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Contributions to the Study of the Comparative Morphology of Teeth and Other Relevant Ichthyodorulites in Living Supra-Specific Taxa of Chondrichthyan Fishes

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Contributions to the Study of the Comparative Morphology of Teeth and Other Relevant Ichthyodorulites in Living Supra-Specific Taxa of Chondrichthyan Fishes I I BULLETIN DE L'lNSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BlOLOGIE, 64 : 165-207, 1994 BULLETIN VAN HET KONINKLJJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN, BlOLOGIE, 64: 165-207, 1994 Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites in living supra-specific taxa of Chondrichthyan fishes Editor : M. STEHMANN Part B : Batomorphii No. la : Order Rajiformes- Suborder Rajoidei- Family: Rajidae Genera and Subgenera: Anacanthobatis (Schroederobatis), Anacanthobatis (Springeria), Breviraja, Dactylobatus, Gurgesiella (Gurgesiella), Gurgesiella (Fenestraja), Malacoraja, Neoraja and Pavoraja by J.HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT and M. STEHMANN Abstract General introduction Part B of this series, comprising the Batomorphii, is initiated with Unlike the principally much more lively interest in taxa of the Rajoidei. The tooth morphology of representatives of nine genera and subgenera is described and illustrated by SEM-photo­ shark teeth, never before received teeth of batoid fishes graphs. An adapted terminology is introduced for the description of (sawfishes, guitarfishes, rays and skates) a similar rajoid teeth, because their tooth morphology differs markedly in degree of attention. In contrast to the majority of sharks, various aspects from th at in selachians. A differential diagnosis for by far most of the batoid species possess very small and conclusions on rajoid odontology, as well as an odontological key teeth closely set in plate-like bands on the jaws. The will be given in a forthcoming issue dealing with the last rajoid taxa. Key-words : Elasmobranchii - Batomorphii - Rajoidei - Odontology - only exceptions are myliobatoid rays with only one row Descriptive terminology. of greatly enlarged teeth (Aetobatus), or the median one of only few tooth rows enlarged in a similar way (Myliobatis, Pteromylaeus, Aetomylaeus, Rhinoptera), Resume to form massive crunching tooth plates. For a long time, there have also been many more fossil records and La deuxieme partie (B) de cette serie concerne Ies Batomorphii. Dans publications on teeth of sharks than on those of batoids. ce premier fascicule l'odontologie de neuf taxons supraspecifiques de Rajoidei est decrite et illustree (cliches MEB). Une terminologie This was mainly due to the difference in size, that adaptee a Ia problematique de leur odontologie est proposee, car celle­ means the tiny batoid teeth were either often over­ ci differe grandement de celle des selaciens. Diagnose differentielle, looked, or not recognised as such, and they were much conclusions systematiques et clef de determination odontologique less obvious and characteristic than shark teeth. With seront presentees dans le fascicule qui cloturera I' etude de ce groupe. Mots-cles : Elasmobranchii - Batomorphii - Rajoidei - Odontologie - regard to living species, sharks are easier available and Terminologie descriptive. also received always more attention of researchers, because many are dangerous to man, and last but not least are jaws and individual teeth of sharks certainly Kurzfassung more spectacular as trophies and collectors items. As a consequence, existing descriptions and illustrations of Teil B dieser Serie, der die Batomorphii umfassen wird, beginnt mit teeth of living batoids are usually rather superficial and der Beschreibung und Illustrierung durch REM-Photos der Zahnmor­ concerning the general type of dentition and eventual phologie einer Gruppe von Rajoidentaxa. Auf Grund erheblicher Unterschiede in der Zahnmorphologie bei Selachiern und Rajoiden sexual dimorphism rather than the detailed complete wird eine entsprechend angepa13te odontologische Terminologie fUr tooth morphology. Based on the principles of descrip­ die Rajoidei vorgestellt. Eine zusammenfassende Di fferentialdiagnose tion and illustration, including root structures and vascu­ und Bewertung zur Odontologie der Rajoidei, sowie ein odontolo­ larization system, introduced for sharks in Part A, Part gische BestimmungsschiUssel werden zum Abschlul3 der Rajoiden in einem spiiteren Beitrag folgen. B of our odontological series will provide the same kind Schiisselworter : Elasmobranchii - Batomorphii - Rajoidei - Odonto­ of consistent and detailed information on the morpho­ logie- Terminologie Zahnmorphologie. logy of batoid teeth. We begin Part B with a first I I 166 J. HERMAN, M. HOVESTADT-EULER, D.C. HOYESTADT and M. STEHMANN contribution on tooth morphology of rajoid skates, the will form the basis for the taxonomic arrangement most specious, systematically most problematical, and