Chondrichthyes) Worldwide and New Occurrences from Colorado and Texas Wayne M

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Historical Biology: An International Journal of Paleobiology Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/ghbi20 Systematics and occurrences of Edestus (Chondrichthyes) worldwide and new occurrences from Colorado and Texas Wayne M. Itano a , Karen J. Houck b & Martin G. Lockley b a Museum of Natural History, University of Colorado, 1995 Dartmouth Ave, Boulder, CO, 80305, USA b Dinosaur Tracks Museum, University of Colorado Denver, PO Box 173364, Denver, CO, 80217, USA Published online: 02 Aug 2012.

To cite this article: Wayne M. Itano , Karen J. Houck & Martin G. Lockley (2012) Systematics and occurrences of Edestus (Chondrichthyes) worldwide and new occurrences from Colorado and Texas, Historical Biology: An International Journal of Paleobiology, 24:4, 397-410, DOI: 10.1080/08912963.2012.658569 To link to this article: http://dx.doi.org/10.1080/08912963.2012.658569

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Systematics and occurrences of Edestus (Chondrichthyes) worldwide and new occurrences from Colorado and Texas Wayne M. Itanoa*, Karen J. Houckb1 and Martin G. Lockleyb2 aMuseum of Natural History, University of Colorado, 1995 Dartmouth Ave, Boulder, CO, 80305, USA; bDinosaur Tracks Museum, University of Colorado Denver, PO Box 173364, Denver, CO, 80217, USA (Received 13 January 2011; ﬁnal version received 15 January 2012)

The species of Edestus Leidy (Chondrichthyes, Edestidae) are divided into two groups: those close to Edestus minor Newberry and those close to Edestus heinrichi Newberry and Worthen. The occurrences of Edestus worldwide (North America, Britain and Russia) are reviewed. In North America, Edestus occurs in the Illinois Basin and in the western United States, including South Dakota, Colorado and Texas, but not in marine beds of the same age in the Appalachian Basin. The absence of Edestus in the Appalachian Basin supports the idea that Edestus required access to the open sea, even if its remains are sometimes found in shallow or marginal marine deposits. The chronostratigraphic range of Edestus is from the middle Atokan through the Desmoinesian (Middle Pennsylvanian). All Atokan and lower Desmoinesian occurrences are from the E. minor group. The ranges of the E. minor group and the E. heinrichi group overlap in the upper Desmoinesian. We report two teeth of E. minor from the latest Atokan, in the Minturn Formation at McCoy, Eagle County Colorado. This is the ﬁrst report of the genus from the southern Rocky Mountain region. We report some teeth of Edestus from Texas for the ﬁrst time. Keywords: Edestus; Chondrichthyes; sharks; Pennsylvanian; Colorado; Texas

Introduction probably represents a ventral whorl of E. heinrichi because Edestus Leidy 1856b is a genus of Carboniferous it is relatively slender and resembles the whorl J-4 chondrichthyans known almost entirely from their tentatively identified by Zangerl and Jeremiah (2004) as symphysial teeth. The crowns, which are roughly ventral. A partially decomposed skull of a juvenile triangular and have serrated edges, are attached to Edestus, specimen FMNH PF2204, preserved in shale and elongated bases, which project posteriorly. Behind the visible by X-ray photography, shows the two whorls in crowns, the bases are open and hollow, so that the newest place and protruding from the front of the head (Zangerl (most posterior) tooth can fit into the trough of the next and Jeremiah 2004, Figure 1). (older) tooth. The oldest (most anterior) teeth were shed. As many as 10 teeth might be present in a tooth whorl at any given time. Figure 1 shows a tooth whorl of Edestus Institutional abbreviations heinrichi Newberry and Worthen 1870. Figure 2 indicates DMNH, Denver Museum of Nature and Science, Denver, the way in which the teeth fit together (Eastman 1902a, CO, USA; TMM, Texas Memorial Museum, University of Figure 7). In Edestus, the tooth bases project posteriorly, Texas, Austin, TX, USA; FMNH, Field Museum of while in some other edestoids, such as Helicoprion Natural History, Chicago, IL, USA; USNM, National Downloaded by [Laurentian University] at 21:18 02 October 2013 Karpinsky 1899, they project anteriorly. Each individual Museum of Natural History, Washington, DC, USA; possessed two tooth whorls, one in the upper jaw and the UMMP, University of Michigan Museum of Paleontology, other in the lower jaw. An associated pair of whorls, Ann Arbor, MI, USA; OUZC, Ohio University Zoological preserved nearly in their natural position with respect to Collections, Athens, OH, USA; ACM, Beneski Museum of one another, was described by Hay (1912). That specimen Natural History, Amherst College, Amherst, MA, USA; is the holotype of Edestus mirus Hay 1912. The upper BHIGR, Black Hills Institute of Geological Research, Hill (dorsal) and lower (ventral) whorls differ in shape, the City, SD, USA. ventral one being more slender and more curved than the dorsal one. Two small teeth resembling those of the genera Campodus de Koninck 1844 or Orodus Agassiz 1843 were Materials and methods associated with the specimen and may be part of the We refer to Ginter et al. (2010) for taxonomic assignments dentition of the same animal (Hay 1912, p. 36). Figure 1 of genus and higher. The North American stages of the

*Corresponding author. Email: [email protected]

ISSN 0891-2963 print/ISSN 1029-2381 online q 2012 Taylor & Francis http://dx.doi.org/10.1080/08912963.2012.658569 http://www.tandfonline.com 398 W.M. Itano et al.

