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BULLETIN DE L' INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BIOLOGIE, 72: 5-45, 2002 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT YOOR NATUURWETENSCHAPPEN, BIOLOGIE, 72: 5-45, 2002
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 B: Batomorphii 4: Order Torpediniformes Family Narcinidae - Subfamily N arcininae Genera: Benthobatis, Diplobatis, Discopyge and Narcine, Subfamily Narkinae Genera: Bengalichthys, Crassinarke, Heteronarce, Narke, Tem,era, and Typhlonarke, Family Torpedinidae Subfamily Torpedininae- Genus: Torpedo- Subgenus: T.(Tetronarke) and T. (Torpedo) and Subfamily Hypninae- Genus: Hypnos.
by J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT & M. STEHMANN
Abstract Kurzfassung
Part B of thi s series, the Batomorphii is continued with th e descrip Mit der Beschreibung und Illustrierung durch RE M-Photos der ti on and illustrati on of the tooth morphology (SEM -photographs) Zahnmorphologie der Ordnung Torpediniformes und additionell er with additional informati on about the tooth vascularisati on of th e Information der Zahnvaskulari sation wird Teil B dieser Seri e (die orderTorpediniformes. Differenti al diagnosis and a classification of Batomorphii) fortgesetzt. Auf Bas is odontologischer Merkmale the torpediniform taxa from the odontological point of view are also werden ferner eine zusammenfassende Differentialdiagnose und presented. It closes the entire seri es on sharks, rays and skates and eine Bewertung der Kl assifikation prasentiert. Nach 16 Jahren wird holocephalans after 16 years. The authors and editor are planning to dieser Seri e tiber Haie, Rochen, und Holocephalen abgeschlossen. enhance the contents of th e series with the publication of additional Di e Authoren und der Editor tiberl egen den Inhalt dieser Serie mit short notices including their personal reflections on the systematics u.a. ihren personlichen "Ansichten der Systematik Rezenter of th e whole group concern ed. Chondrichtyern vom Standpunkt der Odontologie. Key words: Elasmobranchii -Batomorphii - Torpediniformes Schtisselworter: El as mobranchii- Batomorphii - Torpediniformes Odontology. - Odontologie.
Resume Part B: Batomorphii- Order: Torpediniformes Ce fascicul e complete Ia parti e B de cette serie par Ia description et !'illustration (cli ches MEB) de Ia morphologie dentaire des Introduction Torpediniformes. Diagnose differentiel! e, description de leur vascu lari sati on dentaire et conclusions generales concern ant les Torpe Part B of this series, the Batomorphii , is continued with rep dini fo rmes y sont formuh~es .ll cloture cette etude initiee il y a quel resentatives of all supraspecific taxa of the Torpediniformes. ques 16 ans. Les auteurs et leur editeur auront acoe ur de completer The classification of Nelson (1994) is adapted, with eleven cet ensemble par Ia publication de quelques notes additi onnell es genera Benthobatis, Crassinarke, Diplobatis, Discopyge, inclu ant leurs reflexions personnelles concernant Ia systematique Heteronarce, Hypnos, Narcine , Narke, Temera , Torpedo and du groupe concern e. Typhlonarke. Bengalichthys was not mentioned in this classi Mots-des: Elasmobranchii - Batomorphii -Torpedini formes - fication, however, its tooth morphology is here additionall y Odontologie. described, illustrated and compared. Nelson (1994) divided I I
6 J. HERMAN, M. HOVESTADT-EULER , D.C. HOVESTADT & M. STEHMANN the Torpediniformes into two families: the Torpedinidae and Heteronarce garm.ani Narcinidae. He subdivided further the Torpedinidae into the ISH 225-1975 9 219 mm TL subfamiies Torpedininae, with the genus Torp edo, and Hypninae, with the genus Hypnos. Gill (1862) subdivided Heteronarce m.ollis the genus Torpedo into two subgenera: T(T01pedo) and ISH 254-1975 0 196 mm TL T(Tetronarke). The tooth morphology of representatives of both subgenera will be described, illustrated and compared, Hypnos monopte1ygium additionall y. Nelson (1994) also subdivided the Narcinidae into the two subfamilies: Narcininae, with the genera Coli. Hartman 9 265 mm TL Benthobatis, Dip/abatis, Discopyge and Narcine , and Narkinae, with the genera Crassinarke, Heteronarce, Narke, Hypnos subnigrum Temera and Typhlonarke. Coli. Herman 0 140 mm TL The nominal species li sted of each genus or subgenus are af Coli. Herman 9 270mm TL ter MOULD (1999). Cappetta (1988, along with the description and illustration of Narcine brasiliensis a new fossil torpediniform taxon based on tooth morphology, ISH 1856-1968 0 249 mm TL also described and illustrated the tooth morphology of most ISH 1856-1968 9 540 mm TL of the extant genera. The taxa are here redescribed and illus trated by SEM-photographs according to the standard of this Narke dipte1ygia series, with additional infonnation about the tooth vasculari Coli. Herman 9 160 mm TL zation. ISH 8-1 961 9 163 mm TL A differential diagnosis and conclusions on the classification of the torpediniformes are given from the odontological Temera hardwicki point of view. However, being aware of dealing with one complex of characters only, the odontological results will be Coli. Heiman 0 105 mm TL presented here only, and we leave it to following revising authors to incorporate also odontological points of view in a Totpedo (Tetronarke) sp. full systematic review with eventual possible taxonomic and Coli. Heiman o 253 mm TL nomenclatorial changes. The complete bibliographical refer Coli. Herman o 310 mm TL ence for each genus, subgenus and species here described Coli. Hem1an o 315 mm TL will be given in the descriptive section, respectively, and not Coli. Herman o 340 mm TL be repeated under literature references. Coli. Herman o ? mmTL Along with every description of the tooth morphology, the Coli. Hem1an 9 328 mm TL vascularization of the teeth will be described and illustrated. Coli. Herman 9 340mm TL Coli. Hem1an 9 ? mmTL
Material Totpedo (Tetron.arke) nobiliana Coli. Hovestadt 9 620 mm TL The following 61 species of 17 genera and 2 subgenera were Coli. Hovestadt 9 800 mm TL examined for this study: Coli. Hovestadt 9 890 mm TL Coli. Hovestadt 9 1050 mm TL Bengalichthys im.pennis Coli. Herman 9 ? mm TL Coli. Heiman o 160 mm TL Torp edo (Totpedo) torpedo Benthobatis moresbyi Coli. Herman o 350mm TL Coli. Heiman o 215 mm TL Coli. Herman o 350mm TL Coli. Herman o 380 mm TL Crassin.arke dormitor Coli. Herman o 380 mm TL UMTF 20718 o 190 mm TL Coli. Herman o 385 mm TL Coli. Hem1an o 405 mm TL Diplobatis om.mata Coli. Hennan o 410 mm TL Coli. Hennan 9 250mm TL Coli. Hem1an 9 345 mm TL MNHN uncat. 9 150 mm TL Coli. Herman 9 360mm TL
Discopyge tschudii T01pedo (Torpedo) marmorata ISH 883-1979 o 357 mm TL Coli. Hovestadt o 150 mm TL Coli. Herman o 400 mm TL Coli. Hovestadt o 160 mm TL ISH 1632-1966 9 165 mm TL Coli. Hovestadt o 160 mm TL ISH 1632-1966 9 307 mm TL Coli. Hovestadt o 190 mm TL I I
Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 7
Coil. Henn an 3 385 mm TL VASCULARIZATION Coli. Hovestadt <( 120 mm TL Coli. Hovestadt <( 185 mm TL The teeth show the holaul acorhi zid root type with an elon gated, well developed pulp cavity, from whi ch the vascular Coil. Hovestadt 2 250 mm TL Coil. Hovestadt 2 260mm TL tubes of the circumpulpar dentine radiate into crown and Coli. Hovestadt 2 290mm TL root. Osteodentine and inner lateral forami na are absent. (See textfigure 1). Coli. Hennan 2 350 mm TL Coil. Hovestadt 2 360 mm TL Coli. Hovestadt 2 370mm TL Coli. Herman 2 430 mm TL Coil. Hovestadt <( 450mm TL Coil. Hovestadt 2 450mm TL Coli. Hovestadt 2 500 mm TL Coil. Hovestadt 2 500mm TL Textfi gure I. Coli. Hovestadt 2 620mm TL Be111hobatis tooth histological croos section Typhlonarke aysoni Coli. Hetman 3 200mm TL MALES AND FEMALES Additionall y, the illustrations of CAPPETTA (1988) were The crown is divided by sharp mesial and distal cutting edges used. into an inner and outer face. The outer face shows an arched, rounded crown base, largely overhanging the root. Both , me sial and distal cutting edges are of about equal size and Description of the odontological characters arched similarly, each joining seamlessly with the crown base. In about the middle of the crown both cutting edges Family: Narcinidae turn upward to form a nan-ow cusp, being half as high as the Subfamily: Narcinae length of a cutting edge. The cusp becomes only slightly and gradually lower in lateral and posterior teeth. The outer sur The Narcininae comprise the genera Benthobatis, face is sli ghtly concave. Generally, the smooth inner face is Diplobatis, Discopyge and Narcine. concave, but the mid-secti on is convex. Inner and outer orna mentation are absent. The basal surface