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A Lower Carboniferous xenacanthiform shark from Australia

Susan Turnerab; Carole J. Burrowb a School of Geosciences, Monash University, Victoria 3800, Australia b Queensland Museum, Queensland, Australia Online publication date: 21 March 2011

To cite this Article Turner, Susan and Burrow, Carole J.(2011) 'A Lower Carboniferous xenacanthiform shark from

Australia', Journal of Vertebrate Paleontology, 31: 2, 241 257 To link to this Article: DOI: 10.1080/02724634.2011.550359 URL: http://dx.doi.org/10.1080/02724634.2011.550359

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Journal of Vertebrate Paleontology 31(2):241257, March 2011 2011 by the Society of Vertebrate Paleontology

ARTICLE

A LOWER CARBONIFEROUS XENACANTHIFORM SHARK FROM AUSTRALIA

SUSAN TURNER*,1,2 and CAROLE J. BURROW2 School of Geosciences, Monash University, Victoria 3800, Australia; 2 Queensland Museum, Geosciences, 122 Gerler Road, Hendra, Queensland 4011, Australia, sue.turner@qm.qld.gov.au; carole.burrow@gmail.com
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ABSTRACTA new Early Carboniferous (Mississippian, mid-Visean) chondrichthyan, Reginaselache morrisi, n. g., n. sp., from non- or marginal marine sandy mudstones of the Tetrapod Unit of the mid-Visean (330 Ma = top Holkerian/basal Asbian) Ducabrook Formation, northwest of Springsure, central Queensland, is referred to the order Xenacanthiformes. The taxon is represented by robust diplodont teeth with multicristate cusps, a prominent rounded coronal button, and a horseshoe-shaped labial boss. Rare spine fragments from the type locality, and a partial lower jaw from a site close by are also tentatively referred to the taxon. Reginaselache morrisi was a medium-sized, ca. 1 m long shark with numerous teeth, probably feeding on smaller paleoniscoid and other shes and/or invertebrates. Analysis of the teeth and comparison with those of other Carboniferous and later Paleozoic xenacanthiforms shows that the tooth cusp morphology is closest to those of Triodus Hampe and Bohemiacanthus Schneider. A restricted cladistic analysis of the xenacanthiforms with outgroups Leonodus Mader, Phoebodus St John and Worthen, and Antarctilamna Young supports the family Diplodoselachidae Hampe as a clade comprising just two genera, Diplodoselache and Reginaselache.

INTRODUCTION The xenacanthiforms were a successful group of freshwater and marginal marine sharks during the mid- to late Paleozoic and early Mesozoic; articulated specimens including embryonic and juvenile animals are known (Hampe, 2003; Soler-Gijon, 2004). Evidence from older deposits is accumulating to show that the clade was common in shallow-water marine, marginal, and possible freshwater environments (although see Schultze, 2009) and that at times, they had a global, possibly equatorial, distribu tion (e.g., Glikman, 1964; Wurdig-Maciel, 1975; Hampe, 1993, 2003). From the earliest descriptions (e.g., Agassiz, 18331844), xenacanthiforms have been characterized by possession of a distinctive dentition of teeth with two main cusps on a relatively large rounded to subrectangular base, the so-called diplodont type from Diplodus, this original taxonomic name now being defunct (Zidek, 1993; Turner, 1997; Hampe, 2003). Xenacanthiform shark remains, especially teeth, are commonly found in Carboniferous to Permian strata around the world (Hampe, 1993:g. 1); possible older records have been found in recent years, especially in the Southern Hemisphere. The teeth are now dened as having a tricuspidate crown, with two large outer cusps and one or more small or weak (or rarely absent) median cusps, and a single lingually extended base. The dorsal side of the base bears a single coronal button (sometimes apical button or torus). On the labial rim of the base there is a single basal protuberance, the boss or heel. The terminology used within this paper is based on Hampe (1991, 1994:72, g. 8). The earliest articulated Northern Hemisphere xenacanths are found in the Lower Carboniferous (Visean) (e.g., Dick, 1981), whereas the youngest known are from the Triassic of Europe, U.S.A., India, and Australia (e.g., Turner et al., 2008; Ginter et al., 2010). They have been particularly well studied from coal measure basins in Britain (Hampe, 2003) and Germany (e.g.,
*Corresponding

author.

Schneider et al., 1988, 2000) and less so from North America, with teeth now used frequently in biostratigraphical studies in economically rich basins (e.g., Schneider, 1996, and see below). Hampe (2003:191) considered that xenacanthiforms had a practically simultaneous appearance in early Palaeozoic deposits of marine as well as freshwater environments, listing the Early Car boniferous (Visean) Diplodoselache Dick, 1981, from Scotland as the oldest taxon. However, he did not discuss older Australian and North American records of isolated teeth (e.g., Turner, 1993), with one (Turner, 1982; Burrow et al., 2010) referred to Xenacanthus and other examples thought to be cf. Diplodoselache, but with some resemblance to those of Hagenoselache Hampe and Heidtke, 1997, from Germany (Turner, 1993). Previously Hampe (1993) had doubted the existence of Devonian xenacanthiforms, although a cladistic analysis by Hampe and Long (1999:g. 6) based on teeth, spines, and scales concluded that Antarctilamna Young, 1982 (mid-Devonian, E Gondwana), was the sister group of the clade. Antarctilamna is now placed in its own family and considered to be a senior synonym of Wellerodus, known from isolated mid- to Late Devonian teeth in the U.S.A. and Spain (Turner, 1997; Ginter et al., 2006, 2008, 2010). A series of xenacanthiform taxa in the Late Devonian to Early Carboniferous of the Southern Hemisphere (Silva Santos and Salgado Carvalho, 1970; Mensah, 1973; Oelofsen, 1981; Turner, 1982, 1993; Long and Young, 1995; Garvey and Turner, 2006) shows the teeth are found typically in association with a fauna including hybodontiform sharks (e.g., Dick, 1976; Turner, 1993), gyracanthids (Turner et al., 2005), and often tetrapods (Turner et al., 1996). The stepwise stratigraphical and geographical distribution of xenacanthiforms seems to be related to the LaurentianGondwana collision (see below). Only a few Early Carboniferous xenacanthiform taxa have been formally described based on teeth, with others noted in open nomenclature. Specimens found in the 19th century were assigned to Diplodus minutus (gured by Agassiz, 1843), and Diplodus parvulus, Dicentrodus (Cladodus) bicuspidatus, and Xenacanthus (Pleuracanthus) elegans described by

