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 contribu­ systematics, was the genus Raja step by step split up tions of Part B. It will therefore be commented on and further by more recent authors into a number of genera, explained in the introduction to each order, suborder, or and subgenera of some of these. The family Rajidae was family of the batoids, which classification or revision even subdivided into few more families to separately '' Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 167 include skates with quite extraordinary features : Ana­ investigate the tooth morphology of all accepted, even­ canthobatidae VON BONDE & SWART, 1923; Arhyncho­ tually considered, even questionabie supraspecific taxo­ batidae BIGELOW & SCHROEDER, 1953; Pseudorajidae nomic units, with the aim to offer additional arguments BIGELOW & SCHROEDER, 1954; Gurgesiellidae DE BUEN, for the stabilization of classification and perhaps shed 1959; Crurirajidae HULLEY, 1972. MCEACHRAN (1984) light on open questions from the odontological point of finally summarized relevant revisional results published view. by authors mentioned above and obtained by himself Along with every description of the tooth morphology, plus coauthors in a cladistic analysis. This resulted in the vascularization of the teeth will be described his definition of two groups of the Rajoidei equalling and illustrated. The method of makillg the vasculari­ family rank, of which one (the Rajidae) comprehends zation visible follows STENSIO, which was kindly the majority of rajoid genera and subgenera, whereas brought to our attention by Dr. Talimaa, Institute of the other one remained unnamed. However, on the one Geology, Vilnius (personal communication) : com­ hand could he not consider all supraspecific taxa recog­ pletely submerged in anise oil, teeth become totally nized at that time (e.g. Atlantoraja, Rhinoraja, Ri01·aja transparent and internal structures visible in transmittent and subgenera of Anacanthobatis missing), because his light. classification was largely based on anatomjcal structures of neurocranium, scapulocoracoid and claspers, and All supraspecific rajoid taxa taken into consideration for such information was not equally available for all rele­ an odontological investigation, availability of the requi­ vant taxa. On the other hand was his classification con­ red study material given, in this and following issues of cept not unanimously accepted because, e.g., of the our series (Pmt B) are listed below : heterogeneity of diagnostic characters it was based on. Family ANACANTHOBATIDAE : In addition, further supraspecific rajoid taxa were resur­ genus Anacanthobatis VON BONDE & SWART, 1923 rected and established since by authors. McEACHRAN & with four subgenera (after HULLEY, 1973) MIYAKE (1990a) updated the existing information with a Anacanthobatis VON BONDE & SWART, 1923, revised, though still incomplete working hypothesis on Springeria BIGELOW & SCHROEDER, 1951, Sino­ phylogenetic interrelationships of rajoid skates. The batis HULLEY, 1973, and Schroederobatis HuL­ same authors (MCEACHRAN & MIYAKE, 1990b) pre­ LEY, 1973. sented, as an appendix to a zoogeographical and bathy­ Family ARHYNCHOBATIDAE : metrical analysis for skates, a list of the nominal species genus Arhynchobatis WAITE, 1909. of rajoid skates, with assignment to supraspecific taxa Family CRURIRAJIDAE: according to their updated classification. Few species, genus Cruriraja BIGELOW & SCHROEDER, 1948. however, had here been reallocated to other genera wit­ Family GURGESIELLIDAE : hout a specific explanation for such alterations. Several genus Gurgesiella DE BUEN, 1959. groups of species also remained with unresolved assign­ Family PSEUDORAJIDAE : ment in both recent papers by McEACHRAN & MIYAKE genus Pseudoraja BIGELOW & SCHROEDER, 1954. (1990a+b) due to lacking investigational results : RAJOIDEI (Group I = unnamed family rank, after STEHMANN ( 1986) had grouped the about 45 species of McEACHRAN, 1984, and McEACHRAN & MIYAKE, Bathyraja into three morphotypes related to depth distri­ 1990a+b): Sympterygia MOLLER & HENLE, 1841; bution (shallow, deep water, and transitional morpho­ Psammobatis GONTHER, 1870; Arhynchobatis WAITE, types) but drew no taxonomic conclusions. McEACHRAN 1909; lrolita WHITLEY, 1937; Pavoraja (Pavoraja) & MIYAKE (1990a+b) considered two groups of species, WHITLEY, 1939; Raja (Rioraja) WHITLEY, 1939; Rhino­ namely the North Pacific and Amphi-American Raja raja ISHIYAMA, 1952; Pseudoraja BIGELOW & SCHROE­ Assemblages, unresolved within their classification DER, 1954; Bathyraja lSHIYAMA, 1958 (incl. three mor­ concept and said, that also Bathyraja, lrolita, Pavoraja, photypes after STEHMANN, 1986); Notoraja ISHIYAMA, Pseudoraja, and Rhinoraja require further research. 1958; Raja (Atlantoraja) MENNI, 1972; Pavoraja (A) Finally, YEARSLEY & LAST (1992) described a third sub­ sensu McEACHRAN, 1984; Pavoraja (lnsentiraja) genus within Pavoraja, namely P. (lnsentiraja). YEARSLEY & LAST, 1992. Although the phylogenetic interrelationships within the RAJOIDEI (Group II = family RAJIDAE, after Me Rajoidei are still not fully elucidated, and despite thus EACHRAN, 1984, and McEACHRAN & MIYAKE, still existing uncertainties in the recentmost account by 1990a+b) : Raja (Raja) LINNAEUS, 1758; Raja ( Diptu­ Mc EACHRAN & MIYAKE (1990a) on rajoid classification, rus) RAFINESQUE, 1810; Raja (Amblyraja) MALM, 1877; is the latter concept the most up to date and complete Raja (Leucoraja) MALM, 1877; Dactylobatus BEAN & one available. It will be used therefore for our conside­ W EED, 1909; Anacanthobatis VON BONDE & SWART, ration of supraspecific rajoid taxa below family level to 1923 (incl. four subgenera after HULLEY, 1973, see be studied odontologically, regardless their final ranking above); Breviraja BIGELOW & SCHROEDER, 1948; Cruri­ as subgenera, genera, tribes and/or subfamilies. It has raja BIGELOW & SCHROEDER, 1948; Raja (Okamejei) been the principle from the beginning of this series to ISHIYAMA, 1958; Gurgesie/la (Gurgesiella) DE BUEN, ''

168 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT and M. STEHMANN

1959; Malacoraja STEHMANN , 1970; Raja (Rajella) ISH 641-1986 o 277 mm TL STEHMANN, 1970; Raja (Rostroraja) HULLEY, 1970; ISH 641-1986 o 187 mm TL Gurgesiella (Fenestraja) McEACHRAN & COMPAGNO, Pavoraja (A) asperula 1982; Neoraja McEACHRAN & COMPAGNO, 1982; 'North ISH uncatalogued 2 520 mm TL Pacific Raja Assemblage'; 'Amphi-American Raja ISH uncatalogued 2 314 mm TL Assemblage'. ISH uncatalogued o 427 mm TL NOTE, that part of the supraspecific taxa under both ISH uncatalogued o 339 mm TL groups of RAJOIDEI are duplicated from the the first five families above due to our consideration of the various concepts of classification. Terminology

The first group of genera and subgenera, of which we The odontological characters of batoids differ widely present here our odontological results, is listed below from Selachii and therefore, a modified descriptive ter­ under Material. minology for Rajoidei is proposed as used in the descriptions below.