Figure 1. Ventral symphysial tooth whorl of E. heinrichi (cast) in lateral view. Anterior end on the right. Fourth tooth crown from posterior end is restored.

Pennsylvanian Period (Morrowan, Atokan, Desmoinesian) environment of the Minturn Formation in the vicinity of are used preferentially, with British, European and Russian McCoy has been described previously (Chronic and stages and substages converted into the North American Stevens 1958; Houck 1997; Itano et al. 2003). Figure 4 is a ones insofar as is possible. generalised stratigraphic column. Nineteen numbered The E. heinrichi cast in Figure 1 was obtained from the stratigraphic units were recognised by Chronic and Black Hills Institute of Geological Research, Hill City, SD, Stevens (1958). During the Middle Pennsylvanian, this USA. It is ﬁgured here with permission from the BHIGR. The original is from the Anna Shale Member, Carbondale Formation, Kewanee Group, Randolph County, Sparta, IL, USA. The Carbondale Formation is upper Desmoinesian. The specimens from the Minturn Formation were collected by members of the Western Interior Paleonto- logical Society (Denver) and are reposited at the Denver Museum of Nature and Science. Only surface collection was used. In the course of this study, some Edestus specimens were located in the collections of the Texas Memorial Museum and are described and ﬁgured here, with such provenance information as is available.

Locality and stratigraphy The area around McCoy, Eagle County, CO, where the Minturn Formation specimens of Edestus were found, is Downloaded by [Laurentian University] at 21:18 02 October 2013 shown in Figure 3. The stratigraphy and depositional

Figure 3. (A) Map of Colorado, showing the town of McCoy, near the boundary of Routt and Eagle Counties; and (B) enlarged Figure 2. Teeth of a single tooth whorl of Edestus heinrichi, view of the area around the towns of McCoy and Bond. separated to show the way in which they ﬁt together (after The Edestus specimens DMNH 61071 and 61072 were found at Eastman 1902a, Figure 7). Total horizontal extent is 16 cm. Localities 1 and 2, respectively. Historical Biology 399

1874), are also present. Other taxa include the dermal denticle Petrodus patelliformis M’Coy 1848 and dorsal ﬁnspines belonging to the species Ctenacanthus buttersi St. John and Worthen 1883, ‘Ctenacanthus’ furcicarinatus Newberry 1875 and Acondylacanthus nuperus St. John and Worthen 1883, and to the genera Physonemus M’Coy 1848 and Bythiacanthus St. John and Worthen 1875 (two morphotypes). The most abundant of the denticle and spine remains are fragments of C. buttersi ﬁnspines. At present, none of the denticle or spine taxa can be linked with certainty to any of the species on the basis of teeth.

Systematics of Edestus Here we accept the generic assignments of Ginter et al. (2010), who list 11 species of Edestus. Some of these species may prove to be synonymous with one another, since they were based on single or few specimens, with little knowledge of individual variation, ontogeny or possible differences between teeth of the dorsal and ventral whorls. Figure 4. Generalised stratigraphic section of the Minturn Formation at McCoy, CO. The Units 1–19 were defined by The type species is Edestus vorax Leidy 1856b, which Chronic and Stevens (1958). The horizons where Edestus is based on a very incomplete specimen having little remains were found are marked. stratigraphic data: The specimen is most probably from the carboniferous series, and it was obtained by William S. Vaux, Esq., from area was on the eastern margin of the Central Colorado an itinerant showman, who found it at Frozen Rock, Basin, on the western side of the Ancestral Front Range Arkansas River, 20 miles below Fort Gibson, in the Indian uplift. Fusulinds found in Units 3, 5 and 7 date these beds Territory. (Leidy 1856b) to the latest Atokan to Desmoinesian (Houck 1997). The type specimen was figured and described more fully in a later publication (Leidy 1856a). The authenticity of the reported locality, which is near Muskogee, OK, has been Summary of the Minturn Formation chondrichthyan questioned by Branson (1963, 1964) who believed, on assemblage preservational and lithological grounds, that the type The Minturn Formation of central Colorado contains the specimen was collected from Illinois. However, Koenig most diverse Pennsylvanian chondrichthyan assemblage (1965) felt that there was insufficient evidence to change known in the southern Rocky Mountain region. Prelimi- Leidy’s record.

Downloaded by [Laurentian University] at 21:18 02 October 2013 nary reports, listing the chondrichthyan teeth, ﬁnspines Other species of Edestus from the Pennsylvanian of and denticles, have appeared (Lockley 1984; Itano et al. North America have been described, including E. minor 2011). Itano et al. (2003) described the chondrichthyan Newberry 1866, E. heinrichi Newberry and Worthen 1870, ﬁnspines and denticles from the Minturn Formation and Edestus giganteus Newberry 1889, E. crenulatus Hay also reviewed the record of Carboniferous chondrichth- 1910, E. serratus Hay 1910 and E. mirus Hay 1912. yans known from Colorado and neighbouring states. According to Branson (1964), E. giganteus is a junior The most common elements of the Minturn chon- synonym of E. vorax. Edestus protopirata Trautschold drichthyan assemblage are teeth of Petalodus ohioensis 1879, E. karpinskyi Missuna 1908, Edestus minusculus Safford 1853 and Glikmanius occidentalis (Leidy 1860). Hay 1910 and E. kolomnensis (Lebedev 2001b) have been Rare elements of the assemblage, aside from Edestus, are described from the Pennsylvanian of Russia, near Moscow. teeth of cf. Gilliodus peyeri Zangerl 1981, cf. Agassizodus Edestus triserratus Newton 1904 and Edestus pringlei variabilis (Newberry and Worthen 1870), Janassa sp. cf. Watson 1930 have been described from the Pennsylvanian J. unguica Eastman 1903b, Peripristis semicircularis of Britain. (Newberry and Worthen 1866) and Polyrhizodus sp. cf. Some species originally placed in Edestus, including P. carbonarius St. John and Worthen 1875. Holocephalian Helicoprion davisi (Woodward 1886), Toxoprion lecontei tooth plates, including Deltodus angularis Newberry and (Dean 1897), and Lestrodus newtoni (Woodward 1917), Worthen 1866 and Lagarodus specularis (Trautschold have since been transferred to other genera. Of these 400 W.M. Itano et al.

