BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BIOLOGIE, 59: 101-157, 1989 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN, BIOLOGIE, 59: 101-157, 1989

Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites in living supraspecific taxa of Chondrichtyan fishes Editor : M. STEHMANN

Part A: Selachii. No.3: Order: Squaliformes Families: Echinorhinidae, Oxy:riotidae and Squalidae

by J. HERMAN, M. HOVESTADT-EULER and D.C. HOVESTADT

Abstract Erlaubt eine direkte generische Zuordnung und einige taxonomische Anpassungsvorschliige. Die Ergebnisse stiitzen Auffassungen friiherer Autoren, dass die Echinorhinidae eine eigene Ordnung darstellen ; die The teeth of nearly all supraspecific taxa of squaliformes are described Oxynotidae odontologisch mit den Squalidae ubereinstimmen ; die and illustrated by SEM photos. The tooth morphology allows a direct Squalidae-Gattungen in 8 Unterfamilien gruppiert werden Konnten, assignment to all these taxa and some proposals of taxonomic von denen 4 bisher nicht definierbar waren; Centroselachus GARMAN, adjustments. 1913 findet sich bestiitigt und sollte wieder errichtet werden ; odontolo­ The results support opinions of other authors, that Echinorhinidae gisch finden sich keine signiftkanten Unterschiede zwischen Cirrhigaleus represent a separate order, Oxynotidae coincide odontologically with und Squalus, Lepidorhinus und Centrophorus, Scymnodon und Squalidae ; Squalid genera could be grouped into 8 subfamilies, 4 of Scymnodalatias, ebensowenig Squaliolus sowie Heteroscymnoides which could not be defined hitherto; Centroselachus GARMAN, 1913 is einerseits und Euprotomicrus anderseits ; den 3 Somniosus-Arten confirmed and should be revalidated ; significant odontological waren 2 Untergattungen zuzuordnen : Somniosus (Somniosus) und differences do not exist between Cirrhigaleus and Squalus, Lepidorhinus Somniosus (Rhinoscymnus). Ein morphologischer Schliissel zur gene­ and Centrophorus, Scymnodon and Scymnodalatias, neither between rischen Bestimmung wird gegeben. Squaliolus and Heteroscymnoides and Euprotomicrus on the other Schliisselworter: Elasmobranchii- Selachii- Squaliformes- Echinorhi­ hand; 2 subgenera appear to be justified for the 3 species of Somniosus : nidae - Oxynotidae - Squalidae - Odontologie. Somniosus (Somniosus) and Somniosus (Rhinoscymnus). An odontological key is given for all these supraspecific taxa. Introduction Key-words: Elasmobranchii- Selachii- Squaliformes -Echinorhinidae - Squalidae- Odontology. Major revisions of shark systematics and squaloids in Resume particular were published by Bigelow & Schroeder (1957), Bass, d'Aubrey and Kistnasamy (1976) and

Les dents de Ia plupart des taxons supraspecifiques des squaliformes Compagno (1973, 1977 and 1984). sont decrites et figurees (cliches MEB). La morpho Iogie dentaire permet The teeth of Squaliformes were the subject of a study to une identification generique ou subgenerique ainsi que Ia formulation compare fossil teeth by Ledoux (1970, 1972), who d'un certain nombre de propositions de reajustements taxonomiques. described and figured several genera, as well as Casier Dans !'ensemble, ces resultats concordent avec !'opinion d'autres chercheurs. (1961). Ainsi, les Echinorhinidae representeraient un ordre distinct ; les Also Welton (1973) described and figured several Oxynotidae ne se distinguent odontologiquement pas des Squalidae. genera in an unpublished thesis. Les divers genres de Squalidae pourraient etre regroupes en 8 sous­ In this fascicle, will be described and illustrated the familles, dont 4 non encore definies. Centroselachus GARMAN, 1913 est revalidable. II n'existe aucune teeth of all living squaliform families and genera difference odontologique significative entre Cirrhigaleus et Squalus, belonging to this order, with the exception of Mollis­ Lepidorhinus et Centrophorus, Scymnodalatias et Scymnodon, pas quama parini DOLGANOV, 1984, which was not avai­ plus qu'entre Squaliolus, Heteroscymnoides et Euprotomicrus. lable for examination. Les 3 especes de Somniosus peuvent etre reparties au sein de 2 sous­ genres: Somniosus (Somniosus) et Somniosus (Rhinoscymnus). Also Zameus squamulosus, a genus which is in discus­ Une cle odontologique est proposee pour Ia distinction de tous ces sion (Taniuchi & Garrick, 1986), was not available for taxons. examination. Mots-clefs : Elasmobranchii- Selachii- Squaliformes- Echinorhinidae - Oxynotidae - Squalidae - Odontologie. The ordering of genera and species is according to the grouping of taxa based on the present result. Kurzfassung The nomenclature of taxa is according to Compagno ( 1984). The original literature reference of each specific Die Kieferziihne fast aller squaliformen supraspezifischen Taxa werden taxon will be given in the descriptive part, respectively beschrieben und mit REMPhotos illustriert. and not be repeated in the bibliography. 102 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

Systematics and materials The following 904 specimens belonging to 48 species were examined : 1 female ORDER: SQUALIFORMES, Aculeola nigra 3 males sensu COMPAGNO (1984) Centrophorus acus 1 male Centrophorus granulosus 18 males 28 females Centrophorus lusitanicus 2 females This order comprises the families Echinorhinidae, Centrophorus uyato 3 males 1 female Oxynotidae and Squalidae, which will be described Centrophorus squamosus 15 males 47 females and illustrated in detail below. Centroscyllium fabricii 4 males 51 females With the addition of Mollisquama parini DOLGANOV, Centroscyllium granulosum 18 males 24 females 1984, that was not yet included, these families comprise Centroscyllium nigrum 3 males 3 females the following genera : Centroscyllium ritteri I male Family : ECHINORHINIDAE GILL, 1872 Centroscymnus coelolepis 47 males 47 females Genus : Echinorhinus BLAINVILLE, I816 Centroscymnus crepidater 25 males 48 females type species : Echinorhinus brucus Centroscymnus owstoni I female Cirrhigaleus barbifer 1 female Dalatias licha 22 males 40 females Family: 0XYNOTIDAE Deania calcea 48 males 45 females Genus: Oxynotus RAFINESQUE, I810 Deania histricosa 1 male 1 female type species: Oxynotus centt·ina Deania profundorum 2 males 6 females Echinorhinus bntcus 2 males 3 females Family: SQUALIDAE BLAINVILLE, I816 Echinorhinus cookei 2 males 1 female Genus: Aculeola DE BUEN, 1959 Etmopterus bullisi 1 male 2 females type species : Aculeola nigra Etmopterus hillianus 6 males 12 females Genus: Centrophorus MULLER & HENLE, I837 Etmopterus lucifer 2 males 2 females type species : Centrophorus granulosus Etmoptems polli 1 male 2 females Genus: Centroscyllium MULLER & HENLE, 1841 Etmopterus princeps 5 males 4 females type species : Centroscylliumfabricii Etmopterus pusillus 2 males 3 females Genus: Centroscymnus BOCAGE & CAPELLO, 1864 Etmopterus schultzi 3 males I female type species : Centroscymnus coelolepis Etmopterus spinax 45 males 61 females Genus: Cirrhigaleus TANAKA, 1912 Etmopterus virens 2 females type species : Cirrhigaleus barbifer Euprotomicroides zantedeschia 1 male 1 female Genus: Dalatias RAFINESQUE, I810 Euprotomicrus bispinatus 1 male 2 females type species : Dalatias licha Heteroscymnoides marleyi 1 female Genus : Deania JORDAN & SNYDER, 1902 Isistius brasiliensis I male 3 females type species : Deania calcea Oxynotus bmniensis l male Genus: Etmopterus RAFINESQUE, 1810 Oxynotus caribbaeus 1 male type species : Etmoptems spinax Oxynotus cenu·ina 13 males I6 females Genus: Euprotomicroides HULLEY & PEN RITH , 1966 Oxynotus paradoxus 4 males 2 females type species : Euprotomicroides zantedeschia Scymnodalatias albicauda 1 female Genus : Euprotomicrus GILL, 1864 Scymnodon obscurus 2 males 2 females type species : Euprotomicrus bispinatus Scymnodon ringens 18 males 30 females Genus : Heteroscymnoides FOWLER, 1934 Somniosus microcephalus 7 males 18 females type species : Heteroscymnoides marleyi Somniosus pacificus 1 male 2 females Genus: lsistius GILL, 1864 Somniosus rostratus 1 male 4 females type species : Isistius brasiliensis Squaliolus laticaudus l male 4 females Gensu: Mollisquama DoLGANOV, I984 Squalus acanthias 36 males 45 females type species : Mollisquama parini Squalus asper 1 female Genus : Scymnodalatias GARRICK, 1956 Squalus blainvillei 4 males 18 females type species : Scymnodalatias sherwoodi Squalus cubensis 1 male Genus : Scymnodon BOCAGE & CAPELLO, 1864 type species : Scymnodon ringens There is a remarkable morphological difference in Gensu: Somniosus LESUEUR , 181 8 tooth arrangement between Echinorhinidae, Oxynoti­ type species : Somniosus microcephalus dae and Squalidae. The lower tooth ftles are very Genus: Squaliolus SMITH & RADCLIFFE, 1912 compressed in Oxynotidae and Squalidae (with the type species : Squaliolus laticaudus exception of the genera Centroscyllium and Aculeola) , Genus : Squalus LINNAEUS, 1758 resulting in interlocking of the teeth. This interlocking type species : Squalus acanthias is demonstrated by positioning the outer mesial part of I'

Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 103

each tooth behind the inner distal part of the previous Scymnodon ringens 24125 14115 tooth. This phenomenon is even present in both upper Somniosus microcephalus 20124 24/ 29 and lower jaws in some genera of the Squalidae. These Somniosus rostratus 30 161 18 teeth have therefore a particular depression on outer Squaliolus laticaudus 10111 9 mesial and inner distal part of the root. Interlocking is Squalus acanthias 11 115 11 /12 neither existing in upper, nor lower teeth of Echinorhi­ * =or more nidae. Several degrees of dignathic heterodonty are shown Remarkable is the extremely large number of tooth within the genera ofthe order SQUALI FORMES. This is rows in Centroscylliwnfabricii and the extreme reduc­ demonstrated by slightly smaller upper teeth (particu­ tion of tooth rows in Oxynotus, Squaliolus, Hetero­ larly in juvenile specimens) in Echinorhinus, Centroscyl­ scymnoides and Euprotomicrus. lium, Aculeola, Squalus and Cirrhigaleus ; smaller, longitudinally more compressed upper teeth in Deania In a remark, COMPAGNO (1984: 28, 29) suggested to and Centrophorus ; smaller, longitudinally, still group the genera of the families Squalidae and Oxynoti­ stronger compressed upper teeth in Oxynotus and dae into one family with the following subfamilies : Somniosus; much smaller, longitudinally very strongly Family : SQUALIDAE compressed upper teeth in Centroscymnus, Dalatias, Subfamily : Etmopterinae Isistius, Etmopterus, Scymnodon, Euprotomicrus, Genus : Etmopterus Squaliolus, Heteroscymnoides and Euprotomicroides. Genus : Aculeola Several of the genera have either symphysial, or Genus : Centroscyllium parasymphysial, or pseudosymphysial teeth in upper and or lower jaws. The lower jaw teeth are considered Subfamily : Squalinae as particularly important for this phenomenon, because Genus : Squalus they are not affected by nostrils and eyes. A lower Genus : Cirrhigaleus symphysial tooth is present in Oxynotus, Dalatias, Genus : Centrophorus lsistius, Euprotomicroides and Echinorhinus (in Echi­ Genus : Deania norhinus it may only be the relict of a true symphysial tooth row). Parasymphysial teeth were observed in Subfamily : Dalatiinae Etmopterus, Aculeola, Centroscyllium, Squalus, Cen­ Genus : Dalatias trophorus, Deania, Somniosus and Centroscymnus. Genus : Heteroscymnoides Pseudosymphysial teeth were observed in species of the Genus : Euprotomicroides genera Squalus (± 5%) and Centroscymnus (+ 40%). Genus : Euprotomicrus This phenomenon appears to be of no further use and is Genus : Isistius presented as additional information only. Genus : Squafiolus The tooth counts are also very remarkable within the Subfamily : Somniosinae genera of the SQUALIFORMES. We noted the following Genus : Somniosus counts in upper and lower jaw halfs : Genus : Centroscymnus upper lower Genus : Scymnodalatias Aculeola nigra 30 30 Genus : Scymnodon Centroscyllium fabricii 32151 33152 Genus: Oxynotus Centroscymnus coelolepis 24133 17/18 Centroscymnus crepidater 16126 14118 Although no further arguments nor comments were Centrophorus granulosus 18119 14116 presented for this grouping, odontology now also Centrophorus squamosus 15119 13132 allows grouping into subfamilies. In general, this Dalatias licha 16119 819 grouping is similar to that of COMPAGNO (1984) but Deania calcea II I 17 12116 differs in some particular cases. Echinorhinus brucus 10114 9/13 Tooth morphology offers important phylogenetic infor­ Etmopterus spinax 11 I 16 13120 mation based on types of root vascularisation. In the Euprotomicrus bispinatus 9 I 11 10/11 first issue of this series (1987), however, the authors Euprotomicroides zantedeschia 14 I 15 17 have declared their intention not to draw nomenclato­ Heteroscymnoides marleyi 6!7 11 1 12 rially valid conclusions from their odontological results, Isistius brasiliensis 15I 17 12115 respectively. Being aware of dealing with one complex Jsistius plutodus 10 I 14 819 of characters only, they will only present this according Oxynotus cenlf·ina 5I 6 4 odontological conclusions and leave it to following Scymnodalatias afbicauda 25* 15 revising authors to incorporate also odontological Scymnodafatias sherwoodi 16* 16 points of view for a full systematic review with possible (GARR ICK, 1956) taxonomic and nomenclatorial changes. I I 104 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

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Fig. I. Main tooth types of the Squaliformes. Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 105

I I

Descriptions of the odontological morphotypes Oxynotus centrina (LINNAEUS, 1758) Pl. 2, T. pl. 2 Echinorhinus brucus BONNATERRE, 1788 Squalus centrina LINNAEUS, 1758, Syst. Nat., ed. 10, 1 : Pl. I, T. pl. 1. 233.