Material and terminology

For this fascicle the following 34 specimens of 10 spe­ cies were examined : Anacanthobatis (Schroederobatis) americanus ISH 260-1981 2 353 mm TL ISH256-1981 o 314mmTL ISH 260-1981 o 273 mm TL c Anacanthobatis (Schroederobatis) melanosomus ~ IRSNB uncatalogued 2 270 mm TL u

Anacanthobatis (Springeria) longirostris basal ISH 3570-1979 2 660 mm TL crown ornamentation ISH 3583-1979 2 308 mm TL apron ISH 3583-1979 o 465 mm TL &~~~~L_ basal Breviraja spinosa root ornamentation basal ISH 3595-1979 2 305 mm TL root stem median groove root lobe ISH 3520-1979 2 180 mm TL ISH 3600-1979 o 300 mm TL Profile Outer view ISH 3657-1979 o 200 mm TL Dactylobatus armatus Textfigure 1 - Odontological terminology for rajoid teeth. ISH 3582-1979 2 685 mm TL IRSNB uncatalogued 2 160 mm TL ISH 3582-1979 o 270 mm TL Ward (1984) was the first to introduce a specified ter­ Gurgesiella (F enestraja) plutonia minology for describing fossil rajoid teeth. However, ISH 3542-1979 2 267 mm TL further odontological characters for rajoids became ISH 3522-1979 o 232 mm TL known during these investigations, and which are added ISH 3580-1979 2 147 mm TL by modifying Ward's (1984) original drawing (text­ ISH 3522-1979 o 232 mm TL figure 1). ISH 3580-1979 o 164 mm TL The root of the teeth is formed as a stem, often ex­ Gurgesiella (Gurgesiella) dorsalifera hibiting a bilobed base with a basal groove. Basal ISH 1931-1968 2 526 mm TL ornamentation may be present on the edges of the root ISH 1948-1968 o 417 mm TL lobes. In addition, root coating may be present in the ISH 1943-1968 o 383 mm TL upper root part. This coating appears to be enameloid, Malacoraja senta but its nature needs further examination. ISH 251-1985 2571mmTL The crown shape varies in occlusal outer view from ISH 8-1969 2 291 mm TL semi-circular, semi-oval, triangular, to quadrangular or ISH 1029-1982 o 570 mm TL even trapezoid. The basal crown rim is mostly rounded, . ISH 1029-1982 o 366 mm TL with sometimes an inner uvula or/and outer apron pre­ Neoraja caerulea sent. The shape and size of the crown cusp, if present, ISH 175-1983 2 277 mm TL also varies from a small eccentric cone to a elongate or ISH 647-1986 2 239 mm TL lanceolate cusp, with or without an outer and/or distal ,,

Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 169 and mesial cutting edges. Sometimes the cusp is VASCULARIZATION replaced by a transversal keel, dividing the crown in an inner and outer part. A basal outer crown ornamentation The teeth show an adapted kind of holaulacorhizy or may be present varying from some coarse costules to a secondary hemiaulacorhizy, with a large pulp cavity in finer reticulation. the root area, from which the vascular tubes of the circumpulpar dentine radiate into crown and root part. Osteodentine was not observed, and inner lateral fora­ Description of the odontological characters mina are absent.

Genus : Anacanthobatis VON BONDE & SWART, 1923 FEMALES

So far 11 species are described. The genus Anacantho­ The crown base is semi-circular in occlusal occlusal batis was subdivided into four subgenera by HULLEY view. Inner and outer faces are divided by a bluntly ( 1973, namely Anacanthobatis, Schroederobatis, Sino­ rounded transversal keel. Uvula, apron or ornamentation batis, Springeria, based on seven species known at that absent. The inner face is slightly concave, the outer one time and accepted by HULLEY. Only the subgenera slightly convex. Schoederobatis and Springeria will be presented here. The root stem is very low and oval to almost circular in cross-section. Although the root base gradually widens in all directions, it is not bilobed, and the root base outer Subgenus: Schroederobatis HULLEY, 1973 margin is weakly undulated. A basal groove is poorly developed, absent, or sometimes replaced by an inner Monotypic with type species A. americanus BIGELOW & and/or outer foramen. Root coating is absent. SCHROEDER, 1962. MALES Anacanthobatis (Schroederobatis) americanus BIGELOW & SCHROEDER, 1962. The crown base of males is semi-circular to semi­ (Plates : 1, 2 & 3; textfigure 2) quadrangular in occlusal view. The anterior teeth bear a well developed cusp at inner crown edge, which is Anacanthobatis americanus BIGELOW & SCHROEDER, directed inward, oblique distally, but with cutting edges 1962. Bulletin of the Museum of comparative Zoology, curving upward. Harvard College, 128 ( 4) :217. Lateral teeth show a higher cusp, which is stronger oblique distally with cuttinge edges curving stronger HETERODONTY upward. The commissural teeth possess a lower cusp, which is not directed distally but upright. Apron, uvula The dentition is gradient monognathic heterodont. or ornamentation absent. The inner surface is concave, Sexual heterodonty is shown in adults only by the pre­ the outer one flat to slightly convex. sence of a cusp on teeth of males, whereas females have The root stem is very low and oval to almost circular in a transversal keel. cross-section. Although the root base gradually widens Only teeth of males present ontogenetic heterodonty by in all directions, it is not bilobed, and the root base outer bearing a small cone without cutting edges in juveniles, margin .is weakly undulated. A basal groove is poorly which grows to a distinct cusp in maturing specimens. developed, absent or sometimes replaced by inner and/or outer foramina. Root coating is absent.