genera, only Lestrodus Obruchev 1953 seems close to Even a fragmentary tooth crown can usually be assigned to Edestus, but differs in having a much more tightly curved one of these groups, even if it is not possible to assign it to a whorl and shorter tooth bases. species. Some species are distinguished by morphological The type specimen of L. newtoni was found in the upper differences, sometimes rather minute, that could represent Millstone Grit, West Yorkshire, England. It was found in individual variation or different ontogenetic stages, and so association with the ammonoid index fossil Glyphioceras may not prove to be valid. There may be as few as two valid reticulatum (Phillips 1836). This ammonoid was originally species of Edestus. For example, the tooth whorls of designated Goniatites reticulatus; it is now designated E. giganteus ¼ E. vorax might belong to particularly Reticuloceras reticulatum. According to Ramsbottom et al. mature individuals of E. heinrichi (if so, the name E. vorax (1978, pl. 1), this places it in the upper part of the would have priority), and E. minusculus might be based on Kinderscoutian Substage of the Namurian Stage. This is in a tooth of a juvenile E. minor. The E. minor group includes the middle Morrowan (Davydov et al. 2005, p. 227) and all of the species included in the genus Edestodus Obruchev hence is somewhat older than any known Edestus sensu 1953 by Russian workers (Obruchev 1964; Lebedev 2001a, Ginter et al. (2010). The L. newtoni type specimen is also 2001b). The E. heinrichi group includes the species important because, aside from including one largely transferred to the genus Protopirata Trautschold 1888 by complete tooth whorl of at least eight teeth, it includes at Russian workers (E. protopirata, E. heinrichi and E. least one tooth that was interpreted by Woodward (1917, karpinskyi) and also some retained by them in Edestus (E. p. 2) as being from the opposing dentition. This tooth is vorax and E. giganteus). evidence that Lestrodus, like Edestus, had two opposing tooth whorls. Some small teeth resembling Campodus lateral teeth were associated with the specimen (Woodward Geographic and stratigraphic distribution of Edestus 1917, pl. 1, Figures 8, 9, 9a and 10). North American occurrences Without attempting a thorough review, we can say that Koenig (1965) reviewed the occurrences of Edestus in the the species of Edestus fall into two groups based on their mid-continent of the United States. These occurrences tooth morphology. The first group includes those that are were in Illinois, Indiana, Iowa, Oklahoma, and Missouri. similar to E. heinrichi, the best-known species. Zangerl and All occurrences were in the Desmoinesian. Of those Jeremiah (2004) refer to this as the vorax-serratus- occurrences that could be placed stratigraphically with crenulatus-heinrichi species group in which the species more precision than undifferentiated Desmoinesian, all but are difficult to distinguish. To this group we add one were in the Marmaton Group, the remaining one being E. protopirata and E. karpinskyi. The second group in the Cherokee Group. Koenig’s terminology is based on includes E. minor, E. triserratus, E. pringlei, E. minusculus, the Missouri lithostratigraphic sequence, the Marmaton E. mirus and E. kolomnensis. We will refer to these two Group being the upper part of the Desmoinesian Series and groups as the E. heinrichi group and the E. minor group. The the Cherokee Group being the lower part. The Marmaton tooth crowns of the E. heinrichi group are roughly Group occurrences include E. minor and members of the triangular, with an apical angle of around 70–90 degrees. E. heinrichi group, including E. heinrichi, E. crenulatus Tooth crowns of the E. minor group are narrower relative to and E. giganteus. The sole definite Cherokee Group their height. Figure 5 shows a sketch of a single tooth of E. occurrence listed by Koenig was E. minor in Warrick minor found by St. John and published by Eastman (1902c).

Downloaded by [Laurentian University] at 21:18 02 October 2013 County, IN (Collett 1872, p. 202–203). Collett lists this as If a tooth is oriented so that the margin between the crown E. vorax, but as noted by Koenig, it is clear from the and the base is horizontal, then a line drawn from the context that he was misled by an error in Newberry and anterior-most point of the base to the apex of the crown Worthen (1870, pl. 1, Figure 2), where a tooth whorl of points posteriorly for a tooth of the E. heinrichi group and E. minor was mislabelled as E. vorax. Newberry later anteriorly for a tooth of the E. minor group (Compare corrected this error (Newberry 1889, p. 225). Figures 1 and 2 for the former case to Figure 5 for the latter). There are many specimens of Edestus in museum and private collections. Most of these are from the late Desmoinesian of Illinois and Indiana. See, for example, Zangerl (1981, Figures 100F, 100G, 101A–D) for ﬁgures of some specimens in the Field Museum of Natural History, and Zangerl and Jeremiah (2004) for information on some Edestus tooth whorls in a private collection. We will not attempt to list these specimens unless they Figure 5. Single tooth of Edestus minor. From Eastman (1902c, document new occurrences. To the list given by Figure 3), based on a sketch by St. John. Mirror reversed for Koenig (1965), we add the following occurrences in easier comparison with Figures 1 and 2. North America: Historical Biology 401