Squalus brucus BONNATERRE, 1788, Tabl. encyclop. This species shows a strong dignathic heterodonty and method. trois reg. Nat., Ichthyol., Paris : 11. a weak gradient monognathic heterodonty. The root is The species shows a very weak dignathic heterodonty always anaulacorhizid. in having parasymphysial teeth with poorly developed The dimensions of the lower teeth are subcentimetrical, mesial cusplets in the lower jaw. Also the gradient those of the upper teeth plurimillimetrical. monognathic heterodonty is weak. The ontogenetic heterodonty is very strong by monocuspid teeth in Upper jaw. . juveniles and multicuspid teeth in adults. The dimen­ The teeth are compressed, having a more or less sions of the teeth are hemicentimetrical to subcentime­ vertically elongated crown and root. trical in range. The crown is twice as high as the crown base width. The The inner part shows a slightly convex crown surface, symphysial tooth has a symmetrically triangular crown. forming a weak angle with inner root surface. The The lateral teeth show a crown becoming increasingly outer part shows a less convex crown surface with an asymmetrical, less high, and broader based toward the indented crown-root junction. commissure. Both mesial and distal cutting edges are The crown is multicuspid with a principal cusp strongly smooth. inclined toward the commissure. One to three mesial The inner face is strongly convex. There is no trace of cusplets arise from the arched mesial cutting edge of the an uvula. The outer face of the crown is weakly convex, principal cusp on the anterior and lateral teeth, all and a convex and broad apron extends nearly to the directed strongly toward the symphysis. The mesial root base. The apron is not overhanging the root but cusplets diminish to remain as a serration only on forms one part with it. A true outer or inner ornamenta­ posterior teeth. The parasymphysiallower teeth have tion is absent. one small, poorly developed mesial cusplet. Only one The root is elongated and has a more or less rectangular distal cusplet is present on almost all teeth, that is appearance. From the crown-root junction downward, directed toward the commissure, disappearing in the the root is narrowing and the basal edges are slightly crown base in posterior teeth. On the anterior teeth a ro nded. The inner face is flat, showing a median poorly developed second distal cusplet can be present. aperture near the crown-root junction ; sometimes a There is no trace of any outer, or inner crown poorly developed median vertical groove is present, ornamentation. which is more distinct on the lateral and posterior teeth. The root is anaulacorhizid, rectangular with sometimes A deep media-basal sulcus is always present. rounded mesial and distal ends. On the outer face well The outer face shows on both sides of the apron three to developed vertical protuberances are present near the four main foramina. At the basal edge, several randomly crown-root junction. An outer mesial and distal depres­ scattered foramina are present. sion is present. Numerous foramina are scattered over the whole width of the depressed crown-root junction. Lower jaw. The inner face of the root shows one two three median The teeth are strongly compressed and interlocked. The apertures near the crown-root junction, from which triangular crown is as high as the crown base width. poorly developed vertical grooves run to the root base. The crown is high and symmetrically triangular in the The surface of the inner face is pierced with randomly symphysial tooth but becomes increasingly asymmetri­ scattered, vertically elongated foramina. cal and less high in lateral teeth. It consists of a bluntly rounded, cusplet-like knob at each basal edge, conti­ Family: Oxynotidae GILL, 1872 nued upward as strong serration, which becomes reduced in upper crown half and finally disappears This family is monotypic with Oxynotus RAFINESQUE, totally toward the apex. A true ornamentation is 1810 ; type species Oxynotus centrina. absent. The slightly convex inner crown face shows a broad, flat uvula, that is rectangular on the symphysial tooth Genus: Oxynotus RAFINESQUE, 1810 and becomes more oblique on teeth toward the commissure. The genus Oxynotus comprises four species : Oxynotus On the inner face there is a large aperture at each side of paradoxus FRADE, 1929, Oxynotus caribbaeus CER­ the uvula and one large median aperture below the VIGNON, 1961 , Oxynotus bruniensis (OGILBY, 1893) basal depression. A short, broad, media-basal sulcus is and Oxynotus centf·ina (LINNAEUS, 1758). present. The inner surface is flat, and sometimes one or -0\0

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Text plate 1. Echinorhinus brucus (BONNATERRE, 1788). Female 228 em. t.l., Senegal. Upper and lower tooth set: x 1.5. Isolated teeth are macrophotographics of Mr. K. Jansen, Oudenbosch, the Netherlands. Upper tooth: x 3, lower teeth x 4. I' Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 107 two foramina appear above the sulcus. A weak inner The crown base overhangs the crown-root junction. distal depression is due to interlocking of the lower The inner crown-root junction is depressed. teeth. The anaulacorhizid root is strongly compressed. The outer face shows a large aperture at each side of the The inner face of the root is low and strongly convex apron. Near the junction of the apron and root, some longitudinally, more or less supporting the uvula. vertical protuberances are present. Several foramina are present along the crown-root Two to four foramina surround the sulcus, and several junction. There is a large central-basal aperture present, scattered foramina are on the outer basal edge of the that affects also the root base. The root base is root. A weak outer mesial depression is also due to thickened centrally throughout its length. The outer interlocking. face is about twice as high as the inner face. Two or three apertures with several foramina· are present on Family: Squalidae BLAINVILLE, 1816 both sides of the apron.

This family comprises eighteen genera as listed in the Genus: Cirrhigaleus TANAKA, 1912 introduction. This genus is monotypic with the type species Cirrhiga­ Genus : Squalus LINNAEUS, 1758 leus barbifer.

After COMPAGNO, 1984, this genus comprises eight Cirrhigaleus barbifer TANAKA, 1912 species : Squalus asper MERRETI, 1973, Squalus Pl. 4. blain vi/lei (Risso, 1826), Squalus japonicus IsHIKAWA, 1908, Squalus megalops (MACLEAY, 1881), Squalus Cirrhigaleus barbifer TANAKA, 1912, Fig. Descr. Fish. me/anurus FOURMANOIR & RIVATON, 1979, Squa/us Japan, 9: 151. mitsukurii JORDAN & SNYDER, 1903, Squalus rancureli FOURMANOIR & RIVATON, 1979 and type species This species shows a very weak dignathic heterodonty Squalus acanthias LINNAEUS, 1758. in having slightly shorter upper teeth. A gradient monognathic heterodonty is rather obvious in antero­ Squalus acanthias LINNAEUS, 1758 lateral teeth, which becomes broader toward the com­ Pl. 3, T. pl. 3. missure. Sexual heterodonty could not be examined, because only teeth of a female were available. Squalus acanthias LINNAEUS, 1758, Syst. Nat., ed. 10, Both, upper and lower teeth are interlocked, presenting 1: 233. ~ weak inner distal, and a strong outer mesial depres­ SIOn. This species shows a very weak dignathic heterodonty The dimensions of the teeth are hemicentimetrical in in having slightly shorter upper teeth. A gradient range. monognathic heterodonty is rather strong in antero­ The crown has a distally strongly inclined principal lateral teeth becoming broader toward the commissure. cusp, which is mesio-basally rounded. Its long mesial Sexual heterodonty is indicated by a sigmoidal mesial cutting edge is weakly arched, while its short distal cutting edge in males, contrasting an arched mesial cutting edge joins a low distal blade in a notch. cutting edge in females. Serrations are absent on the cutting edges of the Both upper and lower teeth are interlocked, presenting principal cusp are absent. a weak inner distal but a strong outer mesial depression. Except for some short basal costules, a true inner or The dimensions of the teeth are hemicentimetrical in outer ornamentation is absent. range. On the inner face, the principal cusp is weakly convex. The crown has a distally strongly inclined principal A well developed, short central-basal uvula is present. cusp, which is mesio-basally rounded. Its long mesial On the outer face, both principal cusp and distal blade cutting edge is arched or sigmoidal, while its short distal are weakly convex, showing a well developed central­ cutting edge joins a low distal blade in a notch. Weak basal apron, that strongly overhangs the root reaching serrations are sometimes present on the cutting edges of its base. This apron is broad and more or less triangular the principal cusp. Except for some short basal with a rounded basal edge. The crown base overhangs costules, a true inner or outer ornamentation is absent. the crown-root junction. The inner crown-root junction On the inner face, the principal cusp is weakly convex. is depressed. A well developed, short central-basal uvula is present. The anaulacorhizid root is very compressed. On the outer face, both principal cusp and distal blade The inner face of the root is low and strongly convex are weakly convex, showing a well developed central­ longitudinally, more or less supporting the uvula. basal apron, that strongly overhangs the root, and Several foramina are present along the crown-root extends to the base of the root. This apron is narrow junction. There is a large central-basal aperture present, and more or less rectangular with a rounded basal edge. that affects also the root base. The root base is '' 108 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

Text plate 2. Oxynotus centrina (LtNNAEUS, 1758). Female 31 em. t.l., Sete, France, Mediterranean Sea (left)Jemale 36 em. t.l. , Dakar, Senegal (centre) and male 43 em. t.l., Dakar, Senegal (right). All magnifications x 8. Lower jaw: Female 46 em. t.l. x 18 (Diagraph of the first nonjunctionalfile).

Text plate 3. Squalus acanthias (LtNNAEUS, 1758). Female 90 em. t.l., Belgium. Upper jaw x 3; lower jaw x 10 (Diagraph of the outer most functional file). (') 0 :::I s. cr' ~ ...... 5' :::I en ...... 0

