II BULLETIN DE L' INSTITUT ROYAL DES SClENCES NATURELLES DE BELGIQUE, BIOLOGlE, 73: 5-26, 2003 BULLETIN VAN HET KONINK LIJK BELGISCH INSTITUUT VOOR NATU URWETENSCHAPPEN, BIOLOGlE, 73: 5-26, 2003 Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites in living supraspecific taxa of Chondrichthyan fishes Editor: M. STEHMANN Part A: Selachii. Addendum to 1: Order Hexanchifonnes Family Hexanchidae, 2: Order Carcharhiniformes, 2a: Family Triakidae, 2b: Family Scyliorhinidae, 2c: Family Carcharhinidae, Hemigaleidae, Leptochariidae, Sphyrnidae, Proscy lliidae and Pseudotriakidae 3: Order Squaliformes: Family Echinorhinidae, Oxynotidae and Squalidae. Tooth vascularization and phylogenetic interpretation. by J. HERMAN, M. HOVESTADT-EULER & D.C. HOVESTADT Abstract Kurzfassung In the early issues of thi s seri es, tooth vasculari zation was not in cor­ Die innere Geftiss- und Nervenstruktur (Yaskulari sierung) war in porated in the descriptions. This addendum provides the illustra­ den anfa ngli chen Ausgaben dieser Seri e ni cht Bestandteil der Zahn­ ti ons and descriptions of the tooth vascul ariza ti on of tlie taxa in Part beschreibungen. Diese Erganzungsausgabe bringt die Abbildungen A: No. I Hexanchidae, No.2a Triakidae, No.2b Scyliorhinidae, und Beschreibungen zur inneren Zahn-Yersorgungsstruktur der No.2c Proscylliidae, Hemi galeidae, Pseudotriakidae, Lepto­ Taxa in Teil A: Nr.l Hexanchidae, Nr.2a Triakidae, Nr.2b chariidae and Carcharhinidae and No.3 Squalidae of thi s series. Scyliorhinidae, Nr.2c Proscylliidae, Hemigaleidae, Pseudotriaki­ Moreover, tooth vascularization in combination with morphology dae, Leptochariidae und Carcharhinidae und Nr. 3 Squalidae di eser and fo ssil records is validated for interp retation of phylogenetic Seri e. Ferner wird dokumentiert 1 dass die Yaskul ari sierung der Zah­ interrelationships, and a hypothetical systemati c review is given fo r ne, in Kombination mit Zahnmorphologie und fossiele Funde, von the taxa concerned. Bedeutung ist fur die Interpretation phylogenetischer Yerwandt- . schaftsbeziehungen. Hi erzu wird eine hypotheti sche Obersicht zur Key-words: Elasmobranchii - Selachii - Hexanchiformes - entsprechenden Systematik der betroffenen Taxa prasentiert. Carcharhiniformes - Squaliformes - tooth vascul arization - phylogenetic hypothesis. Schliisselworter: El asmobranchii- Selachii - Hexanchiformes ­ Carcharhiniformes - Squaliformes - Zahnvaskul ari sierung - phylogenetische Hypothesen. · Resume Dans les premiers fascicules de cette serie, Ia vascul ari sati on den­ Addendum to Part A: No.1 Hexanchidae, No.2a taire ne fut pas associee a Ia description de Ia morphologie dentaire. Triakidae, No.2b Scyliorhinidae, No.2c Proscylliidae, Cet addendum fournit description et illustration de Ia vascul arisa­ Hemigaleidae, Pseudotriakidae, Leptochariidae and tion dentaire des taxa concern ant les fasci cul es sui vants de Ia Partie Carcharhinidae, and No.3 Squaliformes. A: N°. I Hexanchi dae, N°.2a Triakidae, W.2b Scyliorhinidae, W2c. Proscylliidae, Hemigaleidae, Pseudotriakidae, Leptochariidae and Carcharhinidae, and N°.3 Squalidae. En outre, Ia vascularisation dentaire en combinaison avec Ia morphologie et Jes donnees General introduction paleontologiques est proposee comme element interpretatif paten­ ti el des li gnees phylogenetiques. Un examen systematique des taxa In the earl y issues on sharks of this seri es of Part A, No. I concernes base sur les presentes observations'hi stologiques permet Hexanchidae, No.2a Triakidae, No.2b Scyli orhinidae, No.2c d' avancer certain es hypotheses. Proscylliidae, Hemigaleid ae, Pseudotri akidae, Leptocha­ Mots-cles : Elasmobranchii - Selachii - Hexanchiformes - rii dae and Carcharhinidae and No.3 Squali dae, tooth vascu­ Squaliformes vascul arisation dentaire hypotheses lari zation was not in corporated in the descriptions. Thi s phylogenetiques . addendum provides with their tooth vascul ari zation with a I I 6 J. HERM AN, M. HOVESTADT-EULER, D.C. HO VESTADT & M. STEHMANN schematic drawing and a description of the tooth vasculariza­ Genus Hexanchus RAFINESQUE, 1810 tion of the species concerned, as far as materi al was available Hexanchus griseus (BONNATERRE, 1788) for this research. Additionally, the Sphyrnidae are included as also carcharhiniform taxa. M aterial was available of the The entire midsection of the tooth shows a band of relatively following taxa: Hexanchidae: Heptranchias, Hexanchus and large osteons, from which vertical, more or less parallel ca­ Notorhynchus, Tri akidae: Furgaleus, Galeo rhinus, Hen·ri­ nals pierce the lower root part and end at the inner face of the triakis, Hypogaleus, !ago, Mustelus, Scylliogaleus, Triakis, root as an opening to the vascul ar