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 2, 2011 Turner et al. (1999) interpreted the poorly bedded and sorted sediment with comminuted plant remains characterizing QML1117 as a possible storm deposit. Latterly, Parker and Webb (2008) ascribed the Tetrapod Unit at QML1117 to their LimeFlake Facies, deposited during a storm-induced ood event onto the tidal channel oor in an estuarine environment. Taphonomically, the vertebrate remains occur in beds about 1 m in thickness and although sometimes concentrated in one area (see comments below), specimens appear to be randomly oriented. In addition to vertebrates, branchiopods (Leaia), plant macrofossils of the Lepidodendron ora (White, 1972), and a rich palynoora indicative of mid-Visean age occur within the formation (Veevers, 2000). From a nearby GSQ drill core, Playford (1977) recorded prolic miospores of Granulatisporites frusulen tus, a taxon found in the Famennian to Visean of Australia, and based on the palynoora gave a Visean age (Jones, 1996; Jones et al., 2000). The Ducabrook Formation is dated as mid-Visean (ca. 333335 Ma), equivalent to basal Asbian (V3b) (Heckel and Clayton, 2006), based on recognition of the Anapiculatisporites largus Assemblage (Jones and Truswell, 1992; Jones et al., 2000). The general faunal assemblage and paleoenvironments of the deposits closely resemble the tetrapod-bearing sequences in southeastern Scotland (Wood and Rolfe, 1985; Turner et al., 1999). MATERIALS AND METHODS The vertebrate locality, QM L1117, consists of grey-green friable to indurated sandstone with small to microscopic orangebrown rounded pebbles. The site was reported to the Queensland Museum in late 1995 and a preliminary eld trip conrmed the presence of tetrapods as well as sh taxa (Thulborn et al., 1996). Xenacanthiform teeth were rst found scattered on the surface and collected by Mr. Morris Hawkins in an area that we subsequently called Morriss Mound. The teeth were reported to museum staff during the rst major eld trip at Easter 1996 when samples of all rock types exposed and bones and teeth were collected from the surface (Turner et al., 1996, 1999). Material collected on this and subsequent eld trips has produced more than 120 xenacanthiform teeth and rare spine fragments from surface collecting, excavation of the main bone-bearing horizon, as well as breakdown of rock with subsequent picking in the laboratory, smaller specimens, broken cusps, and separate bases mostly being found in sieved residues. The jaw fragment QM37521 was collected in 2004 by Carl and Joan van der Smissen from a locality south of QM L1117, adjacent to Snake Range National Park. Manual preparation and freeing of exposed xenacanthiform teeth from the sediment was done using sharp needles. As the cusps are particularly vulnerable, specimens were often coated with paraloid or Mowitol. To uncover teeth in the sandy mudstone, blocks were rst immersed in water, then cooked in a microwave oven for 23-minute sessions. Muddy residue was decanted, washed through a 0.1-mm sieve, dried, then picked with brush or needles under a binocular microscope. Scanning electron microscope (SEM) images of the uncoated holotype tooth were produced using an Hitachi TM-1000 Tabletop ESEM; other teeth were coated with platinum and imaged using a Jeol JSM6300F Scanning Electron Microscope; ground thin sections were imaged using an Olympus BX-50 transmission microscope and DP-12 imaging system; larger images were taken using an Olympus SZ40 dissecting microscope and DP-12 imaging system or a Nikon Coolpix 995; gures were compiled using Adobe Photoshop. SYSTEMATIC PALEONTOLOGY Class CHONDRICHTHYES Huxley, 1880 Subclass ELASMOBRANCHII Bonaparte, 1832 Order XENACANTHIFORMES Berg, 1940 Family DIPLODOSELACHIDAE Hampe, 2003

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Traquair (1881, 1888, 1903; see Hampe, 1999, 2003). Dick (1981) described the articulated Diplodoselache woodi, and Hampe et al. (2006) reported on Dicentrodus sp. teeth from Iowa, U.S.A. Turner (1982, 1993) and Lebedev (1996) recorded xenacanthiform Diplodoselache-like teeth in the late Tournaisian to early Visean of Australia and Tournaisian of Russia. A single spine was described as Xenacanthus tocantinsensis Silva Santos and Salgado Carvalho, 1970, from the Lower Carboniferous Poti Formation of northern Brazil, then the oldest xenacanthiform. Turner (1982:g. 8K; Burrow et al., 2010:g. 6S) illustrated an older tooth from the basal Carboniferous of Queensland. The Middle Paddock site (mid-Visean Ducabrook Formation) in central Queensland, eastern Australia, has yielded the rst Carboniferous tetrapod remains from the Southern Hemisphere (Thulborn et al., 1996; Warren and Turner, 1999, 2004; Warren, 2007; see these references for locality information and maps); macro- and microremains of shes (Turner et al., 1999); gyracanthid acanthodian remains (Turner et al., 2005); scales and bones from paleoniscoids, rhizodonts (Turner et al., 1996; Johanson et al., 2000; Parker et al., 2005); lungsh (Turner et al., 1999; Kemp, 2001); and shark material including Ageleodus teeth, hybodont spines, and possible scales, as well as the xenacanthiform remains described below. This paper describes the new xenacanthiform material from Middle Paddock, comparing it with teeth from the earlier Narrien and Raymond formations of Queensland as well as with taxa from elsewhere. Institutional AbbreviationsANSP, Academy of Natural Sciences Museum, Philadelphia; NHMUK PV P, The Natural History Museum, London; FMNH PF, Field Museum of Natural History, Chicago; QMF, Queensland Museum Fossil collection, Hendra, Brisbane.

GEOLOGICAL SETTING The Drummond Basin in east-central Queensland constitutes a discrete Upper DevonianLower Carboniferous depositional entity, the mostly undeformed sediments of which are exposed over about 25,000 km2 (Veevers, 2000:gs. 79, 260). In Late Devonian to Early Carboniferous (Mississippian) times it was an intermontaine basin, receiving up to 12,000 m of mainly uviatile and lacustrine sediments via a northward drainage system (Olgers, 1972) with shallow-marine incursions from the northeast (Veevers et al., 1984; Day et al., 1983). Within the basin, apart from the one tetrapod-bearing member mentioned above, sh have been found in marginal to non-marine sediments in the Upper Telemon, Raymond, Ducabrook, Star of Hope, and Bulliwallah formations. Turner (1982, 1993) and Fox et al. (1995) estimated the age of the Raymond Formation vertebrate-bearing strata as early to mid-Visean. The younger Bulliwallah Formation is exposed in the northern basin (Turner and Cook, 1999; Burrow, 2004; Turner et al., 2005), with the contemporaneous Ducabrook Formation exposed mainly in the southern part. The new xenacanthiform elements were found in the Tetrapod Unit and age-equivalent rocks of the Ducabrook Formation, a thick (>2100 m) sequence of interbedded sandstones and mudstones, predominantly khaki-brown and olive-green in color, with occasional beds of conglomerate, tuff, oolitic limestone, and strings of algal nodules and vertebrate debris (Mollan et al., 1969; Olgers, 1972). The sediments probably accumulated in shallow marginal, lagoonal, uviatile, and lacustrine settings on a oodplain, and, before discovery of tetrapod material at the Middle Paddock locality QML1117 in 1995, had yielded paleoniscoids (Elonichthys) (Hill and Woods, 1964; Turner and Long, 1987) and occasional acanthodian spines (Gyracanthides) (Turner et al., 2005). Some material described here was obtained from correlatable beds on the adjacent Mowbray Station.

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FIGURE 1. Reginaselache morrisi, gen. et sp. nov., holotype, tooth QMF39642. AC, laterolingual view, with B, closeup view of cusp showing wavy, branching cristae, and C, closeup of heavily pitted button and base surface; D, E, labiobasal view, with closeup of median cusp in E; FH, occlusal view, with closeup of larger curved cusp and small median cusp in G, and of button in H. All scale bars equal 1 mm.

RemarksHampe (2003) revised the diagnosis for xenacanthiform families, unifying the basal genera (Diplodoselache, Dicentrodus, Lebachacanthus, Orthacanthus, and Hagenoselache) in one family, the Diplodoselachidae. His revised diagnosis did not include tooth characters, and his cladistic analysis of the order shows the group as a grade rather than a clade (Hampe, 2003:g. 24). A new cladistic analysis (see below) indicates that Diplodoselache and Reginaselache form a poorly supported sister group to other xenacanthiforms. Genus REGINASELACHE, gen. nov. Type SpeciesReginaselache morrisi DiagnosisAs for type and only species. EtymologyFrom the Latin, regina, a queenin allusion to its place of discovery, Queensland, and selache, a shark. REGINASELACHE MORRISI, sp. nov. (Figs. 14AM, 5, 8A) cf. Diplodoselache woodi: Turner et al., 1996:69A. Xenacanth: Turner et al., 1999:178. Xenacanths: Johanson et al., 2000:168. A new species of xenacanthidid: Warren and Turner, 2004:155. A new xenacanthiform: Turner et al., 2005:964. Shark: in part, Parker and Webb, 2008:table 2. HolotypeTooth QMF39642 (Figs. 1, 8A). Other Referred MaterialsParatypes are ca. 120 teeth, isolated cusps, and thin sections of teeth including gured specimens QMF3959939618, 3962139623, 39648, 39649 (Figs. 2AV, 3, 4AM, 5); ungured specimens QMF37392,

QMF3962739629, 3963339641, 39643, 3964539647; and teeth including QMF39644 from nearby Mowbray Station. Other materials including spine fragments, scales, and a lower jaw are tentatively assigned to this species (see below). Type Locality and StratigraphyMiddle Paddock, QM L1117 (details of site kept by the Queensland Museum), Ducabrook Station near Springsure, central Queensland; Tetrapod Unit, Ducabrook Formation; Mississippian, mid-Visean (see above). DiagnosisXenacanthiform with presumed dorsal spine, and numerous small to medium-sized teeth 1.58.5 mm high, slightly asymmetrical with one of two major cusps being thicker than the other; all cusps with subcircular parabasal section and no lateral carinae; two lateral (main) cusps with ve to six wavy cristae on labial side of cusp and up to 20 cristae in total; cristae extending from cusp apex down almost to cusp/base interface, some branching towards base; median cusp not greater than half main cusp height, set slightly labiad; angle between teeth and base ca. 100 ; large rounded high coronal button separated from cusps, positioned in center of dorsolingual base surface, occasionally connected to region of median cusp by narrow ridge; several foramina, fewer than ve, around base of coronal button; base extends lingually beyond coronal button; base robust, circular to heartshaped or subtriangular outline; basal tubercle prominent and more or less equal to height of base, ventrally expressed as a wide horseshoe shape; basal surface with central depression, often with wide groove leading to midpoint of lingual edge; vascular canal foramina large, with no uniform pattern except around edge of basal concavity or lingual to labial boss surface; one large foramen usually in center of basal depression, with less than ve other foramina around edge of central concavity.