Textfigure 2 Schroederobatis tooth: histological cross-section. Textfigure 3 - Springeria tooth :histological cross-section. '' 170 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT and M. STEHMANN

Subgenus: Springeria BIGELOW & SCHROEDER, 1951 Genus : Breviraja BIGELOW & SCHROEDER, 1948

Originally described as a genus for S. folirostris , the This genus comprises the following species :B. clarama­ taxon was ranked as subgenus of Anacanthobatis by culata, B. colesi, B. markeli, B. nigriventralis, B. schroe­ HULLEY (1973), who assigned two species: A. foli­ deri and B. spinosa. The latter one is the type species. rostris (BIGELOW & SCHROEDER, 1951) as type species and A. longirostris BIGELOW & SCHROEDER, 1962. Breviraja spinosa BIGELOW & SCHROEDER, 1950 (Plates : 7 to 11 ; textfigure 4) Anacanthobatis (Springeria) longirostris BIGELOW & SCHROEDER, 1962 Breviraja Spinosa BIGELOW & SCHROEDER , 1950, Bulle­ (Plates : 4, 5 & 6; textfigure 3) tin of the Museum of comparative Zoology, Harvard College, 103 (7) : 400. Anacanthobatis longirostris BIGELOW & SCHROEDER, 1961. Bulletin of the Museum of comparative Zoology, HETERODONTY Harvard College, 128 (4): 223, figs 17 and 18. The dentition is gradient monognathic heterodont. HETERODONTY Sexual heterodonty is shown in adults only by the pre­ sence of a rather high cusp on anterior teeth of males, The dentition is gradient monognathic heterodont. which becomes lower and broader on commissural Sexual and ontogenetic heterodonty absent. teeth. Females have a lower cusp, with a small central cone. Ontogenetic heterodonty is documented by much VASCULARIZATION lower cusps on teeth of juveniles.

The teeth show an adapted kind of holaulacorhizy or VASCULARIZATION secondary hemiaulacorhizy, with a large pulp cavity, from which the vascular tubes of the circumpulpar The teeth of this genus have an adapted kind of holaula­ dentine radiate into crown and root. Osteodentine was corhizy, in that the vascular tubes of the circumpulpar not observed, and inner lateral foramina are absent. dentine radiate into crown and root part from a large pulp cavity in the root. Osteodentine was not observed, FEMALES AND MALES and inner lateral foramina are absent.

The crown base is more or less oval in occlusal view. FEMALES Anterior teeth bear a low cusp with mesial and distal cutting edges, which are replaced by a transversal keel The crown base of females is semi-triangular in occlusal on lateral and commissural teeth. The crown rim has a view. Inner and outer faces are divided by blunt mesial sharp, irregularly shaped ridge of costules around the and distal cutting edges. An apron or ornamentation are outer face. The outer surface of the crown is slightly absent, but a poorly developed inner uvula is present. convex, the inner one is concave. The inner face shows a slightly concave surface, the An apron or ornamentation are absent. However, a outer one is slightly convex. small, poorly developed uvula is present. The root stem is very low and is ovally to almost The root stem is low and is ovally shaped in cross­ cit:cularly shaped in cross-section. The root base section. The root base gradually widens in all directions, strongly widens in all directions, is bilobed, and the root ·is bilobed, and has a smooth root base edge. A well base edge is weakly undulated. A well developed basal developed basal groove is present, which is deeper than groove is deep and encloses a rather large central aper­ wide and encloses one or two small central apertures. ture. Root coating is absent. Some teeth even possess two basal grooves. Root coating is absent. MALES

The crown base of males is semi-triangular in occlusal view. The anterior and lateral teeth bear a well deve­ loped cusp, which more or less directed inward and slightly oblique distally [see above]. Both mesial and distal cutting edges curve weakly [which way?]. The cusp is lower on teeth toward the commissure. An apron or ornamentation is absent, but a poorly developed inner uvula is present. The inner surface is concave, the outer Textfigure 4 - Breviraja tooth :histological cross-section. one flat to slightly convex. Contributions tci the study of the comparative morphology of teeth and other relevant ichthyodorulites 171

The root stem is very low and oval to almost circular in inward, and the mesial and distal cutting edges are blunt cross-section. The root base strongly widens in all direc­ and rounded. The outer surface is slightly convex, the tions, is bilobate, and shows a weakly undulated root inner one strongly concave. base outer margin. A well developed basal groove is An apron or ornamentation are absent. However, a well deep and encloses a large central aperture. Root coating developed uvula is present on the inner crown part [? is absent. surface]. The root stem is moderately high and oval to semi­ triangular in cross-section. The root base strongly Genus : Dactylobatus BEAN & WEED, 1909 widens in all directions, presenting a well developed bilobation with smooth or slightly undulating root base The genus is monotypic with the type species Dactylo­ outer margin. A well developed, relatively broad but batus armatus. MCEACHRAN & MIYAKE (1990b : 326) shallow basal groove encloses an extremely large cen­ list D. clarki as a second species but give no explanation tral aperture. The basal groove may be closed on poste­ for the reallocation of Raja clarki. rior teeth, so that only one or two large foramina remain. Root coating is present half way down the root Dactylobatus armatus BEAN & WEED, 1909 stem. (Plates : 12 & 13 ; textfigure 5)

Dactylobatus armatus BEAN & WEED, 1909. Proceed­ Genus : Gurgesiella DE BUEN, 1959 ings of the United States National Museum 36 : 459. This genus was subdivided into two subgenera G. HETERODONTY (Gurgesiella) and G. (Fenestraja) by McEACHRA N & COMPAGNO ( 1982) The dentition is gradient monognathic heterodont. Al­ though an adult male is not known so far, the well deve­ Subgenus: Fenestraja MCEACHRAN & COMPAGNO, 1982 loped cusp on teeth of the adult female, here described, makes it plausi ble, that sexual, as well as ontogenetic This subgenus comprises the species : G. atripinna, G. heterodonty is not to be expected. cubensis, G. ishiyamai, G. mamillidens, G. plutonia (type species), G. sibogae and G. sinusmexican.us. VASCULARIZATION

The teeth of this genus show an adapted kind of holaula­ corhizy or secondary hemiaulacorhizy, in that vascular strings penetrate the crown section from a rather high, narrow pulp cavity. Both, pulp cavity and vascular string have tubes of the circumpulpar dentine radiating into root and crown. Osteodentine was not observed, and inner lateral foramina are absent. Textfigure 6 - Fenestraja tooth : histological cross-section. FEMALES

The crown base is more or less circular in occlusal view. Gurgesiella (Fenestraja) plutonia (GARMAN, 1881) The well developed, elongated crown is directed (Plates : 14 to 17; textfigure 6)

Raja plutonia GARMAN , 1881. Bulletin of the Museum of comparative Zoology, Harvard College, 8 : 236.

