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Early Jurassic Selachians from the Hasle Formation on Bornholm, Denmark

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Early Jurassic Selachians from the Hasle Formation on Bornholm, Denmark Early Jurassic selachians from the Hasle Formation on Bornholm, Denmark JAN REES Rees, J. 1998. Early Jurassic selachians from the Hasle Formation on Bornholm, Den- mark. - Acta Palaeontologica Polonica 43,3,439452. A selachian fauna from the early Pliensbachian (Uptonia jamesoni to Prodactylioceras davoei zones) is recorded from the Hasle Formation on the island of Bornholm. Three hybodont sharks, Hyb,odus reticulatus, H. delabechei and Lissodus hnsleensis sp. n., and three neoselachians, Synechodus occultidens, Paraorthacodus sp. andAgaleus dorseten- srs, are recognized in the fauna. The enameloid ultrastructure of teeth of A. dorsetensis was examined and found to be comprised of at least two layers, confirming a neosela- chian affinĘ, based on the overall tooth morphology. The palaeogeographical and stratigraphical records of the previously described species are extended. Plienslachian selachian faunas are not well investigated and the descńbed species in the Hasle fauna are all known from Sinemurian deposits elsewhere. K e y w o r d s : Selachians, hybodonts, neoselachians, palaeobiogeography, palaeoeco- logy, Pliensbachian, Early Jurassic, Bornholm, Denmark. Jan Rees pan.Rees@ geol.lu.se], Department of Geology, Lund University, Sólvegatan 13, SE-223 62 Lund, Sweden. Introduction Early Jurassic selachian faunas are dominatedby two groups, the euselachianhybo- donts and the neoselachianpalaeospinacids. The earliest certain records of hybodont sharks are from the Visóan, Early Carboniferous(Duffin 1985).They may originate, howevel as far back as the mid-Devontan(Zangerl 1981).fiie latestrecords are from the Maastrichtian, Late Cretaceous (Cappetta et al. 1993). The most characteristic externalfeatures of a hybodont shark are the large dorsal fin spines and, in malesoone or two pairs of cephalic spines, the latter featurebeing exclusive to the group. They inhabitedrivers, estuariesand lagoons as well as coastal marine environments.Fluvi- atile environments appear to be more frequently inhabited in the Early Cretaceous, or even the Late Jurassic, onwards.The decreaseof hybodontsin marine environments was probably due to overwhelmingcompetition with the rapidly diversĘing neosela- 440 Jurassic selachians : REES Fig. 1. A. General location of the island of Bornholm. B. The coastal cliff section with the type locality of the Hasle Formation marked with a star. chians. The enameloidof hybodont teethconsists of only one layer (single-crystallite enamel of Reif 1973), whereas most neoselachians have three layers making the teeth more resistant to breakage.The simple enameloid structurein hybodont teeth may explain why hybodontsrarely developeda cutting-typedentition with labio-lingually flattenedteeth. Palaeospinacid sharks are an extinct grrup of neoselachians,possibly ranging from the Early Triassic (Thies 1982). The earliest certain records are from the Ladinian, Middle Triassic, of Canada (Johnset ąl.1997). The latestoccuffences are from the Late Palaeocene (Cappetta et aI. 1993).Externally, palaeospinacid sharks are characterisedby att elongatedbody with a subterminalmouth and two dorsal fin spines (Cappetta 1987). The vertebral centra are calcified and there are many diagnostic cranial features,as well as an amphistylic jaw suspension(see Maisey 1985). Palaeospinacid teeth form a clutching type dentition and the root of the teeth is at a pseudopolyaulacorhize stage(Cappetta 1987).They have been the subject of a heateddebate in recent years (Thies I99I, 1993;Cappetta 1987, 199f; Duffin & Ward 1993). The most important result of this controversy came in 1993 when Duffin & Ward, in a paper on the palaeospinacidsof Lyme Regis, concluded that the type species of Palaeospinax, P. priscus, is a nomen dubium since its holotype consists of a single row of vertebrae,lacking generic or specific characters.Species previously referred to PąIaeospinax were reassignedto either of the two remaining valid genera in the family, Paraorthacodus and Synechodus. Geology and biostratigraphy The island of Bornholm (Fig. 1) is part of the Fenno-Scandian Border Znne and constitutesa fault block mosaic (Gravesen et al. 1982). Sporadic outcrops of Early Jurassic strataare exposedin the south-westernpart of the island and are dominatedby ACTA PALAEONTOLOGTCA POLOMCA (43) (3) 441 deltaic and lacustrine deposits.However, due to an early Pliensbachian transgression (Hallam 1975),the limonitic sandstonesand siltstonesof the Hasle Formation were primarily deposited in a shallow marine environment. The dominating sedimentary structuresof the formation include hummocky cross-stratificationand thin, poorly sorted,gravel laminae' representingerosional