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FIGURE 2. Reginaselache morrisi, gen. et sp. nov., teeth. AE, QMF39599 in laterocrown, lateral, labiolateral, laterobasal, and closeup of cusp tip views; FJ, QMF39600 in labial, lateral, linguolateral, labiolateral, and closeup of cusp tip views; KN, QMF39601 in crown, closeup of median cusp base, anterocrown, and laterocrown views; O, P, QMF39602 in anterocrown and crown views; Q, QMF39603 in crown view; R, QMF39604 in laterocrown view; S, T, QMF39605 in basal and lateral views; U, V, QMF39606 in basal and laterobasal views. Scale bar equals 1 mm in AD, FI, K, M, NV, and 0.1 mm in E, J, L. All images are SEMs except F, which was taken in normal light before the tooth was extracted from the matrix and the median cusp lost; the image was equalized in Photoshop for better contrast.

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FIGURE 3. Reginaselache morrisi, gen. et sp. nov., teeth. AD, QMF39607 in laterolabiobasal, basal, labiobasal, and lateral views, respectively; E, F, QMF39608 in occlusal and basal views; G, H, QMF39609 in occlusal and basal views; I, J, QMF39610 in occlusal and basal views. All scale bars equal 1 mm; sketches B, D not to scale. All drawings Dr. S. Turner.

Etymologymorrisi, for Mr. Morris Hawkins of central Queensland, the rst collector of the material.

Description Tooth MorphologyThe teeth range in size from 1.5 to 8.5 mm high and wide. Nearly all teeth are tricuspid and slightly asymmetrical, with one main cusp slightly larger and thicker than the other (Figs. 1F, 2C, K, O, Q, 3A, C). Preservation varies from good with sharp cristae on the cusps to very worn (Fig. 2Q) with almost smooth shiny cusp surfaces, and color varies from light brown to orange to dark brown to black. The smaller lateral cusp is straighter, almost vertical to the base, with the larger cusp curving slightly away laterally. Isolated broken cusps, which when only one is detached is usually the shorter of the two, are identiable by shape and surface ornament. The small median cusp, rarely up to half the height of the larger cusps (Figs. 1, 2BC, F, 3G) and usually present through the tooth size range but often broken off, is set slightly labiad to the main cusps. The teeth bear numerous (820) cristae on the two main cusps, extending almost to their bases, with no lateral carinae. The cusps are usually subcircular in cross-section (Figs. 2K, Q, 4A, D), but can be ovoid to subrhombic. The large prominent rounded button on the dorsolingual basal surface is separated from the cusps in most specimens (Figs. 1, 2A, G, K, P, Q, R); in a few, there is a narrow median shaft connecting the coronal button to the median cusp. The button is up to 2.5 mm high, with a small number of foramina around its rim (Figs. 1H, 2A, Q). The base extends lingually beyond the button and, labiolingually, the outline is ovoid to heart-shaped (Figs. 2K, M, S, 3A, B) with a rounded rim and a few large foramina. The ventral undersurface of the base is concave, with in most specimens a large centrally placed foramen opening into the concavity (Fig. 2TV). On the mid-ventral rim of the labial surface there is a prominent basal boss with a distinctive horseshoe shape that extends into the central basal depression (Figs. 2D, SV).

The holotype QMF39642 (Figs. 1AH; 8A) is 3.0 mm deep labiolingually, 6.0 mm high, and 3.0 mm wide, with both lateral cusps intact but fractured and 0.7 mm of smooth median cusp preserved (Fig. 1E). Each lateral cusp has ca.1012 cristae, several of which branch close to the cusp base (Fig. 1B). The coronal button is rounded and heavily pitted, as is the rest of the upper surface of the base (Fig. 1C, H). A typical tooth QMF39599 (Fig. 2AE) is 4.5 mm deep, 4.5 mm high, and 4.0 mm wide at the base of the cusps, with a rounded coronal button, horseshoe-shaped basal boss, ovoid base outline, and a medial notch on the lingual edge. Although multiply fractured, both main cusps and a central cusp are preserved. Main cusps have ca. 16 well-preserved, wavy vertical cristae, and worn tips; the central cusp is relatively robust and 1.5 mm high. One of the smallest teeth QMF39601 (Fig. 2KN) is 1.5 mm high and wide, and 2 mm deep labiolingually. The main cusp tips and median cusp have broken off; all three have a circular parabasal section. Cristae are sharp, with each main cusp having ca. 10 extending down to the cusp base. One relatively large foramen opens out on the lingual face of the round coronal button. QMF39607 (Fig. 3AD) has at least six, probably seven to eight, ne but prominent cristae on the cusps, with one ending about half way up almost coalescing with another crista. Cristae can be of slightly different length, and not always parallel (e.g., Figs. 2B, H, K, 4B, C, F, HJ, M), with some more sinuous. QMF39614 (Fig. 4E) is a 2.5-mm-high cusp from a mid-sized tooth, also with sharp cristae extending almost to cusp base. QMF39615 (Fig. 4FG) is an isolated cusp from a small tooth, with sharp cristae but a worn and rounded tip. Largest teeth are 5 mm wide, 55.5 mm labiolingual base length, and 5 mm high, e.g., QMF 3960839610 (Fig. 3EH). QMF39608 (Fig. 3E, F) has one cusp base intact, with one broken and a part-depressed fracture, and a small, elongated central cusp. This specimen shows clearly the basal and side conguration, the coronal button separated from the cusps, high and rounded and placed in the center of the dorsolingual base. The main cusps are subrectangular, and ve or more cristae on the labial surface of the broken cusp that do not quite reach to the

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FIGURE 4. Reginaselache morrisi, gen. et sp. nov., tooth cusps and possible scales. A, B, loose cusp QMF39611 in apical and lateral views; C, cusp QMF39612 in lateral view; D, broken cusp QMF39613 in apical view; E, loose cusp QMF39614 in lateral view; F, G, loose cusp QMF39615 in lateral and closeup views; H, loose cusp QMF39616 in lateral view; I, J, loose cusp QMF39617 in lateral and apicolateral views; KM, broken, squashed cusp QMF39618 in ?lingual, apicolateral, and apical views. ?Reginaselache morrisi N, scale QMF39619, in crown view; O, scale QMF39620, in lateral view. Scale bar equals 1 mm in AE, KM, and 0.1 mm in FJ, N, O.