of the crown shows a rather, broad, slightly Genus: Benthobatis ALCOCK, 1898 rounded crown rim at the outer margins, with inner and outer parts joining in blunt mesial and distal angles. The crown Natural hi story notes from H.M. Indian marine survey ship root junction li es in a relatively deep depression in the center "Investi gator", Commander T.H. Heming, R.N., command of the crown's basal surface. ing. Series II, 25. Annals and Magazine of Natural History The hi gh, natTow holaulacorhi zid root is more or less oval in (ser.7) 2 (8): 136-156. cross-section, slightly oblique toward the rear of the tooth, The genus comprises three named species: B. Kreffti (re and slightly diverges at the root base. The root base presents cently described by Rincon, Stehmann & Vooren, 2001), B. a well-developed, deep and narrow median groove, with a marcida and the type species B. moresbyi. A fourth, yet relatively large aperture in the outer central base of the undecribed species from Taiwanes.e waters was mentioned groove. Root coating is present at about one third of the up by Carvalho ( 1999). per part, all around the root. Inner and outer foramina are absent. Benthobatis moresbyi ALCOCK, 1898 (Plates: 1 and 2 ) Genus: Diplobatis BIGELOW & SCHROEDER , 1948 Benthobatis moresbyi ALCOCK, 1898 Natural history notes from H.M.Indian marine survey ship "Inves ti gator", Com New genera and species of batoid fishes. journal of M arine mander T.H. Heming, R.N., commanding. Series II, 25. An Research 7: 543-566. nals and Magazine of Natural History (ser.7) 2 (8): 136-156. This genus comprises the species D. colombiensis, D. guamachensis, D. pictus and the type species D. ommata. HETERODONTY
The dentition is gradient monognathic heterodont with lat Diplobatis ommata (JORDAN & BOLLMAN, 1890) eral and posterior teeth becoming generally smaller and the (Plates: 3 and 4) cusp slightly more inclining toward the commissure. Lacking female and juvenile specimens sexual and ontoge Discopyge ommata JO RDAN & BOLLMAN, 1890 Description neti c heterodonty cannot not be evaluated. of new species of fishes collected at the Galapagos Islands . I 8 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT & M. STEHMANN
and along the coast of the United States (1887-1888). In: Sci Genus: Discopyge HECKEL, 1846 entific results of explorations by the U.S. Fish commission steamer Albatros. Proceedings of the United States national Ichthyologie. In: Tschudi 1846 Untersuchungen tiber die Museum 12 (770): 149-183 (1889) 1890. Fauna Peruana. Scheitlin & Zollikofer, St. Gallen. 1844-1846 in 12 parts Fauna Peru. HETERODONTY The genus is monotypic with the type,species D. tschudii.
The dentition is gradient monognathic heterodont with lat eral and posterior teeth becoming lower and the cusp tip Discopyge tschudii HECKEL, 1846 slightly inclined toward the commissure. (Plates: 5 to 9) Sexual heterodonty is absent. Lacking juvenile material on togenetic heterodonty cannot not be evaluated. Torpedo tschudii HENKEL, 1846 Ichthyologie. In: Tschudi 1846 Untersuchungen i.iber die Fauna Peruana. Scheitlin & VASCULARIZATION Zollikoje1; St.Gallen. 1844-1846 in 12 parts Fauna Peru. The teeth show the holaulacorhizid root type with an elon gated, well developed pulp cavity, from which the vascular HETERODONTY tubes of the circumpulpar dentine radiate into crown and The dentition is gradient monognathic heterodont with lat root. Osteodentine and inner lateral foramina are absent. (See eral an~ _ posterior teeth becoming lower and the cusp slightly textfigure 2) inclined toward the commissure. Sexual heterodonty is absent. Ontogenetic heterodonty is presented by less developed, lower cutting edges and cusp in juveniles.
VASCULARIZATION
The teeth show the holaulacorhizid root type with an elon gated, well developed pulp cavity, from which the vascular Textfigure 2. tubes of the circumpulpar dentine radiate into crown and Diplobatis tooth histological root. Osteodentine and inner lateral foramina are absent. (See cross section textfigure 3)
MALES AND FEMALES
The crown is divided by sharp mesial and distal cutting edges Textfigure 3. into an inner and outer face. The outer face shows an arched, Discopyge tooth histological convex crown base, more or less overhanging the root. Both, cross section mesial and distal cutting edges are of about equal size and blade-like, each joining the crown base in a blunt angle. In about the middle part of the crown both cutting edges turn MALES AND FEMALES upward, after lowering as a blunt notch, to form a nanow, elongated cusp, about as high as the length of a cutting edge. The crown is divided by sharp mesial and distal cutting edges The cusp becomes only slightly and gradually lower and into an inner and outer face. The outer face shows an arched, more oblique distally in lateral and posterior teeth. The outer convex crown base, more or less overhanging the root. Both, surface is slightly concave. Generally, the smooth inner face mesial and distal cutting edges are of about equal size and is concave, but the mid-section is convex. Inner and outer blade-like, each joining the crown base in a blunt angle. In ornamentation are absent. about the middle part of the crown both cutting edges turn The basal surface of the crown shows a rather, broad, slightly upward, after lowering as a blunt notch, to fonn a nanow, rounded crown rim at the outer margins, with inner and outer elongated cusp being about as high as the length of a cutting parts joining in blunt mesial and distal angles. The crown edge. The cusp being only slightly and gradually lower in root junction lies in a relatively deep depression in the centre lateral and posterior teeth. The outer surface is slightly con of the crown's basal surface. cave. Generally, the smooth inner face is concave, but the The nanow holaulacorhizid root is more or less oval in cross mid-section is convex. Inner and outer ornamentation are ab section, slightly oblique toward the rear of the tooth, and di sent. verges at the root base. The root base presents a well-devel The basal surface of the crown shows a rather, broad, slightly oped, moderately broad, shallow median groove, with a rela rounded crown rim at the outer margins, with inner and outer tively large aperture in the outer central base of the groove. parts joining in blunt mesial and distal angles. The crown Root coating is present at about one third of the upper part, root junction lies in a relatively deep depression in the centre all around the root. Inner and outer foramina are absent. of the crown 's basal surface. II
Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 9
The high, narrow holaulacorhizid root is more or less oval in arched, each joining seamlessly with the crown base. At cross-section, slightly oblique toward the rear of the tooth, about one third of the crown both cutting edges turn upward and diverges at the root base. The root base presents a well to fonn a cusp being half as high as the length of a cutting developed, moderately broad, shallow median groove, with a edge. The cusp becomes lower gradually and more oblique in relatively large aperture in the outer central base of the lateral and posterior teeth. The outer surface is slightly groove. Root coating is present at about one third of the concave. Generally, the smooth inner face is concave, but the upper part, all around the root. Inner and outer foramina are mid-section is convex. Inner and outer ornamentation are absent. absent. The basal surface of the crown shows a rather, broad, slightly rounded crown rim at the outer margins, with inner and outer Genus: Narcine HENLE, 1834 parts joining in blunt mesial and distal angles. The crown root junction lies in a relatively deep depression in the centre Uber "Narcine", eine neue Gattung elektrischer Rochen of the crown's basal surface. nebst einer Synopsis der elektrischer Rochen. Berlin: 1-44. The high, narrow holaulacorhizid root is more or less oval in The genus comprises the species N. bicolor, N. brasiliensis, cross-section, slightly oblique toward the rear of the tooth, N. brevilaniata, N. brunnea, N. entemedor, N. firma, N. and slightly diverges at the root base. The root base presents indica, N. lingula, N. maettlata (type species), N. nigra, N. a well-developed, deep and narrow median groove, with a prodorsalis, N. rierai, N. tasmaniensis, N.timlei, N. relatively large aperture in the outer central base of the vermiculatus and N. westralensis. Lacking tooth material of groove. RoRF coating is present at about one third of the up the type species N. brasiliensis was used instead for descrip per part, all around the root. tion and illustration.
