber dyspnoid harvestmen, Eotrogulus fayoli Thev- from the Eocene Braunkohl of Geiseltal in enin, 1901 was reported from the Museum National J.A. Dunlop. A new Carboniferous harvestman 69 Figs 12. Echinopustulus samuelnelsoni gen. et sp.n., a new spiny harvestman from the Upper Carboniferous Coal Measures of western Missouri, USA. 1 part; 2 counterpart. Scale: 2 mm. Ðèñ. 12. Echinopustulus samuelnelsoni gen. et sp.n., íîâûé øèïàñòûé ñåíîêîñåö èç âåðõíåêàðáîíîâûõ óãëåé çàïàäíîãî Ìèññóðè, ÑØÀ. 1 îñíîâíàÿ ÷àñòü; 2 îáðàòíûé îòïå÷àòîê. Ìàñøòàá: 2 ìì. 70 EUROPEAN ARACHNOLOGY 2003 dHistoire Naturelle (MNHN) in Paris, but could Prosoma not be traced in the collections during a recent The carapace is subtriangular, but the ante- search [A. Rage, pers. comm.]. Similarly, the type of rior tip is, unfortunately, not preserved. Exca- Trogulus longipes should be in the Gieseltal Muse- um in Halle, Germany, but could not be traced [M. vation of the anterior end failed to reveal struc- Hellmund, pers. comm.]. The fossil material was tures such as eyes (which remain equivocal) or, also compared to extant harvestmen in the zoological for example, the characteristic bilobed hood collections of the Museum für Naturkunde, Berlin. seen in modern trogulid harvestmen. The cara- pace expresses a weak division or sulcus to- Results and discussion wards its posterior end. Although not a separate sclerite, this corresponds to the metapeltidium or last thoracic tergite sensu Giribet et al. [1999: Morphological interpretation character 21]. The median band and ornament The fossil (Figs 14) is preserved in a nod- here have been noted above. The mouthparts ule as a part and counterpart. Both show the and pedipalps are not preserved. The leg series dorsal surface of the animal, respectively, in is incomplete and only the proximal podomeres positive and negative relief. The latter pre- are preserved. They give the impression of serves more details of the morphology and or- slender, rather gracile legs in which the femora namentation. Descriptions are of the appear- widen distally; see the leg 1 femur
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