cusp/base interface. There is a short apron below the shiny part of the cusp, which presumably was covered in life with epithelium. On QMF39609 (Fig. 3G, H), which measures 6.5 mm 7.5 mm, the base is rather worn, exhibiting a medial groove in the basal concavity. The midpoint of the lingual edge can be deeply notched with a corresponding groove on the underside of the base leading into the central depression (e.g., QMF39608; Fig. 3F). QMF39610 (Fig. 3I, J) has a well-preserved tumid base and a large asymmetric, tumid labial boss; both cusps have wellpreserved ornament but broken tips. Tooth HistologyThin sections (Fig. 5) show that the cusps are composed of an outer relatively shiny orthodentine layer with very ne dentine tubules branching near the outer surface (Fig. 5DF) and an inner core of trabecular dentine ( = osteodentine). Although Hampe (2003) stated that all xenacanthid teeth lack enameloid, a thin outer layer of clear tissue appears to be present in Reginaselache morrisi in some areas (Fig. 5A, B, DG). How-

ever, the layer is not birefringent under polarized light and hence lacks the oriented crystals that typify enameloid (and enamel) and thus this tissue is akin to that called durodentine (e.g., Gross, 1967). The basal tissue is trabecular dentine with a few large canals that penetrate the base with large foramina. The coronal button is also formed of trabecular dentine with large, mainly transverse canals, and is separated from the basal tissue by a thin layer of orthodentine (Fig. 5C, H, I). Tooth PathologyHampe (1997) discussed anomalous growth forms in xenacanthiform teeth, and rare teeth of Reginaselache show evidence of one or two minute cusps between the median and lateral cusps (Fig. 2I, Q, R), indicative of minor developmental anomalies. One isolated cusp (Fig. 4KM) seems to have been crushed before fossilization. Hampe (1997) did not report wear on cusp tips in xenacanthiform genera, whereas several teeth of Reginaselache have cusp tips that have been rounded off and worn down, often with the cristae also worn off well

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FIGURE 5. Reginaselache morrisi, gen. et sp. nov., thin sections of teeth. A, QMF39648, cross-section of lateral tooth tip under blue light excitation, showing growth lines in orthodentine; B, QMF39623, slightly more basal cross-section of a tooth cusp; C, D, I, vertical labiolingual section of tooth QMF39621 showing C, the whole section, D, closeup in polarized light of segment indicated by arrow in C, I, closeup of the structure of the button; EG, cross-section of tooth cusp base QMF39622, E showing nearly the whole section, F a closeup of section with a crista, and G the same closeup in polarized light; H, QMF39649, horizontal section of base, labial edge at bottom, under blue light excitation, showing osteodentine/trabecular dentine with orthodentine layer around lateral and labial margins of the lateral cusps, but not in the base of the central cusp?. Arrows indicate approximate levels of cross-sections. Scale bar equals 1 mm in C, E, 0.1 mm in B, D, I.

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 2, 2011 other xenacanthiform elements are from a different species, because we have not found the spines and other material in articulation with the teeth, we have only tentatively assigned these other elements to R. morrisi. For the purpose of analysis, however, we have treated all as one taxon (see below). MaterialsScales QMF39619, 39620 (Fig. 4N, O); dorsal spine fragments QMF39624, 39625, 39630, 39631 (Fig. 6 AJ), and 39632 from QM L1117; the posterior half of a lower jaw QMF37521 (Fig. 7) from a locality near the Snake Range NP.

below the tip (e.g., Figs. 2E, J, N, 4F, J). These teeth have not been broken either mortem or postmortem, during burial, or during discovery or preparation but seem to have undergone wear during life (see discussion below). ?REGINASELACHE MORRISI (Figs 4N, O, 67) RemarksAlthough all the teeth recovered from the localities are deemed to be from one species, and it is unlikely that the

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FIGURE 6. ?Reginaselache morrisi, gen. et sp. nov., n spine fragments. AC, spine tip QMF39624.a in lateral (leading edge up), trailing edge, and lateral (leading edge down) views; D, longitudinal section QMF39624.b through one side of a more proximal fragment of the same spine (leading edge up, through bases of denticles); E, transverse section QMF39624.c through another more proximal fragment; F, G, spine tip QMF39625 in lateral (leading edge up) and trailing edge views. HK, spine QMF39631, preserved as proximal leading or trailing edge fragmented piece (removed from matrix) plus mid to distal segment, and impression of distal tip in matrix (H); I, proximal end of main segment; J, closeup of denticles; K, distal end of main segment. Scale bar equals 1 mm in AE, 10 mm in FK. Abbreviation: dr, denticle row.

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FIGURE 7. ?Reginaselache morrisi, gen. et sp. nov., partial lower jaw QMF37521. A, lateral view, anterior to left; B, medial view; C, occlusal view. Scale bar equals 10 mm.

Description ScalesScales found at the type locality that might be from Reginaselache morrisi are robust, thick-based rhombic scales similar to those of Holmesella Gunnell, 1931. They are ca. 1.5 mm wide and high, often with only the tumid base and a worn remnant of the crown preserved. Two scales only have been recovered with well-preserved crowns (Fig. 4N, O). Histology is the same as that in Holmesella (Zangerl, 1968:g. 2G) and Protacrodus (Gross, 1973:gs. 27, 28), with a crown having centrifugally arranged, apposed growth zones composed of orthodentine, on an acellular lamellar bone base. Spine MorphologyAll fragments are symmetrical and have rough, narrow, closely spaced ridges running longitudinally. QMF39624.a (Fig. 6AC) and QMF39625 (Fig. 6F, G) are distal tip fragments each ca. 20 mm long, with a double row of ca. 1 mm denticles along the presumed posterior or trailing edge, closely spaced at seven denticles per centimeter in each row. Only the bases of the denticles are preserved, with all projecting apices broken off. The largest preserved specimen is QMF39631 (Fig. 6HK), with an estimated length of 100 mm. The spine fragments comprise the mid 50 mm, plus the impression of one side of the more distal 5 mm, and the trailing or leading half of the proximal end (Fig. 6H). The rounded impression of the distal end indicates the distal spine tip was broken off and worn down in life, or at least before fossilization. The spine is laterally attened; maximum depth 10 mm and width 5 mm, both near the middle of the spine, which tapers towards distal and proximal ends (Fig. 6I, J). Narrow, irregularly arranged but longitudinal ridges separated by narrower grooves cover the whole spine. Two rows of smooth, recurved pointed denticles extend for the distal-most 15 mm, oriented posteriorly (cf. life position of spines) along each side of the trailing edge; some denticles are preserved (Fig. 6K), but most are represented only by their bases, with the rest broken off during collection/preparation.

Spine HistologyExternal longitudinal ridges and denticles are formed of osteodentine; no dentine tubules are visible in the bases of denticles cut through in longitudinal section (Fig. 6D). Small fragments broken off from QMF39624 show that the internal structure is wholly composed of a highly trabecular osteodentine (Fig. 6D, E). There is no evidence of an outer enameloid or orthodentine layer. RemarksAssuming that all fragments are from the same species, the largest spines must have been at least 100 mm long in life, denticulated only along the distal quarter or less. Given the family assignment, the dorsal spine in Reginaselache would have been placed anterior to the dorsal n rather than on the back or top of the head as in more advanced xenacanthidids (e.g., Soler Gijon, 1997:g. 6). Lower Jaw FragmentSpecimen QMF37521 (Fig. 7AC) is the posterior two-thirds of a robust, heavily calcied Meckels cartilage; total length is 105 mm. At the anterior fracture surface the jaw is 25 mm deep, and 30 mm at its deepest just anterior to the shelf that is presumed to demark the lower limit of the dental lamina. The ventral edge is slightly convex and thickened, curving more tightly posteriorly and forming a ange behind the circular articular cotylus. A shallow groove for the adductor muscle extends from the anterior limit of the cotylus to the buttress running up from the ventral edge, at the front of the ange. The dorsal edge of the jaw bulges upwards in front of the cotylus to form an articulatory surface, a feature referred to by Ginter and Maisey (2007) as a mandibular knob, which slotted into a quadrate concavity on the palatoquadrate. The dorsal edge attens out in front of the knob to form a slightly concave shelf medially, under which the ceratohyal is presumed to have lain. The shelf fades out near the end of a low ridge that extends forward from the shelf along the middle of the medial face of the jaw. This ridge probably represents the base of a shallow, featureless dental groove. No evidence is preserved of the position of the dental families on the jaw. COMPARISON Hampe (1999, 2003) revised the British Lower Carboniferous taxa, enabling a detailed assessment of the relationship between Diplodoselache, the new Australian taxon, and other xenacanth taxa. Thus we can compare Reginaselache morrisi with the oldest known articulated xenacanthiform Diplodoselache woodi from the contemporaneous Visean Lower Oil Shale Group of Scotland (Fig. 11). Diplodoselache woodi has numerous small bicuspid teeth (Dick, 1981:g. 12) of the type rst gured by Agassiz (Diplodus minutus in 18331844:v. 3, table 22b, gs. 68; listed as nomen nudum because the specimens were incomplete and are now lost) from the Burdiehouse Limestone (Hampe, 2003). Agassiz did not record a median cuspule in these teeth. The Middle Paddock teeth show the same size range (18 mm) as those described by Agassiz. Diplodus was recorded also from younger horizons including the Blackband Ironstone of the Burghlee (or Borough Lee) Limestone (Limestone Coal Group) and the South Parrot Coal (Upper Limestone Group B-P1) (e.g., Traquair, 1881, 1903, 1905) and Gilmerton (Dick, 1981), but there is no clear idea given in these papers of cusp ornament or of basal conguration; such characters were rarely identied by earlier workers. The somewhat similar teeth from the Visean Borough Lee Ironstone, for which Traquair (1881:36) erected Diplodus parvulus, have a median cuspulea small boss or knob which is frequently divided or notched into several minute rounded lobules, a carina that runs up the labial margin of the base ending on this knob, and tooth cusps that also bear carinae or ridges towards their apices (Hampe, 2003:g. 6). This taxon, which Hampe (2003) reassigned to Diplodoselache parvulus, is recorded from several sites in eastern Scotland, including Borough Lee and Pitcorthie (Hampe, 2003). Hampe