HETERODONTY

The dentition is gradient monognathic heterodont. Sexual heterodonty is presented in adults only by a rather elongated cusp on anterior teeth of males, which becomes lower and broader and loses [do you mean that?] mesial and distal cutting edges on teeth toward the commissure. Females also have a well developed but lower cusp on anterior, lateral and commissural Textfi gure 5- Dactylobatus tooth :histological cross-section. teeth, which gradually become lower [where ?] and I I

172 J. HERMAN, M. HOVESTADT-EULER, D.C. HOYESTADT and M. STEHMANN

possess mesial and distal cutting edges. Ontogenetic Subgenus: Gurgesiella DE BUEN , 1959 heterodonty is only documented in juvenile males, teeth of which have a lower cusp with mesial and distal This subgenus comprises the species: G. atlantica, G. cutting edges. dorsalifera (type species) and G.furvescens.

V ASCULAR1ZATION Gurgesiella (Gurgesiella) dorsalifera McEACHRAN & COMPAGNO, 1980 The teeth show an adapted kind of holaulacorhizy in (Plates : 18 to 21; textfigure 7) that semi-parallel tubes of the circumpulpar dentine radiate from the vascular string into crown and root Gurgesiella (Gurgesiella) dorsalifera McEACHRAN & from a large and high pulp cavity in the root. Osteoden­ COMPAGNO, 1980. Archiv fiir Fischereiwissenschaft, 31 tine was not observed, and inner lateral foramina are (1): 1-14 figs. 1-8. absent. HETERODONTY FEMALES The dentition is gradient monognathic heterodont. The crown base is semi-oval in occlusal view. Inner and Sexual heterodonty is present in adults only, in that outer faces are not strictly divided, and mesial and distal males have a rather elongated cusp on anterior teeth, cutting edges are absent. A semi-central cone is present which becomes lower and broader on teeth toward the on the crown. An apron, uvula and ornamentation are commissure. Females also have a well developed but absent. The inner face shows a slightly concave surface, lower cusp on anterior, lateral and commissural teeth, the outer one is slightly convex. which gradually become lower. Ontogenetic hetero­ The root stem is moderately high and more or less oval donty is only documented in juvenile males, teeth of in cross-section. The root widens at the base [see above] which have a low cusp only. but is not bilobated. The root base edge is weakly un­ dulated. A basal groove is overgrown, and only enlarged VASCULARIZATION outer and inner apertures remain. Root coating is only present on the crown-root junction. The teeth show an adapted kind of holaulacorhizy, in that a broad canal runs vertically into the crown area MALES from a large but low pulp cavity in the root. The semi­ parallel tubes of the circumpulpar dentine spread out The crown base is semi-circular to semi-triangular in from this canal into crown and root. Osteodentine was occlusal view. The anterior teeth have a well developed, not observed, and inner lateral foramina are absent. broad based but narrow and elongated cusp, more or less directed inward and slightly oblique distally [see FEMALES above]. Lateral and commissural teeth have a much lower cusp. Both, mesial and distal cutting edges are The crown base is semi-oval in occlusal view. Inner and absent in anterior but present in lateral and commissural outer faces are divided by sharp mesial and distal teeth. The outer crown surface is strongly convex on cutting edges. An apron or ornamentation are absent, anterior but rather plain on lateral and commissural but a well developed inner uvula is present. The inner teeth. An apron, uvula and ornamentation are absent. face shows a slightly concave surface, the outer one is The root stem is moderately high and oval to almost slightly convex. circular shaped in cross-section. The root base strongly The broad root is moderately high and more or less oval widens in all directions but is not bilobed. A basal in cross-section. The root widens at the base but is not groove is overgrown, and only enlarged outer and inner bilobate. The root base edge is weakly waved [see apertures remain. Root coating is only present on the above]. Only remains of a basal groove are one or two crown-root junction. enlarged outer and inner apertures. Root coating is present on upper third of the root stem.

MALES

The crown base is semi-circular in occlusal view. The anterior and lateral teeth bear a well developed cusp, which is more or less directed inward and slightly oblique distally. Both, mesial and distal cutting edges curve weakly. A poorly developed outer cutting edge is Textfi gure 7 - Gurgesiella tooth : histological cross-section. present on the upper part of the outer crown surface. '' Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 173

The cusp is lower on posterior teeth but has well deve­ FEMALES loped cutting edges. The outer crown surface is rather plain, and an outer cutting edge is absent. An apron or The crown base is semi-circular in occlusal view. Inner ornamentation are absent, but a well developed inner and outer faces are not strictly divided. The poorly uvula is present. developed cusp is reduced to a cone, without mesial or The root stem is moderately high and oval to almost distal cutting edges. The inner face is rather concave, circular in cross-section. The root base strongly widens the outer is convex. An apron or ornamentation are [see above] but is not bilobed. A basal groove is over­ absent, but a poorly developed uvula is present. grown and only inner and outer apertures remain. Root The root stem is moderately high and more or less oval coating is present on the upper third of the root stem. in cross-section. The root base gradually widens in all directions and is bilobed with weakly · undulated root base outer margins. A basal groove is poorly developed, Genus: Malacoraja STEHM ANN, 1970 absent, or sometimes overgrown with only inner and/or outer foramina remaining. Root coating is present on the This genus comprises 3 species : M. kreffti, M. senta and upper half of the root stem. M. spinacidermis (type species under junior synonym Raja mollis). MALES

Malacoraja senta (GARMAN, 1885) The crown base is semi-circular in occlusal view. The (Plates : 23 & 24; textfigure 8) anterior teeth bear a well developed cusp, which is directed inward and slightly oblique distally but lacks Raja senta GARMAN, 1885. Proceedings of the United true mesial or distal cutting edges. States National Museum, 8 : 43. The cusp is higher, less oblique distally on lateral teeth, and gradually becomes lower and more oblique on teeth closer to the commissure. The inner surface is concave, the outer one is flat to slightly convex, and both surfaces are not strictly divided. An apron or ornamentation are absent but a poorly developed uvula is present. The root stem is moderately high and more or less oval in cross-section. The root base gradually widens in all directions and is bilobed with weakly undulated root base outer margins. A basal groove is poorly developed, absent or sometimes overgrown with only inner and/or Textfigure 8 - Malacoraja tooth : histological cross-section. outer foramina remaining. Root coating is present on the upper half of the root stem.