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0 Text plate 4. Etmopterus spinax (Lt N NAEUS, 1758). Female 55 em. t.l., S. Lousy, N. E. Atlantic (upper jaw x 8). Lower jaw x 8 ( Diagraph of -\0 the first non-functional file). '' 110 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT thickened centrally throughout its length. The outer The high inner face of the root is fait, and a small face is a little higher than the inner face. One or two central basal sulcus gives a pseudo bilobate appearance apertures with some foramina are present at both sides to the anaulacorhizid root. A small central aperture is of the apron. present near the crown-root junction. The low outer face of the root is divided by the central Genus: Etmopterus RAFINESQUE, 1810 basal sulcus. Several large foramina are scattered along the crown-root junction. After CoMPAGNO (1984), this genus comprises sixteen Lower jaw species: Etmopterus baxteri GARRICK, 1957, Etmopte­ The lower teeth are always strongly interlocked, presen­ rus brachyurus SMITH & RADCLIFFE, 1912, Etmopterus ting deep inner distal and outer mesial depressions bullisi BIGELOW & SCHROEDER, 1957, Etmopterus affecting both, crown and root. The root is anaulacorhi­ decacuspidatus CHAN, 1966, Etmopterus gracilispinis zid. KREFFr, 1968, Etmopterus granulosus (GUNTHER, The crown has a very strongly inclined triangular 1880), Etmopterus hillianus (POEY, 1861), Etmopterus principal cusp reaching sometimes beyond the distal lucifer JORDAN & SNYDER, 1902, Etmopterus polli root end in anterior teeth. The distal blade is short and BIGELOW & SCHROEDER, 1953, Etmopterus princeps joins the principal cusp in a notch. The cutting edges COLLETT, 1904, Etmopterus pusillus (LOWE, 1839), are always smooth. Etmopterus schu/tzi BIGELOW & SCHROEDER, 1953, The inner face of the crown is slightly convex. A flat Etmopterus sentosus BASS, d'AUBREY & KISTNASAMY, uvula is present, covering also the upper central and 1976, Etmopterus unicolor (ENGELHARDT, 1912), mesial part of the root. There are two particular Etmopterus villosus GILBERT, 1905, Etmopterus virens depressions on the uvula base. Inner ornamentation is BIGELOW & SCHROEDER, 1953 and the type species absent. Etmopterus spinax LINNAEUS, 1758. The outer crown face is more or less flat, showing a flat apron reaching half of the root's height and covering the upper central and distal part of the root. Etmopterus spinax (LINNAEUS, 1758) The uvula-root junction is mesio-distally straight. An Pl. 5, T. pl. 4. outer ornamentation is absent. The root is more or less rectangular. Squalus spinax LINNAEUS, 1758, Syst. Nat., ed. 10, 1 : The inner face is flat with a strong distal depression. 233. There are three apertures present along the uvula-root junction, two under both mesial and distal lower ends This species is extemely dignathic heterodont and of the uvula and one at deeper level in the central part. gradient monognathic heterodont in both upper and The outer face of the root shows a strong mesial lower teeth. The root is always anaulacorhizid. depression. There are two apertures present, one under the mesial lower end of the uvula and one in the centre Upper Jaw of the root. Along the uvula-root junction, there are up The upper teeth are always multicuspid in having one to 8 smaller foramina present. principal cusp flanked by one to two cusplets. The teeth are not interlocked. The dimensions of the teeth are Centroscyllium M ULLER & HENLE; 1841 plurimillimetrical in range. The crown has an erect, slender principal cusp with a This genus comprises (after COMPAGNO, 1984) the weak basal constriction. Both, mesial and distal cutting following six species : Centroscyllium fabricii (REIN­ edges therefore have a sigmoidal appearance. One or HARDT, 1825), which is the type species, Centroscyllium two cusplets always flank the principal cusp. If a second granulatum GUNTHER, 1887, Centroscyllium kamoha­ cusplet present, it is only one third as high as the first rai ABE, 1966, Centroscyllium nigrum GARMAN, 1899, cusplet. The first cusplet can reach half the height of the Centroscyllium ornatum (ALCOCK, 1889) and Cen­ principal cusp. The cusplets show the same basal troscyllium ritteri JORDAN & FOWLER, 1903. constriction as the principal cusp. They become lower toward the commissure and disappear on the posterior Centroscyllium fabricii (RHEINHARDT, 1825) teeth. Pl. 6, T. pl. 5. On inner face, the principal cusp and cusplets, are weakly convex. Spinax fabricii REINHARDT, 1825, Overs. K. danske Uvula and inner ornamentation are absent. Vidensk. Selsk. Forh., (1824-1825): 3. On outer face, the principal cusp and cusplets, are flat or slightly convex. This species is almost homodont but shows a gradient A strong basal ornamentation is present on principal weak monognathic heterodonty by reduction of tooth cusp and cusplets by partly branched vertical custules. size toward the commissure. Sexual heterodonty is not Apron is absent. existing. The teeth are plurirnillimetrical in range. I I Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites Ill -N

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Textplate6. Somniosus microcephalus (BLO CH & SCHNEIDER, 1801). Female470 em. t.l., N. Lousy, N.E. Atlantic. Upper jaw x 1; lower jaw x 0.7. Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 113

The crown has an erect, slender and triangular principal shows a strong ornamentation. Long vertical costules cusp. flanked by a pair of also erect and slender but reach half way up the principal cusp, or higher. A lower mesial and distal cusplets.The height of both reticulated, secondary basal ornamentation is present cusp lets is more than half of the height of the principal between these costules. Apron is absent. cusp on anterior and antero-lateral teeth, but only The root is hemiaulacorhizid but there is no true about half the height on latera-posterior teeth. Both, bilobation. The mesial and distal basal ends are distal and mesial cutting edges of principal cusp and rounded. cuplets are smooth. The principal cusp becomes lower The inner face shows a broad shallow pseudo-groove in and broader based toward the commissure. lower half of the root and a large central aperture near The inner face of principal cusp and cusp lets is weakly the crown-root junction. Some randomly scattered convex, and they have a strong ornamentation. Vertical foramina are present at the basal edge of the root. costules cover the basal half of the principal cusp and The outer face shows up to six foramina along the the entire length of the cusplets. Distinct reticulated, crown-root junction. The basal part of the inner secondary ornamentation is present between the basal pseudo-groove is visible. costules, uvula is absent. The outer face of principal cusp and cusplets is flat to Genus: Somniosus LESUEUR, 1818 slightly convex and shows the same ornamentation with an extension. Vertical costules reach half the This genus comprises the three species Somniosus height of the principal cusp and are all over the cusp lets rostratus (RISSO, 1826), Somniosus pacificus BIGELOW and basal reticulation as on inner face. & SCHROEDER, 1944, and Somniosus microcephalus The root is hemiaulacorhizid and bilobate. The root (BLOCH & SCHNEIDER, 1801 ), of which the latter is the lobes are rounded at the mesial and distal basal ends. type species. We will also describe, illustrate and discuss The inner root face shows a broad, deep central Somniosus rostratus (RISSO, 1826), because of the pseudo-groove in lower half of the root and a large discussion amongst several authors on the species central aperture near the crown-root junction. At the existing. basal edges of each root lobe, numerous randomly scattered foramina are present. Somniosus microcephalus (BLOCH & SCHNEIDER, The outer face shows ten or more foramina along the 1801) crown-root junction. The basal part of the inner Pl. 8, 9, T. pl. 6, 7. pseudo-groove is visible. Squalus microcephalus BLOCH & SCHNEIDER, 1801, Syst. Ichthyol., Berlin : 135. Genus: Aculeola DE BUEN, 1959 This species shows a strong dignathic heterodonty and The type species of this mono typic genus is Aculeola a weak gradient monognathic heterodonty. nigra. The dimensions of the lower teeth are centrimetrical and those of the upper teeth are hemicentimetrical in Aculeola nigra DE BUEN, 1959 range. Pl. 7. Upper jaw. Aculeola nigra DE BUEN, 1959, Bol. Mus. Nac. Hist. The teeth are slightly compressed, crown and root are Nat. Santiago, Chile, 27(3): 180. vertically elongated. The crown is twice as high as the crown base width. The This species is almost homodont but shows a gradient teeth have an irregularly shaped, though more or less weak mono gnathic heterodonty by reduction of tooth symmetrical erect crown, that is slightly inclining size toward the commissure. Sexual heterodonty is toward the commissure on latera-posterior teeth. Both present in that females have a slightly lower, broader mesial and distal cutting edges are undulated but not based crown with a more intensely reticulated outer serrated. ornamentation. The teeth are plurimillimetrical in The inner face is strongly convex. Uvula and ornamen­ range. tation are absent. The crown has an erect, slender and symmetrical The outer face is slightly convex. A prominent, principal cusp. Mesial and distal cusplets are absent, vertically elongated apron is present, forming one part and the cutting edges are smooth. with the root. Near the crown-root junction, some The principal cusp becomes slightly lower, a little vertical protuberances are present. The pseudoholaula­ broader based and curves stronger inward near the corhizid root is elongated and divided into two root comrrussure. lobes. Both root lobes have a more or less rectangular The inner face of the principal cusp is convex and appearance. The basal edges are slightly rounded. without ornamentation. Uvula is absent. The inner face is flat, showing a broad median groove, Outer face of principal cusp is slightly convex, and that includes a large central foramen with five or six 114 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

Text plate 7. Somniosus microcephalus (BLOCH & SCHNEIDER, 1801). Male 176 em. t.l., E. Oreille, N.E. Atlantic. Upper jaw x 2; lower jaw x 2. (Diagraph of the first non-functional file).

Text plate 8. Somniosus rostratus (Risso, 1826). Male 109 em. t.l., W Hebrides, N.E. Atlantic. Upper jaw x 3; lower x 3. I I

Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 115 smaller foramina. The root is slightly depressed mesio­ The outer face is weakly convex. Apron is absent. A distally. Some scattered foramina are present on the strong, reticulated basal ornamentation is present, in root lobes. which a secondary much finer reticulated ornamenta­ The outer face shows on both sides of the apron several tion is included. randomly scattered foramina. The basal part of the The anaulacorhizid root is short and slightly constricted inner median groove is visible. from below the crown-root junction downward. The basal ends are slightly rounded. Lower jaw. The inner face is flat, showing a large central foramen The teeth are very compressed, with a strong distally below the crown-root junction. Below this central inclined principal cusp. The mesial cutting edge is foramen a second one is present from which a broad slightly arched, while the distal cutting edge joins the sulcus runs to the central basal end of the root. distal blade in a notch. The height of the crown is half to The outer face is slightly concave and shows two or a quarter of the crown base width. Both mesial and three large central foramina. Also two or three small distal cutting edges are smooth. A true ornamentation scattered mesial and distal foramina are present. The on the crown is absent. basal part of the inner median groove is visible. The slightly convex, inner face of the crown shows a small, convex uvula, forming one part with the root. Lower jaw. The outer face shows a rather short, triangular apron, The teeth are very compressed, with a triangular, also forming one part with the root. The outer surface is distally inclined principal cusp. The mesial cutting edge a little more convex than the inner one. is straight and the distal cutting edge joins the distal The vertically expanded anaulacorhizid root has roun­ blade in a notch. Both mesial and distal cutting edges ded mesio- and distal-basal ends and is constricted and distal blade are smooth. There is no true omamen-· from just below the crown-root junction toward the tation on the crown. The height of the crown has the basal root end. This constriction becomes much stron­ dimensions of the crown base length. ger toward the commissure. The slightly convex inner face of the crown has no true On the inner face there is a large aperture situated uvula. The outer face shows a rather short, rectangular distally besides the uvula. This aperture includes three apron, forming one part with the root. large foramina. A broad, shallow median sulcus is The vertically elongated anaulacorhizid root has roun­ present. Several foramina are randomly scattered on ded mesial and distal basal ends and is slightly the flat inner face of the root. Interlocking has caused a constricted from just below the crown-root junction strong inner distal depression. downward. Three large apertures are present on the The outer face shows some large apertures at each side inner face, situated on the central, distal and mesial of the apron. Near the junction of the apron and root, parts along the crown-root junction. some vertical protuberances are present. From the central aperture, a broad shallow sulcus runs Interlocking has resulted in a mild outer mesial to the central basal end of the root. One or two small depression. foramina are present in this sulcus. Interlocking has led Somniosus rostratus (RISSO, 1826) to a strong inner distal depression. Pl. 10, T. pl. 8. On the outer face, the apron is flanked by a pair of large apertures. Scymnus rostratus RISSO, 1826, Hist. nat. pnnc1p. Five to six foramina are present along the crown-root prod. Europe Merid., Paris 3 Poissons : 138. junction. The basal part of the sulcus is visible. This species also shows a strong dignathic heterodonty Interlocking has resulted in a mild outer mesial and a weak gradient monognathic heterodonty. The depression. root, however, is always anaulacorhizid. The dimensions of the lower teeth are hemicentimetrical Genus: Centrophorus MULLER & HENLE, 1837 and those of the upper teeth are plurirnillimetrical in range. The genus Centrophorus comprises (after COMPAGNO, 1984) the following eight species: Centrophorus acus Upper jaw. GARMAN, 1906, Centrophorus granulosus (BLOCH & The teeth are more or less compressed with the root SCHNEIDER, 1801), Centrophorus harrissoni McCUL­ short and the crown vertically elongated. LOCH, 1915, Centrophorus lusitanicus BOCAGE & The triangular, symmetrical crown is three times as CAPELLO, 1864, Centrophorus moluccensis BLEEKER, high as the crown base width and slightly inclining 1860, CentrophorusniaukangTENG, 1959, Centropho­ toward the commissure on latera-posterior teeth. Both, rus squamosus (BONNATERRE, 1788), Centrophorus mesial and distal cutting edges are smooth. tessellatus GARMAN, 1906, and Centrophorus uyato The inner face is weakly convex. Uvula is absent. A (RAFINESQUE, 1810). weak ornamentation is presented by some basal cos­ The type species is Centrophorus granulosus (BLOCH & tules. SCHNEIDER, 1801). 116 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

Text plate 9. Centrophorus granulosus (BLOCH & SCHNEIDER, 1801). Male 86 em., t.l. Nice, Mediterranean Sea. Upper jaw x 3. Female 97 em. t.l. Motril, Mediterranian, Diagraph of the lower functional file (x 5).