Subfamily: Narkinae Narcine brasiliensis (OLFERS, 1831) (Plates: 10 and 11) The Narkidae comprise the genera Crassinarke, Hetero narce, Narke, Temera and Typhlonarke. Torpedo brasiliensis OLFERS, 1831 Die Gattung Torpedo in ihren naturhistorischen und antiquarischen Beziehungen erli:iutert. Berlin: 1-35. Genus: Crassinarke TAKAG I, 195 1
HETERODONTY Sur Ia nouvelle raie torpille, Crassinarke dormitor, gen. et sp. nov., appartenant a Ia sous-famille Narkinee. Journal of the The dentition is gradient monognathic heterodont with lat Tol~:yo University of Fisheries 38 (1): 27-34. eral and posterior teeth becoming lower and the cusp slightly This genus is monotypic with the type species C. dormitor. inclined toward the commissure. Sexual heterodonty is absent. Ontogenetic heterodonty is presented by less developed, lower cutting edges and cusp in Crassinarke dormitor TAKAGI, 1951 juveniles. (Textplate I; no SEM illustration of isolated teeth because the VASCULARIZATION roots are too badly preserved)
The teeth show the holaulacorhizid root type with an elon Crassinarke dormitor TAKAGI, 1951 Sur Ia nouvelle raie gated, well developed pulp cavity, from which the vascular torpille, Crassinarke dormitor, gen. et sp. nov., appartenant a tubes of the circumpulpar dentine radiate into crown and Ia sous-famille Narkinee. Journal of the Tokyo University of root. Osteodentine and inner lateral foramina are absent. (See Fisheries 38 (1): 27-34. textfi gure 4) HETERODONTY
The dentition is gradient monognathic heterodont with lat eral and posterior teeth becoming lower and the cusp slightly Textfigure 4. inclined toward the commissure. Narcine tooth histological Lacking a female and juvenile specimen sexual nor ontoge cross section neti c heterodonty could not be evaluated.
VASCULARIZATION MALES AND FEMALES The teeth show the holaulacorhizid root type with an elon The crown is divided by sharp mesial and distal cutting edges gated, well developed pulp cavity, from which the vascular into an inner and outer face. The outer face shows an arched, tubes of the circumpul par dentine radiate into crown and rounded crown base, overhanging the root. Both, mesial and root. Osteodentine and inner lateral foramina are absent. (See distal cutting edges are of about equal size and sinilarl y textfigure 5) I I
10 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT & M. STEHMANN
Textplate I . Crassinarke donnitor TAKAGI , 1951. Upper and lower dentitions of a male 19cm l.l., off Japan. Enl argement circa 20. '' Contributions to the study of the comparati ve morphology of teeth and other relevant ichthyodorulites 11
VASCULARIZATION
The teeth the show ho1aulacorhizid root type with an elon gated, well developed pulp cavity, from which the vascular Textfi gure 5. tubes of the circumpulpar dentine radiate into crown and Crassinarke tooth histological root. Osteodentine and inner lateral foramina are absent. (See cross section textfi gure 6)
MALES AND FEMALES
The crown is di vided by sharp mesial and distal cutting edges into an inner and outer face. The outer face shows a crown Textfigure 6. base that forms a blunt angle in the center, and overhangs the Heteronarce tooth histological root. Both, mesial and distal cutting edges are of about equal cross section size and curve upward to form a well-developed cusp; each joins the crown-base in a blunt angle. Lateral and posterior teeth become graduall y lower and asymmetrical. The outer MALES AND FEMALES surface is slightly concave. Generally, the smooth inner face The crow'n is divided by sharp mesial and distal cutting edges is concave, but the mid-secti on is convex. Inner and outer into an inner and outer face. The outer face shows a slightly ornamentation are absent. arched, rounded crown base, overhanging the i·oot. Both, The basal surface of the crown shows a rather, broad, slightly mesial and distal cutting edges are about equal size and simi rounded crown rim at the outer margins, with inner and outer larly arched, and continuously with the crown base. At about parts joining in blunt mesial and distal angles. The crown one third of the crown both cutting edges turn upward to root j unction lies in a relatively deep depression in the centre form a cusp, half as high as the length of a cutting edge, that of the crown's basal surface. lowers gradually in lateral and posterior teeth. The outer sur The narrow holaul acorhi zid root is more or less oval in cross face is slightly concave. Generally, the smooth inner face is section, slightly oblique toward the rear of the tooth, and concave, but the mid-section is convex. Inner and outer orna slightly diverges at the root base. The root base presents a mentation are absent. well-developed, deep and narrow median groove, with a The basal surface of the crown shows a rather, broad, slightly relatively large aperture in the outel' central base of the rounded crown rim at the outer margins, with inner and outer groove. Root coating is present at about one third of the parts joining in blunt mesial and distal angles. The crown upper part, all around the root. Inner and outer foramina are rootjunction lies in a relatively deep depression in the centre absent. of the crown's basal surface. The moderately high, narrow holaulacorhizid root is more or less oval in cross-section, slightly oblique toward the rear of Genus: Heteronarce REGAN, 192 1 the tooth, and diverges at the root base. The root base pres ents a well-developed, deep and narrow median groove, with New fishes from deep water off the coast of Natal. Annals an aperture in the outer central base of the groove. Root coat and Magazine of Natural Histmy, Series 9, 7 ( 4 1): 4 12-420. ing is present at about one third of the upper part, all around The genus comprises the species H. bentuviai, H. garmani the root. (type species), H. mollis and H. prabhui. Part of the dentition of a male of Heteronarce mollis is also illustrated on plate 13. Heteronarce garmani REGAN, 192 1 (Plate: 12; see also plate: 13) Genus: Narke KA UP, 1826
Heteronarce garmani REGAN 1921 New fishes from deep Beitrage zu Amphibiologie und Ichthyologie. Isis 19 ( 1): 87- water off the coast of Natal. Annals and Magazine ofNa tural 90. History, Series 9, 7 (41): 4 12-420. The genus comprises the type species N. capensis and N. dipte1ygia and N. japonica. Lacking tooth material of the HETERODONTY type species N. dipte1ygia was used instead for description and illustration. The dentition is gradient monognathic heterodont with lat eral and posterior teeth becoming lower and the cusp slightly inclining toward the commissure. Narke dipte1ygia (BLOCH & SCHNEIDER, 1801) Only a male specimen was at hand for this study, but female (Plate: 14) teeth were ill ustrated by CAPPETTA (1988), so that absence of sexual heterodonty could not be confirmed. Lacking juvenile T01pedo dipte1 ygia BLOCH & SCHNEIDER 1801 M .E. material ontogenetic heterodonty could not evaluated. BLOCH II Systema Ichtyologiae iconibus ex ill ustratum. Post I I
12 J. HERMAN, M. HOYESTADT-EULER, D.C. HOYESTADT & M. STEH MANN
obitum auctoris opus inchoatum absolvit, correxit, interpola Genus: Temera G RAY, 1836 vit Jo. Gottlieb Scheider, Saxo. Berolini. Sumtibus Austoris Impressum et Bibliopolio Sanderiano Commissum. Systema Descri ption of twelve new genera of fish, discovered by /chtylogiae: 1-584, 110 pis. General Hardwicke in India, the greater part in the British Museum. Zoological Miscellany I: 4-9. HETERODONTY The genus is monotypic with the type species T hardwicki.