HETERODONTY

The dentition is gradient monognathic heterodont. Sexual heterodonty is present in adults only, in that teeth of males have a well developed, elongated cusp, which is a low, more or less central cone on teeth of females only. Only males present ontogenetic heterodonty by a more ovally shaped crown in occlusal view, with a distinct Textfigure 9 - Neoraja tooth : histological cross-section. cusp, which is still a small cone in juveniles, while in females only a more ovally shaped crown in occlusal view is significant for the ontogenetic heterodonty. Genus : Neoraja McEACHRAN & COMPAGNO, 1982

VASCULARIZATION This genus comprises the species N. africana, N. caeru­ lea (type species), N. carolensis and N. stehmanni. The teeth show an adapted kind of holaulacorhizy or secondary hemiaulacorhizy, in that the vascular tubes of Neoraja caerulea (STEHMANN, 1976) the circumpulpar dentine radiate into crown and root (Plates : 25, 26 & 27; textfigure 9) from a large and high pulp cavity in the root. Osteoden­ tine was not observed, and inner lateral foramina are Breviraja caerulea STEHMANN, 1976. Archiv fi.ir Fisch­ absent. wissenschaft, 27 (2): 97-114 I I

174 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT and M. STEHMANN

HETERODONTY developed, absent, or sometimes overgrown with only inner and/or outer foramina remaining. Root coating is The dentition is gradient monognathic heterodont. present on the upper half of the root stem. Sexual heterodonty is presented in adults only, in that males show a well developed, elongated cusp on teeth of the anterior and antero-lateral rows. The lateral and Genus: Pavoraja WHITLEY, 1939 commissural teeth of males and females show a low, more or less central cone and hardly sexual heterodonty. This genus was subdivided by McEACHRAN (1984) into Males present ontogenetic heterodonty in anterior teeth, two subgenera Pavoraja (Pavoraja) and Pavoraja (A), which only have cusps with a small cone in juveniles. the latter still unnamed. YEARSLEY & LAST (1992) intro­ duced a third subgenus P. (lnsentiraja). VASCULARIZATION Subgenus : Pavoraja (A), sensu McEACHRAN (1984) The teeth present an adapted kind of holaulacorhizy, in that from a large, but low pulp cavity in the root section This subgenus comprises the species P. asperula (type several vascular canals enter the crown section. From species) and P. spinifera. these canals the the vascular tubes of the circumpulpar dentine radiate into the crown section. Osteodentine was Pavoraja asperula (GARRICK & PAUL, 1974) not observed, and inner lateral foramina are absent. (Plates : 28, 29 & 30; textfigure 10)

FEMALES Bathyraja asperula GARRICK & PAUL, 1974. Journal of the Royal Society of New Zealand, 4 (3) : 363, figs 8 The crown base of anterior and commissural rows of and 10. females is semi-circular in occlusal view. The crowns in lateral rows are more ovally-shaped in occlusal view. Inner and outer faces are not strictly divided. The poorly developed cusp is reduced to a cone, without mesial or distal cutting edges. The inner face shows a rather concave surface, while the outer one is convex. An apron, ornamentation, or a true uvula are absent (an uvula may develop on anterior teeth). The root stem is moderately high and more or less oval in cross-section. The root base gradually widens in all directions and is bilobed with multilobed root base outer margins. A extremely shallow, basal groove is poorly developed, absent, or sometimes overgrown with only inner and/or Textfigure I 0 - Pavoraja tooth : histological cross-section. outer foramina remaining. Root coating is present on the upper part of the root stem.

MALES HETERODONTY

The crown base of anterior and commissural teeth is The dentition is gradient monognathic heterodont. semi-circular in occlusal view. Tooth crowns in lateral Sexual heterodonty is presented in adults only, in that rows are more oval in occlusal view. The anterior teeth males bear an elongated cusp on anterior teeth, which bear a well developed cusp, which is broad based with a becomes lower on teeth toward the commissure. nan·ow, elongated, inward directed cone lacking true Females only possess a low semi-central cone. Onto­ mesial or distal cutting edges. genetic heterodonty is documented only in juvenile The cone of the cusp strongly reduces in height in males, which have a low cusp. lateral teeth and further gradually in commissural teeth. The inner surface is concave, the outer one is flat to VASCULARIZATION slightly convex. They are not strictly divided. An apron or ornamentation are absent, but a poorly developed The teeth show an adapted kind of holaulacorhizy or uvula is present. secondary hemiaulacorhizy, in that a large pulp cavity in The root stem is moderately high and more or less oval the root section, from which semi-parallel tubes of the in cross-section. The root base gradually widens in all circumpulpar dentine radiate into crown and root part directions and is bilobed with multilobed root base outer [see above]. Osteodentine was not observed, and inner margins. An extremely shallow, basal groove is poorly lateral foramina are absent. Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 175