Text plate 10. Centrophorus squamosus (BONNATERRE, 1788). Male 92 em. t. l. , S. W. Rockall, N.E. Atlantic. Upper jaw x 4; lower jaw : Diagraph of the functional file (x 5). Formerly Lepidorhinus squamosus (BONNATERRE, 1788). 11

Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 117

Centrophorus granulosus (BLOCH & SCHNEIDER, 1801) situated mesially aside the apron. It is the origin of a PI. II, T. pl. 9. broad, shallow sulcus running to the central root end. Squa/us granu/osus BLOCH & SCHNEIDER, 1801, Syst. Few small foramina are on the distal region of the Ichthyol. : 135. crown-root junction. On the mesial part, one large foramen and two or three minor foramina are present. This species shows a strong dignathic heterodonty and The inner distal depression is only present in the region a weak gradient monognathic heterodonty. The root is of the crown-root junction as a result of interlocking. always anaulacorhize. The outer face of the root is flat with some small ridges The dimensions of the lower teeth are hemicentimetrical near the apron. Along the apron, one or two major and those of the upper teeth are plurimillimetrical in mesial apertures are present near the crown-root range. junction and eight to ten minor foramina. Two or three Upper jaw major apertures with some minor ones are on the distal The teeth have a triangular principal cusp and a small part near the crown-root junction. The mesial depres­ distal blade joins the distal cutting edge in a notch. The sion is well developed as a result of interlocking. principal cusp inclines slightly toward the commissure Genus: Deania JORDAN & SNYDER, 1902 on lateral and posterior teeth. Both distal and mesial cutting edges are smooth and slightly sigmoidal. A The genus Deania comprises (after COMPAGNO, 1984) constriction from below the crown-root junction down­ the four species: Deania calcea (LOWE, 1839), Deania ward is stronger developed mesially. The mesial and histricosa GARMAN, 1906, Deania profundorum distal basal ends are rounded. (SMITH & RADCLIFFE, 1912) and Deania quadri­ The inner face of the crown is convex and is about as spinosum (mcGULLOCH, 1915). The type species IS high as the crown base width. A small central basal Deania calcea. uvula is present. The outer face is slightly convex and narrow. A Deania calcea (LOWE, 1839) vertically elongated apron is present, reaching almost Pl. 13, Tpl. 11. the central root end. The inner face of the root has a depressed crown-root junction. The root becomes Acanthidium calceum LOWE, 1839, Trans. Zool. Soc. strongly convex directly below this depression, forming London, 3(1) : 19. a longitudinal ridge. A large central aperture is present This species shows a strong dignathic heterodonty and on or just below the ridge and is the beginning of a a weak gradient monognathic heterodonty. The root is broad, shallow sulcus running to the central root end. always anaulacorhizid. A few small foramina are on the crown-root junction The dimensions of lower and upper teeth are plurimilli­ and a large foramen is present in the mesial region. metrical in range. The outer face of the root is flat with some small ridges near the apron. Along the apron, one major mesial Upper jaw aperture is present near the crown-root junction and The teeth have a triangular principal cusp and a small another one on the mesio-central root end. Also one or distal and mesial blade both joining the principal cusp two minor foramina are present along the apron. in a notch. The principal cusp inclines slightly toward the commissure on lateral and posterior teeth. Both Lower jaw distal and mesial cutting edges are smooth. The teeth have a strong, distally inclined principal cusp, The inner face of the crown is convex and its height is and a small distal blad joins the distal cutting edge in a about equal to the crown base width. A small central notch. The arched mesial cutting edge is slightly basal uvula is present. serrated, while the distal one and the distal blade are The outer face is slightly convex, and a triangular, smooth. A constriction from below the crown-root vertically elongated apron with a rounded end is junction downward is stronger developed mesially. The present, reaching almost the central root end. mesial basal end is almost rectangular, while the distal The inner face of the root has a depressed crown-root basal end is rounded. junction. Directly below this depression the root The inner face of the crown is convex, and its height is becomes strongly convex, forming a longitudinal ridge. about half of the crown base width. A small central A large central aperture is situated on the ridge. A basal uvula is present. second aperture is present below this aperture, which is The outer face is slightly convex, and a narrow, the origin of a broad, shallow sulcus running to the vertically elongated apron is present reaching half way central root end. Two to four small foramina are on the toward the central root part. crown-root junction at both sides of the central The inner face of the root has a depressed crown-root aperture. junction. The root becomes strongly convex directly The outer face of the root is flat. A major mesial and below this depression, forming a longitudinal ridge. A distal aperture is present along the apron near the large semicentral aperture is present on the ridge, crown-root junction. 'I

118 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

Text plate 11. Deaniacalcea(LOWE, 1813). Female 107 em. t.l., W. Hebrides, N.E. Atlantic. Upper jaw x4; lower jaw: Diagraph ofthe first non-functional file (x 5).

Text plate 12. Centroscymnus coelolepis BOCA GE & CAPELLO, 1864. Male 85 em. t.l., N. Bille Bailey, N.E. Atlantic. Upperjaw x 5; lowerjaw : Diagraph of the first non-functional file (x 6). Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 119

Lower jaw Upper jaw The teeth have a strong, distally inclined principal cusp The teeth have an elongated, basally constricted, and a small distal blade joining its distal cutting edge in narrow principal cusp. Cusplets are absent. The mesial a notch. The mesial and distal cutting edges are and distal cutting edges are slightly sigmoidal. The root smooth. The low root has a constriction from below the is bilobate. The inner face of the crown is convex. crown-root junction downward, that is stronger mesial­ Ornamentation and uvula are absent. ly. The mesial basal end is narrower rounded than the The outer face of the crown is flat to slightly convex. distal basal end. Apron is absent. The crown base is strongly depressed, The inner face of the crown is convex, and its height is developing a mesial and distal vertical ridge on anterior about half of the crown base width. A small central teeth. On lateral teeth, from the "eye"-position toward basal uvula is present. the commissure, the basal depression is absent and The outer face is slightly convex, and a triangular, mesial and distal ridges join in a pseudo-apron. A true basally rounded apron is present reaching almost the ornamentation is absent. central root end. The inner face of the root is flat, showing one large The inner face of the root has a depressed crown-root central aperture near the crown-root junction and one junction. Directly below this depression the root on the basal apart of the root. becomes strongly convex, forming a longitudinal ridge. The outer face of the root has a strong central root Below the apron, a large central aperture is present on depression. One or two mesial and distal apertures are the ridge. A second aperture appears below this present. aperture, forming the origin of a broad, shallow sulcus running to the central root end. A few small foramina Lower jaw are on the distal region of the crown-root junction. One The teeth are strongly compressed. The slightly sigmoi­ large foramen and three to six minor foramina are dal principal cusp is strongly inclined toward the present on the mesial part. commiss4re, joining the distal blade in a notch. The The inner distal depression is only present in the region crown is half as high as the crown base width. The of the crown-root junction. Two or three major rectangular root is constricted from the crown-root apertures are on the distal part near the crown-root junction downward and is vertically elongated. The junction. Interlocking has led to a strong mesial mesial and distal root ends are slightly rounded. depression. The inner face of the crown is flat to very weakly convex. The basal part is slightly depressed. A true uvula is absent. Genus: Centroscymnus BOCAGE & CAPELLO, 1864 The outer face of the crown is also weakly convex. A vertically elongated apron is present, that has a fair The genus comprises (after CoM­ Centroscymnus constriction at half of its length. The apron is not PAGNO, 1984) the following eight species : Centroscym­ overhanging the root but forms one part together with nus coelolepis .SOCAGE & CAPELLO, 1864, Centroscym­ it. (BOCAGE & CAPELLO, nus crepidater 1864, Centro­ The inner face of the root shows near the crown-root scymnus cryptacanthus REGAN, Centroscymnus 1906, junction a large central and large mesial aperture. One REGAN, macracanthus 1906, Centroscymnus owstoni or two distal foramina are also present near the crown­ GARMAN, 1906 and Centroscymnus plunketi (WAITE, root junction. A second large central aperture is 1900). present, in the middle of the root forming the origin of a The type species is Centroscymnus coelolepis. Additio­ broad, shallow sulcus running toward the central root nally (BOCAGE & CAPELLO, Centroscymnus crepidater base. A short distal depression is only present directly 1864) was examined. Because so many remarkable below the distal blade, as a result of interlocking. differences were discovered between both species, also The outer face of the root shows a very large mesial will be described below, Centroscymnus crepidater aperture and one or two smaller ones, as well as two or along with comments on its generic status. three large distal apertures along with one or two smaller ones. The latter are vertically arranged. The Centroscymnus coelolepis Boc AG E & CAPELLO, 1864 basal part of the sulcus is visible. The mesial depression Pl. 14, T. pl. 12. is well developed.

Centroscymnus crepidater (BO CAGE & CAPELLO, 1864) Centroscymnus coelolepis, BocAGE & CAPELLO, 1864, Pl. 15, T. pl. 13. Proc. Zoo!. Soc. London, 24 : 263. Centrophorus crepidater BOCAGE & CAPELLO, 1864 This species has a strong dignathic heterodonty and a Proc. zoo!. Soc. London, 24 : 262. weak gradient monognathic heterodonty. The root is anaulacorhizid. Both upper and lower teeth are pluri­ This species has a strong dignathic heterodonty and a millimetrical in range. weak gradient monognathic heterodonty. The root is ''

120 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

Text plate 13. Centroscymnus crepidater (BOCAGE & CAPELLO, 1864). Female84 em. t.l., Porcupine, N.E. Atlantic. Upper jaw x4; lower jaw : Diagraph of the first non-functional file (x 6).

Text plate 14. Scymnodon ringens B OCAGE & CAPELLO, 1864. Female 102 em. t.l., W. Hebrides, N.E. Atlantic. Upper jaw x 3 ; lower jaw: Diagraph of the first non-functional file (x 4). Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 121

anaulacorhizid. Both, upper and lower teeth are pluri­ This species has a strong dignathic heterodonty and a millimetrical in range. weak gradient monognathic heterodonty. The root is anaulacorhize. The upper teeth are hemicentimetrical Upper jaw and the lower teeth centrimetrical in range. The teeth have an elongated, narrow principal cusp. Cusp lets are absent. The mesial and distal cutting edges Upper jaw are slightly sigmoidal The root is quadrangular. The The teeth have an elongated, narrow principal cusp. inner face of the crown is convex. Uvula is absent. An Cusp lets are absent. The mesial and distal cutting edges ornamentation is presented by short vertical costules. are slightly sigmoidal. The root is quadrangular. The The outer face of the crown is flat to slightly convex. principal cusp inclines toward the commissure in Apron is absent. The crown base is depressed with a latera-posterior teeth. The inner face of the crown is mesial and distal vertical ridge on anterior teeth. On convex. Uvula is absent, as well as an inner ornamenta­ lateral teeth, from the "eye"-position toward the com­ tion. The outer face of the crown is flat or slightly missure, the basal depression is absent and mesial and convex. Apron is absent. The crown base is depressed, distal ridges join in a pseudo-apron. A true ornamenta­ with a mesial and a distal vertical ridges. An outer tion is absent, but some scattered basal costules are ornamentation is presented by some scattered basal present. costules on anterior and antero-lateral teeth and well The inner face of the root joins the crown surface in an developed vertical basal costules on lateral and commis­ angle. One large central aperture is present near the sural teeth. crown-root junction. The inner face of the root joins the crown surface in an The outer face of the root has a central root depression. angle. Apertures or foramina are absent. A short, Three or four central apertures are present. narrow and deep basal sulcus is present. Lower jaw The outer face of the root has a central root depression. The teeth are strongly compressed. The sigmoidal Some randomly scattered foramina are present on principal cusp is inclined toward the commissure, but latera-posterior and commissural teeth. its distal cutting edge tends to be almost vertical and joins the distal blade in a notch. The rectangular root is Lower teeth narrowing from just below the crown-root junction The teeth are strongly compressed. The triangular downward, becoming more oblique toward the com­ crown is as high as the crown base wide. missure. The sigmoidal principal cusp is inclined toward the The inner face of the crown is flat to very weakly commissure but its distal cutting edge tends to be convex. The basal part is slightly depressed. A true almost vertical and joins the distal blade in a notch. The uvula is absent. rectangular root is narrowing from below the crown­ The outer face of the crown is also weakly convex. A root junction downward becoming more oblique to­ short quadrangular apron is present. The apron is not ward the commissure. overhanging the root but forms one part with it. The The inner face of the crown is flat to very weakly crown is as high as the crown base width. convex. The basal part is slightly depressed. A true The inner face of the root is strongly convex near the uvula is absent. crown-root junction, forming a longitudinal ridge. A The outer face of the crown is also weakly convex. A central and mesial aperture is situated on the ridge. short quadrangular apron is present. The apron is not Sulcus is absent. A distal depression, as result of inter­ overhanging the root but forms one part with it. locking, is hardly developed. The inner face of the root is strongly convex near the The outer face of the root shows numerous foramina, crown-root junction, forming a longitudinal ridge. A mainly surrounding the apron. The upper mesial and central and mesial aperture is situated on the ridge. A distal foramina are much larger than the basal ones. A short basal sulcus is present. mesial depression, as result of interlocking, is hardly A distal depression, as result of interlocking, is poorly developed. developed. The outer face of the root shows numerous foramina, Genus: Scymnodon BOCAGE & CAPELLO, 1864 mainly surrounding the apron. The upper mesial and The genus Scymnodon comprises the three species distal foramina are more or less vertically arranged and Scymnodon ringens BOCAGE & CAPELLO, 1864, Scym­ horizontally expanded, being much larger than the nodon obscurus (VAILLANT, 1888) and Scymnodon basal ones. The mesial depression is poorly developed. squamulosus (GUNTHER, 1877). The type species is Scymnodon ringens. Genus: Scymnodalatias GARRICK, 1956 Scymnodon ringens BOCAGE & CAPELLO, 1864, Proc. The genus Scymnodalatias comprises the four species zool. Soc. London, 24: 263. Scymnodalatias albicauda TANIUCHI & GARRICK, Pl. 16, T. pl. 14 1986, Scymnodalatias oligodon KUKUJEV & KONOVA- I I

