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Acanthorhachis, a New Genus of Shark from the Carboniferous (Westphalian) of Yorkshire, England

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Acanthorhachis, a New Genus of Shark from the Carboniferous (Westphalian) of Yorkshire, England Geol. Mag. 151 (3), 2014, pp. 517–533. c Cambridge University Press 2013 517 doi:10.1017/S0016756813000447 Acanthorhachis, a new genus of shark from the Carboniferous (Westphalian) of Yorkshire, England ∗ ∗ DAVID M. MARTILL †, PETER J. A. DEL STROTHER‡ & FLORENCE GALLIEN ∗ School of Earth and Environmental Sciences, University of Portsmouth, Portsmouth PO1 3QL, UK ‡PJDS Consulting Ltd., Clitheroe BB12 7JA, UK (Received 12 January 2013; accepted 15 May 2013; first published online 8 July 2013) Abstract – An association of diverse hollow spines and dermal denticles (ichthyoliths) from the Carboniferous (Westphalian) of Todmorden, Yorkshire, England are attributed to a new genus of enigmatic shark that may lie close to Listracanthus Newberry & Worthen, 1870. Scanning electron microscopy shows that denticle morphology is highly variable, but forms a morphocline including elongate multi-spined elements as well as robust dome-like stellate denticles and recurved spinose elements. Histological analysis suggests an absence of enameloid. Continuous variation of form between elongate multi-cusped spines to boss-like circular denticles shows that all previously described Palaeozoic species of Listracanthus are probably junior synonyms of the type species L. hystrix Newberry & Worthen, 1870. The status of Listracanthus as a surviving ‘Lilliputian’ taxon after the Permian extinction is questioned. Although the new specimen has affinities with Listracanthus, significant differences in the form of the posterior spines on elongate denticles warrants its placement in the new genus Acanthorhachis gen. nov. The family Listracanthidae is erected to accommodate Listracanthus and Acanthorhachis. Keywords: Acanthorhachis gen. nov., Carboniferous, Chondrichthyes, dermal denticles, England. 1. Introduction Listracanthus. Listracanthus has also been considered present in the Triassic of Canada where its occurrence Listracanthus Newberry & Worthen, 1870 is a ‘form’ has been considered a part of the so-called Lilliputian taxon representing an enigmatic chondrichthyan known recovery fauna after the Permian mass extinction only from its mostly small but elaborate spines and event (Mutter & Neuman, 2009). Although enigmatic, a few dermal denticles (ichthyoliths). It ostensibly Listracanthus was considered by Patterson (1965) to be ranges from the Carboniferous to the Triassic (Mutter a junior synonym of the holocephalan Deltoptychius, & Neuman, 2006) and was first described from but recent workers accept that Listracanthus represents Carboniferous strata in Illinois, USA (Newberry & an early, although somewhat unusual, elasmobranch Worthen, 1870). Elsewhere in the United States it has (e.g. Mutter & Neuman, 2006). Here we describe a been reported from Ohio, Indiana, Missouri and Kansas new specimen of elasmobranch with associated spines (Newberry, 1873, 1875; Newberry & Worthen 1870; and denticles that resembles Listracanthus but displays Hibbard, 1938; Chorn & Reavis, 1978; Schultze et al. a wide variety of dermal denticle morphotypes as well 1982; Hamm & Cicimurri, 2005), while in Europe as several significant differences in spine structure from the genus has been reported from the Carboniferous the holotype of Listracanthus. of England (Stobbs, 1905; Edwards & Stubblefield, Institutional abbreviations. UALVP – Laboratory 1948), Scotland (Woodward, 1891, p. 149), Belgium for Vertebrate Paleontology – University of Alberta – (Woodward, 1891; Koninck, 1878; Derycke et al. Edmonton – Canada; YPM – Yale Peabody Museum – 1995), Germany (Bolton, 1896; Schmidt, 1950) and USA; NHMUK – Natural History Museum – London. the Czech Republic (Lang, 1979; Štamberg & Lang, 1979). In Asia Listracanthus has been reported from the Carboniferous of northern China (Lu et al. 2002, 2005) 2. Locality and also from the Permian of the Ural region of Russia (Ivanov, 2005). Turner (1993) reported Listracanthus Material examined in this study was collected by one in the Carboniferous of Queensland, Australia (the of the authors (PdS) on the east side of an old colliery first occurrence in the southern Hemisphere) on the spoil dump on Todmorden Moor, Calderdale District, basis of a single spine, but several dermal denticles West Yorkshire (NGR SD 89292 24865) (Fig. 1a). This figured by her as ‘hybodontid’ also resemble those of spoil dump was generated from mine workings of the Sandy Road Colliery belonging to the East Lancashire Brick Co. Ltd., abandoned in 1966. It appears to be located close to the boundary between the Millstone †Author for correspondence: [email protected] Grit and the Lower Coal Measures (Wright et al. Downloaded from