The dentition is gradient monognathic heterodont with lat eral and posterior teeth becoming lower and the cusp slightly Temera hardwicki GRAY, 1836 inclining toward the commissure. (Plate : 15) Only a female specimen was at hand for this study, but male teeth were illustrated by CAPPETTA ( 1988), so that the Temera hardwicki GRAY, 1836 Description of twelve new absence of sexual heterodonty could not be confirmed. On genera of fish, discovered by General Hardwicke in India, togenetic heterodonty is presented by less developed, lower the greater part in the British Museum. Zoological Miscel cusps in juveniles. lany I: 4-9.
VASCULARIZATION HETERODONTY
The teeth show the holaul acorhizid root type with an elon The dentition is gradient monognathic heterodont, with gated, well developed pulp cavi ty, from which the vascular lateral·and posteri or teeth lowering toward the commissure. tubes of the circumpulpar dentine radiate into crown and Lacking material of j uveniles and full-grown females, root. Osteodentine and inner lateral foramina are absent. (See ontogenetic or sexual heterodonty could not be determined. textfigure 7) However, the low crown with a tranverse keel instead of a cusp indicates absence of sexual heterodonty.
V ASCULARIZATJON
The teeth show an adapted, holaulacorhizid root type with a relati vely small pulp cavity. The vascular tubes of the cir cumpulpar dentine radiate into crown and root. Osteodentine Textfigure 7. and inner lateral foramina are absent. (See textfigure 8) Narke tooth hi stological cross secti on
MALES AND FEMALES
The crown is divided by sharp mesial and distal cutting edges Textfigure 8. into an inner and outer face. The outer face shows an arched, Temera tooth hi stological cross rounded crown base with an apron-like bulge, overhanging section the root. Both, mesial and distal cutting edges are somewhat irregularly straight, more or less of equal size; each joins the MALES AND FEMALES crown base in a blunt angle, and both merge in an apex, giv ing the outer face a triangular appearance, that slightly low In occlusal view, the Grown is mesio-distally broad and ex ers gradually in lateral and posterior teeth. The outer surface hibits an inward bent, relatively high transverse keel, which is slightly concave. Generally, the smooth inner face is con is often flattened by abrasion, and lowers gradually in lateral cave, but the mid-secti on is convex. Inner and outer orna and posterior teeth. It divides the crown into an inner and mentati on are absent. outer part. The outer and inner margins of the crown are more The basal surface of the crown shows a rather, broad, slightly or less equally arched, and both margins join in sharp mesial rounded crown rim at the outer margins, with inner and outer and distal angles. The smooth inner face is concave with a parts joining in blunt mesial and distal angles. The crown weak bulging of the upper central part. The outer face's up root j unction lies in a relati vely deep depression in the centre per part is concave, changing into an apron-like lower part, of the cromn's basal surface. that is strongly convex and sometimes presenting a central The moderately high, narrow holaulacorhizid root is more or depression. Inner and outer ornamentation are absent. The less oval in cross-section, slightly oblique toward the rear of basal surface of the crown shows a broad, slightly convex the tooth, and diverges at the root base. The root base pres crown rim at the outer margin, gradually nanowing to half its ents a well-developed, deep median groove, with an aperture width at the inner part. The crown-root j unction lies in a shal in the outer central base of the groove. Root coating is low depression in the centre of the crown's basal surface. present at about one third of the upper part, all around the The high, narrow holaulacorhizid root is more or less oval in root. cross-section, slightly oblique toward the rear of the tooth, and diverges at the root base. The root base presents a well- I I
Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 13 developed, median groove, with an aperture in the outer cen Generally, the smooth inner face is concave, but the mid tral base of the groove. Root coating is present at about one section is slightly convex. Inner and outer ornamentation is third of the upper part, all around the root. absent. The basal surface of the crown shows a rather, broad, slightly rounded crown rim at the outer margins, with inner and outer Genus: Typhlonarke WAITE, 1909 parts joining in blunt mesial and distal angles. The crown root junction lies in a relatively deep depression in the center Scientific results of the New Zealand government trawling of the crown's basal surface. expedition 1907. Records of the Canterbury Museum 1: 41- The generally small, narrow holaulacorhizid root is more or 156. less oval in cross-section, slightly oblique toward the rear of The genus comprises the type species T aysoni and T the tooth, and diverges at the root base. The root base pres tarakea. ents a well-developed, narrow median groove, with an aper ture in the outer central base of the groove. Root coating is present at about one third of the upper part, all around the Typhlonarke aysoni (HAMILTON, 1902) root. (Plates: 17 and 18)
Astrape aysoni HAMILTON, 1902 Notice of an electric ray Family: Torpedinidae new to the fauna of New Zealand, belonging to the genus Subfamily: Hypninae Astrape. Transactions and Proceedings of the New Zealand Genus: Hypnos DUMERIL, 1852 Institute 34: 224-226 Monographie de Ia famille des Torpediniens, ou poissons HETERODONTY plagiostomes electriques, comprenant Ia description d 'un genre nouveau, de trois especes nouvelles, et de deux especes The dentition is gradient monognathic heterodont with lat nommees dans le Musee de Paris, mais non encore decrites. eral and posterior teeth becoming lower and the cusp slightly Rev. Magazine de Zoologie (ser.2) 4: 176-189. inclined toward the commissure. The genus comprises the type species: H. monoptel)'gium Only a male specimen was at hand for this study, but a female and H. subnigrum. was illustrated by CAPPETTA (1988), so that absence of sexual heterodonty is confirmed. Lacking juvenile material ontogenetic heterodonty could not be· examined. Hypnos monoptel)'gium SHAW & NODDER , 1794 (Plates: 19 and 20) VASCULARIZATION Lophius monoptel)'gium SHAW & NODDER, 1794 Naturalist's The teeth show the holaulacorhizid root type with an elon Miscellany, or colored figures of natural objects drawn and gated, well developed pulp cavity, from which the vascular described from nature. Naturalist's Miscellany, London: un tubes of the circumpulpar dentine radiate into crown and numbered pages. root. Osteodentine and inner lateral foramina are absent. (See textfigure 9) HETERODONTY
The dentition is gradient monognathic heterodont with lat eral and posterior teethbecoming lower and the cusp slightly inclined toward the commissure. Sexual and ontogenetic heterodonty are absent.
Textfigure 9. VASCULARIZATION Typhlonarke tooth hi stological cross section The teeth show the holaulacorhizid root type with an elon gated, well developed pulp cavity, from which the vascular . tubes of the circumpulpar dentine radiate into crown and MALES AND FEMALES root. Osteodentine and inner lateral foramina are absent. (See textfigure 10) The crown is divided by sharp mesial and distal cutting edges into an inner and outer face. The outer face shows an arched, MALES AND FEMALES rounded crown base with an apron-like bulging, overhanging the root. Both, mesial and distal cutting edges are of about The tri-cuspid crown is structured by sharp mesial and di stal equal size and blade-like, each joining with the crown base in cutting edges at each sid e of the cusps dividing the crown a blunt angle. At about one third of the crown both cutting into an inner and outer face. The outer face shows an almost edges turn upward to form a cusp, half as or equally as hi gh straight to slightly arched crown base, overhanging the root. as the length of a cuttin g edge and lowering gradually in lat The extremely elongated cusps are constricted at their base, eral and posterior teeth. The outer surface is sli ghtly concave. and the central one can be more th an seven times as hi gh as I I
14 J. HER MAN, M. HOVESTADT-EULER, D.C. HOVESTADT & M. STEH MA NN
Descriptiones Diagnostices. Annals of the Lyceum of Natu ral History of New York 7 (32): 367-4 13.
