BULLETIN DE L'INSTITUT ROYAL DES SCIENCES NATURELLES DE BELGIQUE, BIOLOGIE, 60: 181-230, 1990 BULLETIN VAN HET KONINKLIJK BELGISCH INSTITUUT VOOR NATUURWETENSCHAPPEN, BIOLOG IE, 60: 181-230, 1990

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

Part A: Selachii. No. 2b :· Order: Carcharhiniformes - Family: Scyliorhinidae by J. HERMAN, M. HOVESTADT-EULER & D.C. HOVESTADT

Abstract authors always restricted their illustations to simplified drawings. The tooth morphology of all the genera and subgenera of the Scyliorhinidae, A careful examination was initiated of the teeth of almost with the unique exception of Pentanchus (not available for examination), all living genera of the Scyliorhinidae, comprising (after is described and illustrated. CoMPAG NO, 1988) the subfamilies Atelomycterinae with Several systematic considerations are given. The odontology of the genera Centroscyllium and Aculeo/a, which are the genera Atelomycterus and Aulohalaelurus, Schroede­ rather aberrant among the Squaliforms, is compared with that of all the richtyinae with the single genus Schroederichthys, Scylio­ Scyliorhinidae taxa presently described. rhininae with the genera Scyliorhinus, Poroderma and Key-words: Elasmobranchii, Selachii, Scyliorhinidae, odontology. Cephaloscyllium, and Pentanchinae, which is devided into the tribe Galeini with the genera Asymbolus, Cephalurus, Galeus, Parmaturus, Haploblepharus, Holohalaelurus and Resume: Halaelurus, and the tribe Pentanchini with the genera Pen­ tanchus and Apristurus. The latter genus comprises the A !'exception de celle du genre Pemachus (non disponible), Ia morphologie brunneus-, indicus-, laurussonii-, longicephalus-, manis-, des dents des genres et des sous-genres des Scyliorhinidae est decrite et profundorum-, platyrhynchus-, riveri-, sibogae- and spon­ figuree. giceps-groups. The Halaelurus has the two subgenera Diverses considerations systematiques sont formulees. Halaelurus and Bythaelurus. L 'odontologie des genres Centroscyllium et Aculeola, si isoles au sein des Teeth of the genus Pentanchus were not available for Squaliformes, est comparee avec celle de tous les taxa de Scyliorhinidae presentement decrits. examination and could not be described nor illustrated. Mots-clefs : Elasmobranchii, Selachii, Scyliorhinidae, odontologie. CoMPAGNO (1988) indicates differences between the type species of Cephalurus and specimens from off Peru. The teeth of one of these specimens were examined, described and illustrated. Kurzfassung

Die Zahnmorphologie aller Gattungen und Untergattungen der Scyliorhini­ dae werden beschrieben und illustriert, mit Ausnahme von der Gattung Pentachus, die zum untersuchen nicht verfilgbar war. Description of the odontological morphotypes Mehrere systematische Erwagungen werden vorgeschlagen. Die Odontologie der Gattungen Centroscyllium und Aculeola, abweicht ORDER : CARCHARHINIFORMES, von den Squaliformes, werden verglichen mit allen hier beschr iebenen sensu COMPAGNO (1988) Scyliorhinidentaxa. Schliisselworter : Elasmobranchii, Selachii, Scyliorhinidae, Odontologie. This order comprises, after CoMPAGNO (1988), eight fami­ lies : Triakidae, Proscylliidae, Leptochariidae, Scyliorhini­ dae, Pseudotriakidae, Sphyrnidae, Carcharhinidae and Introduction Hemigaleidae. Besides the Scyliorhinidae, the tooth mor­ phology of which will be described and illustrated in this The teeth of the Scyliorhinidae, in general, were occasio­ fascicle, the remaining families include the following sub­ nally described but never properly illustrated. Previous fami lies, tribes and genera : 182 J. HERMAN, M. HOYESTADT-EULER & D.C. HOYESTADT