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FIGURE 8. Drawings of teeth from Reginaselache and other xenacanthiform genera, showing specimens from type species where possible; lingual views unless noted. A, Reginaselache morrisi holotype, lateral view; B, Diplodoselache woodi (Hampe, 2003:g. 4a); C, Dicentrodus bicuspidatus (Hampe, 2003:g. 8a); D, Hagenoselache sippeli, lateral view and coronal view of isolated base (Hampe and Heidtke, 1997:g. 4B, C); E, Lebachacan thus senkenbergianus (Soler-Gijon, 1997:g. 2 in part; as far as we are aware, no detailed drawings of Lebachacanthus teeth have been published); F, Wurdigneria obliterata, lingual and lateral views (Richter, 2005:g. 10C, D); G, Triodus sessilis (Heidtke et al., 2004:g. 6b); H, Orthacanthus gibbosus (Hampe, 2003:g. 9b); I, Xenacanthus laevissimus (Hampe, 2003:g. 15a); J, Bohemiacanthus sp. (after Schneider, 1988:g. 1); K, Plicatodus plicatus (Hampe, 1997:g. 1). All scale bars equal 10 mm.

et al. (2006:g. 3) gave the stratigraphic ranges of Scottish Early Carboniferous xenacanthiform taxa (cf. Fig. 11). Diplodoselache woodi teeth (Dick 1981:g. 12) have a centrally placed coronal button and a small central cusp, with main

FIGURE 9. Cladistic analysis of outgroups Leonodus, Antarctilamna, and Phoebodus, and xenacanthiform genera represented by teeth plus other elements. As only 11 taxa were thus included, an ACCTRAN branch-and-bound search was undertaken using PAUP 4.0b10 (Swofford, 2002), all 19 characters unordered and of equal weight; multistate taxa were treated as uncertainty. The single shortest most parsimonious tree is 40 steps, CI = 0.7750, RI = 0.7632. Changes at nodes are listed in Appendix 2.

cusps that are laterally compressed, sharply carinate but otherwise smooth (Fig. 8B); the Ducabrook teeth differ in all these characters. Of the other non-xenacanthid genera, Dicentrodus differs from Reginaselache by having extremely asymmetrical teeth with a at base and divergent main cusps with a lanceolate cross-section and nely serrated edges (Hampe et al., 2006; Figs. 8C, 10E, F); Orthacanthus teeth always have a minute median cusp, some serrations on the main cusps, and a basal tubercle without a concave depression (e.g., Hampe, 2003; Fig. 8H); Lebachacanthus Soler-Gijon, 1997, teeth have massive, serrated main cusps (Hampe, 2003; Fig. 8E); and Hagenoselache teeth are small (less than 3.5 mm) with their median cusp 3/4 or 4/5 the height of the main cusps, and the angle between crown and base 120125 (Hampe and Heidtke, 1997; Fig. 8D). The other described xenacanthiform taxa from Gondwana is the younger Permian Wurdigneria (Richter, 2005; Fig. 8F); the teeth are robust but have smooth cusps with only a lateral carina and no cristae. The older xenacanthiform teeth from the nearby Narrien Range (Turner, 1993:g. 4CG) resemble more closely those of Hagenoselache sippeli Hampe and Heidtke, 1997, from the Namurian B of Germany. The teeth of Reginaselache show some of the diagnostic characters of the Xenacanthidae as revised by Hampe (2003), most closely resembling those of Triodus (and Bohemiacanthus Schneider, 1996, if this is a valid genus) but differing in having slightly sinuous cristae that extend from the tip to close to the base of the main cusps, and in the histological structure of coronal button and underlying base. The Permian xenacanthid Plicatodus is the only other xenacanthiform to have wavy cristae on the main cusps, but its teeth have a at base, many nutrient foramina, and a small at coronal button (Hampe, 2003). The spine fragments referred here to Reginaselache are compatible with assignment to a xenacanthiform shark in having a double row of posterior denticles, lacking a posterior groove, and being wholly composed of osteodentine. The rough ridges forming the outer surface of the spines are comparable to the bark-like ornamentation described for Diplodoselache woodi (Hampe, 2003:199, g. 4k) rather than the smoother and/or more

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FIGURE 10. Late Devonian and Early Carboniferous xenacanthiform teeth, some not yet assigned to species. AD, Diplodoselache-like tooth QMF13391, Tournaisian Telemon Formation, Narrien Range, central Queensland (ST drawings) in A, lingual view; B, occlusal view; C, lateral view; D, labiobasal view. EH Dicentrodus sp. 1A FMNH PS-14028a, Heimstra Quarry, near Delta, Iowa (ST drawings), in E, lingual view; F, lateral views; G, labial view; H, basal view. I, xenacanthiform tooth ANSP 23232, mid-Famennian of Red Hill, Pennsylvania, in labial view. J, K, Diplodoselache? antiqua (Lebedev 1996:g. 9b, d), Tournaisian of Tula, Moscow Basin. L, M, JHQ-09 xenacanthid? tooth FMNH (PF15378) from the mid-Visean Upper St Louis Formation, Heimstra Quarry, Delta, Iowa. Scale bar equals 0.1 mm in L, M; 0.5 mm in EG; 1 mm in AD, H, I.

nely ribbed Xenacanthus and Orthacanthus spines (Soler-Gijon, 2000; Soler-Gijon and Siebert, 2001). The dorsal spine on Diplodoselache woodi (NMS 1974.51.4), which lies near the front of the dorsal n approximately half way between the level of the pectoral and pelvic ns, is estimated to have been 75 mm long with a maximum width of 9 mm (Dick, 1981); the spine on Reginaselache was longer and more slender. Hampe (2003:231) grouped the more basal xenacanthiforms in the family Diplodoselachidae based on the autapomorphic character of a dorsal spine which always has a rounded cross-section and a ventrally arranged double row of denticles. Based on the denticle pattern, Reginaselache groups with his diplodoselachids rather than the xenacanthids, which have laterally extending posterior denticle rows, but differing from all other xenacanthiforms in having a laterally attened cross-section plus posterior denticle rows. The lower jaw (Fig. 7) is typical xenacanthiform, with the secondary articulation anterior to the articular cotylus shared only by xenacanthiforms and modern elasmobranchs (Ginter and Maisey, 2007). The dental furrow is smooth as in Xenacanthus, rather than subdivided by shallow grooves as in some other groups of Paleozoic sharks (Dick, 1981). The uncompressed, 3D preservation of QMF37521 gives an unequalled view of the structure of a xenacanthid Meckels cartilage. The proportions of the jaw, by comparison with those of Diplodoselache (e.g., NMS.GY.1974.51.4 in Dick, 1981:g. 2), indicate that the whole sh would have been over a metre in length. The scales tentatively assigned to Reginaselache morrisi are of the kind found posterior to the pelvic region on Diplodoselache