Torpedo (Tetronarke) nobiliana BONA PARTE, 1835 (Pl ates: 2 1 and 22)
Textfig ure I 0. Torpedo nobiliana BONAPARTE 1835 Iconografia della fauna Hypnos tooth hi stological cross itali ca perle quattro cl assi degli animali vertebrati. III Pesci. sec ti on -Roma Iconografi a 3 (12- 14): 1-824.
HETERODONTY its base width, the mesial and di stal cusps can be up to fi ve times as hi gh as their base width. The latter ones slightly The dentition is gradi ent monognathic heterodont with lat curve away from the central cusp. The three cusps are posi eral and posteri or teeth becoming lower and the cusp inclined ti oned next to each other, in occlusal view, the central one to ward the commissure. stands slightly backwards and is slightly more bent in ward. Sexual and ontogeneti c heterodonty are absent. The teeth only diminish in their general size and the three cusps onl y slightly become more oblique distall y in lateral VASCULARI ZATION and posteri or teeth. The outer surface is sli ghtly concave. The teeth show the holaul acorhi zid root type with an elon Generall y, the inner face of the cusps is strongly convex and gated, well developed pulp cavity, from which the vascular from the cusps, the inner face protrudes almost hori zontall y tubes of the circumpulpar dentine radi ate into crown and inward to a central basal depression at the inner margin . Inner root. Osteodentine and inner lateral fo ramina are absent. (See and outer orn amentati on is absent. textfigure 11 ) The basal surface of the crown has a rather, broad, sli ghtly rounded crown rim at the outer margins, and is more or less quadrangul ar with blunt mesial and distal angles. The crown root junction lies in a relati vely nan ow depression in the cen tre of the basal crown's surface. The low, broad holaul acorhi zid root is slightly oblique to ward the rear of the tooth , and di verges at the root base. The root base presents a well-developed, deep and very broad Textfig ure II . medi an groove, with an aperture in the outer central base of Tetronarke too th his tological the groove and sometimes one or several scattered small er cross secti on apertures. Root coating is present at about one third of the upper part, all around the root. MALES AND FEMALES
The crown is di vided by sharp mesial and distal cutting edges Subfamily: Torpedininae into an inner and outer face. The outer face shows an upward Genus: Torpedo HOUTTUYN, 1764 arched, rounded crown base, overhanging the root. Both, mesial and distal cutting edges are of about equal size, each Natuurlyke historie of uitv.oeri ge beschryving der dieren, joining seamlessly with·the crown base. They curve abruptly pl anten en mineraalen, volgens het samenstel van den Heer upward to form a broad-based, elongated cusp and j oint Linnaeus. Met nauwkeurige afbeeldingen. 3 vols. in 37 parts. apex, more than ones as hi gh as the length of each cutting Natuurlyke histori e, Amsterdam 1761-1 785 edge. The cusp becomes lower graduall y in lateral and poste The genus was subdivided by Gill ( 1862) in to the two rior teeth. The outer surface is sli ghtly concave, with a basal subgenera T (Torpedo) and T (Tetronarke). The former com central depression, giving the impression of mesial and di stal pri ses the species T anderson.i, T alexandriensis, T basal lobes. Generall y, the smooth inner face is concave, but bauchotae, T fuscomaculata, T marmorata, T mackayana, the mid-secti on is convex, the mesial and dis tal lower lobe T panthera, T sin.uspersici, T pan.thera, T torpedo (type like parts protruding in wardly. Inner and outer ornamenta species), and another new species from the Gulf of Aden ti on are absent. (CARVA LH O, M. , Stehmann , M .F.W. , MAN ILO, L.G. in press. The basal surface of the crown has a rather, broad, sli ghtly T(Tetronarke) compri ses the species T california, T rounded crown rim at the outer margins, and is more or less fairchildi, T microdiscus, T nobiliana (type species), T quadrangul ar with blunt mesial and distal angles. The crown puelcha, T semipelagica, T tokionis and T tremens. root j unction lies in a relati vely nan ow depression in the cen tre of the basal crown's surface. Subgenus: To 1pedo (Tetronarke) GILL, 1862 The low, broad holaul acorhizid root is sli ghtly oblique to ward the rear of the tooth , and diverges at the root base. The A nalyti cal synopsis of the Order of Squali and revision of the root base presents a well-developed, deep and very broad nomenclature of the genera: Squalorum generum Novorum median groove, with an aperture in the outer central base of \I
Contributions to the study of the comparati ve morphology of teeth and other relevant ichthyodorulites 15
the groove and sometimes one or several scattered smaller The basal surface of the crown has a rather, broad, sli ghtly apertures. Root coating is present at about one third of the rounded crown rim at the outer margins, and is more or less upper part, all around the root. quach:angul ar with blunt mesial and distal angles. The crown root junction lies in a relati vely naiTow depression in the cen tre of the basal crown's surface. Subgenus: Totpedo (Torpedo) HOUTTUYN , 1764 The low, broad holaulacorhi zid root is sli ghtly oblique to ward the rear of the tooth, and diverges "at the root base. The Natuurlyke histori e of uitvoeri ge beschryvin g der dieren, root base presents a well-developed, deep and very broad pl anten en mineraalen, volgens het samenstel van den Heer medi an groove, with an aperture in the outer central base of Linnaeus. M et nauwkeurige afb eeldingen. 3 vols. in 37 parts. the groove and sometimes one or several scattered smaller Natuurlyke hi stori e, Amsterdam 1761-1785 apertures. Root coatin g is present at about one third of the upper part, all around the root. Torp edo (Torpedo) to1pedo L! NNAEUS , 1758 (Plates: 23 to 26) Family: not defined Raia torp edo LI NNAEUS , 1758 Systema naturae per regna tri a Genus: Bengalichthys ANNANDA LE, 1909 naturae, secundum classes, ordinus, genera, species, cum characteribus, di fferentii s, synonymis, Jocis. Tomus I Editio Report on the fishes taken by the Bengal fi sheri es steamer decima, ref01mata. Holmiae: 1-824. Golden Crown Part I , Batoidei. Memoirs of th e Indian Museum '2.' ( I ): 1-60. HETERODONTY The genus is monotypic with the type species B. impennis.
The dentition is gradient monognathic heterodont with lat eral and posteri or teeth becoming lower and the cusp inclined Bengalichthys impennis ANNANDALE, 1909 toward the commissure. (Pl ate: 16 ) Sexual and ontogenetic heterodonty are absent. Bengalichthys impennis ANNANDALE, 1909 Report on the VASCULARI ZATION fishes taken by the Bengal fisheri es steamer Golden Crown The teeth show the holaul acorhizid root type with an elon Part I , Batoid ei. Memoirs ofthe Indian Museum 2 ( 1): 1-60. gated, well developed pulp cavity, from whi ch the vascular tubes of the circumpulpar dentine radi ate into crown and HETERODONTY root. Osteodentine and inner lateral foramina are absent. (See The dentition is gradient monognathic heterodont, with lat textfigure 12) eral and posteri or teeth lowering toward the commissure. Lacking materi al of juveniles and full-grown fe males, on togenetic and sexual heterodonty could not be detennined.