Family : Proscylliidae Family : SCYLIORHINIDAE GILL, 1862 Subfamily : Proscylliinae Proscyllium HILGENDORF, 1904 This family includes, after CoMPAGNO (1988), the fol­ Eridacnis SMITH , 1913 lowing subfamilies : Ctenacis CoMPAGNO, 1973 Subfamily : Golluminae Atelomycterinae Gollum CoMPAGNO, 1973 Schroederichtyinae Family : Leptochariidae Scyliorhininae Leptocharias SMITH, Pentanchinae in MuLLER & HENLE, 1838 Family : Pseudotriakidae The subfamily Atelomycterinae· comprises the genera : Pseudotriakis BRITO CAPELLO, 1867 Atelomycterus GARMAN, 1913, type species: Scyllium mar­ Family : Triakidae moratum Subfamily : Triakinae Aulohalaelurus FowLER, 1934, type species : Catu.lus Triakis MOLLER & HENLE, 1838 labiosus Mustelus LINCK, 1790 Scylliogaleus BouLENGER, 1902 The subfamily Schroederichthyinae comprises the genus : Subfamily : Galeorhininae Schroederichthys SPRINGER, 1966, type species: Schroede­ Tribe : Iagini richthys maculatus Hemitriakis HERRE, 1923 WHITLEY, Furgaleus , 1951 The subfamily Scyliorhininae comprises the genera : COMPAGNO, 1973 Gogolia Cephaloscyllium GILL, 1862, type species: Scyllium lati­ !ago _ . COMPAGNO & SPRI NGER, 1971 ceps Tribe : Galeorhinini Poroderma SMITH , 1837, type species : Squalus africa nus Galeorhinus BLAIN VILLE, 1816 Scyliorhinus BLAINVILLE, 1816, type species : Scyliorhinus . , Hypogaleus SMITH, 1957 canicula Family : Sphyrnidae Eusphyra GILL, 1862 The subfamily Pentanchinae comprises the tribes : Sphyrna RAFINESQUE, 1810 Galeini, including the genera : Family : Carcharhinidae Asymbolus WHITLEY, 1939, type species: Scyllium anale Subfamily : Galeocerdinae Cephalurus BIGELOW & ScHROEDER, 1941 , type species: Galeocerdo MOLLER & HENLE, 1837 Subfamily : Scoliodontinae Catulus cephalus Gale us Gale us melasto­ Scoliodon MOLLER & HENLE, 1837 RAFINESQUE, 1810, type species : Subfamily : Carcharhininae mus Tribe: Rhizoprionini Halaelurus GILL, 1862, type species: Halaelurus burgeri Loxodon MOLLER & HENLE, 1838 Haploblepharus GARMAN, 1913, type species: Haploble­ Rhizoprionodon WHITLEY, 1929 pharus edwardsii Tribe : Isogomphodontini Holohalaelurus FowLER, 1934, type species: Scyliorhinus Isogomphodon GILL, 1862 reg ani Tribe : Carcharinini Parmaturus GARMAN, 1906, type species: Parmaturus Carcharhinus BLAINVILLE, 1816 polisus Glyphis AGASSIZ, 1843 Pentanchini, including the genera : Prionace CANTOR, 1849 Apristurus GARMAN , 1913, type species : Scyliorhinus indi­ Nasolamia CoMPAGNO & GARRICK , 1983 cus Lamiopsis GILL, 1862 Pentanchus SMITH & RADCLIFFE, 1912, type species: Pen­ Negaprion WHITLEY, 1940 tanchus profundicolus Tribe : Triaenodontini Triaenodon MOLLER & HENLE, 1837 The following 136 specimens belonging to 28 species were Family : Hemigaleidae examined: Subfamily : Hemigaleininae Apristurus maderensis 2 males Hemigaleus BLEEKER, 1852 Apristurus laurussoni 15 males 9 females Paragaleus BuDKER, 1935 Asymbolus analis 2 males 1 female Chaenogaleus GILL, 1862 Atelomycterus marmoratus 2 males Subfamily : Hemipristinae Aulohalaelurus labiosus 1 male 1 female Hemipristis AGASSIZ, 1843 Bythaelurus canescens male Cephaloscyllium isabellum male 1 female Cephaloscyllium. su.fflans male Cephalurus cephalus male Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 183