Illinois In eastern San Saba County near Bend, the basal Smithwick is 60 feet above the top of the limestone ... Some E. heinrichi teeth were reported from the Anna Shale Member of the Carbondale Formation in southern On the other hand, according to Grayson and Merrill Illinois (Grenda 1974). Jillson (1949) noted that a single (1991, p. 53), regarding the ‘Bend Dump’ locality: tooth of E. minor, USNM 14790, from the Number 5 Coal The formation boundary is drawn at the top of the highest at Springﬁeld, IL (Carbondale Formation), collected by prominent, resistant, carbonate layer. This designation is St. John, was in the collections of the US National more or less arbitrary but is determined by the weathering Museum of Natural History. From Jillson’s description, characteristics of the two lithic types; ... this is probably the same one shown here in Figure 5. The contact between the Marble Falls and the Smithwick Formations, which is conformable, is in the Atokan, but the age varies according to location, being older in the Kentucky eastern part of the Llano region than in the western part (Groves 1991; Erlich and Coleman 2005). The presence of A tooth whorl of E. minor with seven teeth has been the conodont Neognathodus atokaensis Grayson 1984 in reported from south-eastern Muhlenberg County in western the lower Smithwick Formation at the Bend Dump locality Kentucky above the Number 9 Coal in the Carbondale (Grayson et al. 1985) places this bed in the middle Atokan Formation (Jillson 1949). A tooth whorl of E. heinrichi with (Barrick et al. 2004). N. atokaensis is also found in South seven teeth was found at the same horizon in Union County Guizhou, South China, where it straddles the Bashkirian– in western Kentucky (Corgan 1995). A particularly large Moscovian boundary (Wang et al. 2011). According to tooth whorl belonging to the E. heinrichi group was found Davydov et al. (2005, p. 227), the base of the Moscovian is in a coal mine in Webster County, western Kentucky, in a coincident with the base of the Westphalian C. marine shale above the Springﬁeld (western Kentucky No. One incomplete tooth of E. minor from the Smithwick 9) coal (Desmoinesian) (Bauguess 2011; Greb et al. 2011) Formation (Atokan) of Mason County, TX, and several A fragment of a second, smaller, tooth whorl was found in incomplete E. minor teeth from the Strawn Group close association with the large whorl. The second tooth (Desmoinesian), in San Saba County, TX, were found in whorl is direct evidence that sharks of the E. heinrichi the collections of the Texas Memorial Museum. These group had two tooth whorls, similarly to E. mirus, which specimens are discussed in the ‘Systematic Paleontology’ belongs to the E. minor group. section. A tooth from the Finis Shale (Upper Pennsylvanian) at the Lost Creek Reservoir Spillway, Coleman County, TX, Texas labelled Edestus sp. (McKinzie 2003, p. 115, Figure 5-8), clearly does not belong to Edestus. The tooth base is Cummins (1890, p. 149) reported E. vorax from the curved, tapers distally and is not fused to the neighbouring Carboniferous, on the west side of the Colorado River, bases in the series. It may belong to the genus ‘just above the limestone, in the black shale’, near Bend, Campyloprion Eastman 1902b. TX. The limestone referred to is the highest one in the section. Evidently this is the same specimen reported by Cummins (1891, p. 392) as E. minor ‘in the black shale Michigan

Downloaded by [Laurentian University] at 21:18 02 October 2013 about one mile west of the town of Bend, in the southeast corner of San Saba County’. The confusion between E. A single occurrence of E. minor is noted at Six Mile Creek, vorax and E. minor, which are easy to distinguish, was Shiawassee County, in the Saginaw Group (Kelly 1936, likely caused by the previously noted error in Newberry p. 11). This specimen is also noted by Dorr and Eschman and Worthen (1870, pl. 1, Figure 2). The whereabouts of (1970, p. 398) with specimen number UMMP 14411. this specimen are unknown. E. minor, possibly the same According to current terminology, the bed in which the specimen, was reported later in faunal listings of the specimen was found is in the Verne Limestone Member of Smithwick Formation but without any details (Moore the Saginaw Formation. The age is late Atokan (Landing 1919, p. 229, 234; Plummer and Moore 1921, p. 220). The and Wardlaw 1981). age of the bed in which Cummins found the E. minor tooth is of some importance, since this may be the earliest occurrence of Edestus in North America. From Cum- Kansas mins’s stratigraphic description, the bed is in either the Hamm and Cicimurri (2005) report three tooth fragments lowermost Smithwick Formation or the uppermost Marble of Edestus sp. cf. E. heinrichi from the Lake Neosho Shale Falls Formation, depending on the deﬁnition of the units. Member of the Altamont Limestone of Montgomery According to Plummer (1950, p. 78), this bed would be in County, KS. The Altamont Limestone is in the Marmaton the uppermost Marble Falls Formation: Group. The tooth crowns compare well to E. heinrichi. 402 W.M. Itano et al.