VASCULARIZATION
The teeth show an adapted, holaul acorhizid root type with a relatively small pulp cavity. The vascul ar tubes of the cir Textfigure 12. cumpulpar dentine radiate into crown and root. Osteodentine Torpedo tooth hi stological and inner lateral fo ramina .fire absent. (See textfigure 13) cross secti on
MALES AND FEMALES
The crown is divided by sharp mesial and distal cutting edges into an inner and outer face. The outer face shows a rounded, upward arched crown base, overhangin g the root. Both, me Tex tfigure 13. sial and di stal cutting edges are of about equal size and con Bengalicluhys tooth tinuous with the crown base. They curve from abruptly up hi stological cross section ward to fo nn a broad-based, elongated cusp and join t apex being twice or m ore as hi gh as the length of each cutting MALES AND FEMALES edge. The cusp becomes lower graduall y in lateral and poste rior teeth. The outer surface is sli ghtly concave, with a basal T he crown is div ided by sharp mesial and distal cutting edges central depression, whjch gives the impression of mesial and into an inner and outer face. The outer face shows an arched, distal basal Jobes. Generall y, the smooth inner face is con rounded crown base overhanging the root. Both, mesial and cave, but the mid-section is convex, with the mesial and distal cutting edges are stTa ight, more or less o f equal size, di stal lower lobe-like parts protruding in ward. Inner and and j oining the crown base in an angle. Both cutting edges outer orn amentation are absent. j oin in an apex, givin g the outer face a triangul ar appearance, It
16 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT & M. STEHMANN which lowers gradually in lateral and posterior teeth. The hys are straight, those of Crassinarke slightly curved upward outer surface is slightly concave. Generally, also the smooth and those of Narke irregularly, more or less undulated. inner face is concave. Inner and outer ornamentation are Benthobatis, Diplobatis, Discopyge, Heteronarce, Narcine absent. and Typhlonarke all possess a central cusp, which is more or The basal surface of the crown shows a rather, broad, slightly less resulting from the shape of the mesial and distal cutting rounded crown rim at the outer margins, with inner and outer edges. With the exception of BenJhobatis and Heteronarce , parts joining in mesial and distal angles. The crown-root with upward curving cutting edges, cutting edges of the re junction lies in a relatively deep depression in the centre of maining taxa are more or less blade-like. The cusp of the basal crown's SUiface. Benthobatis is nanow, whereas that of Heteronarce is The moderately high, narrow holaulacorhizid root is more or broader and more massive. The cutting edges of Benthobatis less oval in cross-section, slightly oblique toward the rear of are irregularly shaped, but smooth in Heteronarce. The me the tooth, and diverges at the root base. The root base pres sial and di stal blades of Diplobatis, Discopyge and Narcine ents a well-developed median groove, with a large aperture join the cusp base in a notch. Those of Typhlonarke are con in the outer central base of the groove. Root coating is tinuous with the cusp base. The notches of Diplobatis are present at about one third of the upper part, all around the shallow and the cusp is low, whereas in Discopyge and root. Narcine, the notches are deeper and the cusp is high to mod erately high. Generally, the cusps of Discopyge are lower than those of Narcine. However, the tooth morphology dif Differential diagnosis ferences between Diplobatis, Discopyge and Narcine are mi nor and may vary within these taxa. The most significant tooth morphology characters are sum marized in table 1. Generally for Torpedinifmmes, the most characteristic part is the outer face of the crown, particularly Conclusions the absence or presence of a cusp, its shape and the shape of the mesial and distal cutting edges. Further, the type of root is From the odontological point of view the Torpediniformes significant: a narrow and high or moderately high or low and can be divided into two major groups by their typical root broad root. Hypnos, T(Tetronarke) and T(T01pedo) share a type. Hypnos, T(Tetronarke) and T (Torpedo) have a low low and broad root. The tricuspid crown is typical for broad root. All remaining taxa possess a narrow and high or Hypnos. There are minor odontological differences between moderately high root. T(Tetronarke) and T(Torpedo) only, which may only be interspecific variation. · Group 1 Among the remaining taxa with a high to moderately high, Hypnos and Torpedo are considered as separate evolutionary narrow root, only Temera possesses a tranverse keel. developments. The odontological differences between Bengalichthys, Crassinarke and Narke share a more or less T(Tetronarke) and T (Torpedo) are minimal and do not sup triangularly shaped crown. The cutting edges of Bengalicht- port retaining two separate subgenera.
Table 1. General morphologically significant characters - Species Crown Root
Bengalichthys Cusp triangular Cutting edges straight Moderately hi gh and narrow
Benthobatis Cusp narrow and moderately hi gh Cutting edges curved upward Moderately hi gh and narrow
Dip/abatis Cusp narrow and moderately high Cutting edges blade-like Moderately hi gh and narrow
Discopyge Cusp narrow and high Cutting edges blade-like Moderately hi gh and narrow
Crassinarke Cusp triangularly Cutting edges sli ghtl y curved upward Moderately hi gh and nmTow
He/eronarce Cusp relatively broad Cutting edges curving upward Moderately hi gh and narrow
Na rcine Cusp narrow and hi gh Cutting edges blade-like Moderately hi gh and nan·ow
Narke Cusp triangular Cutting edges straight Moderately hi gh and narrow
Temera Transversal keel high Low and narrow
Typhlonarke Cusp relatively broad Cutting edges blade-like, curving upward Low and narrow
H)pnos Tticuspid Low and broad
To1pedo (Tetronarke) Cups hi gh Cutting edges blade-like, curving upward Low and broad
Torpedo (TIJ/p edo) Cups hi gh Cutting edges blade-like, curving upward Low and broad ''
Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 17
Crown unicuspid Group 1 Torpedo Root low and broad . Hypnos Crown tricuspid ..
.. Cutting edges curved upward Benthobatis Torpedinifonnes Heteronarce
,---- Diplobatis Cutting edges blade-like t-- Discopyge ... t-- Narcine ~ Typhlonarke Root high and narrow
Group 2 Cusp triangular r- Bengalichthys Crassinarke .....____ Narke Transverse keel Temera
Group 2 Acknowledgements Thi s group is subdivided into three separate evolutionary lines: The authors thank Prof. Dr. H. Wilkens, Zoologisches Subgroup l Temera with a transverse keel and more or less Institut und Zoologisches Museum der UniversiUit Hamburg dasyatid appearance. (ZMH), Dr. H. Ishihara, Museum of the Tokyo University of Subgroup 2 Bengalichthys, Crassinarke and Narke with Fisheri es (MTUF), Mr. B. Seret, Antenne XXX , Musee their more or less triangul arly shaped crown. Nati onal d'Histoire Naturtelle, Paris (NMHN) , Dr. D. Nolf, Subgroup 3 Benthobatis, Diplobatis, Discopyge, Hetero Institut Royal des Sciences naturelles de Belgique, Brussels narce, Narcine and Typhlonarke with their particular cusp (IRSNB) and Mr. C. Hartman, Ryan's Falls (Australia) for and mesial and distal cutting edges. permission to examine specimens at their disposal. The Within thi s subgroup Benthobatis and Heteronarce are dif SEM -photographs were taken by J.Cillis, Institut Royal des ferent by the shape of mesial and distal cutting edges, as well Sciences naturelles de Belgique, Brussels. as Dip/abatis, Discopyge, Narcine and Typhlonarke are char acterized by the shape of mesial and distal blades and cusp. Probably due to the small size of the torpediniform micro Bibliography teeth, there are only few fossil records known. However, their tooth morphology shows similarity with the dasyatids, CAPPETTA, H. 1988 Les Torpediniform es (Neoselac hii , which particularly is documented in Temera. Not onl y the Batomorphii) des phosphates du Maroc. Observations sur Ia denture transversal keel, but the whole tooth morphology has similar des ge nres ac tu els. Tertiary research I 0 (I): 21- 52. ity with dasyatids, and the Torpediniformes may well have CARVALHO, M. 1999 A synopsis of the deep-sea ge nu s Benthobatis deri ved from the Dasyatidae. ALCOCK , with a redescri pti on of the type species Benthoba/is The Jack of sexual heteronty appears to be a more advanced moresby ALCOCK, 1898 (C hondri chth yes, Torpedini fo rm es , stage of development. Narcinidae). /n: Sere!, B. & Sire, 1.- Y. ( eds): Proceedings of th e 5th The vascularization of the teeth of all torpediniform taxa is Indo-Pacific Fish Conference, Nourmea, 1997. Paris. Societe principall y the same. The orthodont type, Jacking osteoden fran caise d 'Jchthyo/ogie: 23 1-255. tine in the root, is also an indicati on of an advanced develop CARVALHO, M. , STEHM ANN, M.F.W. , MAN ILO, L.G. (in press) Tor ment. pee/a adenensis, a new species of elec tri c ray from the Gulf of Aden, with co mm ents on nominal species of Torpedo from the Western In dian Ocean , Arabian Sea and adjacent areas (Chond ri chth yes: Torped ini formes: To rpecl inidae. American Museum of Novilates. I'
18 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT & M. STEHMANN .. GILL , T. 1862 Analytical synopsis of the Order of Squali and revi Symphysial sions of the nomenclature of the genera: Squalorum Generum Teeth at junction of both halves of a jaw. Novorum Descriptiones Di agnosticae. Annals of the Lyceum for Natural History of New York, 7 (32): 367-4!3. Transverse Outer/inner direction of a row. MOULD, B. 1997 Classification of the recent elasmobranchii. A Classification of the Living Sharks and Rays of the World. Unpublished Database accessible under
NELSON, J.S. !994. Fishes of the World n~ ed.). Jolm Wiley & An-, Hemi-, Hoi- and Polyaulacorhizid Sons, Inc. New York: p.57-65 Concerning their vascularization, E.Casier(l947) recognized RINCON, G., STEHMANN, M. & VOOREN, C.M. 2001 Results of the and described four phylogenetically significant root types research cruises of FRY "Walther Herwig" to South America. within the orthodont histotypes of elasmobranch teeth . LXXIV. Benthobatis kreffti n.sp. (Chondrichthyes, Torpedini formmes, Narcinidae), a new deep-water electric ray from off South Anaulacorhizid Brasil and the third species of the genus. Archives of the FisheiJ' Vascularization through scattered foramina of equal size on Marine Research. 49 (!): 45-60. both outer and inner faces, (e.g. Hexanchidae).