122 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

LENKO, 1988, Scymnodalatias garricki KUKUJEV & short basal sulcus is present. KONOVALENKO, 1988 and Scymnodon sherwoodi A distal depression, as result of interlocking, is poorly (ARCHEY, 1921) ; Type species: Scymnodon sherwoodi. developed. The outer face of the root shows numerous Because the type species was not available in time for foramina, mainly surrounding the apron. The upper examination, Scymnodalatias albicauda was used for mesial and distal foramina are more or less vertically description, which seams to have a similar tooth arranged and horizontally expanded, being much morphology. larger than the basal ones. The mesial depression is poorly developed. Scymnoda/atias a/bicauda TANIUCHI & GARRICK, 1986 Genus : Isistius GILL, 1864 Pl. 17, T. pl. 15. This genus comprises the species Isistius plutodus Scymnoda/atias a/bicauda, TANIUCHI & GARRICK, GARRICK & SPRINGER, 1964 and lsistius brasiliensis 1986, Japanese J ourn. of Ichthyology 33(2) : 119-134. (QUOY & GAIMARD, 1824), of which the latter is the type species. This species has a strong dignathic heterodonty and a weak gradient monognathic heterodonty. The root is lsistius brasiliensis (QuoY & GAIMARD, 1824) anaulacorhizid. The upper teeth are hemicentrimetrical Pl. 18, T. pl. 16. and the lower teeth centrimetrical in range. Scymnus brasiliensis QvoY & GAIMARD, 1824, Zoolo­ Upper jaw gie, voy. Uranie et Physicienne, 1817-20 : 198. The teeth have an elongated, narrow principal cusp, that has a slight constriction near the crown base. This species has a strong dignathic heterodonty and a Cusp lets are absent. The mesial and distal cutting edges weak gradient monognathic heterodonty. The root is are strongly sigmoidal. The root is bilobated on anaulacorhizid. The lower teeth are strongly inter­ anterior and lateral teeth becoming quadrangular locked. The upper teeth are plurimillimetrical and the toward the commissure. The principal cusp inclines lower ones are centrimetrical in range. toward commissure in latera-posterior teeth. The inner face of the crown is strongly convex. Uvula is absent, as Upper jaw well as inner ornamentation. The outer face of the The teeth have an elongated principal cusp. Cusplets crown is slightly convex. Apron is absent. The crown are absent. The mesial and distal cutting edges are base is depressed, with a mesial and distal vertical ridge. slightly curved. The root is pseudobilobated by a deep Outer ornamentation is absent. sulcus, having quadrangular pseudo-lobes. The root is The inner face of the root joins the crown surface in an constricted just below the crown-root junction down­ angle. Apertures or foramina are absent. A short, ward. The crown is slightly distally inclined in lateral narrow and deep basal sulcus is present. and posterior teeth and 50% higher than the width of its The outer face of the root has a central root depression. base. Some randomly scattered foramina are present on The inner face of the crown is slightly convex. Uvula is latera-posterior and commissural teeth. absent, as well as inner ornamentation. The outer face of the crown is flat to slightly convex and Lower teeth a triangular, basally rounded apron is present. Outer The teeth are strongly compressed. The height of the ornamentation is absent. triangular crown equals the crown base width. The inner face of the root shows a longitudinal ridge The slightly arched principal cusp is inclined toward the near the crown-root junction, on which a large central commissure, joining the distal blade in a notch. The aperture is present. A narrow mesial depression is rectangular root is narrowing from below the crown­ present below this ridge. A short, broad and deep basal root junction downward becoming more oblique to­ sulcus is present. ward the commissure. The outer face of the root has three foramina near the The inner face of the crown is flat to very weakly crown-root junction : two foramina flanking the apron convex. The basal part is slightly depressed. A true and one central foramen below the apron. The central uvula is absent. basal sulcus is also present on the outer face. The outer face of the crown is also weakly convex. A relatively narrow, short quadrangular apron is present. Lower jaw The apron is not overhanging the root but forms one The strongly compressed teeth are always interlocked, part with it. presenting deep, vertically long, inner distal and outer The inner face of the root is strongly convex near the mesial depressions. Distal and mesial blades are absent. crown-root junction, forming a longitudinal ridge. A The crown inclines slightly toward the commissure in central and mesial aperture is situated on the ridge. A latera-posterior and posterior teeth. I I

Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 123

Text plate 15. Scymnodalatias albicauda (TANJUCHI & GARRICK, 1986). Sex unknown97.2 em t.l., Hokitika Canyon, New Zealand. Upper jaw: uncompletefile x 3; lower jaw: Diagraph x 4 (The commissural tooth is missing).

Text p late 16. Isistius brasiliensis (QuaY & GAIMA RD, 1824). Upper jaw: Female 41 em t.l., Dakar, Senegal x 5. Lower jaw: Diagraph x 6, Female 24 em t.l. Hawai Island. I I

124 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

The inner face of the crown is flat to very weakly Lower jaw convex and is as high as the crown base width. Uvula is The strongly compressed teeth are always interlocked, absent. presenting deep, vertically long, inner distal and outer The outer face of the crown is also weakly convex. A mesial depressions. Distal and mesial blades are present vertically long quadrangular apron covers the central on the symphysial tooth. The mesial blade is absent on and distal parts of the root. The apron is not overhan­ the other teeth. The crown inclines slightly toward the ging the root but forms one part with it. commissure in latera-posterior and posterior teeth. The The inner face of the root is vertically long and very flat, principal cusp is over twice as high as its base width. showing a central aperture and another smaller distal The root has a slightly rounded base with rounded foramen with sometimes a small mesial foramen near basal ends. the crown-root junction, a large, elongated central The inner face of the crown. shows a slightly convex aperture is present. A distal depression, as result of principal cusp. Uvula is absent. interlocking, is well developed. The outer face of the The outer face of the crown is also weakly convex. A root shows a central aperture near the crown-root broad, vertically long rounded apron covers the central junction, that is the origin of a vertically long central and distal parts of the root. The apron is not overhan­ depression. A smaller distal aperture is present also ging the root but forms one part with it. near the crown-root junction. A well developed mesial The inner face of the root is vertically long and very flat, depression is present as a result of interlocking. showing a central aperture flanked by one smaller mesial and one distal foramen near the crown-root Genus: Euprotomicroides HULLEY & PENRITH, 1966 junction. The distal depression, as a result of inter­ locking, is well developed. The type species of this monotypic genus is Euprotomi­ The outer face of the root shows a large central aperture croides zantedeschia HULLEY & PENRlTH, 1966. with two or three mesial and distal foramina along the crown-root junction. A well developed mesial depres­ Euprotomicroides zantedeschia HULLEY & PENRlTH, sion is present as a result of interlocking. 1966 Pl. 19, T. pl. 17. Genus: Euprotomicrus GILL, 1864

Euprotomicroides zantedeschia HULLEY & PENRlTH, The type species of this monotypic genus is Euprotomi­ 1966, Bull. mar. Sci., 16(2) : 222. crus bispinatus (QVOY & GAIMARD, 1824).

This species has a strong dignathic heterodonty and a Euprotomicrus bispinatus (QUOY & GAIMARD, 1824) weak gradient monognathic heterodonty. The root is Pl. 20, T. pl. 18. anaulacorhizid. The lower teeth are strongly inter­ locked. Both upper and lower teeth are plurirnillimetri­ Scymnus bispinatus Quoy & GAIMARD, 1924, Zoolo­ cal in range. gie, Voy. Uranie et Physicienne, 1817-20: 197. This species has a strong dignathic heterodonty and a Upper jaw weak gradient monognathic heterodonty. A weak The teeth have a very elongated, narrow principal cusp, sexual heterodonty is present in lower teeth by a slightly that is symmetrical on the anterior positions and sigmoidal mesial cutting edge of the principal cusp in sigmoidal on lateral and posterior positions. Cusplets males and an arched mesial cutting edge in females. are absent. The root is anaulacorhizid. The lower teeth are strongly The root is pseudobilobated by an arched root base, interlocked. Both upper and lower teeth are plurirnilli­ having rounded pseudo-lobes. The crown is more than metrical in range. five times higher than the width of its base. The inner face of the crown is very convex. Uvula is Upper jaw absent, as well as an inner ornamentation. The teeth have a very elongated, narrow principal cusp, The outer face of the crown is convex. Apron and outer that is constricted just above the crown-root junction. ornamentation are absent. Mesial and distal cutting edges therefore are sigmoidal. The inner face of the root shows a convex crown-root Cusplets are absent. The teeth become lower and junction, on which a large central aperture is present. broader based toward the commissure. Downward, the surface is flat, sometimes showing a The root is pseudobilobated by an arched root base, short, shallow basal sulcus with one or two foramina in having rounded pseudo-lobes. The lateral teeth have a it. shorter distal pseudo-lobe. The crown is over three The outer face of the root has a central aperture with times as high as the width of its base. two or three mesial and distal foramina along the The inner face of the crown is very convex. Uvula is crown-root junction. The central part between the absent. An inner ornamentation is present by fine, root's pseudo-lobes is depressed. short basal costules. ''

Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 125

Text plate 17. Euprotomicroides zantedeschia HULLEY & PENRITH, 1966. Male 41.6 em. t.l., S. W Atlantic. By courtesy of Dr. M. STEHMANN, I.S.H. Hamburg, Germany (x 5). The outer face of the crown is convex. The basal Lower jaw constriction has caused a mesial and a distal ridges. The The strongly compressed teeth are always interlocked, crown-root junction is strongly arched and the crown presenting deep, vertically long, inner distal and outer has formed an apron, which is divided into a mesial and mesial depressions. Distal and mesial blades are present a distal pseudolobe, that both are vertically long and on the symphysial tooth. The mesial blade is absent on narrow. the other teeth. The crown inclines slightly toward the The inner face of the root is flat with a slight central, commissure in latera-posterior and posterior teeth. The basal depression showing a large central aperture near principal cusp is over twice as high as its base width. the crown-root junction. One or two foramina are Mesial and distal cutting edges are sigmoidal. The root present on the root's pseudo-lobes. has an arched base with rounded basal ends. The outer face of the root has a strong central The inner face of the crown shows a slightly convex depression, in which a very large central aperture is principal cusp. Uvula is absent. present. Both pseudo-lobes have a central foramen The outer face of the crown is also weakly convex. The asides the apron. crown base is strongly arched; the apron is consequently I I

126 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

divided into a mesial and a distal pseudo-lobe. On The outer face of the root has a strong central lateral teeth the mesial apron is narrower than the distal depression, in which a very large central aperture is one. Some basal, vertical costules are present. present. Both pseudo-lobes may have a central foramen The inner face of the root is vertically long and very flat, aside the apron. showing a central aperture. A distal depression, as a result of interlocking, is well developed. Lower jaw The outer face of the root shows a very large central The strongly compressed teeth are always interlocked, aperture. A foramen is present also at the junction of presenting deep, vertically long, inner distal and outer the pseudo-lobes and the root. A well developed mesial mesial depressions. Distal and mesial blades are present depression is present as a restJlt of interlocking, in on the symphysial tooth. The mesial blade is absent on which one or two foramina are present along the the other teeth. The crown inclines slightly toward the crown-root junction. commissure in latera-posterior and posterior teeth. The principal cusp is over twice as high as its base width. Genus: Squaliolus SMITH & RADCLIFFE, 1912 Mesial and distal cutting edges are sigmoidal. The root has an arched root base with rounded basal ends. The type species of this genus is Squaliolus laticaudus The inner face of the crown shows a slightly convex SMITH & RADCLIFFE, 1912. Squa/iolus a/iae TENG, principal cusp. Uvula is absent. 1959 has recently been revaiidated from synonymy The outer face of the crown is also weakly convex. The with Squaliolus laticaudus by SASAKI & UYENO (1987). crown base is strongly arched, presenting an apron divided into a mesial and a distal pseudo-lobe. On Squaliolus laticaudus SMITH & RADCLIFFE, 1912 lateral teeth, the mesial pseudo-lobe is narrower than Pl. 21. the distal one. The inner face of the root is vertically long and very flat, Squaliolus laticaudus SMITH & RADCLIFFE, 1912, showing a large central aperture. A distal depression, as Proc. U. S. nat. Mus., 41 : 684. result of interlocking, is well developed. The outer face of the root shows a very large central This species has a strong dignathic heterodonty and a aperture. Another foramen is present at the junction of weak gradient monognathic heterodonty. A weak pseudo-lobes and root. A well developed mesial depres­ sexual heterodonty is present in lower teeth by a slightly sion is present as a result of interlocking, in which one sigmoidal mesial cutting edge of the principal crown in or two foramina are present along the crown-root males and a straight mesial cutting edge in females. The junction. root is anaulacorhizid. The lower teeth are strongly interlocked. Both upper and lower teeth are plurimilli­ Genus: Heteroscymnoides FOWLER, 1934 metrical in range. The type species of this monotypic genus is Heteroscym­ Upper jaw noides marleyi FOWLER, 1934. The teeth have a very elongated, narrow principal cusp, that is constricted just above the crown-root junction. Heteroscymnoides marleyi FOWLER, 1934 Mesial and distal cutting edges therefore are sigmoidal. Pl. 22. Cusplets are absent. The teeth become lower and Heteroscymnoides marleyi FOWLER, 1934, Proc. Acad. broader based toward the commissure. nat. Sci. Philad., 85 : 239. The root is pseudobilobated by an arched root base having rounded pseudo-lobes. The lateral teeth have a This species has a strong dignathic heterodonty and a shorter distal pseudo-lobe. The crown is over three weak gradient monognathic heterodonty. The mesial times as high as its base width. cutting edge of the lower teeth is almost straight, The inner face of the crown is very convex. Uvula is reaching beyond the crown base and joining with it in a absent. An inner ornamentation is present by fine, short rounding. short basal costules. The root is anaulacorhizid. The lower teeth are strongly The outer face of the crown is convex. The basal interlocked. Both upper and lower teeth are plurimilli­ constriction has caused a mesial and distal ridge. The metrical in range. crown-root junction is strongly arched and the crown has formed an apron, which is divided into a mesial and Upper jaw distal pseudo-lobe, that both are vertically long and The teeth have a very elongated, narrow principal cusp, narrow. that is constricted just above the crown-root junction. The inner face of the root is flat, with a slight central, Mesial and distal cutting edges therefore are sigmoidal. basal depression, showing a large central aperture near Cusplets are absent. The teeth become lower and the crown-root junction. On the pseudo-lobes one or broader based toward the commissure. two foramina are present. The root is pseudobilobated by an arched root base, l ''

Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 127 having rounded pseudo-lobes. The lateral teeth have a locked. The upper teeth are subcentimetrical and the shorter distal pseudo-lobe. The crown is over three lower ones centimetrical in range. times as high as its base width. The inner face of the crown is very convex. Uvula is Upper jaw absent. An inner ornamentation is present by fme, The teeth have an elongated principal cusp, that is short basal costules. broad based and constricted just above the crown base. The outer face of the crown is convex. The basal Cusplets are absent. The principal cusp is always constriction has caused a mesial and a distal ridge. The slightly directed toward the commissure, still more so in crown-root junction is strongly arched, and the crown lateral and posterior teeth. The root is pseudobilobated has formed a mesial and a distal apron, that both are by a deep sulcus, having slightly rounded pseudo-lobes. vertically long and narrow. The crown height is twice or more the crown base The inner face of the root is flat with a slight central, width. basal depression, showing a large central aperture near The inner face of the crown is slightly convex. Uvula is the crown-root junction. One or two foramina are absent. Also an inner ornamentation is absent. present on the pseudo-lobes. The outer face of the crown is slightly convex. Apron The outer face of the root has a strong central and outer ornamentation are absent. depression, in which a very large central aperture is The inner face of the root shows a low longitudinal present. Both pseudo-lobes can have a central forame­ ridge near the crown-root junction, on which a large na side the apron. central aperture with a mesial and a distal foramen is present. A short, broad and deep basal sulcus originates Lower jaw just below the central aperture, in which a second The strongly compressed teeth are always interlocked, central foramen is present. presenting deep, vertically long, inner distal and outer The short outer face of the root sometimes has one or mesial depressions. Distal and mesial blades are present two foramina scattered below the crown-root junction. on the symphysial tooth. The mesial blade is absent on The central basal sulcus is also present on the outer the other teeth. The crown inclines slightly toward the face. commissure at latera-posterior and posterior teeth. The principal cusp is over twice as high as its base width. Lower jaw Mesial and distal cutting edges are sigmoidal. The root The str ngly compressed teeth are always interlocked, has an arched base with rounded basal ends. presenting deep, vertically long, inner distal and outer The inner face of the crown shows a slightly convex mesial depressions. Small distal and mesial blades are principal cusp. Uvula is absent. present at the symphysial tooth. The lateral teeth have a The outer face of the crown is also weakly convex. The small distal blade only, that becomes more distinct on crown base is strongly arched, presenting an apron posterior teeth. The crown inclines slightly toward the which is divided into a mesial and a distal pseudo-lobe. commissure in lateral and posterior teeth. On lateral teeth the mesial pseudo-lobe is narrower Mesial and distal cutting edges and distal blade are than the distal one. serrated. This fine serration is very regular, reaching The inner face of the root is vertically long and very flat, almost the apex of the principal cusp and half way on showing a central aperture. A distal depression, as a the cutting edge of the distal blade. result of interlocking, is well developed. The rectangular root is vertically long and pseudo­ The outer face of the root shows a very large central bilobated with rounded pseudo-lobes. The root becomes aperture. Another foramen is present at the junction of more oblique in lateral and posterior teeth. pseudo-lobes. A well developed mesial depression is The inner face of the crown is flat to very weakly present as a result of interlocking, in which one or two convex and as high as the crown base width. Uvula is foramina are present along the crown-root junction. absent. Genus: Dalatias RAFINESQUE, 1810 The outer face of the crown is also weakly convex. Vertically long mesial and distal aprons are present. The type species of this monotypic genus is Dalatias The aprons are not overhanging the root but form one licha (BONNATERRE, 1788). part with it. Ornamentation is absent. The inner face of the root is vertically long and very flat, Dalatias licha (BONNATERRE, 1788) showing a central aperture and a mesial and a distal Pl. 23, T. pl. 19. foramen. Just below the central aperture, a vertically Squalus licha BONNATERRE, 1788, Tab!. encyclop. long, broad, deep sulcus divides the root into two method. trois reg. nat., Ichthyol., Paris : 12. pseudo-lobes. The outer face of the root shows the sulcus, running This species has a strong dignathic heterodonty and a from the crown-root junction between both pseudo­ weak gradient monognathic heterodonty. The root is lobes of the apron to the basal end. Foramina are anaulacorhizid. The lower teeth are strongly inter- absent. I I

128 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

Text plate 18: Euprotomicrus bispinatus (Quoy & G AIMARD, 1824). Female 22 em. t. l., Indian Ocean. Upper jaw x 10 ; lower jaw x 10.

Text p late 19. Dalatias lich a(BONNATERRE, 1788). Female84 em. t.l. , W. Hebrides, N.E. Atlantic. Upper jaw x4; Lowerjaw: Diagraph of the functional file. (x 5). I I

Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 129

Both pseudo-lobes have irregularly scattered depres­ 3. - Lower teeth : root narrowing, apron flanked by two large SIOns. apertures, short but broad uvula and a long vertical sulcus - Upper teeth : numerous foramina scattered around the apron, sulcus very long and broad (text fig. 1, fig. 3) Differential diagnosis Somniosus (microcephalus-form) Squaliform and hexanchiform teeth are derived from Somniosus the anaulacorhizid root type. Their compressed and (rostratus-form) vertical expanded shape is the result of a long evolution, which has led to several forms of vascularisation and 4. - Lower teeth: apron broadly rounded, surrounded by several foramina, root rectangular and equally broad developed the tooth types as displayed in Hexanchiform - Upper teeth: uvula absent, short sulcus (text fig. I, fig. 4) and Squaliform species. · Oxynotus The following remarkable features were noted during our examinations : 5. - Lower teeth: apron flanked by several horizontally expanded -Both species of Echinorhinus show an hexanchiform foramina that are arranged in a vertical row, respectively, or numerous small foramina randomly arranged around the root type of jaw teeth, and their odontology does not apron give any evidence of a relationship with Squaliformes. - Upper teeth : entire tooth well developed and elongated with -There is hardly any morphological difference in teeth outer basal ridges on the principal cusp, root pseudobiloba­ of the genera Squalus and Cirrhigaleus, nor is there any ted (text fig. I, fig. 5) evidence in their external morphology to maintain two . Scymnodon Scymnodalatias separate genera. Centroscymnus -A similar situation exists for Squaliolus, Euprotomi­ (coelolepis-form) crus and Heteroscymnoides, all with one or two species l:uproto1nicroides only. Despite their strong dignathic heterodonty and Centroscymnus sexual heterondonty, there is not any morphological (crepidater-form) evidence to maintain three seperate genera. 6. - Lower teeth : apron divided into a mesial and a distal part, - The genera Scymnodon and Scymnodalatias are uvula absent odontologically very similar. We examined one species - Upper teeth : crown elongated and outer root centre of the genus Scymnodalatias, but the minor differences depressed with either one large, or up to three smaller in tooth morphology are covered by the natural foramina (text fig. I, fig. 6) variability within the genus Scymnodon. . Dalatias lsistius - The morphological differences between teeth of Squaliolus Somniosus microcephalus and Somniosus rostratus l:uprotomicrus are that extreme, that each could represent a separate . Heteroscymnoides supraspecific taxon. Upper teeth C. coelolepis C. crepidater - Lower teeth : extremely broad apron covering the central root pseudobilobated root rectangular and distal root parts sulcus present sulcus absent mesial and distal foramina central foramen only 7. - Upper teeth : branched root and multicuspid crown (text fig. Lower teeth I, fig. 7) apron long apron short and broad l:tmopterus foramina horizontally foramina surrounding expanded and situated aside the apron 8. - Lower and upper teeth almost homodont : elongated crown, the apron unicuspid (text fig. I, fig. 8) sulcus present sulcus absent . Aculeola root ends rounded root ends angled The degree of vacularisation in squaliform tooth roots has led to several typical tooth types. (see text-plate 1). Conclusions These tooth types will be described below and the genera grouped accordingly into 8 subfamilies: Despite its relative dignathic homodonty, the strong 1. - Lower and upper teeth homodont : crown low, apron and ontogenetic heterodonty, the absence of an anal fm and uvula well developed and root low (text fig. I, fig. I) 5 pairs of gill slits only, all in contrast to the hexanchi­ . Squalus . Cirrhigaleus form genera, the genus Echinorhinus based on odonto­ logical features does not have a relationship to squali­ 2. - Lower teeth : similar to type I but higher root ; uvula less forms but rather to hexanchiforms. However, the developed and sulcus longer differences to both orders are otherwise extreme, and - Upper teeth : principal cusp erect, uvula absent, root therefor we rather tend to follow the palaeontologists rectangular and apron fl anked by two foramina (text fig. I, fig. 2) PFEIL ( 1983) and CAP PElT A ( 1987) in their proposal to . Deania place Echinorhinidae in a separate order: Echinorhini­ . Centrophorus formes . 130 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

Odontologically the genus Oxynotus has the typical in general show odontological similarities with the characteristics of Squalidae, and thus no additional family Scyliorhinidae. These will be discussed in the arguments are offered to support some authors in forthcoming contributions on scylicirhinid odontology. holding a separate familie Oxynotidae. Key to the squalid genera based on odontological Odontologically no differences exist between Cirrhiga­ characters : leus and Squalus. 1. Principal cusp blade-like and broad based : Only minor odontological differences exist between a- Uvula present: Euprotomicrus, Squaliolus and Heteroscymnoides, Sulcus absent which thus might be congeneric. - Apron and uvula cover the whole upper part of the root Etmopterus Subdivision of the genus Somniosus into two subgenera (lower teeth) (Pl. 5). has often been proposed by authors. The present Sulcus short odontological result supports this view as expressed by - Apron narrow and short QUERO ( 1976) and COMPAGNO ( 1984: 102), to group S. Uvula small microcephalus and S. pacificus into one subgenus Crown strongly directed distally Somniosus (Somniosus) and S. rostratus into another . Squalus (upper and lower teeth) subgenus Somniosus (Rhinoscymnus). (Pl. 3). - Apron broad The species of Centroscymnus share a characteristic Uvula broad external morphology. However Centroscymnus crepi­ Crown erect, with serrated cutting edges dater differs so markedly in its tooth morphology from Oxynotus (lower teeth) its congeners, that GARMAN's (1913) introduction of a (Pl. 2). separate genus Centroselachus for this species appears Sulcus long justified. Crown strongly directed distally - Uvula very small Although Centroscymnus and Scymnodon have the Root always broader than high . Deania same type of root vascularisation, all their odontologi­ (lower teeth) cal characters differ that extreme that generic distinction (Pl. 13). should remain. - Uvula narrow and small Width and height of root base about equal However the distinction of Scymno_don and Scymno­ . Centrophorus (lower teeth) dalatias by odontology is of minor importance only (Pl. II). and therefore both may be congeneric. - Uvula broad and short Root base narrower than the height of the root Lepidorhinus squamosus has been reassigned to the a-Apron short genus Centrophorus by COMPAGNO (1984). This deci­ Somniosus (Somniosus) sion is supported by odontological characters as (lower teeth) illustrated for C. squamosus in the present paper (Pl. (Pl. 8, 9). 12, Tpl. 10). b-Apron long Centroscymnus The degree of development of vascularisation in tooth (lower teeth) (Pl. 14). roots is a significant indication of phylogenetic relation­ A pair of large foramina flanking the apron ships and whould allow to group the squalid genera in Somniosus subfamilies. COMPAGNO (1973) had subdivided the (Rhinoscymnus) Squalidae, as then understood by him, into four (lower teeth) subfamilies but withdrew this concept (1984 : 29) (Pl. 10). Several horizontally expanded foramina flanking the apron pending his further completion of work on squaloid Scymnodon morphology, and he considered separate families for (lower teeth) Echinorhinus and Oxynotus. Though based on other (Pl. 16). characters, CoMPAGNo's (1973) concept of squalid subfamilies is generally in accordance with the present b-Uvula absent odontological results to group squalid genera into 8 Short apron flanked by two main foramina subfamilies as indicated in the diffential diagnosis - Root high, strongly narrowing from just below the crown-root junction downward above. . Centrophorus (upper teeth) Remark : Centroscyllium fabriccii and Aculeola nigra (Pl. II). I I

Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 131

- Root low, almost equally wide from just below the crown-root - Cusplets present junction downward Etmopterus (upper teeth) . Deania (pl. 5). (upper teeth) - Cusplets absent (Pl. 13). - Outer crown base slightly depressed Long apron surrounded by foramina Crown strongly constricted at base - Crown height twice the crown base width. Root long Root mesio-distally wide Slightly rounded root lobes Euprotomicroides . Dalatias (lower teeth) (upper teeth) (Pl. 19). (Pl. 23). - Crown height equals the crown base width - Outer crown base not depressed a-Root low Crown not constricted Centroselachus Root mesio-distally narrow, with quadrangular root lobes /sistius (lower teeth) (upper teeth) (Pl. 15). (Pl. 18). b-Root very high - Outer crown base strongly depressed, with a large central Oxynotus foramen (upper teeth) Root mesio-distally narrow with rounded root ends Euprotomicrus (Pl. 2). (upper teeth) Long and broad apron, divided into a mesial and a distal part (Pl. 20). - Sulcus long apron dividing from crown-root junction downward Basal ridge on the outer part of the principal cusp present Cutting edges serrated - Root bilobate One broad, outer mesial and another distal foramen present . Dalatias Centroscymnus (lower teeth) (upper teeth) (Pl. 23). (Pl. 14). - Sulcus short - Root more or less quadrangular Crown triangularly shapes a-Sulcus present Apron divided at apron base only Outer mesial and distal foramina absent (some outer scattered foramina present on posterior teeth) lsistius Scymnodon (lower teeth) (upper teeth) (Pl. 18). (Pl. 16). - Sulcus absent b-Sulcus absent Three to four outer central foramina present Apron divided in lower half only Root shape quadrangular Euprotomicrus Centroselachus (lower teeth) (upper teeth) (Pl. 20) (Pl. 15). Apron prominent and long Basal ridge on the outer part of the principal cusp present Somniosus - Root bilobate (Somniosus) One broad, outer mesial and another distal foramen present Centroscymnus (upper teeth) (upper teeth) (Pl. 8, 9) (Pl. 14). Apron absent - Root more or less quadrangular Fine reticulated ornamentation present on the outer crown base a- Sulcus present Somniosus Outer mesial and distal foramina absent (some outer scattered foramina present on posterior teeth) ( Rhinoscymnus) Scymnodon (upper teeth) (upper teeth) (Pl. 10) (Pl. 16). b-Sulcus absent Three to four outer central foramina present 2. Principal cusp vertically long and slender : Root shape quadrangular Root hemiaulacorhizid Centroselachus - Cusplets present (upper teeth) Centroscyllium (Pl. 15). (upper and lower teeth) Basal ridge on the outer part of the principal cusp absent (Pl. 6). - One or two outer central foramina and two or three mesial and - Cusplets absent distal foramina present . Acu/eo/a Root more or less bilobate (upper and lower teeth) Euprotomicroides (Pl. 7). (upper teeth) Root anaulacorhizid (Pl. 19) 132 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

Acknowledgements Zealand, Wellington, for the permission to examine specimens at their disposal. Dr. M. STEH MANN provi­ We whould like to thank Dr. J.L.V. COMPAGNO, ded specimens of I.S.H., Hamburg. formerly San Francisco State University, California : We also whould like to thank Captain P. G UEGUEN for Dr. J .P . GOSSE, Institut Royal des Sciences naturelles allowing us to collect specimens during his voyages. de Belgique, Brussels ;·G.H. BURGESS, Florida State We would also like to thank Miss F. LADEUZE, Museum, Mrs. S.L. JEWETT, United States National F.N.R.S., Brussels, for the careful correction of the Museum of Natural History, New York; Dr. W.N. galleys. EscHMEYER, California Academy of Sciences, San The SEM-photographs were taken by J. C!LLIS, Institut Francisco ; Dr. M. BoESEMAN, Rijksmuseum van Royal des Sciences naturelles de Belgique, Brussels and Natuurlijke Historie, Leiden ; Dr. M.L. BAUCHOT, printed by H. STOUT, Brussels. The macro-photographs Musee National d'Histoire naturelle, Paris; B. SERET, as well as the diagraphs were realized by our friends G. formerly 0RSTOM, Dakar ; A. WHEELER, British BROGNIET, J . DEGREEF and M. VALLE, Brussels. Museum of Natural History, London ; Dr. G.S. Exceptions are mentioned in the legends of the plates. HARDY and A. STEWART, National Museum of New

Bibliography DOLGANOV, V.N., 1984, A new shark from the family Squalidae caught on theNaska submarine Ridge. Zoological BASS, A.J., D'AUBREY, J.D. and KISTNASAMY, N., 1976, Journal. 63(10) : 1589-1591. Sharks of the east coast of southern Africa. 6. The families GARMAN, S., 1913, The Plagiostomia. (Sharks, skates and Oxynotidae, Squalidae, Dalatiidae and Echinorhinidae. rays). Memoir Museum comparative Zoology, Harvard. 36. Investigational Report Oceanographic Research Institute, 515 pp. Durban, (45) : 103 pp. LEDOUX, J.-C., 1970, Les dents des Squalides de Ia Medi­ BIGELOW, H.B. and SCHROEDER, W.C., 1957, A study of tefi·anee occidentale et de !'Atlantique Nord-Ouest africain. the sharks of the suborder Squaloidea. Bulletin Museum Vie et Milieu. Ser. A. Biologie marine. 21(2a) : 309-362. comparative Zoology, Harvard, 117 (1) : 150 pp. LEDOUX, J.C., 1972, Les Squalidae (Euselachii) miocenes CAPPETTA, H., 1987, Chondrichtyes 2. Mesozoic and des environs d'Avignon (Vaucluse). Documents laboratoire cenozoic elasmobranchii. Handbook of Paleoichthyology Geologie Lyon: 52: 133-175. (ed. H.P. Schultze). Vol. 38 : 193 pp. PFEIL, F. H., 1983, Zahnmorphologische Untersuchungen in COMPAGNO, L.J.V., 1973, Interrelationships of living elas­ rezenten und fossilen Haien der Ordnungen Chlamydose­ mobranchs. In : Interrelationships of fishes. (ed. P.H. lachiformes und Echinorhiniformes. Palaeoichthyologica. 1 Greenwood, R.S. Miles and C. Patterson). Journal Linnean : 1-315. Society (Zoology), 53, suppl. 1, 15-61. QUERO, J.-C., 1976, Somniosus bauchotae sp. nov. (Selachii, COMPAGNO, L.J.V., 1977, Phyletic relationships of living Squalidae, Scymnorhininae) espece nouvelle de !'Atlantique sharks and rays. American Zoologist, 17(2): 302-322. N.E. Revue Travail Institut Peches Maritimes, Nantes. COMPAGNO, L.J.V., 1984, FAO species catalogue. Vol. 4. 39(4) : 455-469. Sharks of the world. An annotated and illustrated catalogue T ANIUCHI, T. and GARRICK, J.A.F., 1986, A new species of of shark species known to date. Part 1. Hexanchiformes to Scymnodalatias from the Southern Oceans, and comments Lamniformes. FAO Fisheries Synopsis. (125). Vol. 4. Part 1: on the other squaliform sharks. Japanese Journal ofIcht hyo­ 24-129. Part 2: 564. logy. Vol. 33(2): 119-134.

Glossary (also applying to previous issues of this series) Holaulacorhizid Vascularisation through many small foramina An-, Hemi-, Hoi- and Poly- aulacorhizid concentrated in a median groove running from Considering their vascularisation, E. Casier (1947) outer to inner face. recognised and described 4 phylogenetically charac­ (like Carcharhinidae, Lamnidae, R ajidae etc.) teristic root forms of elasmobranch teeth : Polyaulacorhizid Anaulacorhizid Vascularisation through many small foramina V ascularisation through scattered foramina of concentrated in several grooves running parallel equal size on both outer and inner faces. from outer to inner face and near crown-root (like Hexanchidae) junction on both inner and outer faces. (like Hemiaulacorhizid Myliobatidae, etc.) Vascularisation through a median and 1 or 2 Anterior lateral foramina on inner face. Tooth positions close to junction of left and 1ight (like Squatinidae and Orectolobidae) jaw part. Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 133

Apron Posterior Expansion of the central part of the outer crown Tooth positons toward end of jaw. base. Row Basal Tooth row from inner face to outer face of jaw Bottom face concerned. (transversally). Commissural Striae Tooth positions near the end of jaw. Vertical ridges running from crown base toward Costule apex. Short, vertical ridges sometimes present on inner Sulcus and/ or outer crown base. Groove developed by the main vascularisation Dignathic string leading from root base to the main foramen Heterodont by having different tooth morphology on analacorhizid root types. · in upper and lower jaw. It differs from the median groove of the holaulacor­ Distal hizid root type and the parallel grooves of the Tooth row from symphysis toward end of jaw. polyaulacorhizid root type, in which several forami­ (longitudinally) na are concentrated, respectively. Heterodonty Symphysial Different tooth morphology within a tooth file. Teeth at junction of both halfs of a jaw. Homodonty Uvula Equal tooth morphology within a tooth file. Lobate extension of the inner crown base. Inner face View from inside the mouth. Lateral STEHMANN M., Tooth positions half way along the jaw. Aussenstelle Ichthyologie Median keel des Instituts fiir Seefischerei, Transversal ridge dividing crown into inner and cf o Zoologisches Museum outer face. der UniversiHit Hamburg, Mesial Martin-Luther-King-Platz 3, Toward junction (symphysis) of left and right jaw D-2000 Hamburg 13, parts. Fed. Repl. of Germany. Outer face View from outside the mouth. Panisymphysial HERMAN J., First anterior tooth row if a symphysial tooth row Service Geologique is absent de Belgique, Pseudoapron Rue Jenner 13, Apron-like vertical ridges that appear sometimes B-1040 Brussels, Belgium. on lateral and posterior teeth Pseudosymphysial HOVESTADT-EULER M. One of the parasymphysial tooth rows becomes and HOVESTADT D.C., the character of a symphisial tooth row (sym­ Merwedelaan 6, metry). This phenomenon is sometimes present in NL-4535ET Terneuzen, spectes. The Netherlands. I I 134 J. HERMAN, M. HOVESTAD-EULER and D.C. HOVESTADT

Compositions of the plates

As far as possible, for all the genera and subgenera, one plate of the complete tooth file of upper and lower jaw and another plate with SEM-photographs or macro­ photographs of isolated teeth are presented. Both plates have the same composition : the upper part shows always the upper jaw or teeth and the lower part the lower jaw or teeth. The outer and inner view of the isolated teeth (macro­ or SEM-photographs) always. are symmetrically pre­ sented. Only the outer view of the jaw halfs or tooth files is presented, because it is our intention to show the number of tooth rows, their dispositions and the variations in form and size of the teeth. There are two techniques of photographing used : diagraphs (the tooth file used as negativ) and composition of photo­ graphs that partly overlap every previous one tot form one complete jaw half. The choice ofleft or right jaw half depends of quality of preservation of the specimens's tooth files only. The upper scale bar refers to the upper teeth, the lower scale bar to the lower teeth. Photographs of posterior and commissural teeth mostly are enlarged and have their own scale bar.

Legend

s = Symphysisial position ps = Parasymphysial position a = Anterior position al = Antero-lateral position 1 = Lateral position lp = Latera-posterior position p = Posterior position c = Commissural position Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 135

(.) c.

-.] ~ E u

0

-..: ~ ~ 0 s::: -<:) ~ -s:: c; s --i:: ~ ~ .... <:u ~ Cll ::::5 <:i ~ ~ ~ E' 1o ~ -..:'- <.)E ~ ~ ~ ~ ~ ;:;;} :~J -.~ ~

!- :I: tTl ;;o 3:: _>z

~ :I: 0 < tTl (/)-l > 0 c:m r tTl "' "'c.;::> 0 h :r: 0 < tTl (/)-l > -l0

1 t mm l p ~ a

Plate 2. Oxynotus centrina (L !NNAEUS, 1758). Female 46 em. t.l., Dakar, Senegal. (only the lower commissural tooth refers to the large scale). Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 137

en ] c.