Chorn and Reavis (1978, p. 5) state that Edestus and Russia Cladodus are known from a locality of the Little Osage The Russian occurrences of Edestus are presented in a Member of the Fort Scott Limestone (Marmaton Group, chart by Lebedev (2001a, p. 93–95), which lists upper Desmoinesian) in Bourbon County, KS. occurrences according to subdivisions of the Russian Platform substages for each of the species E. minusculus, E. kolomnensis, E. protopirata, E. karpinskyi, and an Missouri undetermined species resembling E. minor. The ranges Two tooth fragments that appear to be E. minor are figured are somewhat tentative, since all of the species are on a website (Sutton 2011). They were found in the Lake known by only a few specimens (one to five in a bed), Neosho Shale Member of the Altamont Limestone, near and some specimens were collected long ago, without St. Louis, MO. A photograph of these teeth, taken from the good stratigraphic control. To summarise the data, website, has appeared in print (McKinzie 2003, p. 116, various species have overlapping ranges, with none Figure 5-9). being earlier than middle Myachkovian and none being later than early Krevyakian. Thus, the Russian occurrences all fall within the late Desmoinesian (Davydov South Dakota et al. 2005, p. 227). Cicimurri and Fahrenbach (2002) reported a single We note that Hay (1910) erred in describing the type fragmentary tooth of Edestus sp. from the upper Minnelusa specimen of E. minusculus as being from the lowermost Formation, Meade County, SD. The narrow shape of the Permian, Artinskian Stage. The specimen in question was crown is enough to identify this as E. minor. The associated called Edestus cf. minor by Karpinsky (1899), who stated conodonts indicate a middle-to-late Desmoinesian age. that it was from the quarry at Myachkova, near Moscow, in the zone of the brachiopod Spirifer mosquensis. Hay confused the age of the Edestus specimen with that of Oklahoma Helicoprion, which was described in the same publication. Figure 10A of Mapes and Chaffin (2003) shows a tooth The range of Spirifer mosquensis, which was originally fragment that appears to be a large specimen of E. minor.It designated Choristites mosquensis Fischer de Waldheim is from the Atoka Formation (Atokan) near Bromide, OK 1825, is from the Tsninian Substage of the Moscovian (October 2011 emails from R.H. Mapes). At present, the age Stage to the Krevyakian Substage of the Kasimovian Stage cannot be determined more precisely than undifferentiated (Wagner et al. 1996, p. 94, Figure 26). This is equivalent to Atokan. It is reposited in the Ohio University Zoological the late Atokan through latest Desmoinesian (Davydov et al. Collections with specimen number OUZC 5833. 2005, p. 227). The brachiopod zone is consistent with, but broader than, the range given for E. minusculus by Lebedev (2001a, p. 93), which is from late Myachkovian to Colorado early Krevyakian. Two incomplete teeth of E. minor have been found in latest Atokan beds in the Minturn Formation at McCoy, Britain Eagle County, CO. They are discussed in the ‘Systematic E. triserratus is based on a single tooth, found in a marine Paleontology’ section.

Downloaded by [Laurentian University] at 21:18 02 October 2013 band 16.5 metres below the Twist Coal at Nettlebank, North Staffordshire (Newton 1904). According to Stobbs (1905), the Twist Coal at North Staffordshire is equivalent Wyoming to the Gin Mine Coal. According to Ramsbottom et al. Two tooth crown fragments from the Permian of the middle (1978, pl. 2 and 3), the marine band below the Gin Mine Phosphoria Formation of Wyoming were assigned, Coal is the Aegiranum Marine Band (standard name). questionably, to Edestus by Branson (1933). Without The Aegiranum Marine Band marks the base of the more complete remains, such as a part of the tooth base, Westphalian C, which is also the base of the Bolsovian they cannot be assigned to Edestus with any conﬁdence. (Guion et al. 2000, p. 261). This places it in the middle They probably belong to some other edestoid, such as Atokan (Davydov et al. 2005, p. 227). Helicoprion. Three teeth referred to a new species E. pringlei were found in a marine band at depths of 514 metres in No. 1 shaft and 537 metres in No. 2 shaft at the Cefn Coed Occurrences outside North America Colliery, near Neath, South Wales (Watson 1930). These Occurrences of Edestus outside North America were not levels correspond to the Cefn Coed Marine Band reviewed by Koenig (1965). Deﬁnite occurrences are only (Ware 1930), which is equivalent to the Aegiranum from Russia and Britain. Marine Band. Thus, both E. triserratus and E. pringlei are Historical Biology 403