Cephalurus sp. (SE-Pacific) I female HEEMSTRA, 1979, Aprisrurus federovi DoLGANOV, 1985, Ga!eus me!astomus 14 males 18 females Aprisrurus herklorsi (FowLER, 1934), Apristurus indicus Galeus murinus 7 males 9 females (BRAUER, 1906), Apristurus investigatoris (MISRA, 1962), Galeus polli I male I female Apristurus japonicus NAKAYA, 1975, Apristurus kampae Halae!urus biirgeri I female T AYLOR, 1972, Apristurus !aurussoni (SAEMUNDSSON, Halaelurus nata!ensis male 1922), Apristurus !ongicephalus NAKA YA, 1975, Apristu­ Haplob!epharus edwardsii male rus macrorhynchus (TANAKA, 1909), Apristurus maderen­ Ho!oha!aelurus regani male sis CADENAT & MAUL, 1966, Aprisrurus manis (PRINGER, Parmaturus me!anobranchius female 1979), Apristurus microps (GILCHR IST, 1922), Apristurus Parmaturus pi!osus male nasutus DE BuEN, 1959, Apristurus parvipinnis SPRINGER Parmaturus xaniurus male female & HEEMSTRA, 1979, Apristurus pinguis DENG, XIONG & Poroderma africanum female ZAHN, 1983, Apristurus p!atyrhynchus (TAN AKA, 1909), Poroderma pantherinum male Apristurus profunc!orum (GooDE & BEAN, 1896), Apristu­ Scyliorhinus canicula 5 males 9 females rus riveri BIGELOW & ScHROEDER, 1944, Apristurus sal­ Scyliorhinus capensis 1 female danha (BARNARD, 1925), Apristurus sibogae (WEBER, Scy!iorhinus stellaris 7 males 6 females 1913), Apristunts sinensis CHu & Hu, 1981, Apristurus Scyliorhinus torazame I male spongiceps (GILBERT, 1895), Apristurus stenseni (SPRIN­ Schroederichthys chilensis I male GER, 1979), Apristurus verweyi (FowLER, 1934). Schroederichthys maculatus male The type species of this genus is Scy!iorhinus indicus BRAUER, 1906 by original designation. However, teeth of The teeth of the species of the family Scyliorhinidae, the type species were not available for examination, and generally, have a well developed slender principal cusp, therefore Apristurus laurussoni will be used instead, for flanked by up to three cusplets. illustration and description of the tooth morphology. The root is rarely holaulacorhizid, more often secondary hemiaulacorhizid, which is demonstrated by a partly closed median groove, and mostly secondary anaulacorhizid, by Apristurus laurussoni (SAEM UNDSSON, 1922) a fully closed median groove. Because fossil ancestors of (Plates I, _2, 3, 4) this family mostly have holaulacorhizid roots with a strongly developed median groove, the root types of living Scyllium laurussoni SAEMUNDsso , 1922, Vidensk. Meddr Dansk Scyliorhinidae can not be hemiaulacorhizid nor anaulacor­ Naturhist. Foren. Kobenhavn., 74 : 173. hizid sensu stricto (CASlER, 1947), but these types of hemiaulacorhizy or anaulacorhizy are formed by syncre­ The teeth of this species have a rather slender and elonga­ tion of the median groove. Therefore, we follow CAPPETTA ted principal cusp, which is more distinct in upper teeth. ( 1987) and use the terms secondary hemiaulacorhizy and The principal cusp is strongly inclined toward the commis­ secondary anaulacorhizy for this type of root. sure and is commonly fl anked by two or (in lateral teeth) three also slender and elongated cusplets. The cusplets HETERODONTY closest to the principal cusp reach half the height of the principal cusp or even more in lower lateral teeth. A weak dignathic heterodonty can be present by relatively The root varies between holaulacorhizid and secondary higher cusplets and a shorter principal cusp in lower teeth. anaulacorhizid, showing two root lobes, that are relatively A weak sexual heterodonty is sometimes present by a long and narrow, forming an angle at the root base. The slightly higher principal cusp and more cusplets for fema­ dimensions of the teeth are plurimillimetrical in range. les. A strong ontogenetic heterodonty is often present by The outer face of the principal cusp and cusplets is flat broader cusplets and a much shorter, less convex principal to very weakly convex, presenting well developed striae, cusp in teeth of juvenile specimens. A gradient monogra­ that run from crown base toward the apex of the principal phic heterodonty is always present by little broader lateral cusp and cusplets. Although those striae are rather regular teeth with a more inclined or oblique principal cusp. The in shape, they are sometimes slightly sigmoidal. They lateral teeth and those next to the commissure are mostly reach a maximum densisty of six striae on the principal smaller than the anterior ones. cusp in anterior teeth, decreasing to two on the lower lateral one. Striae on cusplets are always less distinct than on the principal cusp. Genus : Apristurus GARMAN, 191 3 Reticulated ornamentation is present on the crown base between the striae, which is more distinct on posterior M any authors have studied thjs genus but there is till now teeth. much uncertainty about the validity of its species. T he The inner face of the principal cu p and cusp lets is strong ly most recent revision is of CoMPAGNO (1988), who included convex, also presenting striae, which are a little less dis­ the following 27 species in this genus : tinct than the outer ones. Apristurus at/anticus (KoEFOED, 1932), Apristurus brun­ They al o run from crown base toward the apex of the neus (GILBERT, 1892), Apristurus cm~ u tus SPRINGER & principal cusp and cusplets in a more or less regular way. 184 J. HERMAN, M. HOVESTADT-EULER & D.C. HOVESTADT