(Dick, 1981:g. 11c). If Reginaselache morrisi also had more fragile, thin-based scales similar to those found on the rest of body of Diplodoselache (Dick, 1981:g. 10), it is unlikely that they would have survived the turbulent depositional environment represented in the type locality. DISCUSSION Zidek (1993) recognized dental morphology as a valid criterion for distinguishing xenacanthiform species but was doubtful regarding the usefulness in dening all genera. Schneider (1996), Hampe and Heidtke (1997), and Hampe (2003), for example, have made it clear that full descriptions of the morphological (Fig. 8) and histological characteristics of teeth do enable denition of genera. We agree with Schneider (e.g., 1988) that cusp characters can aid in analysis of taxa in combination with tooth base characters. Even though Hampe and Heidtke (1997) stressed the number of cristae as an important factor relating Hagenoselache sippeli to Triodus carinatus, Hampe (2003) placed the latter in the Xenacanthidae and the former in the Diplodoselachidae. Despite some overall similarity, the illustrated teeth of Hagenoselache have few cristae on the cusps, and those only close to the tip, so that Reginaselache teeth differ in that feature. Perhaps the specic tooth morphology, particularly ornament, depends on diet at least as much as on phylogenetic relationships. We undertook a cladistic analysis (Appendix 1, Table 1, Fig. 9) of xenacanthiform genera known from more than isolated teeth,

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 2, 2011 the button area is composed of the same trabecular osteodentine as forms the rest of the base. In the resulting cladogram, the xenacanthiforms are united by ve characters (see Appendix 2), and within the xenacanthiforms, Diplodoselache and Reginaselache form a sister group to the other genera, although the node supporting their relationship is only poorly supported (Appendix 2), with the single synapomorphy being the coronal button composition. Nevertheless, we tentatively refer Reginaselache to the family Diplodoselachidae, with Diplodoselache as the only other genus included. Our analysis indicates that Hagenoselache, Plicatodus, Triodus, and Xenacanthus form a better-supported clade. The node uniting the other xenacanthiform genera is more strongly supported, as is the family Xenacanthidae (Appendix 2). Paleobiogeography In Australia we now have found xenacanthiform teeth including Reginaselache and cf. Diplodoselache (Fig. 11) in the Early Carboniferous (e.g., Turner, 1982, 1993; Jones et al., 2000) from at least four horizons spanning the basal Tournaisian to mid-Visean in the Drummond Basin; a small tooth from one of the basal beds in the Telemon Formation is shown here (Fig. 10AD). Small (microscopic <5 mm) xenacanthiform teeth have also been noted in the Famennian Red Hill fauna of Pennsylvania (S.T., pers. observ., 2001; Fig. 10I); in the possible D/C boundary beds of Manseld, Victoria (Long in Garvey and Turner, 2006); and in the basal Tournaisian of Horton Bluff, Nova Scotia (S.T., pers. observ.). Turner (1993) described the chondrichthyan assemblage of the Raymond Formation, underlying the Ducabrook Formation, which contains a closely similar fauna to that from the Scot tish Lower Carboniferous (early to mid-Visean) at least at the generic level and compared the xenacanthiform teeth to those of Diplodoselache as illustrated by Dick (1981). Based on a survey of North American, some 50 Scottish, and the Queensland sites, she dened an Ageleodus-xenacanthiform-hybodontiform (Tristichius) (AXT) assemblage, which appears throughout nonmarine (including marginal to freshwater) facies. Fischer et al. (2010) recognized a similar facies association in the coal basins of Europe. Lebedev (1996) disputed the identity of the Raymond Formation assemblage and considered that scales called hybodontoid by Turner (1993) and the Ageleodus teeth were all specialized denticles belonging (with the associated xenacanthiform teeth) to an Early Carboniferous xenacanthiform shark. No articulated xenacanth material has been recovered as yet to support this hypothesis, although branchial denticles in some of Fritschs (1889, 1890) xenacanths do resemble Ageleodus. However, the strong basal growth and histological structure of Ageleodus and another of the unusual hand-like teeth, Cynopodius, mitigate against their not being teeth (e.g., Garvey and Turner, 2006). Nevertheless, it is possible that most of the scales gured

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FIGURE 11. Stratigraphical ranges of Early Carboniferous xenacanthiform taxa (cf. Hampe and Ivanov 2007:g. 5; extra data from: Garvey and Turner, 2006; Hampe et al., 2006:g. 3; Ginter et al., 2010; boundary dates from Ogg et al., 2008).

using tooth, spine, jaw, and scale characters to test the current ideas of relationship. We revised and extended the list of these characters from the analysis by Hampe (2003:g. 24), in some instances reversing character polarity based on our interpretation that the diplodont, rather than the cladodont, tooth form is plesiomorphous for xenacanthiform lineage (Appendix 1). The Devonian genera Leonodus Mader, 1986, Antarctilamna Young, 1982, and Phoebodus St John and Worthen, 1875, which also have diplodont teeth and have often been compared with xenacanthiforms (e.g., Ginter, 2004), were used as outgroups. Botella et al. (2005, 2009) investigated the histological structure of the cusps on Leonodus teeth; Botella (pers. comm., 2010) has advised that

TABLE 1. Character matrix for selected xenacanthiform and outgroup taxa. 1 Leonodus Antarctilamna Phoebodus Diplodoselache Hagenoselache Lebachacacanthus Orthacanthus Plicatodus Reginaselache Triodus Xenacanthus 0 1 1 2 2 2 2 2 2 2 2 2 0 1 3 1 2 1 1 2 1 2 2 3 0 0 1 1 0 2 2 0 0 0 1 4 0 0&2 0&1 0&2 0 2 2 1 1 0 0&2 5 1 0 0&1 1 1 1 1 1 1 1 1 6 0 0 0 0 ? ? 0 0 0 1 1 7 0 1 1 0 0 0 0 0 0 0 0 8 2 0 0 2 2 2 3 2 1 2 2 9 0 2 0 2 1 ? 1 1 0 0 1 10 0 1&2 1&2 2 2 2 2 2 2 2 2 11 0 0 2 0 1 1 1 1 0 1 0&2 12 0 0 0 0 1 0 0 1 0 1 0 13 0 ? 0 1 0 ? 0 2 1 2 0 14 ? 0 ? 0 ? 0 1 1 ? 1 1 15 0 0 ? 1 ? 1 1 2 1 2 2 16 0 0 ? 1 ? 1 2 2 1 2 2 17 0 0 ? 0 ? 0 1 1 ? 1 1 18 ? 0 ? 1 ? 1 1 1 1 1 1 19 0 0 0 0 1 1 ? ? ? 1 ?

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FIGURE 12. Lower Carboniferous xenacanthiform sites marked by solid triangles on an Early Carboniferous (Visean; 340 Ma) paleogeographical reconstruction (after Metcalfe, 1996) with estimated extent of southern glaciation (cf. Jones et al., 2000); Australia: Middle Paddock, Ducabrook Formation in the Drummond Basin, central Queensland; Snowy Plains Formation, Manseld Group, Victoria; Africa: Sekondi Formation, Ghana; Europe: sites in the U.K., mainly Scotland; eastern U.S.A. and Canada: sites in New Brunswick and Nova Scotia along northern Gondwana margins; see text for stratigraphical and geographical detail.

by Turner (1993) might belong to a diplodoselachid shark, when compared with those illustrated by Dick (1981). Lebedev (1996) had discovered teeth in the D/C boundary beds in Russia, which he named ?Diplodoselache antiqua, and these do have smooth cusps more like the type from Scotland than the teeth from Queensland. Further study of these taxonomic points will be considered elsewhere when further material is available. Interestingly, the youngest record for a diplodoselachid has been reported from the Permocarboniferous lacustrine (lagoonal?) limestone of Guardia Pisano basin, Sardinia (Fischer et al., 2010). Other isolated teeth from the Mississippian (mid-Visean, Holkerian) Upper St Louis Formation of Heimstra Quarry, near Delta, Iowa (e.g., Bolt, 1990), resemble those of Orthacanthus in having highly symmetrical smooth cusps with serrated edges (S.T., pers. observ., 1997; Fig. 10E, F), but compare most closely with those of Dicentrodus bicuspidatus from slightly younger beds in Scotland (Hampe, 2003); Hampe et al. (2006) assigned the American teeth to Dicentrodus sp. 1A, the oldest xenacanthid found in North America. The only other Mississippian xenacanthiform is also from Heimstra Quarry (FM collection), which John Bolt (pers. comm. to S.T., 1997) reports as a large, considerably attened specimen with a full set of upper and lower teeth (not been seen by the authors); an isolated tooth from a sample of Upper St Louis Formation bonebed is illustrated here (Fig. 10L, M). Following the climatic highs of the Devonian (e.g., Young et al., 2010), both marine and non-marine/marginal vertebrate records in Australia become increasingly sparse throughout the Carboniferous with the onset of the southern glaciation that predominated until later in the Permian (Jones et al., 2000; Fig. 12). Queensland, at least in Visean times, was still in the temperate region above 60 south (Veevers, 2000; Burrow et al., 2010) so that the Middle Paddock continental to marginal tetrapod and sh fauna, including Reginaselache, was able to ourish. Fielding et al. (2008:g. 3) have discerned the oscillating nature of the long southern glaciation event, and the vertebrate occurrences, including the Ducabrook Formation fauna, support the possibility of later mid-Carboniferous ice age onset in Queensland with milder interglacial times, with a series of assemblages recorded until well into the Pennsylvanian (Burrow et al., 2010).