system outside. Scyliorhinidae: Apristurus, Asymbolus, Atelomycterus, Above the midsection, the osteons become narrow to form a Cephaloscyllium, Cephalurus, Galeus, Halaelurus, Haplo­ reticulated system of more or less horizontal canals. This sys­ blepharus, Holohalaelurus, Parmaturus, Pentanchus, Para­ tem is connected to separate triangular-shaped pulp cavities derma, Schroederichthys and Scyliorhinus, Carcharhinidae: in each mesial cusp let of the crown. From these cavities, nu­ Carcharhinus, Galeocerdo, Glyphis, Lamiopsis, Loxodon, merous tiny parallel canaliculi of the circumpulpar dentine Nasolamia, Negaprion, Prionace, Rhizoprionodon, Scolio­ radiate into the cusplets. don and Triaenodon, Hemigaleidae : Chaenogaleus, Hemi­ galeus, Hemipristis and Paragaleus, Leptocharidae: Lepto­ charias, Sphyrnidae: Eusphyra and Sphyrna, Proscylliidae: Proscyllium, Eridacnis and Ctenacis, Pseudotriakidae: Pseudotriakis, Family Echinorhinidae: Echinorhinus, Oxy­ Textfigure 2: Hexanchus notidae: Oxynotus and Squalidae: Centrophorus, Centros­ cylliwn, Centroscymnus, Dalatias, Deania, Etmopterus, Isistius, Scymnodon, Somniosus, Squaliolus and Squalus. Tooth vascularization is used here for an interpretation of phylogenetic interrelati onships and a hypothetical systematic review is given for the taxa concerned to be taken in account Genus Notorhynchus AYRES, 1855 in future systematic research. Notorhynchus cepedianus (PERON , 1807) The vascularization canals have been made visible by soak­ ing in ani se oil a thin secti on of the tooth thus becoming The entire midsection of the tooth shows a band of relatively translucent. large osteons, from which vertical, more or less parallel ca­ For the terminology used here see HO VESTADT & nal s pierce the lower root part and end at the inner face of the HOVESTADT-EULER (1993). root as an opening to the vascular system outside. Above the mi qsection, the osteons become narrow to form a Part A: No.1 Hexanchidae reti cul ated system of more or less hori zontal canals. This sys­ tem is connected to separate triangularl-shaped pulp cavities in each mesial cusplet of the crown. From these cavities nu­ This family is subdivided into the genera Heptranchias, merous tiny parallel canaliculi of the circumpulpar dentine Hexanchus and Notorhynchus. radiate into the cusplets. Genus Heptranchias RAFINESQ UE, 1810 Heptranchias perlo (BONNATERRE, 1788) The entire midsection of the tooth shows a band of relatively large osteons, from which vertical, more or less parallel ca­ Tex tfi gure 3: nals pierce the lower root part and end at the inner face of the Noto rhynchus root as an openin g to the vascular system outside. Above the midsecti on, the osteons become narrow to form a reticulated system of more or less horizontal canals. This sys­ tem is connected to remains of separate triangular-shaped pulp cavities in each mesial cusplet of the crown. These cavi­ ties are more or less filled with osteodentine. From these Part A: No.2a Triakidae cavities numerous tiny parallel canaliculi of the circum­ pulpar. dentine radi ate into the cusplets. This family comprises the genera Furgaleus, Galeorhinus, Gogo lia, Hemitriakis, Hypogaleus, !ago, Mustelus, Rhinotriacis, Scylliogaleus and Triakis. Tex tfi gure 1: Heptranchias The fo ll owing species were selected to represent the tooth vascularization of these genera: Furgaleus macki, Galeo rhinus galeus, Hemitriakis japanica, Hypogaleus hyugaensis, !ago omanensis, Mustelus mustelus, Rhinotriacis henlei, Scylliogaleus quecketti and Triakis semifasciata. Material of Gogolia was not avail able for hi s­ tological examinati on. '' Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 7 Genus Furgaleus WHITLEY, 195 I Genus Hypogaleus SMITH , 1957 Furgaleus macki (WHITLEY, 1943) Hypogaleus hyugaensis (MIYOS I, 1939) The root shows a coarse, reticulated system of relatively The root shows a fi ne, reti cul ated system of relatively narrow large osteons. Thi s system is connected to a relatively large , osteons that become courser toward the mesial and distal more or less triangularl-shaped pulp cavity in the crown. margin s. The system is connected to a relatively large, more From this cavity, numerous tiny parallel canaliculi of the or less triangular- shaped pulp cavity in the crO\'{ n. From this circumpulpar dentine radiate into the cusp. Strings of par­ cavity, numerous tin y parallel canaliculi of the circumpulpar ticularl y concentrated canaliculi do not extend from the pulp dentine radiate into the cusp. Strings of particularly concen­ cavity toward the mesial cusplets. trated canali culi do not extend from the pulp cavity toward the mesial cusplets. Textfigure 4: Textfigure 7: Furgaleus Hypogaleus Genus !ago COMPAGN O