Their density is rather steady with six striae on the princi­ the inner face of the root into two parts and its central pal cusp and less on cusplets. section is more or less protuberated. Basal ornamentation is absent. Sometimes, one or two foramina are present on each root The outer face of the root has a strong central depression, lobe, on the part toward the crown in the depressions, presenting a line of up to ten well developed f01mina, of fl anking the protuberation of the ridge. Foramina are which the central ones are often merged. absent near the crown-root junction. A foramen is always The inner face of the root shows a ridge, which has about present in the centre of the ridge. the same shape as the angle of the root lobes. It divides Some randomly scattered foramina are usually present on the inner face of the root into two parts, of which the basal the basal face. face is often called the root base. Two or four foramina are present on each root lobe, on the part toward the crown and along the ridge. Foramina Genus : Atelomycterus GARMAN , 191 3 are absent near the crown root junction. A fo ramen is always present just below the centre of the ridge, and is The genus Atelomycterus comprises, after CoMPAGNO sometimes more or less extended toward the root base, (1988), the two species Atelomycterus mac/eayi WHITLEY, forming a partial groove or even sometimes a complete 1939 and Atelomycterus marmoratus (BENNETT, 1830). median one. The type species is Scyllium marmoratum BENNETT, 1830 Some randomly scattered foramina are present on the basal by original monotypy. face.

Ate/omycterus marmoratus (BENNETT, 1830) Genus : Asymbo/us WHITLEY , 1939 (Plates 7, 8)