The last record of a xenacanthiform in the Carboniferous of the southeastern Pangaean landmass (eastern Australia) is that of Reginaselache. The xenacanthiforms presumably then survived elsewhere (further north in equatorial Paleotethys: see Fig. 12) and only reappear in the stratigraphic record of eastern Australia in the Lazarus taxa of the early to mid-Triassic of the Sydney Basin (Turner et al., 2008). There exists a note about xenacanthid teeth from the Lower Carboniferous Sekondi Formation of Ghana, West Africa (Mensah, 1973), but absence of specimen data or illustrations does not allow us to gauge this record as truly xenacanthiform. Attempts (by S.T. and O. Hampe, pers. comm., 1999) to obtain material have so far proved unsuccessful. Elsewhere in the Lower Carboniferous of the former western Gondwana, Xenacanthus tocantinsensis (see above) is represented by only a small 30-mmlong spine. What is the relationship of Reginaselache to younger and better-known Carboniferous and Permian taxa from successive lacustrine (and/or possibly marginal lagoonal) horizons in Stephanian to Asselian basins (see e.g., Werneburg et al., 2007:g. 9)? Similar Triodus species assigned to Bohemiacanthus by Schneider and Zajc (1994) include T. carinatus, T. lauteren sis, T. palatinus, and T. obscurus (Ginter et al., 2010), with the latter three species showing a stratigraphically older to younger decrease in the number of labial and lingual cristae (Schneider, 1996:g. 8). There is still a large temporal gap to ll (see Hampe and Ivanov, 2007:g. 5) and search of suitable horizons in the southern hemisphere should help ll the gaps in the early xenacanthiform history, as has already been shown in Queensland, Brazil, and Ghana. Similarly, further work is needed in Pennsylvania and maritime Canada, which were part of, or closely associated with, the northern Gondwanan shore or drainage systems by the end-Devonian (e.g., Jones et al., 2000; Turner et al., 2005). Phylogenetically, Hampe (2003) placed xenacanthiforms as the sister group of phoebodontids. Ginter (2004) further discussed the relationships of Devonian examples of these clades plus Omalodontiformes. The teeth found in the Famennian of Red Hill (Fig. 10I) with three median cusplets possibly represent a transitional form related to Antarctilamna within the xenacanthiform lineage (cf. Ginter, 2004:g. 3). Hampe (2003) did not take into consideration the southern hemisphere occurrences of xenacanthiform teeth. Even if we accept that Leonodus, Antarctilamna (Wellerodus), and such pre-Famennian teeth and spines are not Xenacanthiformes s.s., the early incidences of xenacanthiforms in the Gondwanan regions (Australia, West Africa, Brazil, U.S.A., and eastern Canada), need to be taken into account (e.g., Turner et al., 2008). Paleobiology Comparison with Diplodoselache, Hagenoselache, and Triodus suggests that an adult Reginaselache morrisi was an animal of around 12 m in length, presumably with smaller juveniles, bearing a high tooth complement in each jaw. Based on other Lower Carboniferous xenacanthiforms (e.g., Dick, 1981; Hampe, 2003), the teeth from the type locality, which come from a small area a few metres square, could be from just one animal or a few individuals (most likely the latter because there are at least three spine tips from the same area), the 18 mm size range reecting the growth and tooth-le size range. However, smaller Reginaselache morrisi teeth in the sample might be from juveniles or sub-adults. The teeth exhibit a range of color and preservation conditions, which suggests that they were not all fossilized at the same time: some have smoother cusps; some broken cusps, which might have resulted from the relatively turbulent paleoenvironment and/or burial event; and some have cusps worn down on the tips only.

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 2, 2011 ther exploration especially in such deposits across the former northern Gondwana is warranted to verify this prediction. The new Australian xenacanthiform was a shallow-water dweller within the seaward part of the large Drummond Basin river system, probably capable of living in marginal and freshwater environments. Reginaselache morrisi was a medium-sized predator well-endowed with a dentition of probably 60 or more similarly sized teeth for feeding (possibly even chewing) on smaller shes or shelled invertebrates, and predictably, as a basal xenacanthiform, with a post-occipital dorsal n spine and heterocercal tail. Tooth cusp wear suggests a novel style of food processing for xenacanths. ACKNOWLEDGMENTS We are grateful to the Hawkins and Cann families, former and present owners of Ducabrook Station, as well as those of neighboring stations, for alerting us to the material and allowing access; to C. and J. van der Smissen (Brisbane) for nding and donating the jaw specimen; A. Warren (Melbourne) for preparation; and M. Brazeau (Berlin) for identifying the fragment as shark. S.T. thanks C. Dufn (London), O. Hampe (Berlin), J. Schneider (Freiberg), and G. D. Johnston (Dallas) for discussions and information on xenacanth teeth; and Hampe for help with histology. E. Daeschler (Philadelphia), S. McLean (Hancock Museum), J. Bolt and W. Simpson (FMNH), and the late C. Patterson (NHM) assisted with and gave permission to use museum collections and specimens. We are grateful to R. van der Kamp for translations; J. Braceeld, J. Ford, C. Northwood, and K. Stumkat for SEM and research assistance; C. Bonde and B. Tangey for preparation, curation, and databasing of specimens; and to all eld work participants. Work was initiated when S.T. was ARC Australian Research Fellow (19952000), nancially assisted 19971999 and 20002002 by ARC grants A39700915 and 00000629 to her and A. A. Warren. The Ian Potter Foundation, Melbourne, and the Field Museum provided travel grants to S.T. to visit the U.S.A. in 1997. We thank the Queensland Museum Board for basic facilities; QM librarians are ever helpful and Geoscience Australia also provided documents. Two referees and editor Dr. Charlie Underwood provided critical comments for the improvement of the manuscript in review. LITERATURE CITED
Agassiz, L. 18331844. Recherches sur les Poissons Fossiles. Imprimerie de Petitpierre, Neuchatel, 1420 pp. Andrews, S. M. 1982. The Discovery of Fossil Fishes in Scotland up to 1845 with Checklists of Agassizs Figured Specimens. Royal Scottish Museum, Edinburgh, 87 pp. Berg, L. S. 1940. Classication of shes, both recent and fossil. Travaux de lInstitute zoologique de lAcademie des Sciences de lURSS 5(2):85517. [Russian and English] ` Blieck, A., G. Clement, H. Blom, H. Lelievre, E. Luksevics, M. Streel, J. Thorez, and G. C. Young. 2008. The biostratigraphical and palaeogeographical framework of the earliest diversication of tetrapods (late Devonian); pp. 219235 in R. T. Becker and W. T. Kirchgasser (eds.), Devonian Events and Correlations. SDS Volume in Honour of M. R. House. Geological Society, London, Special Publication 278. Bolt, J. R. 1990. Mississippian vertebrates from Iowa. National Geographic Research 6:339354. Bonaparte, C. L. 1832. Selachorum tabula analytica. Nuovi Annali della Scienze Naturali (Bologna) 1:195214. Botella, H., P. C. J. Donoghue, and C. Martnez-Perez. 2009. Enameloid microstructure in the oldest known chondrichthyan teeth. Acta Zoologica 90:103108. Botella, H., J. I. Valenzuela-Ros, and C. Martnez-Perez. 2005. Mor fologa e histologa de las escamas de Leonodus carlsi Mader, un condrictio primitivo del Devonico Inferior de la Cordillera Iberica (Espana); pp. 205219 in G. Melendez, C. Martnez-Perez, S. Ros,

