Pohlsepia Mazonensis, an Early 'Octopus' from the Carboniferous of Illinois
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POHLSEPIA MAZONENSIS, AN EARLY `OCTOPUS' FROM THE CARBONIFEROUS OF ILLINOIS, USA by JOANNE KLUESSENDORF and PETER DOYLE ABSTRACT. Pohlsepia mazonensis gen. et sp. nov. from the Mazon Creek Konservat LagerstaÈtte (Carboniferous) of Illinois is an exceptionally preserved soft-bodied fossil coleoid, with well-de®ned body and arms. Lacking an internal shell and possessing eight subequal and two modi®ed arms, Pohlsepia can be compared with both the living cirrate octopods and the decabrachian sepiardarids, both of which lack a well-developed internal skeleton. Given its sac-like body, lack of a well-de®ned head and presence of ®ns, Pohlsepia can be safely compared with modern cirrate octopods. It is the oldest known completely soft-bodied coleoid and as such has great signi®cance with respect to the phylogeny of the group, given that both the octobrachian and decabrachian clades have previously been thought to have evolved in the Jurassic. KEY WORDS: Coleoidea, Octobrachia, Konservat LagerstaÈtte, Mazon Creek, Carboniferous. T HE preservation of cephalopods with the soft parts intact is relatively uncommon in the fossil record, and is common only within the Coleoidea, a group which includes belemnoids and the familiar living squid, cuttle®sh and octopods. The majority of known records are of Mesozoic coleoids, and these have effectively been derived mostly from the Jurassic and Cretaceous Konservat LagerstaÈtten of Europe and the Middle East. A notable exception to this is the Upper Carboniferous Mazon Creek fauna of Illinois, USA, which famously preserves the soft parts of a wide range of phyla within sideritic concretions. Although many cephalopods have been recovered from the Mazon Creek fauna, soft parts are rare even here, and this deposit has yielded specimens of some of the oldest coleoids with soft-part preservation, including the ten-armed Jeletzkya douglassae (Johnson and Richardson 1968; Saunders and Richardson 1979), the origins of which have been much debated, and at least one other possible teuthid form (Allison 1987). The specimen described herein is one of the best preserved cephalopods recovered from the Mazon Creek fauna, and is preserved with arms, indistinct head, and body intact. It is unlike Jeletzkya douglassae, which has been interpreted as a belemnoid, although this is still in debate (D. T. Donovan, pers. comm. 1999), and although it has some morphological similarity to the specimen described by Allison (1987), the excellence of its preservation, and the absence of any shelly material, either externally or internally, provides an important record of great phylogenetic signi®cance. The aim of this paper is, therefore, to describe in detail the specimen, to compare it with known taxa, and to discuss its wider phylogenetic signi®cance. STRATIGRAPHY AND DEPOSITIONAL SETTING The Mazon Creek LagerstaÈtte occurs within the Upper Carboniferous (Westphalian D: Pennsylvanian in North America) Francis Creek Shale Member of the Carbondale Formation in north-eastern Illinois. The Francis Creek Shale was deposited in a deltaic setting where high sedimentation rates effected rapid burial that resulted in the extraordinary fossil preservation within early diagenetic siderite concretions. Two biotic associations occur in this LagerstaÈtte: the Braidwood biota contains terrestrial plants and a variety of freshwater terrestrial organisms; and the Essex biota, which is dominated by a planktonic and nektonic taxa representation of nearshore marine to brackish water conditions (Richardson and Johnson 1971). The distribution patterns of these biotas delineate the limit of marine conditions and the transition to a swamp environment (Richardson and Johnson 1971). The type area for the Essex biota is the former Pit Eleven strip mine of the Peabody Coal Company in Will and Kankakee counties, Illinios (Johnson and Richardson [Palaeontology, Vol. 43, Part 5, 2000, pp. 919±926] q The Palaeontological Association 920 PALAEONTOLOGY, VOLUME 43 1966; Richardson and Johnson 1971). Most Mazon Creek cephalopods, including the specimen described herein, were collected from the Essex biota at Pit Eleven (see Saunders and Richardson 1979). SYSTEMATIC PALAEONTOLOGY Class Cephalopoda Cuvier, 1797 Subclass Coleoidea Bather, 1888 Order ?Cirroctopoda Young, 1989 Genus Pohlsepia gen. nov. Derivation of name. The genus is named in honour of James Pohl of Minnesota, formerly of Belgium, Wisconsin, who collected and donated the specimen to the Field Museum of Natural History. Jim regularly accompanied his father Joe Pohl, noted fossil collector, dairy farmer, and adventurer, on collecting trips to the strip mines of Illinois. Both Jim and Joe have collected many new taxa from the Mazon Creek LagerstaÈtte, which they have shared with the scienti®c community and generously donated to museums. Type species. Pohlsepia mazonensis sp. nov. Diagnosis. Shell absent; mantle subcircular in outline and sac-like; dorsal mantle fused to indistinct head; ten circumoral appendages comprise eight equal