The genus Asymbolus comprises, after CoMPAGNO ( 1988), Scy/lium marmora/Urn B ENNETI, 1830. Memoir of the li fe and the two species Asymbolus vincenti (ZEITZ, 1908) and public Services of Sir Stamford Raffles, London, p. 693 . Asymbolus ana/is (OGILBY, 1885). The type species is Scy//ium anale 0 GILBY, 1995 by original designation. The teeth of this species have a rather broad based but elongated principal cusp in both upper and lower jaws. Asymbo/us ana/is (OGILBY, 1885) The principal cusp is erect on anterior teeth, becoming (Pl ates 5, 6; Pl ate 35, figs. 1 & 2) sli ghtly oblique on lateral and commissural ones. A cusplet is always present at each side, broad based, like the princi­ Scy/lium anale 0 GILBY, 1885. Proceedings of the Linnean Society , pal cusp, and they tend to curve to its apex. New South Wales, 10 : pp. 445-447. The root is always secondary hemiaulacorhizid and shows two root lobes, which are relatively broad and subquadrate, The teeth of this species have a broad based but elongated forming a straight line at the root base. principal cusp. The principal cusp is more or less oblique The teeth are plurimillimetrical in range. toward the commissure and has commonly one also short The outer face of the principal cusp and cusplets is rather and broad based cusplet at each side. A vaguely developed convex, presenting little developed striae that run from second cusplet is present on the extreme edges, next to 1/4 to about 3/4 of the height of the principal cusp and each cusplet. cusplets. These striae are rather irregular in shape. They The anaulacorhizid root shows two root lobes that are reach a maximum density of five to eight striae on the relatively long and narrow, forming an angle at the root principal cusp in both upper and lower teeth. One or two base. striae are always present on the cusplets. The teeth are plurimillimetrical in range. The crown base is slightly depressed near the crown-root The outer face of the principal cusp and cusplets is weakly junction. Ornamentation is absent. convex, presenting well developed, short basal costules The inner face of the principal cusp and cusplets is strongly and poorly developed striae. The crown more or less over­ convex, also presenting well developed striae. hangs the root. They also run from 1/4 tot 3/4 of the height of the principal A basal ornamentation is absent. cusp and cusplets in a more or less irregul ar way. Their The inner face of the principal cusp and cusplets is strongly density is rather steady, of seven to eight on the principal convex, presenting striae. They run from the crown base cusp and of two to three on both cusplets. toward the apex of the principal cusp and cusplets in a Basal ornamentation is absent. more or less regular way. Their density is rather high, of There is a deep depression on both mesial and distal parts about nine to thirteen on the principal cusp and of tluee of the principal cusp, which is caused by two strong root to five on the cusplets. Basal ornamentation is absent. depressions. The outer face of the root is very low and The outer face of the root presents an irregul ar series of divided into mesial and distal root parts by the remains three to six well developed foramina. of a median groove. The inner face of the root shows a ridge, which has about Both mesial and distal root parts present three to four the same shape as the angle of the root lobes. It divides foramina along the crown-root junction. \I

Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 185

The inner face of the root shows a ridge, which is strongly just above the centre of the protuberated part of the ridge. protuberated in the central part. It divides the inner face The relict of a median groove is present on the basal root of the root into two sections. face, running from root base up to half the height of the One or two foramina are present on each root lobe in a basal root face. deep mesial and distal depression. An aperture is present Numerous randomly scattered foramina are present on the just above the centre of the protuberated part of the ridge. basal face of both root lobes. The relict of a median groove is present on the basal root face, running from the root base up to half the height of the basal root face. Numerous randomly scattered foramina are present on the Genus : Cephaloscyllium GILL, 1862 basal face of both root lobes. The genus Cephaloscyllium was described by GILL, 1862 and comprises, after CoMPAG NO 1988), the species Cepha­ Genus : Aulohalaelurus FowLER, 1934 loscyllium fasciatum CHAN, 1966, Cephaloscyllium isabel­ fum (BoNNATERRE, 1788), Cepha!oscy!!ium laticeps The monotypic genus Auloha!aelurus was described by (DUM ER IL, 1853), Cephaloscyllium nascione WHITLEY, FowLER, 1934 as a subgenus of Halaelurus GILL, 1862. 1932, Cephaloscyl!ium silasi (TALWAR, 1974), Cephalos­ cyllium sufflans (REGAN, 1921) and Cephaloscyllium ven­ However, SPRINGER ( 1979) elevated Aulohalaelurus to the (GARMAN, 1880). The type species is generic rank, and was followed by COMPAGNO (1984). The triosum Scyllium laticeps DuMER IL, 1853 by original designation, but was type species of the genus is Catulus labiosus WAITE, 1905 not available for examination, and therefore, Cephaloscyl­ by original designation. lium isabellwn will be used instead, which is very closely related to the type species (CoMPAGNo, 1984, 1988). Aulohalaelurus labiosus (WAITE, 1905) (Plates 9, 10) C ephaloscyllium isabellum (BoNNATERRE, 1788) Catulus /abiosus WAITE, 1905. Records of the Australian (Plates 11 , 12) Museum, 6 (2): 57.