about the same age as the tooth of E. minor reported from Remarks the Smithwick Formation, near Bend, TX (Cummins 1890, Hitchcock (1856) reported and figured, but did not name, 1891). The Cefn Coed Marine Band contains the richest a tooth whorl containing seven teeth. Hitchcock’s marine fauna of the British Westphalian, including corals, specimen was found in Parke County, IN, in a shale brachiopods, echinoids, crinoids, ammonoids and trilo- layer above a coal bank. According to Koenig (1965, bites (Ramsbottom 1952; Ramsbottom et al. 1978). p. 16), the shale is equivalent to the Little Osage A single lateral tooth of Campodus type was associated Formation, in the Fort Scott Subgroup of the Marmaton with the E. pringlei teeth (Watson 1930, Figure 1G). This Group (upper Desmoinesian). Newberry described association, together with the associations of Campodus- E. minor, on the basis of a single tooth crown of the type teeth with E. mirus and L. newtoni as previously same kind (Newberry and Worthen 1866, pl. 4, Figure noted, supports the idea that the dentition of Edestus 24). This tooth is technically the holotype of E. minor. included teeth of this type. However, its whereabouts are unknown, and it does not E. triserratus and E. pringlei are both very similar to seem to have ever been figured again. Hitchcock’s much E. minor. They differ mainly in having serrations (cusplets) more complete specimen, figured by Newberry and that are trifid and bifid (divided into three or two parts), Worthen (1870, pl. 1, Figure 2), but erroneously labelled respectively, rather than being undivided. as E. vorax, is reposited at the Beneski Museum of Natural History, Amherst College, with specimen number China ACM 85 (October 2011 email from K. Wellspring). Part of this figure (Newberry and Worthen 1870) is shown in There is a questionable occurrence from the Ortu the background of Figure 7. Whatever the formal Formation, Qijiagou, Xinjiang Province, China (Cheng designation, the more complete specimen, which has et al. 1996). It is a fragment of a tooth crown showing a been figured many times (Newberry 1888, 1889; von serrated edge. The serrations are atypical for Edestus. No Zittel 1890; Woodward 1891; Dean 1895, 1897; details are given of the base, which is important for Karpinsky 1899; Eastman 1903a; Newton 1904; Obru- differentiating Edestus from other edestoids. The fact that chev 1953, 1964; and others), is effectively the type one side is more convex than the other (Cheng et al. 1996, specimen of E. minor. [Eastman (1903a), who shows p. 704) would seem to exclude it from Edestus, since photographs of both lateral views, seems to be the only Edestus teeth always have bilateral symmetry about a one to figure both sides.] It is interesting to note that the plane containing the serrated edges. It may belong to crown figured by Newberry and Worthen (1866) is more another genus having serrated teeth, like Carcharopsis sharply pointed than those on the tooth whorl of ACM 85 Agassiz 1843. The corals and ammonoids in the Ortu and is very similar to those on the holotype of E. mirus. Formation indicate a late Pennsylvanian age. In fact, the characters of the shape of the tooth crown used by Hay (1912, p. 36) to distinguish E. mirus from Systematic paleontology E. minor would apply equally to the specimen of Newberry and Worthen (1866, pl. 4, Figure 24). If Hays’s Class: Chondrichthyes Huxley 1880 species concept were to be consistently applied, the Subclass: Euchondrocephali Lund and Grogan 1997 specimen reported by Hitchcock (ACM 85) would require Order: Eugeneodontiformes Zangerl 1981 a new name, and E. mirus would probably become a

Downloaded by [Laurentian University] at 21:18 02 October 2013 Superfamily: Edestoidea Hay 1929 junior synonym of E. minor. Family: Edestidae Jaekel 1899 Genus: Edestus Leidy 1856b Edestus minor Newberry 1866 Minturn Formation specimens Figures 6–12 Figures 6–9

[No name]; Hitchcock (1856, text-ﬁgure on p. 229) Edestus minor; Newberry 1866, in Newberry and Material Worthen (1866, pl. 4, Figure 24) Two incomplete teeth, DMNH 61071 and 61072. Edestus vorax; Newberry and Worthen (1870, pl. 1, Figure 2) Edestus minor; Newberry (1889, p. 218, pl. 39, Figures Locality and age 1, 1a) DMNH 61071, Unit 3c of Houck (1997), Minturn Edestus minor; Eastman (1902c, Figure 3) Formation, Section 5, T2S, R83W, Eagle County, CO, Edestus minor; Jillson (1949, p. 12–13, frontispiece collected by D. Nelson. DMNH 61072, Unit 3b of Houck ﬁgure) (1997), Minturn Formation, Section 2, T2S, R84W, Eagle Edestus minor; Zangerl (1981, Figure 100F) County, CO, collected by S. Reinhold. According to 404 W.M. Itano et al.

the posterior edge of the crown are partially preserved, but the tips of all of the cusplets are broken. Figure 7 shows the same tooth on a ﬁgure of E. minor ACM 85 (Newberry and Worthen 1870, pl. 1, Figure 2). The ﬁgure is of natural size. There is a very good match with the third tooth from the anterior (right) end of the tooth whorl, both in terms of the shape of the portion of the crown which is preserved and the shape of the margin between the crown and the base. DMNH 61072 (Figures 8 and 9) is a single tooth, broken so as to expose the interior of the base, but preserving most of the outline of the base and also a small section of the crown, mainly as an external mould. A cast was made of the crown portion by pressing a piece of modelling clay into the specimen. In the resulting cast (Figure 9), the serrations of the posterior crown edge are clearly seen. Also, the orientation of the exposed trabecular dentine of the interior of the tooth crown shows that the apex of the crown was directed forward as Figure 6. Partial tooth of Edestus minor DMNH 61071 from the in the teeth of the E. minor group. The base is not Minturn Formation, CO. (A) Lateral view, anterior end on the diagnostic as to species. The crown morphology places right; (B) coronal view showing broken serrations on posterior this within the E. minor group, and we refer it to E. minor. crown edge; and (C) posterior view. Tooth is coated with The limestone matrix contains spiriferid and other ammonium chloride. Scale bar ¼ 1 cm. brachiopods, crinoid columnals and other crinoid ossicles, Houck (1997), Units 3b and 3c are within the fusulinid and archaeocidarid (echinoid) spines, all indicative of a biozone of Fusulinella famula, F. iowaensis and normal marine environment. F. iowaensis leyi of Ross and Ross (1987), which is latest Atokan. Smithwick Formation specimen Figure 10 Discussion DMNH 61071 (Figures 6 and 7) is a single tooth, missing the distal end of the crown and preserving only the portion Material of the base immediately below the crown. The serrations on One incomplete tooth, TMM 40312-1. Downloaded by [Laurentian University] at 21:18 02 October 2013