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The latter is surprising considering xenacanth sharks have been regarded as purely grasping predators; such pathological wear might represent manipulation of prey or more likely thegosis during life of the teeth in the tooth les. Johnson and Thayer (2009) did note indications of worn cusps in some of their xenacanthiform teeth from the Pennsylvanian of Arizona. In addition, other contemporaneous Early Carboniferous sharks from Scotland, Tristychius (which might also be present in the Narrien Range, cf. Turner, 1993) and Onychoselache, show wear on the tooth cusps to such a degree that Dick (1976) surmised a durophagous diet to produce these strong effects. Our contention, therefore, is that Reginaselache was not a shark that quickly grasped its prey and swallowed, but one that either might have masticated its food to some extent, or experienced opposable wear. The Ducabrook Formation, including beds above and below the Tetrapod Unit, preserves evidence of algal mounds that probably formed in standing lagoons; these could have been a possible source of silica-rich food on which Reginaselache might have grazed occasionally. Another shark in the assemblage, Ageleodus, known only from small hand-like teeth (Turner, 1993), might have been an algal nibbler and in turn been prey for the xenacanth. The spines presumed to be from Reginaselache are compa rable to those of Diplodoselache (Soler-Gijon, 2000), but the largest specimen is longer than that of Diplodoselache woodi (see above). The D. antiqua spine (Lebedev, 1996:g. 9g, h) shows similar lateral compression to that of Reginaselache, but the denticles extend one-third spine length (cf. <1/4 for Reginaselache). The ridges on Reginaselache are more bark-like as in D. woodi and not smooth and regular as in D. antiqua. As in other more primitive xenacanthiforms, the dorsal spine was presumably positioned in the post-occipital re gion (Soler-Gijon, 2004). No growth lines (cf. Soler-Gijon and Siebert, 2001) are seen within the ?Reginaselache spine tip (Fig. 6). Based on the work of Soler-Gijon (2004), the presence of closely spaced well-formed denticles on spines indicates that the animals were post-parturition with erupted spines, possibly from very young individuals. CONCLUSIONS The mid-Visean xenacanthiform shark Reginaselache morrisi, gen. et sp. nov., from the Ducabrook Formation of central Queensland, is a new taxon that has teeth morphologically akin to those of Triodus, Bohemiacanthus, Hagenoselache, and Xenacanthus found in marine and non-marine beds in the Late Carboniferous to Permian of Europe. The new shark is represented by its numerous diplodont teeth, with several associated spine fragments, a partial lower jaw, and scales highly likely to be from Reginaselache. Based on a cladistic analysis, Reginaselache is placed tentatively within the family Diplodoselachidae. However, the teeth share few morphological characters with the type Diplodoselache woodi and those of another purported but unlikely diplodoselachid, Wurdigneria Richter, 2005, although they do resemble other Scottish diplodoselachid taxa, especially Diplodoselache parvulus. Analysis of teeth and comparison with those from other Carboniferous sites worldwide shows close morphological similarities with those of the later CarboniferousPermian from central Europe. Histological structure is identical to that of the Early Carboniferous Diplodoselache. The spines of the two taxa are also similar, although that of Reginaselache is comparatively slender. Where records of older xenacanthiforms occur in the present Southern Hemisphere or in terranes that were part of Gondwana and based on our phylogenetic analysis as well as analysis of associated fauna, we surmise that the Xenacanthiformes originated in Gondwana, inhabiting non-marine/marginal environments. Fur-

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(12) Composition of tooth base (Hampe [11]): trabecular dentine (0); orthodentine (1). (13) Composition of coronal button (expanded from Heidtke et al. [13]): trabecular dentine (0); trabecular dentine dome with orthodentine cap (1); orthodentine (2). (14) Dorsal spine (Hampe [13]): present, posterior to neurocranium (0); present, articulates with neurocranium (1). (15) Spine cross-section (Hampe [14]): nodose ridged, triangular or laterally compressed (0); circular cross-section, double row of denticles ventrally (i.e., along trailing edge) (1) dorsoventrally compressed distally, lateral rows of denticles (2). (16) Spine composition (Hampe [17], Soler-Gijon [9] combined): trabecular dentine with orthodentine outer layer forming ridges (0); trabecular dentine, rough surface texture (1); trabecular dentine, smooth surface texture (2). (17) Adult spine width/length ratio (Hampe [15]): robust, <1:12 (0); slender, >1:12 (1). (18) Meckels cartilage prearticular protuberance (new character): absent (0); present (1). (19) Dermal denticles (Hampe [30]): polyodontode (0); monodontode (1). APPENDIX 2. Apomorphy lists for the nodes in the cladogram in Figure 10; Character numbers correspond to those listed in under the Change column repAppendix 1; the double arrow resents unambiguous changes and the single arrow represents ambiguous changes; all changes are one step; CI = consistency index.
Branch node 20 Leonodus node 20 node 12 Character 2 10 1 5 7 8 9 2 3 11 1 4 15 16 18 13 3 9 4 8 3 9 11 19 14 16 17 2 3 4 12 15 13 4 6 9 3 11 12 8 Steps 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 CI 1.000 1.000 1.000 1.000 1.000 1.000 0.500 1.000 0.500 0.667 1.000 0.500 1.000 1.000 1.000 1.000 0.500 0.500 0.500 1.000 0.400 0.500 0.667 1.000 1.000 1.000 1.000 1.000 0.400 0.500 0.500 1.000 1.000 0.500 0.500 0.500 0.400 0.667 0.500 1.000 Change 1 0 2 0 01 10 0 1 2 0 0 2 1 3 0 1 0 2 02 02 0 1 01 01 0 1 0 1 0 2 2 1 2 1 02 01 0 1 01 0 1 1 2 0 1 1 2 20 20 0 1 1 2 0 2 0 1 0 1 1 0 0 1 1 0/2 1 0 2 3

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APPENDIX 1. Tooth, spine, jaw, and scale characters and character states used in cladistic analysis; comparable characters used by Hampe (2002), Soler-Gijon (2000), and Heidtke et al. (2004) are noted in brackets. (1) Number of tooth cusps (Hampe [1], changed polarity): two (0); more than three (1); three (2). (2) Relative length of lateral versus median cusps (Hampe [2], extra states added): no median cusps (0); median cusp(s) <half height of lateral cusps (1); median cusps shorter but >half height of lateral cusps (2); median and lateral cusps equal height (3); median cusp higher than lateral cusps (cladodont) (4). (3) Cusp cutting edges (carinae) (Hampe [3], muricated edges state dropped): absent (0); smooth (1); serrated (2). (4) Cusp cristae (Hampe [5]): cristae straight (0); wavy vertically oriented cristae (1); absent (2). (5) Labial nutrient foramina (Hampe [6]): present (0); absent (1). (6) Coronal button (new character): separated from cusps (0); close to/contiguous with cusps (1). (7) Outline of tooth base (Hampe [7], polarity reversed): subcircular or labiolingually elongate (0); mesiodistally elongate (1). (8) Prominent basal tubercle (Hampe [9]): absent (0); present, horseshoe-shaped (1); present, circular with concave depression (2); present, circular without depression (3). (9) Nutrient foramina on basal surface (new character): large, 1 or 2 (0); large, >2 (1); small and few (2). (10) Enameloid layer on teeth (Hampe [12], new ancestral polarity state added based on Leonodus): present on upper half of cusp (0); present on whole cusp (1); absent (2). (11) Composition of tooth cusps (Hampe [10], polarity changed): trabecular dentine basal core plus orthodentine (0); orthodentine (1); trabecular dentine (2).

node 12 Antarctilamna node 12 Phoebodus node 20 node 19 (Xenacanthiformes)

node 19 node 13 (Diplodoselachidae) node 13 Diplodoselache node 13 Reginaselache node 19 node 18

node 18 node 17 (Xenacanthidae) node 17 node 16

node node node node node

16 node 15 15 Plicatodus 15 node 14 14 Triodus 14 Xenacanthus

node 17 Orthacanthus