Hemiau'facorhizid General glossary Vascularization through a median groove and 1 or 2 lateral fo ramina on inner face, (like in Squatinidae and Orectolobidae) (applying to all previous issues of this series) Concerning the jaw Holaulacorhizid Vascularization through many small foramina concentrated in a Anterior median groove running from outer to inner face, (e.g. Rajidae) Tooth position close to junction of left and right jaw halves. Polyaulacorhizid Commissural Vascularization through many small foramina concentrated in Tooth posi tion near the end of jaw. several grooves running parallel from outer to inner face, (e.g. Myliobatidae) Dignathic Heterodont by havi ng different tooth morphology in upper and Apron lower jaws. Expansion of the central part of the outer crown base.
File Basal Tooth row from symphysis toward end of jaw. Bottom face concerned.
Heterodonty Inner central ridge Different tooth morphology within a tooth file. There are two Convex protrusion at the upper midsection of the inner crown types of heterodonty: dignathic and monognathic. face.
Homodonty Costules Uniform tooth morphology within a tooth file Short, vertical ridges sometimes present on inner and/or outer crown base. Lateral Tooth positions half way along the jaw. Crown Longitudinal Enamelated tooth part. Symphysial/commissural direction of a tooth file . Distal Monognathic Tooth edge or part toward angle of jaws. Heterodonty within one jaw only. (this can appear as gradient or disjunct) Hi stotype Type of intern al tooth vascularization. Parasymphysial First anterior tooth row, if a symphysial tooth row is absent. Inner face Viewed from inside the mouth. Posterior Tooth positions toward the angle of jaws. Longitudinally Apico"basall y directed structuring on a tooth. Pseudosymphysial One of th e parasymphysial tooth rows placed in the position of Median groove the symphysial tooth row (symmetry). Groove running from the inner root base to the inner crown-root junction, dividing a holaulacorhizid type of root into two root Row lobes. It includes the main foramina of the vasculari zation sys Tooth row from inner face to outer face of jaw. tem. I I
Contributions to the study of the compara ti ve morphology of teeth a nd other relevant ichthyodorulites 19
Mesial Transverse Tooth edge or part toward junction (symphysis) of left and right Mesio-distall y directed. jaw hal ves. Transverse keel Neo-holaul acorhi zy Transverse ridge dividing the crown into inner and outer face. Modification of the hol aul acorhi zi d type of root, combining a shallow med ian groove and an extremely expanded pulp cavity. Uvul a Lobate extension of the inner crown base. Orthodont Histotype of vascul ari zation, by whi ch a tooth is supplied pri maril y by an internal pulp cavity radiating into numerous tiny STEHMANN, canals penetrating the orthodentine layer. M. Institut fUr Seefischerei, Bundesforschungsanstalt fi.ir Fischerei, Osteodont Palmaille 9, Hi stotype of vasculari zation, by whi ch a tooth is suppli ed with D-22767 H amburg, G etm any. out any pulp cavity by scattered tiny cavities and canals pen etrating the osteodentine layer of the root and th e intern al crown HERMAN, J. material. Service Geologique de Belgique. Rue Jenner 13 , Outer face Viewed fro m outside the mouth. B-1 010 Brussels, Belgium.
Pseudo-apron HOVESTADT-EULER, M . Apron-like vertical ridges that appear sometimes on lateral and & HOVESTADT, D.C. posterior teeth. M erwedelaan 6, NL-4535ET Te rneuzen, Pseudo-osteodont The N etherlands. The former pulp cavity of an originally orthodont hi stotype of tooth being fi ll ed secondaril y with osteodentine.
Pulp cavity Cavity inside the tooth from whi ch th e vascul arizati on is spread via canali culi .
Root Non-enamelated tooth part, that forms th e junction with the jaw gum and provides vascul ari zation of th e tooth.
Root coating Coating on the upper part of the root (probably enameli od)
Root stem Root part between the crown base and root lobe section. Composition of the plates
Second ary anaul acorhi zid As far as possible plates of isolated teeth of one juve nile Medi an groove of a holaul acorhi zid type of root totally over (m ale or female) and of both male and female adults are pres grown to fo rm a closed tube internall y connected or merged e nted for each supraspecific taxon. with the pul p cavity The plates have a consistent composition: upper teeth are Secondarily hemiaul acorhi zid presented with their cusps downward and lower teeth w ith Median groove of hol aul acorhi zid type of root overgrown to the ir cusps upward. various extent, converting the median groove to an in tern al The choice of left or ri ght jaw halves illustrated depends on tube, whi ch is merged with the pulp cavity. the preservatio n quality of the specime n's tooth fil es only.