Squa/us isabella B oNNATERRE, 1788. Tableau encyclopedique et The teeth of this species have a rather slender, elongated methodique des trois regnes de Ia terre : Ichthyologie, 6. La principal cusp in both upper and lower jaws. The principal mer du sud. cusp is erect in anterior teeth, becoming slightly oblique in lateral and commissural teeth. One well developed short The teeth of this species have a broad based, relatively cusplet is always present, with a poorly developed second short principal cusp, which is more or less oblique toward one on its extreme mesial and distal parts. The cusplets the commissure. A short and broad based cusplet is present are broad based and tend to curve toward the principal at each side. A vaguely developed second cusplet is present cusp. on the extreme edges, next to each cusplet. The root is always secondary hemiaulacorhizid and shows The root varies between holaulacorhizid and secondary two root lobes, which are relatively broad and subquadrate, anaulacorhizid, showing two root lobes that are relatively fmming a straigth line at the root base. long and narrow and f01m an angle at the root base. The teeth are plurimillimetrical in range. The teeth are plurimillimetrical in range. The outer face of the principal cusp and cusplets is rather The outer face of the principal cusp and cusplets is weakly convex, presenting little developed costules. convex, presenting well developed, short basal costules. Ornamentation is absent. The crown overhangs the root. The inner face of the principal cusp and cusp lets is strongly Some striae are occasional on the principal cusp and cus­ convex, sometimes presenting poorly developed striae. plets, in males only. Basal ornamentation is absent. Basal ornamentation is absent. There is a deep depression on both mesial and distal parts The inner face of the principal cusp and cusplets is strongly of the principal cusp, which is caused by two strong root convex, presenting striae that are more distinct in males. depressions. They run from the crown base to the apex of the principal The outer face of the root is divided into mesial and distal cusp and cusplets in a more or less regular way. Their root sections by the remains of a median groove. Both density is rather high with about fifteen striae on the mesial and distal root sections present one or two foramina principal cusp and five to seven on the cusplets. along the crown-root junction. Basal ornamentation is absent. The inner face of the root shows a ridge, which is strongly The outer face of the root is rather high, presenting an protuberated in the central part. It divides the inner face irregular series of five to eight well developed foramina, of the root into two sections. of which the central ones are often merged. One or two foramina are present on each root lobe in a The inner face of the root shows a ridge similar shaped deep mesial and distal depression. An aperture is present as the angle of the root lobes. It divides the inner face of 186 J. HERMAN, M. HOVESTADT-EULER & D.C. HOYESTADT