Figure 7. Edestus minor tooth DMNH 61071 from the Minturn Formation, CO, superposed on natural-size ﬁgure (Newberry and Worthen 1870, plate 1, Figure 2) of type specimen of E. minor ACM 85. Tooth is coated with ammonium chloride. Anterior end of tooth whorl is on the right. Scale ¼ 1 cm. Historical Biology 405

Figure 8. Tooth of Edestus minor DMNH 61072 from the Minturn Formation, CO (colour online). The coronal side is up. The anterior end is to the left. The crown and the anterior end of the base are mainly preserved as an external mould. Scale ¼ 1 cm.

Locality and age of fusulinid ranges, fusulinid biozones and lithostrati- Smithwick Formation, upper Atokan, Honey Creek, 13 km graphic units (Groves 1991, Figure 3) indicates that the south-west of Mason, Mason County, TX. Smithwick Formation at Zesch Ranch is upper Atokan. The shape of the crown places this tooth in the E. minor group, and we refer it to E. minor. The shape of the margin Discussion between the crown (enamelled) part of the tooth and the base is a variable feature in E. minor. On this tooth, the margin has The label states that this specimen was ‘taken from old a ‘hook’ or concave downward section near the anterior end collections of Geological Survey of Texas’. An older label (left side of Figure 10(a)). This feature is also seen on some has a specimen number 40129-12 that indicates that it was teeth of the holotype of E. mirus (Hay 1912, pl. 1). in a group of fossils that was transferred to the TMM from the Texas Bureau of Economic Geology (J.C. Sagebiel, personal communication). In the Llano region, the age of the Smithwick Formation decreases from east to west (Groves 1991; Erlich and Coleman 2005). Mason is west of Bend, so this tooth is somewhat younger than the one reported by Cummins from the vicinity of Bend. The Mason locality is close to the Zesch Ranch Section (Groves 1991). A chart Downloaded by [Laurentian University] at 21:18 02 October 2013

Figure 9. Cast of crown portion of DMNH 61072. Anterior end is to the right. The margin between the base and the crown (enamelled part) is nearly horizontal. A row of seven serrations is Figure 10. (Colour online) Edestus minor tooth, TMM 40312-1, visible on the posterior edge. On the right, the exposed interior of from the Smithwick Formation, Mason County, TX (colour the tooth has the ‘grain’ of the trabecular dentine oriented along online). (A) Lateral view, anterior end on left; and (B) coronal the main axis of the crown. Scale bar ¼ 1 cm. view, posterior crown edge. Scale bar ¼ 1 cm. 406 W.M. Itano et al.

Figure 11. Edestus minor tooth, TMM 40111-7, from the Figure 12. Edestus minor tooth, TMM 40234-8, from the Strawn Group, San Saba County, TX, embedded in matrix Strawn Group, San Saba County, TX (colour online). (A) Lateral (colour online). Lateral view, anterior end on the right. The view, anterior end on the right; (B) posterior view; and (C) margin between the crown (enamelled part) and the base is nearly coronal view of posterior crown edge. Scale bar ¼ 1 cm. horizontal. Scale ¼ 1 cm. such as ACM 85. Although no scale was given for the Strawn Group specimens sketch of USNM 14790, Jillson (1949, p. 10–11), who examined the specimen, described this tooth as belonging Figures 11 and 12 to a juvenile of E. minor. TMM 40234-8 (Figure 12) is a complete crown, Material preserving part of the base. The posterior edge of the Nineteen incomplete teeth, TMM 40111-7, 40234-8, crown is concave, like the Colorado specimen DMNH 40234-9, 40234-10, 40234-11, 40234-12 and 13 teeth 61071 (Figure 6), which itself is very similar to teeth of the sharing number TMM 40234-1. ACM 85 specimen of E. minor (see Figure 7). The crown is more erect than is typical of E. minor. We consider this

Downloaded by [Laurentian University] at 21:18 02 October 2013 to represent individual variation. Figure 12(c) of TMM Locality and age 40234-8 shows some deviation from bilateral asymmetry Upper Strawn Group, middle Desmoinesian, 5 km west of of the crown, which we consider to be a growth Richland Springs, San Saba County, TX. abnormality, not a characteristic feature of the species. Although these specimens were found long ago, probably as long ago as the 1893 surveys of E. Cope and Discussion W.F. Cummins (J.C. Sagebiel, personal communication), All of these teeth are believed to be from the same locality. they have not been reported in the published literature. TMM 40111-7 (Figure 11) is a nearly complete crown, The locality is the same, or very close to, the Big Uncle preserving part of the base, embedded in a sandstone and Creek Stop in the ﬁeld trip guidebook of Grayson and conglomerate matrix with other fossils, including what Merrill (1991), which is described as being 2.80 km west appears to be the base of a Petalodus tooth. TM 40111-7 and slightly north-west of the centre of Richland Springs. has a relatively straight posterior edge, like the tooth found This ﬁeld trip stop is in the upper Strawn Group. The age, by St. John, USNM 14790 (shown here in Figure 5), in as indicated by conodonts Idiognathodus delicatus, contrast to some other teeth referred to E. minor. This is Neognathodus medadultimus, Neognathodus medexulti- likely an ontogenetic difference, TMM 40111-7 and mus and Gondolella bella, is early to middle Desmoinesian USNM 14790 being small compared to other specimens, (Grayson et al. 1985; Barrick et al. 2004). Historical Biology 407