chosen. geographically most widely distributed group of Like before in Part A on sharks, will the authors also batoids. For the rajoids, as well as later within few here in Part B on Batomorphii not draw any nomencla­ other diverse batoid groups, will the sequence of torially valid conclusions. Being aware of dealing with publication of supraspecific taxa not reflect any certain one complex of characters only, they will present their classification or taxonomic ordering but merely depend odontological results and leave it to following revising on availability of study material and progress made in authors to incorporate also odontological points of view its investigation. If a larger number of supraspecific taxa in a full systematic review with possible taxonomic and within an order, suborder, or even family of batoids nomenclatorial changes. The original reference of each has to be published, due to technical limitations, in specific taxon will be given in the descriptive section, more than one issue of this series, the last contribution respectively, and not be repeated under literature refe­ will provide for each higher taxonomic category, res­ rences. pectively, the summarizing discussion, conclusion, differential diagnosis, and odontological key. So is the present issue on some supraspecific rajoid taxa only the first of several contributions on rajoid tooth mor­ Part B : Batomorphii phology. Systematics and classification of sharks are not yet fully ORDER:RAJIFORMES understood and clarified in the sense, that existing Suborder : Rajoidei systems uniformly would reflect resolved phylogenetic interrelationships. However, considerably more progress Introduction in revisional work has been achieved for this group of chondrichthyan fishes than for the remaining two, Considerable revisional progress within the Bato­ namely Batomorphii and Holocephali. So has CoM­ morphii has been made during the past about 40 years PAGNO (1984) presented a world catalogue of living especially in rajoid skates. Critical regional faunal shark species and somewhat later ( 1988) added a inventories and revisions have played a key role in this thourough revision of the Carcharhinifmmes. Despite process, and this kind of revisonal approach is less sur­ several still existing unresolved problems, these works prising, if the total number of rajoid species is conside­ have offered a good systematic basis for Part A on red. These are about 230, which is nearly 50 % of all sharks of this series, the odontological results of which batoid species known. In addition, unlike sharks and have provided additional arguments for eventually holocephalans, are rajoid skates distributed almost resolving part of still questionable shark taxa and their everywhere in the world oceans from Arctic to Antarctic interrelationships. A similar general revision of the and from shallow coastal to deep waters of several Holocephali is soon to be expected by DIDIER (in press, thousand meters depth. They become scarce in tropical pers. comm.). Quite different is the situation for syste­ coastal regions only or, like in tropical coral reef areas, matics of the Batomorphii, although their number of are missing. Apparently has this group of batoids been species and supraspecific taxa nearly equals that of sufficiently flexible and managed to settle in nearly all sharks; that again documents the by far greater research kinds of marine habitats, resulting in greatly diverse effort invested in sharks. evolutionary lineages of the rajoids. A world catalogue of batoid species is still far from Regional · revisions were, e.g., published by CLARK being completed, because revisional effort has been (1926) for the European rajids, including the Mediter­ quite heterogeneous for a long time regarding the ranean, BIGELOW & SCHROEDER (1953) for the entire various orders and suborders of batoid fishes and distri­ Western North Atlantic batoid fauna, IsHIYAMA (1958) butional ranges of the species. GARMA N (1913) was the for the Western North Pacific Rajidae, MENNI (1972, last investigator having revised "The Plagiostomia" 1973) for the southwestern, STEHMANN (1970) for the (sharks, skates and rays) in total. All following revision­ northeastern, and HULLEY (1970) for the southeastern a) accounts on Batomorphii, or Batoidea only dealt Atlantic rajids. Early authors already erected few, now with individual family taxa (e.g., NORM AN, 1926, for as before valid rajid genera (Sympterygia MOLLER & the Rhinobatidae), single genera, or mainly defined HENLE, 1841; Psammobatis GONTHER, 1870; Dactylo­ geographical faunas. As a consequence, unlike the batus BEAN & WEED, 1909) in addition to the collective Selachii of Part A, we cannot rely on a relatively recent genus Raja LrNNAEUS , 1758. Paralleled by the extension up to date taxonomic arrangement for the supraspecific of character complexes used for the analysis of rajid batoid taxa for publishing the odontological
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