arms and two arms modi®ed as tentacles; lobate ®ns longer than wide and free. Pohlsepia mazonensis gen. et sp. nov. Text-®gure 1 Derivation of name. The new species is named for the well known Mazon Creek LagerstaÈtte where the holotype was found. Diagnosis. As for genus. Type specimen. PE51727, Field Museum of Natural History, Chicago, Illinois, USA. Description of holotype. The specimen (Text-®g. 1) is exceptionally well preserved, providing a ventral view of its low relief, and is represented as a slight colour difference within the dark greyish brown siderite concretion, which is 80 mm long and 50 mm wide. Light coloured features in the concretion to the left posterior (and possibly to the right anterior) of the specimen probably represent ¯uids expressed from the animal after burial. Distinct body, head and arms can be distinguished, as well as a number of internal and delicate external features. The body of the coleoid, which has been compressed dorsoventrally, is subcircular and 35 mm wide at its broadest point, with two distinct and symmetrical ®ns at its anterior. These ®ns are narrow and con®ned to the posterior margin of the coleoid. The mantle appears to have crumpled slightly, causing the posterior ®ns to tilt slightly to the anterior. An internal feature to the posterior of the coleoid may be interpreted as either an ink sac or a gut trace; ink sacs are common in Mesozoic coleoids, and the ¯ask-like form of the trace is reminiscent of these. However, it is unusual for TEXT-FIG. 1. A, Pohlsepia mazonensis, gen. et sp. nov., holotype, in siderite concretion, Upper Carboniferous (Westphalian D), Francis Creek Shale Member, Carbondale Formation, near Braidwood, Will County, Illinois; ´ 1´9. B, drawing of Pohlsepia mazonensis gen. et sp. nov.; ´ 1´9. Abbreviations: e, eye; ef, expressed ¯uid; f, ®n; fu, funnel; is?, ink sac (or gut trace); m, mandibles; ma, modi®ed arm (tentacle); r, radula. KLUESSENDORF AND DOYLE: CARBONIFEROUS `OCTOPUS' 921 922 PALAEONTOLOGY, VOLUME 43 the ink not to be preserved, as the melanin of coleoid ink is stable. A similar feature was also described from Allison's specimen of an unknown coleoid (Allison 1987). The specimen shows no sign of any internal shell or phragmocone. The head is identi®able but indistinct from the body, and possesses mandibular architecture, eyes, a funnel and arms. Mandibles and radula are preserved in the head region as strong impressions. The mandibles are articulated, although crushed and dif®cult to identify, and the radula is preserved in situ between them. Radula are commonly preserved in many of the Mazon Creek cephalopods. However, unlike these, the present specimen has its radula obscured by matrix and is otherwise unidenti®able. A short funnel may be visible at the anterior centre of the head, indicating the ventral aspect of the view, and is distinct and broadly central although no cartilagenous locking apparatus is present. The eyes are preserved as small patches of dark pigment which are spaced on either side of the well-de®ned head, and the dark pigmentation is a typical feature of eyes in Mazon Creek vertebrates. The arm crown is indistinct, although its component arms are clearly circumoral, and both short arms and longer modi®ed arms (tentacles) may be distinguished. Only the right appendages are well preserved (left as seen in the ventral view), and comprise what appears to be four short arms (only three are de®nitely visible) and one longer tentacle. No hooks are present and suckers are not visible. Comparison with Jeletzkya douglassae. Jeletzkya douglassae is a well-known coleoid from the Mazon Creek fauna (Johnson and Richardson 1968; Saunders and Richardson 1979) but may be distinguished from the current specimen on the following characteristics: an arm crown comprising ten subequal arms; distinct arm-hooks; a torpedo-like body shape; and an indistinct shell. These features have led to a general acceptance that this taxon represents the soft-body parts of a belemnoid, although whether a true belemnite or an aulacocerid is still under debate (Doyle et al. 1994; Pignatti and Mariotti 1995; D. T. Donovan, pers. comm. 1999). Pohlsepia mazonensis is distinct from this taxon in the presence of a differentiated arm crown, the absence of hooks and an internal shell, and in the subcircular body shape. Comparison with Allison's specimen. Allison (1987) has described a ten-armed coleoid from the Mazon Creek fauna which he did not name. The specimen is poorly preserved and is elongate, with the head and arms extended. The body is globular and preserves posterior lateral ®ns which are similar in morphology to that of the specimen currently under investigation. No mandibles or radula are preserved or described, but identi®able eyes and possible ink sac or internal gut trace are amongst the features discussed. The ten arms are elongate and lacking their anterior tips. However, it is possible that they are differentiated and in this way are similar to the specimen currently under discussion. In general terms, Allison's specimen may be safely compared with the current one and is in all probability representative of Pohlsepia.