-~ s:: -Sl ~ kl ~ ~- ~ kl .....- ~ <::>" -..<::> ~ <::! t:: ~ a;;- .....,~ :::sv) "l ~ 2:; t:i ~ :.E '§ '-' Ol ::l"' t;l ::l cr (/J rv-i ~ -Sl t:l..., w 00-

~ ::r: tTl

~ .:z

~ ::r: 0 < tTl C/J ;! 0 r, c: r tTl

:l"' 0."' 0 0 ::r: 0 < tTl C/J ;! -l0

0 mm

Plate 4. Squalus barbifer (TANAKA, 1912). Female 82 em. t.l., Sagami Bay, Japan (latera/teeth). Formerly Cirrhigaleus barbifer (TANAKA, 1912). Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 139

(.)

(/) (/) a. a.

<:::i ~ s:: %1 s:: ....~ ~ ~ .,; ~ -'

E<.) 0,

~"" <::s !::: ~ ~ -.~ Vi ;::) Lq ~ ~ ~ ~ .!3 0.. "' "'::l.... !l 0.. s0 I.Il .,..; ~ ..!:! Q..; ,__ ~ 0

~ ::r: tTl :>:' :::: )> .z ;s: ::r: 0 < tTl ps ., ., \. I '1i..,C£.&>ali I I~ )> r,CJ c: r tTl :>:' ""'c. CJ 0 ::r: 0 < tTl (/J ...., )> ....,CJ

ps c

1 l ,;m I

Plate 6. Centroscyllium fabricii (REI NHARDT, 1825). Female 85 em. t.l., N. Bill Bailey, N.E. Atlantic. Upper commissural tooth is 3 times more enlarged than the other teeth. Lower lateral tooth has an abnormal root structure. ps (') 0 ::I ......

0 mm ~ :::1. 1...... ! 0" t:: ...... o· ::I en ...... 0 ...... ::r (1> en ,-< I 12 0... '<: ...... ,0 ...... lr (') ·.\ s0

.>~ I 1'§ ~ ~ ( _ <' "' (1> *l '~, s . 'I 0 ps '"'1 '1:S ::r 0

·~ 0 ~ ...... ,0 ...... (1> (1> ...... ::r .., ::I 0... 0 ...... ::r (1> '"'1 '"'1 (1> ~ ..,< ::I ...... ()' ::r...... ~ '<: liiDI• 0 0... 0

~ (1> en

Plate 7. Aculeola nigra DEBUEN1 1959. Upper photo series: male 44.5.cm. t.l., Peru. Central photo series: male 40 em t.l., Chile. (Details of the micro-ornamentation of the commisural teeth x 90). Lower photo series :female 38 em. t.l., Chile. A ll these SEM-ph otos by courtesy of Mr. F CIGALA-FULGOSI, Parma, italy. -+>-- -.j:::.. IN a ,_ I ~ ::r: tT1

~ "';J> _z

?= ::r: 0 < tT1 (/) ~ 0 r, l'c: tT1

"'P> 0."' 0 0 ::r: 0 < I'T1 (/) ~ ....,0

\ c Ia

Plate 8. Somniosus microcephalus (BLOCH & SCHNEIDER, 1801). Female 470 em. t.l., N. Lousy N.E. Atlantic. Upper teeth x 35; lower anterio-lateraltooth x 2.5; lower commissural tooth x 7. Macro-photographs were taken by K. Jansen, Oudenbosch, The Netherlands. New taxonomic proposition: Somniosus (Somniosus) microcephalus (BLOCH& SCHNEIDER, 1801). (J 0 ,....::::1 !:1. 0" ~ c;·...... ::::1 VJ ,.... 0 ,.... ::r (1> VJ ...... 15. ps ~ I '< ...... ,0 ,.... ::r (1> (") 0 "0s ~ ~ Pl,.... :;::· (1> s 0

~ . .a . -- ·" ;::> .. 0 0 ps '• t ~ I (. '/ 0...... , ,.... ' . ' (1> ,....(1> ;::>

~ 0.. . '· ~~ ,....0 ' J ~ ~ ;::> ' ' (1> ·f ..,

(1> (0 < Pl ,....::::1 ()" ;::>,.... '< 0 0.. 0

,....~ (1> VJ Plate 9. Somniosus microcephalus (BLOCH & SCHNEIDER, 1801). Central part : male 176 em. t.l. , E. Oreille, N.E. Atlantic. Outer parts: female 470 em. t.l. , N. Lousy, N.E. Atlantic. New taxonomic proposition : Somniosus (Somniosus) microcephalus (BLOCH & SCHNEIDER, 1801). -w~ -t

~ ::I: tT1 I "' \ .z~ I $:: ps ::I: 0 < tT1 Cll

-~- s; C) ~- 0 mm ! r, c ·-·-- r tT1

"'!5 c. C) 0 ::r: 0 < a Bl-l )> -l0 ~

ps a

Plate 10. Somniosus rostratus (Risso, 1826). Male 109 em. t.l., W. Hebrides, N.E. A tlantic. The micro-ornamentation of the upper parasymphysialtooth x 90 and x 650, respectively. New taxomic proposition: Somniosus (Rhinoscymnus) rostratus (RISSO, 1826). 'I

Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 145

<:i ~ s::: <::l "'s::: ~ II) ] ~ c. ~ -..:·;;:: ~

-'...; E '-' !'-. 0\ ~ <::l ;::: ~ -.--.:.. <:::> -.00 ~ Q kl :<: tl ~ ~ tl a --1 ~

:­ ;r: tTl ::0 s:: _z> s:: ;r: 0 < tTl Ul ;! r,0 c:: r tTl ::0

="'0. 0 0 ;r: 0 < tTl Ul-l > -l0

a

Plate 12. Centrophorus squamosus (B ONNATERRE, 1788). Male 92 em. t.l., S. W. Rockall. N.E. Atlantic. Formerly Lepidorhinus squamosus (BONNATERRE, 1788). I I

Contributions to the study of the comparative morphology of teeth and other relevant ichtyodorulites 147

-.

I I j I .:,} ~ ~ ~ ki <: '-i' ~·;::: ~ ~ :s: -I ~ ~ '-' t-...... 0 ~ 1:3 12 ~ ...... ~ ...... 00 ::14 a t:i ro

,....

::I: ps '\ A I I tTl - ::0 ~ >- _z

~ ::I: 0 < tTl C/l -1 >- 0 rr, c: r tTl ::0 "' "'0.. 0 0 ::I: 0 < tTl (/) -1 >- 0 -1

c

a 0 rnm : ~ ps c & uppe r -----lcelh

Plate 14. Centroscymnus coelolepis BOCAGE & CA PELLO, 1864. Female 110 em. t.l., N. Bill Bailey, N.E. Atlantic. Both upper parasymphysial tooth and anterior tooth are from a female 102 em . t.l., Nice Mediterranean. (') a0 ::l . ccr' c;·...... en:::3 ...... 0 ...... ::r (1) en c...... 0.. '< 0-., ...... ::r (1) () 0 '0s e; ~ ......

~· s 0 .a ::r 0 0 ~ -.,0 ...... (1) (1) ...... ::r § 0.. 0 ...... ::r (1) '"1 (a ro­ < § ...... ps 0 " ::r ...... 0 mm 1 '< lo •"" • .I 0 0.. 0

...... ~ (1) en Plate 15. Centroscymnus crepidater (BOCAGE & CAPELLO, 1864). Female 84 em. t.l., Porcupine, N.E. Atlantic. Both upper parasymphysial and anterior teeth are from a male 61 em. t.l., N. Bill Bailey, N.E. Atlantic. New taxonomic proposition : Centroselachus crepidater(BOCAGE -~ & CAPELLO,1864) such as proposed by GARMAN(1913). \0 Vl p -0

~ :c Ia tTl ::0 ~ )> .z ps ~ :c 0 lp < tTl {/) ;; 0 1 t w ,:,mw w I m c::: r tTl ::0 "' 0.." 0 0 :c 0 < tTl {/) ;; ....,0

ps

0~ 1 a

Plate 16. Scymnodon ringens SO CAGE & CAPELLO, 1864. Female 102 em. t.l., W. Hebrides, N.E. Atlantic. (Upper teeth). Lower teeth : Male 61 em. t.l., N. Bill Bailey, N.E. At lantic. n 0 ps a ::s s. 0" ..-+c c;· ::s U) ..-+ 0 ..-+ ;::r' ~ U) ..-+c 0... ~ '< 0...... , ..-+ ;::r' ~ () s0 '"d ~ ~ ~· s~ 0 .a ;::r' 0 0 ~ ...... ,0 ..-+ ~ ..-+~ ;::r' § 0... s g0 >-1 w(il g (=i' ;::r' ..-+ '<: bfuJ 0 a 0... 0 >-1 §.: c:; U)

Plate 17. Scymnodalatias albicauda T AN IUCHI & GARRI CK, 1986. Sex unknown 97.2 em. t.l., Hokitika Canyon, New Zealand. New Vl taxonomic proposition: Scymnodon albicauda (TA NIUCHI & G A RRICK, 1986). - LJl I - N L w .!Amw - I

~ :I: tTl ::0 ps ~ )> .z ~ :I: 0 < tTl Vl-l )> 0 r, c:: r tTl ::0 c."';:l 0 0 :I: 0 < tT1 Vl-l )> -l0

s

~ a p c

Plate 18. Isistius brasiliensis (QuaY & GAIMARD, 1824). Female 41 em . t.l., Dakar, Senegal. The parasymphysialtoothfrom a female 24 em. t.l., Hawai Island (2 x more enlarged). () 0 ::l s. 0" ps ~..... c;· ::l Vl ..... 0 ::;..... n Vl ..... ~ 0.. '<

...... ,0 1 ..... t w ..:m. • 1 ::; n () I 0 "0s e; ~ ::::· n s 0 .a ::; 0 0 ~

...... ,0 (t n ::;.....

~ 0.. 0 ::;...... ,n .....n ~ 5 <

.....~ (=i' 1 ::; 0 ;.,m - I d,l !w - ~ 0 0.. 0 ~ .....n Vl Plate 19. Euprotomicroides zantedeschia H ULLEY & PENRITH, 1966. Female (Holotyp e) 17.6 ern. t.l., S. Africa: lower symphysial and upper teeth. (except the lateral one). The lower anterior and lateral teeth, as well as upper lateral are from a male 41.6 ern. t.l., S. W. Atlantic. Vl (IS. H.). -w Ul -~ ( /{\:.:

~'ft ' ::-.: ps ,~,\ a §,~' ...', ~ ~ if.~ ~ ~ ~ 1a a "' · I I I~ ::r: tT1 "'s:: .z~ ~ ::r: 0 < tT1 Ul-l ~ 0m c: r tT1 ., "'::J Q. 0 0 ::r: 0 .._. < DK1 1'/Jiffj/ I I~ ~ -l0

01...-""""'--1 mm 1

s

·-­--..~-

Plate 20. Euprotomicrus bispinatus (Q UO Y & GAIMARD, 1824). Male 20 em. t.l., Mauritius Island. Centre of the plate: Female 11.5 em. t.l., near Ascension Island, Central Atlantic: n 0 :::3 s. o" ...... c c;·

131 ...... 0 ...... ::::> ('1) rn c...... 0.. '< .....,0 ...... ::::> ('1) 0 0 "0s e;

!=?.

<"('1) s 0 .a ::::> 0 0 ~ .....,0 ...... ('1) ('1) ...... ::::> g; 0.. 0 ......

...,1r wCti g; ...... cs· ::::> '<...... 0 0.. 0

...... ~ ('1) rn

Plate 21. Squaliolus laticaudus (SMITH & RAD CLIFFE, 1912). Female 22.7 em. t.l., N.E. Atlantic. By courtesy of Dr. M . Stehmann l.S.H. ,__ Vl Hamburg, Germany. New taxonomic proposition: Euprotornicrus laticaudus (SMITH & RAD CLIFFE, 1912). Vl 0\-Vl

a I I~ ::r: m :;<:! ~ ;J> _z

~ ::r: 0< (llm -l ;J> 0 r, c 0 mm 1 r 1ww __, I m :;<:!

"'0. 0 0 ::r: 0 < (llm -l ;J> -l0

r!l • ps

Plate 22. Heteroscymnoides marleyi (FOWLER, 1934). Female 28.4 em . t.l., S.E. Atlantic. By courtesy of Dr. M. Stehmann J.S.H . Hamburg, Germany. New taxonomic proposition : Euprotomicrus marleyi (FOWLER, 1934). () 0 ::l ...... :::::!. ps 0"' ...... c c;·

~ ...... 0 ...... ::r (1) en ...... c 0 mm 1 0. lo::.::I::JDd '< ...... ,0 ...... ::r (1) (") a0 '"0 ....,!:~) ~ <:• (1) a 0 .a ::r 0 0 ~ ...... ,0 ...... (1) (1) ...... ::r § 0. s 0 ...... ::r (1) ...., @ c:r < § ...... (=i" ::r 0 1 ob.J a «" 0.0 0 ~ ...... (1) en

Plate 23. Dalatias licha (BONN A TERRE, 1788). Female 147 em. t.l. (upper teeth), Female 84 em. t.l. (lower teeth), both from W Hebrides, N.E. Atlantic. -Vl -.1