surfaces(Gravesen et ąl. |98f).Deposi- tion took place at a depth of 10-40 meters on the open shelf, one or two kilometres from the coast line (Surlyk & Noe-Nygaard 1986).The sedimentarystructures show that sedimentation was active during storms that were either frequent or so vigorous that they erodedall fair-weatherdeposits. The age of the formation has been determinedby means of ammonitesto include the Zones of Uptonia jamesoni to ProdacĘIioceras davoei in the early Pliensbachian (Gravesen et al. 1982). Invertebrate fossils are very scarce in the section yielding selachianremains and it has not been possible to make a more precise age determination. Material and methods All fossils describedherein were collected during thę last decadeat the type locality of the Hasle Formation,situated along the coast,south of the harbourin the town of Hasle (Fig. 1B). The strataare exposed along the shore for approximately700 meters.The sandstonesare often too consolidatedto make bulk processingsuccessful. As a result, most of the selachianmaterial in this study was obtainedthrough surfacepicking. tn places, weatheńnghas softenedthe sandstoneallowing some bulk sampling.In total 18 kg were gently wet sieved and the residue above 500 pm was picked. Prior to photography,the large teeth were coated with ammonium chloride, while small teethwere coatedwith gold and photographedusing a SEM. The ultrastructureof the teethwas examined with a SEM after etching the teethwith 5 N hydrochloric acid (HCl) for 10 to 30 secondsand then coating themwith gold (see Reif 1973:p. 23f). Systematic palaeontology Genęral systematics,taxonomy and descriptive terminology for palaeospinacidsfol- low those of Cappetta (1987). Descriptive terminology for Agalerzs follows that of Duffin & Ward (1983)and terminology for hybodontsis modified from Duffin (1985) and shown in Fig. 2. All illustrated specimens are deposited in the Ępe collection of the Division of Historical Geology and Palaeontology,Lund University, Sweden and prefixed LO (Lund Original). The suffix 'T' denotes holotype and 't' denotes other illustrated specimens. Cohort Euselachii Hayo 1902 Superfamily Hybodontoidea Zangerl, 198 L Remarks. - As a result of the extreme rarity of skeletal remains or even associated teeth from single individuals, the interrelationships andtaxonomy of hybodonts is complicated. Arevision of hybodont 44f Jurassic selachians : REES cutting edge veńical ridges centralcusp lateralcusplets crownshoulder labial node ridge crown- root irregular junction foramina 0\000 e00( D rootbuttress labialpeg occlusalcrest F'tr B Fig.2. A generalizedhybodont shark tooth in labial (A) and occlusal @) views, showing descriptiveterms used in the text. taxonomy and systematics is greatly needed, as a few genera may be paraphyletic or even poly- phyletic. This is, however, beyond the scope of this paper and I will consequentĘ follow the current taxonomy. Family Hybodontidae Owen, 1846 Genus Hybodus Agassiz, 1837 Type species; Hybodus reticulatus Agassiz, 1837 from the Sinemurian, Early Jurassic of Lyme Regis, England. Hyb odus reticulatus Agassi z, 1837 Fig.3G-L. Hybodus reticulatus Agassiz 1837: p.50, pl. 22:22-23. Hybodus reticulatus Agassiz, 1837;Woodward 1889:p. 266-268,p1. 10: 16-18. Hybodus reticulatus Agassiz, L837; Malling & GrÓnwa]l 1909:p.f94,pl.II If. Hybodus reticulatus Agassiz, 1837;Maisey 1987:p. 1-39, figs 1:7,9-I3,18. Hybodus reticulatus Agassiz, 1837; Duffin 1993: p. 49 , ftg Id. Hybodus reticulatus Agassiz, 1837;Delsate & Duffin 1993:p. 110, pl. 1: 1 Material. -fI incomplete crowns, LO7953t,LO 7954t and LO 7955t, all others unnumbered. Description. - The teeth have a high central cusp and two to three pairs of lateral cusplets. All teeth are approximately as high as they are wide. The largest tooth measures 11 mm in height. The central cusp and the lateral cusplets are slender with circular to oval cross section. The cusp and the cusplets of lateral teeth are stightly inclined posteńorly. The cusplets vary from being very small elevations of the occlusal crest to reaching approximately one quarter of the height of the central cusp. Ornamentation consists of weak vertical ridges, occasionally bifurcating, covering the crown except for the apex of the central cusp. The cutting edges are moderately strong. The labial and lingual sides are equally convex. Monognathic heterodonĘ in this species is manifested by differen- ACTA PALAEONTOLOGTCA POLONTCA (43) (3) 443 Fig. 3. Hybodusteeth and hybodontfin spine.A. Hybodontfin spine,LO 7963tx 1.8.B-F. H. delabechei Charlesworth,1839 x 1.6.B-D. LO 7956tin labial (B), lingual (C), and occlusal(D) views (stereopair). E, F. LO 7957tinlabial (E) andlingual (F) views.G-L H. reticulatusAgassiz, 1837 x 3.5.G, H. LO 7953t in labial(G) andlingual (H) views.I, J. LO l954tin labial(I) andlingual (J) views.K, L. LO 7955tin lingual (K) and
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