Conclusion be harder to understand if it were adapted to freshwater or We have added some new and little-known occurrences of estuarine environments. It is also consistent with its large Edestus sensu Ginter et al. (2010) and removed some size. One would still expect its remains to be found in erroneous ones. All species of Edestus can be placed in one shallow and marginal marine sediments. However, the of two groups, called here the E. heinrichi group and the possibility that it might have spent most of its time in E. minor group. Some of the species in each group may deeper water would help account for the scarcity of its prove to be synonymous. remains in shallow marine deposits. Edestus has still not Edestus had a wide range geographically but a rather been found in the Pennsylvanian coal mining areas of the narrow range chronologically, about six million years Appalachian Basin, comprising parts of Ohio, Pennsylva- according to the time scale of Davydov et al. (2005, nia, West Virginia, and eastern Kentucky. This is p. 227). No occurrences of Edestus are known below the understandable if the Appalachian Basin was not well middle Atokan (base of the Westphalian C) or above the connected to the deeper marine basins to the west, so that the environment would be less suitable to a large, pelagic latest Desmoinesian (early Krevyakian). [However, an fish. The continued absence of finds of Edestus remains Oklahoma E. minor tooth is not dated more precisely than from the Pennsylvanian marine beds of the Appalachian the undifferentiated Atokan (Mapes and Chaffin 2003 and Basin, over a century after Newberry’s observation, is October 2011 emails from R.H. Mapes).] North American strong evidence that Edestus required access to the open occurrences are found through the entire range from the sea during part of its life cycle. Remains of presumed middle Atokan to the late Desmoinesian. British marine chondrichthyans, such as Petalodus, and marine occurrences are in the earlier part of the range; Russian invertebrates are found in deposits of the Appalachian occurrences are in the later part. All occurrences in the Basin of the same age as the Edestus of the Illinois Basin Atokan and the lower Desmoinesian are of the E. minor (Hansen 1986; Feldmann and Hackathorn 1996). group. Both the E. minor group and the E. heinrichi group The Russian Edestus remains are from the Moscow occur in the upper Desmoinesian. Lestrodus, known only Syneclise, which would have been connected to the from the middle Morrowan (Namurian, Kinderscoutian) of Uralian Ocean to the east during the Pennsylvanian. Britain, may be ancestral to Edestus. Edestus seems to The British occurrences of Edestus are only in the have died out without giving rise to any successor genera. Aegiranum Marine Band, which may have been a In North America, Edestus is widespread, though particularly deep marine incursion, judging by the richness uncommon, in marine chondrichthyan assemblages of of its marine fauna. The occurrence of Edestus in this bed Atokan and Desmoinesian age. It has not yet been reported may indicate a connection to the open sea, possibly brief. from assemblages of this age from Arizona (Elliott et al. The fact that many Edestus remains have been found in 2004) or New Mexico (Lucas and Estep 2000; Ivanov et al. close proximity to coal beds (but in the black shales above 2009), but this may be due to the fact that relatively little the actual coal beds) has led to Edestus being nicknamed the collecting has been done at those localities. ‘coal shark’ (see, e.g., Cocke 2002, p. 127). The association Over a century ago, Newberry (1889, p. 218) noted with coal beds should be seen more or less as an accident. that the geographical distribution of Edestus was ‘some- The Colorado and Texas Edestus finds, including DMNH what peculiar’. He noted that the genus had been found in 61072, embedded in crinoidal limestone, and TMM 40111- the coalfields of Illinois and Indiana, but not in the 7, embedded in arkosic conglomerate containing a coalfields to the east:

Downloaded by [Laurentian University] at 21:18 02 October 2013 Petalodus tooth fragment, are not closely associated with In Ohio and Pennsylvania much more extensive coal deposits, nor are they in black shales. These finds are excavations in the coal rocks and numerous collections consistent with Edestus being a marine pelagic fish which of Carboniferous fossils have been made, but not a trace of sometimes came to shallow water and not, as Newberry Edestus has been found there. Hence we must infer that it thought, an inhabitant of freshwater lagoons. never passed the highlands of the Cincinnati arch, which separated the western from the eastern coal basins. He noted, correctly, that the remains of Edestus were Acknowledgements found in the carbonaceous black shales above coal seams, We thank D. Nelson and S. Reinhold for collecting and but misinterpreted the shale beds as being freshwater donating the Colorado specimens; J.C. Sagebiel of the sediments. He imagined Edestus as inhabiting ‘lagoons of TMM for facilitating the loan of the Texas specimens and water that occupied portions of the coal marshes’. These for providing information on their provenance; N. Larson shale beds are now known to represent marine incursions. of the BHIGR for providing information on the E. heinrichi The geographical distribution of Edestus becomes cast in Figure 1; K. Wellspring of the ACM for providing understandable if Edestus was a pelagic, sea-going fish, at information on the type specimen of E. minor; and least for part of its life cycle. This mode of life would help R.H. Mapes of Ohio University for information on the account for its wide geographic distribution, which would Atokan Edestus tooth from Oklahoma. We also thank two 408 W.M. Itano et al.

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