Stri ae Verti cal ridges running from crown base toward apex. Legend
Sulcus a a nte ri or positi on Groove developed by th e primary vascul ari zation canals lead Ia latera-anterior ing from root base to the main foramina in anaul acorhi zid root I lateral position type. It differs from the medi an groove in whi ch several fo ramina are concentrated of the holaul acorhi zid root type and the lp Latera-posterior parall el grooves of th e polyaulacorhi zid root type, respectively, p posterior positio n in that a sul cus lacks foramin a. c commissural position N 0
~ fli ;>:l :s: )>z :s: :r:: 0 < tTl (/) ~ 0-; m c: r tTI ;>:l 0 0 :r:: 0 < tTl (/)
0 mm 1 ~ 0-; Ro :s: (/)-; tTl :r:: :s: )> z z \ ')
a
)
Plate I . Benthobatis marcida BEAN & W EED, 1909. Male 34.5 em t.l., off Florida. Upper and lower teeth. t t
Contributions to the study of the comparati ve morphology of teeth and other relevant ichthyodorulites 2 1
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0) C ~.:::;. Q a. I I Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 23 .!: 0) ~.... :::"' 0 ...J c-d E "'c: c.."' "- "- 0 -...; E u V) ~"' E L.L.."' 8 0. 00 E E z<( ::E ...l ...l 0 co o<:! z <( Cl ~ 0 0 (3 '€:; ~ 0 0 .::J '€:; ~ C> ~ Q -i 0,) 'til 0:: I I 24 J. HERMAN, M . HOVESTADT-EULER, D.C. HOVESTADT & M . STEHMANN ..c 0 ~... ll) 0. 0. :J >. co"' .. -.; c 2 u0 --' E t.l 0 '<:!" ll) c;; ~ E 0 E '<:!" 00 _j UJ ;,: u UJ ·:c- :::;: """'-s::: \..) .:::: "' 0 ~\..) Q'"' v) ..c ;) ~.... 3: ""0 .....l >. co"' -;::; ec: u0 -...; E u 0 " 26 J. HERMAN, M. HOYESTADT-EULER, D.C. HOYESTADT & M. STEHMANN 0. ..c 0 ~.... Q) 0.. 0.. ;::) >, co~ -;::; ....g u0 -..; E u (<) Q) ~ E Q) E u... E 0 ""00 .s L.U ~ u :r:L.U ""§ -"'u .'::: 0 "'co ~ ..,u Q r--: coQ) 0::: ··- 0. '' Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 27 ~ 0 ~ ;:; ~ 0 ....l ;>.. co"' 03g 0 u -....; E u "'~ "'E E t.I.."' E 0 "' 0 :::: "'"<> <.>~ Ci"' oci <;;"' 0::: .;. I I 28 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTA DT & M. STEHMANN .c 0) ~ '- Cl) 3:: .2 v c "''- Cl) D.. D.. ::::> cci (:5 0:: "' Cl) v 0 c::,...... 0 -5 5 ~ -E (.) E Cl) co c 03 ~ 0 " 0 ""..:::: "'<.; ~ co"' <.;~ cs"' o\ ~ 0::"' II Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 29 // " ( '•, / --' J 03 / ~.... 0 ~ ~ -o c "'.... 0 0.. 0.. ::::> .; I E c ·c"' C/)"' ~ ' • 0 -,_; // § ' r c,; "'L.lJ Ll. ....1 8 - ""~ !:: ~·;::; !3 -<:; -e~ ~ c:i 0 0::"" VJ 0 ~ :r: tT1 a ,~ .... ;;::l ,, s: \ )> ' .z s: "' ...... :r: 0 m< (/) >! tJ_, mI c r m ;;::l tJ 0 0 mm 1 :r: 0 < (/)m >! tJ_, Ro s: (/) ni :r: s: )> z z \ ' a Plate II. Narcine brasiliensis (OLFERS, 183 1). Male 49 em t.l. , off Brasil. Upper and lower teeth. ,, Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 31 ..<:: ;:) ~ ti 3: / ,,47" .3 ~ '"ti 0.. 0.. ::J c;; E '"0 C/J '-- '-'- 0 -..c E u "" C'C ""Q) E C3 E E Q) u. 0\"" £ < c.J UJ ~ ·-§~ ~ ~ "" 0 l::'"' ~ 2 ~"' ::r:: 0i coQ) 0:: I I 32 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT & M. STEHMANN 0 ~....., ..s:? -o"' § i::i 0.. 0.. ::::> '"" E '"0 Cl) t:: 0 - ~ E u l/") 0\ E ~ ro E 2"' ;:::- 0 0\ 0 0 ~ 2 ~ (5 0 ~ "'~ ~ ~ 2 ::t::"' ,....; ~ 0:::"' / ' II Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 33 ..c Q3 ~ ..... (!) :;: .3 -o c ro ..... (!) 0.. 0.. ::J cd -o c ..... ct tt:: 0 ~ E u 00 (!) ""E (!) E u.. E ~ 0 00 c::Z Ll.l 0 Uiz :r: u C/) c: .::;,.~ ".., ~ <:~ ..t (!) c:~ 0 w """' :- a .411 :... \W;Ie.·~~ I lgj ::0 s:: )> _z s:: :r: 0 < tT1 (/) ~ -l0 m l'c tT1 .?" 0 0 :r: 0 < tT1 (/) ~ 0-l Ro s:: (/) -l tT1 :r: s:: )> z z ,.- I ' ~ a 0 mm 1 Plate 15. Temera hardwickii GRAY, 1836. Male 10.5 em t.l., Singapore. Upper and lower teeth. Contributions to the study of the comparative morphology of teeth and other relevant ichthyodoruli tes 35 0 ~..,.... :::: ~ u c ...... ,"' 0. 0. ::::J co "'5 co '0 ::l 0 -....; E u \D .., E ::E E "' ~ 0 0\ u.i ...J < Clz ~ 0 0:::"' I I 36 1. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT & M. STEHMANN c. tl ~... 0 c.. c.. :::::> -oc c;;"' 0 N ~ z0 -oa :§ :S 0 (/)"' "- 0 -...; c u 0 N 0 c;; :2 N' 0 0\ z 0 ~ :E ::s-< 0 ·-Ci "' - ~ s ~" iS -9 ~:::- ~ r..: 0 ~ 0::: c. I I Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 37 ..<:: 0 ~.... 0 ~ 0 .....l -:::) c / cc"' 0 N ~ z0 -oc -;;"' ..<:: :; 0 (/) "- "- 0 E t) 0 E N E co0 :2 01 0 0\ ;i 0 ~ ~ -< 6 0 § "' ..,5' ~ ~ _§ -"' ~ oO 0 0::"" 0. w 00 ~ @ ::0 :s: )> _z :s: :r 0 < tT1 Vl ~ CJ-l I ctT1 r tT1 ?' CJ 0 :r 0 < tT1 Vl ~ .. CJ : ·~ : ..~ ·--.\ I ~. -- I ~ :s: "' :r~ :s: )> z z 0 mm 1 \ Plate 19. Hypnos monopterygius (SH AW & NODDER,1 794). Female 26 em t. l., Port Jackson, Australia. Upper teeth. I I Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 39 Q) ~.... ;;:ll.l 0 ....J cci § VJ :::;) :- ::r: tT1 :s:"" )> _z :s: ~ ,, ... 'i"i. . l::r: a \ I 0 < tT1 til :;! 0 m-l c: r tT1 .?' 0 0 ::r: 0 < tiltT1 :;! 0-l ?? :s: til tri ::r: :s: )> z z p 0 mm 1I L...... I Plate 21. Torpedo (Telronarke) nobiliana BONAPARTE, 1835. Female 89 em t.l. , South England. Upper teeth. '' Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 4 1 .. ..c t5 ~ ..... 0.) 3 0 -l -oc: '. -eo"' ci3 -5 0 (/)"' -~ E :.; cr. ) 00 0.) -;;; E 0.) u.. vi (") 00 E E w ~ -;f. ~.- .?'! a ,..;_:.~.. . :!: :..:.;·__ ,.~ ~.s,;; ~ -l0 mc r tr1 ? 0 '! / /1~~ I~ \ ~ - / I 0 < tr1 ~ (I'J ~ 0-l Rl' $: (I'J :rtTl $: >-z z ·" 7 , 0 mm 1 Plate 23. Torpedo (Torpedo) torpedo (LINNAEUS, 1758). Male 28 em t.l., off Dakar, Senegal. Upper teeth. II Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 43 .c 0) ~ ~ ~ 0 ....l ~ ' Oil -....; E u 00 01 ";::::a ~ 0 :-... @.. ..-- G """:-...~ ~ -.;f 01 3:: :r: 0 ~ ..:1//!•'!'Q-. _r-~, t~'t~ I lrii '~ (/) );! t" / I "''). --. ,~,\ ~ / ::;., ~ . ~~ ' . . \, \ =<:·.~ I lo-l ~ m c: r tr1 ? 0 0 :r: 0 < tr1 (/) );! 0-l Ro 3:: (/) -l tr1 :r: 3:: )>z z -~ '~ 0 mm 1I L.- ..... Plate 25. Torpedo (Torpedo) torpedo (LINNAEUS, 1758). Male 41 em t.l., off Dakar, Senegal. Upper teeth. Central lower part of the plate, radicular view of an uncommon example of the fusion of two dental germinae of commissural teeth...... 1 0__ mm _ I n 0 ;:?. :J. cr a. 0 ::::: til 0 s- ('tl til 2 0. '< .....,0 s- (1> (") 0 3 "0 ...,::>:> g. < (1> 3 0..., "0 :::::- 0 0 ao '< 0...,., CD (1> s- ::>:> :::::0. 0 s- ...,(1> ..., (1> c:n < ::>:> ;:?. c;· :::::- s- '< 0 a # ~ I 1§- • ..., • s CD til Plate 26. Torpedo (To1pedo) torpedo (LINNAEUS, 1758). Male 4 1 em t.l. , off Dakar, Senegal. Lower teeth. Central upper part of the plate, outer view of an uncommon example of the fusion of .p.. two dental germinae of commissural teeth. Vl