FEMALES CLARK, R.S. - 1926. Rays and Skates, a revision of the Euro­ pean species. Scientific Investigations of the Fishery Board of The crown base is semi-oval in occlusal view. Inner and Scotland, 1926 (I) (Text) : 1-66, figs 1-42, pis 1-36. outer faces are not strictly divided and, mesial and distal COMPAGNO, L.J. V. - 1984. FAO species catalogue. Vol. 4. cutting edges are absent. An apron, uvula and ornamen­ Sharks of the world. An annotated and illustrated catalogue of tation are absent. A semi-central cone is present on the shark species known to date. Part I. Hexanchiformes to Lamniformes. FAO Fisheries Synopsis No. 125, vol. 4 (I): cusp. The inner face shows a slightly concave surface, iii-viii, 1-249. the outer one is slightly convex. The inner and outer basal crown rim is irregularly undulated. CoMPAGNO, L.J.V. - 1984. FAO species catalogue. Vol. 4 . The root stem is moderately high and more or less oval Sharks of the world. An annotated and illustrated catalogue of shark species known to date. Part 2. Carcharhiniformes. F AO in cross-section. The root base gradually widens in all Fisheries Synopsis No. 125, vol. 4 (2) : iii-x, 251-655. directions and is bilobed. The root base outer margin in weakly undulated. A wide, shallow, basal groove is COMPAGNO, L.J.V. - 1984. Sharks of the Order Carcharhini­ formes. Princeton University Press, Princeton, New Jersey : poorly developed, absent, or sometimes overgrown with vii-xxi, 3-486, figs 1.1-21.10, pis 1-35. only inner and/or outer foramina remaining. Root coating is present on the upper quarter of the root stem. DIDIER, D.A. (in press). Phylogenetic systematics of extant chimaeroid fishes (Holocephali, Chimaeroidei). Bulletin of the American Museum of Natural History. MALES GARMAN, S.W. - 1913. The Plagiostomia (sharks, skates and The crown base is semi-circular in occlusal view. The rays). Memoirs of the Museum of Comparative Zoology, Har­ vard College, 36 : xiii + 515 pp., 75 pis. anterior and lateral teeth bear a well developed, broad based, narrow and elongated cusp, which is more or less HULLEY, P.A. - 1970. An investigation of the Rajidae of the directed inward and slightly oblique distally. Mesial and west and south coasts of Southern Africa. Annals of the South African Museum, 55 (4): 151-220,21 figs, 13 pis. distal cutting edges are absent. Inner and outer surfaces are convex, and the inner and outer basal crown rims are HULLEY, P.A.- 1973. Interrelationships within the Anacantho­ irregularly undulated. The cusp becomes lower on teeth batidae (Chondrichthyes, Rajoidea), with a description of the lectotype of Anacanthobatis marmoratus VON BoNDE & toward the commissure, the outer crown surface plainer. SwART, 1923. Annals of the South African Museum, 62 (4) : An apron, uvula and ornamentation are absent. 131 - 158. The root stem is high and more or less circular in cross­ section. The root base gradually widens in all directions ISHIYAMA, R. - 1958. Studies on the Rajid Fishes (Rajidae) found in the Waters around Japan. Journal of the Shimonoseki and is bilobed. The root base outer margin in weakly College of Fisheries, 7 (2, 3) : 193-394. undulated. A well developed basal groove encloses a large foramen. Root coating is present on the upper part McEACHRAN, J.D. - 1984. Anatomical Investigations of the New Zealand Skates Bathyraja asperula and B. spinifera, of the root stem. With an Evaluation of Their Classification Within the Rajoid­ ei (Chondrichthyes). Copeia 1984 (1) : 45-58. McEACHRAN, J.D. & COMPAGNO, L.J.V. - 1982. Interrelations­ Acknowledgements hips of and within Breviraja based on anatomical structures (Pisces : Rajoidei). Bulletin of Marine Science, 32 (2) : 399- The authors thank the editor for having provided most 425. of the material for this study. McEACHRAN, J.D. & MIYAKE, T. - 1990a. Phylogenetic Inter­ We also whould like to thank Dr. D. NoLF, Institut relationships of Skates : A Working Hypothesis (Chondrich­ Royal des Sciences naturelles de Belgique, Brussels for thyes Rajoidei). In Pratt Jr., H.L., Gruber, S.H. & Taniuchi, T. permission to examine specimens at his disposal, and (Eds.) Elasmobranchs as Living Resources : Advances in the Miss. F. LADEUZE, F.N.R.S., Brussels, as well as Miss. I. Biology, Ecology, Systematics, and the Status of the Fishe­ SILVA, Jacqmain, Brussels for technical assistance. The ries : Proceedings of the Second United States-Japan Work­ shop East-West Center. NOAA Technical Report NMFS 90 : SEM-photographs were taken by J. CILLIS, Institut 285-304. Royal des Sciences naturelles de Belgique, Brussels, and the photographs printed by G. BROGNET and J. Mc EACHRAN, J.D. & MIYAK E, T.- 1990b. Zoogeography and DEGREEF, Brussels. Bathymetry of Skates (Chondrichthyes, Rajoidei). In Pratt Jr., H. L., Gruber, S.H. & Taniuchi, T. (Eds.) Elasmobranchs as Living Resources : Advances in the Biology, Ecology, Syste­ matics, and the Status of the Fisheries : Proceedings of the Bibliography Second United States-Japan Workshop East-West Center. NOAA Technical Report NMFS 90: 305-326. BIGELOW, H.B. & SCHROEDER, W.C. - 1953. Sawfishes, Gui­ NORMAN , J.R. - 1926. A Synopsis of the Rays of the Family tarfishes, Skates and Rays (pp. 1-514). In : TEE-VAN, J. et al. Rhinobatidae, with a Revision of the Genus Rhinobatus. (Eds). Fishes of the Western North Atlantic. Memoir Sears Proceedings of the Zoological Society of London, 1926 (4) : Foundation for Marine Research, l (2) : I-XV + 588 pp. 94 1-982. 'I

176 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT and M. STEHMANN

MENNI, R.C. - 1972. Raja (Atlantoraja) subgen. nov. y lista Posterior critica de los Rajidae Argentinas (Chondrichthyes, Rajifor­ Tooth posiiions toward the angle of jaw. mes). Revista del Museo de La Plata (N.S.) (Zoologia), 11 : Pseudosymphysial 165-173. One of the parasymphysial tooth rows placed in the MENNI, R.C. - 1973. Rajidae del Litoral Bonarense. I. position of the symphysial tooth row (symmetry). Especies de los generos Raja, Bathyraja y Sympterygia Row (Chondrichthyes). Physis (A) , 32 (85) : 413-439. Tooth row from inner face to outer face of jaw. STEHMANN , M. - 1970. Vergleichend morphologische und Symphysial anatomische Untersuchungen zur Neuordnung der Systematik Teeth at junction of both halves of a jaw. der nordostatlantischen Rajidae (Chondrichthyes, Batoidei). Transversal Archiv fiir Fischereiwissenschaft, 21 (2) : 73-164. Outer/inner direction of a row. STEHMANN, M. - 1986. Notes on the systematics of the Rajid genus Bathyraja and its distribution in the World Ocean. In Uyeno, T. et a/. (eds.) : Indo-Pacific Fish Biology : Pro­ CONCERNING THE TOOTH ceedings of the Second International Conference on Indo­ Pacific Fishes. Ichthyological Society of Japan, Tokyo: 261- 168. An-, Hemi-, Hoi- and Polyaulacorhizid WARD, D.J. - 1984. Additions to the fish fauna of the English Concerning their vascularization, E. Casier (1947) Palaeogene. 5. A new species of Raja from the London Clay. recognised and described 4 phylogenetically signifi­ Tertiary Research, Leiden 6 (2) : 65-68. cant root types within the orthodont histotypes of YEARSLEY, G. & LAST, P.- 1992. Pavoraja (lnsentiraja) laxi­ elasmobranch teeth. pella, a new subgenus and species of skate (Chondrichtyes : Anaul acorhizid Rajoidei) from the western Pacific. Copeia 3 : 839-850. Vascularization through scattered foramina of equal size on both outer and inner faces, (e.g. Hexanchi­ dae). Hemiaulacorhizid General glossary Vascularization through a median groove and 1 or 2 lateral foramina on inner face, (like in Squatinidae (applying to all previous issues of this series). and Orectolobidae) Holaulacorhizid CONCERNING THE JAW Vascularization through many small foramina con­ centrated in a median groove running from outer to Anterior inner face, (e.g. Rajidae) Tooth position close to junction of left and right jaw Polyaulacorhizid parts halves. Vascularization through many small foramina con­ Commissural centrated in several grooves running parallel from Tooth position near the end of jaw. outer to inner face, (e.g. Myliobatidae) Dignathic Apron Heterodont by having different tooth morphology in Expansion of the central part of the outer crown upper and lower jaws. · base. File Basal Tooth row from symphysis toward end of jaw. Bottom face concerned. Heterodonty Costules Different tooth morphology within a tooth file. Short, vertical ridges sometimes present on inner There are two types of heterodonty : dignathic and and/or outer crown base. monognathic. Crown Homodonty Enamelated tooth part. Uniform tooth morphology within a tooth file Distal Lateral Tooth edge or part toward angle of jaws. Tooth positions half way along the jaw. His to type Longitudinal Type of internal tooth vascularization. Symphysial/commissural direction of a tooth file. Inner face Mono gnathic Viewed from inside the mouth. Heterodonty within one jaw only. (this can appear as Longitudinally gradient or disjunct) Apico-basally directed structuring on a tooth. Parasymphysial Median groove First anterior tooth row, if a symphysial tooth row is Groove running from the inner root base to the inner absent. crown-root junction, dividing a holaulacorhizid type ''

Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 177

of root into two root lobes. It includes the main Sulcus foramina of the vascularization system. Groove developed by the primary vascularization Median keel canals leading from root base to the main foramina Transverse ridge dividing the crown into inner and in anaulacorhizid root type. It differs from the outer face. median groove in which several foramina are con­ Mesial centrated of the holaulacorhizid root type and the Tooth edge or part toward junction (symphysis) of parallel grooves of the poly-aulacorhizid root type, left and right jaw halves. respectively, in that a sulcus lacks foramina. Neo-holaulacorhizy Transversal Modification of the holaulacorhizid type of root, Mesio-distally directed. combining a shallow median groove and an extre­ Uvula mely expanded pulp cavity. Lobate extension of the inner crown base. Orthodont Histotype of vascularization, by which a tooth is STEHMANN, M. Institut fUr Seefischerei, supplied primarily by an internal pulp cavity radiat­ Bundesforschungsanstalt fUr Fischerei, ing into numerous tiny canals penetrating the ortho­ Palmaille 9, dentine layer. D-22767 Hamburg, Gennany. Osteodont Histotype of vascularization, by which a tooth is HERM AN, J. supplied without any pulp cavity by scattered tiny Service Geologique de Belgique. Rue Jenner 13 , cavities and canals penetrating the osteodentine B-1 040 Brussels, Belgium. layer of the root and the internal crown material. Outer face HOVESTADT-EULER, M. and HOVESTADT, D.C. Viewed from outside the mouth. Merwedelaan 6, Pseudo-apron NL-4535 ET Terneuzen, The Netherlands. Apron-like vertical ridges that appear sometimes on lateral and posterior teeth. Pseudo-osteodont The former pulp cavity of an originally orthodont Composition of the plates histotype of tooth being filled secondarily with osteodentine. As far as possible plates of isolated teeth of one juvenile (male or female) and of both male and female adults are pre­ Pulp cavity sented for each supraspecific taxa. Cavity inside the tooth from which the vasculariza­ The plates have a consistent composition : upper teeth are pre­ tion is spread via canaliculi. sented with their cusps downward and lower teeth with their Root cusps upward. Non-enamelated tooth part, that forms the junction The choice of left or right jaw halves illustrated depends on with the jaw and provides vascularization of the the preservation quality of the specimen's tooth files only. tooth. Legend Root coating a = anterior position p = posterior position Coating on the upper part of the root (probably ena­ I = lateral position c = commissural positio maliod) Root stem Root pa1t between the crown base and root lobe sec­ tion. Secondarily anaulacorhizid Median groove of a holaulacorhizid type of anaula­ corhizid root totally overgrown to form a closed tube internally connected or merged with the pulp cavity Secondarily Median groove of holaul acorhi zid type of hemiaul a­ corhizid root overgrown to various extent, but Striae Vertical ridges running from crown base toward apex. Terminally groove or pores still open. ._.) -00

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Plate 11 - Breviraja spinosa BIGELOW & SCHROEDER, 1950. Ju venile o i SH 3657-1979, 200 mm tl. , N.W. Atlantic. n 0 q::J & ~ c;· ::J (/) 0 s- (1) (/) 2 0.. '-<

.....,0 s- (1) (l 0 3 "0 § ..... < (1) 3 0 .:a ::r 0 0 0 mm 0.5 (JQ '-< p 0....., (ti (1) .....::r § 0.. 0 s- (1) ...., @ (i' < § ..... ('i ' ::r s- '-< 0 0.. 0 2

~· (/)

...... Plate 12 - Dactylobatus armatus BEAN & W EED , 1909. Adult ~ISH3582- 1979, 685 mm tl., Florida . 00 \0 ...... \0 0

:--- @ Al 3:::

~

3:::

::r: 0

(T)< (/) >! 0 7 (T)

?(T) Al 0 0 ::r: 0

(T)< (/) >! 0 0 m m 0,5 ~ p "'0." 3::: (/) -l (T) ::r:

~

Plate 13 - Dactylobatus armatus BEAN & WEED , 1909. Adult <(ISH 3582-1979, 685 mm tl. , Florida. () 0 ::l ::3. 0" .....c: o· ::l "' 0 s- ..._____ (1)

..... "'c: p_, '< p .....,0 ::r..... (1) () 0 3 '0

~ ::=.