https:/www.cambridge.org/core. Open University Library, on 25 Jan 2017 at 01:11:02, subject to the Cambridge Core terms of use, available at https:/www.cambridge.org/core/terms. https://doi.org/10.1017/S0016756813000447 518 D. M. MARTILL AND OTHERS Figure 1. (Colour online) Locality maps for the Todmorden Acanthorhachis occurrence. (a) Generalized map of central England showing the location of Todmorden with respect to main commercial centres. (b) The Acanthorhachis locality is an abandoned mine dump by the field road between Todmorden and Bacup. (c) Insert showing Google Earth image of abandoned mine dump. 1927) (Fig. 2). Presently there is very little exposure Mary Stopes in 1903 (Briant, 1962). Todmorden of the fossiliferous strata in this region, but the spoil Moor is currently registered as a site of Local dump is well-known for yielding so-called ‘coal balls’, Geological Interest by the West Yorkshire Geology carbonate concretions rich in 3D plant remains (Stopes Trust. & Watson, 1908), and goniatites from the Gasteroceras listeri marine band. There are few accounts of 3. Material and methods vertebrate remains from this locality; an early exception is the report of fish remains by Bolton (1889) who The associated spines and denticles examined in described coelacanth and indeterminate fish bones this analysis were all obtained from a concentration from a locality east of Bacup and an occurrence of in a layer within a dark grey limestone fragment Listracanthus in shales above the Bullion Coal, also measuring c. 160 × 90 × 40 mm. Part of the at Bacup (Fig. 1), but this specimen was lost at the fragment was broken and placed in acetic acid at 7– time of its discovery (Bolton 1896). The abundance of 10 % concentration until all the matrix had dissolved ‘coal balls’ at this locality has made it an important (approximately one month). A second piece was placed historic site, which was collected by palaeobotanist in acetic acid, but with an acid-resistant coating Downloaded from https:/www.cambridge.org/core. Open University Library, on 25 Jan 2017 at 01:11:02, subject to the Cambridge Core terms of use, available at https:/www.cambridge.org/core/terms. https://doi.org/10.1017/S0016756813000447 Acanthorhachis: new shark genus from Yorkshire Westphalian 519 Figure 2. (Colour online) A generalized stratigraphic section for the Westphalian sequence in the vicinity of Todmorden. The insert (arrowed) is a more detailed section seen in the abandoned coal mine that yielded the Acanthorhachis specimen, from Dugdale (1887). Downloaded from https:/www.cambridge.org/core. Open University Library, on 25 Jan 2017 at 01:11:02, subject to the Cambridge Core terms of use, available at https:/www.cambridge.org/core/terms. https://doi.org/10.1017/S0016756813000447 520 D. M. MARTILL AND OTHERS Figure 3. (Colour online) A portion of the specimen yielding the Acanthorhachis denticles after partial dissolution in 10 % acetic acid: (a) looking down onto bedding plane surface (NHMUK P73205a); (b) etched section through the portion (NHMUK P73206b); (c) detail of the distribution of denticles on bedding after etching and (d) before etching showing fresh, unweathered colour of the matrix; (e) weathered surface showing cream colouration; and (f) goniatite cf. Gastrioceras listeri photographed under alcohol. The specimen is seen in sagittal section, showing this was not the original outer surface of the concretion. (paraloid) applied to the sides so that a bedding Images have been manipulated using Corel Photo-Paint plane surface was partially etched to reveal several X5 and CorelDraw X5. large spines without complete separation from the matrix. Acid-resistant residues were rinsed initially in tap water, with a final rinse in distilled water. 4. Geological setting and stratigraphy Suspended clay was decanted off and the remaining insoluble residue contained only dermal denticles and 4.a. Geological setting did not require sieving. A similar process was also The geology of Rossendale and Todmorden Moor is performed on pieces of Dinantian crinoidal limestone described by Aitken (1868) and Wright et al. (1927) from Steeplehouse Quarry, Wirksworth, Derbyshire with a more recent, though general, account provided containing Petrodus denticles for comparative pur- by Aitkenhead et al. (2002). The fossiliferous strata poses. Specimens for electron microscopy analysis are located on the eastern flank of the Rossendale Sub- were mounted on flat aluminium stubs, sputter coated basin of the Pennine Basin and are part of an anticline with gold and examined using a JEOL JSM 6100 associated with the so-called Todmorden Smash Belt, machine. where they dip gently to the SW. Exposures of the Thin-sections were produced from remaining frag- fossiliferous strata are rare on Todmorden Moor due to ments and examined using a petrological microscope extensive cover by Holocene peat deposits, but a
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