(1)< 3 0 mm 0.5 0 .a ::r 0 0 (JQ '< .....,0

~ (1) s- § p_, 0 ::r..... (1) ..., p @ 0 < § ..... (i" ::r s- '< 0 p_, ...,0 c:

~ "'

_. Plate 14 - Gurgesiella (Fenestraja) plutonja GARMAN, 1881. Adult r.?iSH 3542-1979,267 mm tl., Florida. \0 ll 192 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT and M. STEHMANN

~ ~ ~ k.. ~ E: E: ('... \() o\"' ('... 0\ "';' "' ""'lr) ""::r:: ~ C>+ ~ '"'=:! ""<: ....; 00 00 ...... ~- "'( ~ <:>:: "'( c.J ·a0'<1 0 .3 Ill 0. c:i '2 "' ·~ b (/) Q.) c: sQ.) E ~ E v '{/j Q.) bO,_ ::::1 0

0

~ 0'<1 0::: '' Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 193

-~ 0:: ~ ""<: :s: <: ~ s:: s:: "' """' ~ -.0\ C".l' "'lr) "" ~-. "o -::;- "'\:::s ""<: -...: co co-.

\0

194 J. HERMAN, M. HOVESTADT- EULER, D.C. HOVESTADT and M . STEHMANN

~ .§ lt.. -::::: E: !i: ('... ';to E ....., E o;- ('...... ,0\ c:, 00 lr) "" ~ 0+ ~ · ~ \U ;:.. \U ;:...... ,::; .....; 00 .....,00 ;;£ ~ ~ c.J ·roa 0 '5 0.. ---. ro ·c;r b C/) ~ 1':: u..~ '-' ro ·u;~ ~ bl)..... ::s 0

r-

~ ~ 0:: n 0 ::l s. 0"' a~· ::l C/l 0 s- (1)

C/l a 2 c I 1'<0...... ,0 ::r~ (1) () a0 '"0 e; Ill c .

<(1) a 0 mm 0.5 0 -a ::r 0 0 '

c I l (il ~ < Ill ;:?. ()" ::r ..... ::r '< 0 0.. ...,0 §.: ~ (1) C/l

...... Plate 18 - Gurgesiella (Gurgesiella) dorsalifera M cEAC HRAN & COMPACNO, 1980. Adult ':( ISH 1931 -1968, 526 mm tl., Brasil. \0 Vl \0...... 0\

:- :r: rn ;;o z~

~ :r: 0 < ~

~ 0 -;-! rn c: r rn ?' 0 0 :r: 0 0 mm 0.5 < rn Vl >! 0-l "'::l a. ~ Vl-l rn :r: ~z z

Plate 19 - Gurgesiella (Gurgesiella) dorsalifera McEACHRAN & CoMPAGNO, 1980. Adult 9 ISH 1931-1968, 526 mm tl., Brasil. (J 0 =q cr= c ...... c;· en= ...... 0 & (1) C/) 2 0. '-<

0...... , & (1) () 0 3 "0 e; I=» ...... :;;::- (1) 3 0 0 .a mm 0.5 ::r 0 0 (/q '-<

0...... ,

~ (1) & I=» =0. 0 & (1) ..., @ (5'" < § ...... c=;· ::r & '-< 0.0 0..., c ...... ~ (1) C/)

Plate 20 - Gurgesiella (Gurgesiella) dorsalifera M cEAC!-IRAN & COMPAGNO, 1980. Adult o ISH 1948-1968,4 17 mm tl., Brasil. \0 -.J \0 -00

~ ::r: tTl ;:o 3:: ;I>z 3::

::r: 0 < ~ >!

~ tTl ? tTl ;:o n0 ::r: 0 < tTl 0 mm 0.5 en >! ...,0 :» "'0. 3::

en..., tTl ::r: 3:: ;I>z z

Plate 2 1 - Gurgesiella (Gurgesiella) dorsalifera M c E ACHRAN & CoMPAGNO, 1980. Adult o ISH 1948-1968, 417 mm tl., Brasil. I I Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 199

Ci "'1::1 1:: -c t;j ....:] -..... !:': 0 I:: ...... f',. lr)

ti0\ '""';< ...... lr) C"-1 ::r::: ...... {/) 0+ ..... :i "'1::1-«:

iG00 ......

Q:;~ '«: c.:> '--- «< c::v Vl «< ";;' ..... 0 (.) «< c. «l ~

C"'l C"'l 2:l «< 0:: I I

200 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT and M. STEHMANN Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 201

-~ ~ E ~ E "<:( :s: <: ~ !::: " !:: ""'-; 0\ "l " o._' \Q 0\ ""'-; oO ~ ();.- ~ -~ IU ;o. ::s ...... _ lr)'""' 00 00 -<;' ~ 0::; C.) '--" c:C<:l Cl)

"'C<:l - ~ ..... 0 u C<:l -C<:l ~

"'"N ~ C<:l 0:: N 0 N

:- :r: tT1 :;<:) s: ;J> .z s: :r: 0 < tT1 Vl :;! 0 mm 0.5 ;l0 ctT1 r tT1 :;<:) 0 n :r: 0 < ~ :;! 0 -1 "-' "0. s: Vl -1 tT1 :r: s: ;J>

~

a

Plate 25 - Neoraja caerulea (STEHMANN,l976). Adult I( ISH 175-1983, 277 mm tl., E. Rockall. II

Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 203

0 1\

204 J. HERMAN, M. HOVESTADT-EULER, D.C. HOVESTADT and M. STEHMANN Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 205

-.::i ::::: ~ <::i N"' s <:"'

"'0 ~ !:: !:: C) "-1 lr) E ~ E .§ I..J "'~ "1:j "' 0 ~ ~s <:::i I..J .. ::st::: ~ ();. ~ ~ " Cd 0.. I 00 N ~ ro 0: N 0 0\

:--

Ri ;:o $: )> ?-

~ :I: 0 < ['T1 (/) :;! p I 10 a "j'l ['T1 p c r ['T1 JO 0 0 :I: 0 < ['T1 (/) :;! 0-l

""=Q. $: (/) -l ['T1 :I: $: )> 0 z mm 0.5 z

p

Plate 29 - Pavoraja asperula (GARRICK & PAUL, 1974). Adult o ISH uncatalogued specimen 52711111111., New Zealand. I Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 207

"' "' "\:! !;;: (::! ~ ~ N S: <:~ ";::i In 0 t:: !::: 0\ "'"'> "'"'> ~ .§ E '-' E ~ ~ ""'=! ~::s 0() <::l ~ 0 ~ '-> s:: ::s ::t: ~ "a ~·;;:: ~ ;>...... ,::s ...... ~ 0\...... :i Q..," "Cj ::.:: ;;:'-> <>:: (.J..__" ro "3... II) 0.. "'ro ro "2 i3 >ro 0... I 0 ("')

II) C