the root into two sections and its central part is more or Cephalurus sp. cf. cephalus less protuberated. (Plates 15, 16) Two to three foramina are present on each root lobe on the part toward the crown in the depressions, flanking the CoMPAGNo ( 1988) : Sharks of the Order Carcharhiniformes, Prin­ protuberation of the ridge. Foramina are absent near the ceton University Press. Princeton New Jersey. crown-root junction. A central foramen is always present just below the centre of the ridge, which is sometimes The teeth of this species have a rather short, broad based more or less elongated toward the root base, forming partly principal cusp, which is always strongly inclined toward a groove or sometimes even a complete median one. the commissure. One distal cusplet is always present, and Some randomly scattered foramina are usually present on up to three mesial cusplets may also be present. All cus­ the basal face. plets tend to arise from the lower part of the mesial and distal cutting edges of the principal cusp. They have a rather triangular shape and tend to point inwards. Genus : Cephalurus BIGELOw & ScHROEDER, 1941 The root is holaulacorhizid, showing two root lobes that are relatively long and narrow, forming a weak angle at This monotypic genus was erected by BIGELOW & ScHROE­ the root base. The teeth are plurimillimetrical in range. DER (1941) for Catulus cephalus GILBERT, 1892. CoM­ The outer face of the principal cusp and cusplets is flat PAGNO (1984, 1988) cited previous authors regarding to very weakly convex, presenting some poorly developed Cephalurus - like sharks from the Eastern central and striae that are better developed on lateral teeth. (In juve­ SE-Pacific differing from the type species. niles all teeth have better developed striae). Theeth of one of the latter specimens were examined and Costules and ornamentation are absent. are described and illustrated here for comparison with the The inner face of the principal cusp is strongly convex, type species of more Northern Eastern Pacific distribution. also presenting some striae that are slightly more distinct than the outer ones. Cephalurus cephalus (GILBERT, 1892) The outer face of the cusplets is rather flat and has no (Plates 13, 14) striae. Basal ornamentation is absent. The outer face of the root is low, presenting an aperture

Cant! us cephal us GILBERT, 1892. Proceedings of the United States with some irregularly lined foramina. National Museum, 14: 541. The inner face of the root shows a ridge that is angled as the root lobes. It divides the inner face of the root into The teeth of this species have an elongated, slender princi­ two sections. pal cusp, which is always somewhat inclined toward the Two to four foramina are present on each root lobe on commissure. One distal cusplet is always present, and up the section toward the crown and are lined up near the to two mesial cusplets may be present in addition. All ridge. Foramina are absent near the crown-root junction. cusplets tend to arise from the lower part of the mesial A foramen is always present in the centre of the ridge. and distal cutting edges of the principal cusp and have a The basal section is rather concave. rather triangular shape. Some randomly scattered foramina are usually present on The root varies between holaulacorhizid and secondary the basal face. hemiaulacorhizid, showing two root lobes that are relati­ vely long and broad and form a weak angle at the root base. The teeth are plurimillimetrical in range. Genus : Gale us RAFINESQUE, 1810 The outer face of the principal cusp and cusplets is flat to very weakly convex, presenting some poorly developed This genus comprises (after CoMPAGNO, 1988) 11 species : striae, which are more developed in lateral teeth. Galeus arae (NICHOLS, 1927), Galeus at/anticus (VAIL­ Costules and ornamentation are absent. LANT, 1888), Galeus boardmani (WHITLEY, 1928), Galeus The inner face of the principal cusp is strongly convex eastmani (JORDAN & SNYDER, 1904), Galeus murinus (CoL­ and also presents some well developed striae. LETT, 1904), Galeus nipponensis NAKAY A, 1975, Galeus Basal ornamention is absent. piperatus SPRINGER & WAGNER, 1966, Gale us polli CADE­ The outer face of the root is low, presenting some irregu­ NAT, 1959, Galeus sauteri (JORDAN & RI CHARDSON, 1909), larly lined foramina. Galeus schultzi SPRINGER, 1979 and the type species The inner face of the root shows a ridge of about the same Gale u.s melastomus RAFINESQUE, 1810. shape as that the angle of the root lobes. It divides the inner face of the root into two parts. One or two foramina are present on each root lobe on the section toward the Gale us melastomus R AFIN ESQUE, 1810 crown flanking the ridge. Foramina are absent near the (Plates 17, 18) crown-root junction. A foramen is always present in the centre of the ridge. The basal section is rather concave. Caratleri di alcuni nuovi generi e nuove specie di animali (princi­ Some randomly scattered foramina are usually present on palmente di pesci) e pi ante della Scicilia, con vari e osserva­ the basal face. zioni sopra i medisimi. Palermo, pt. I : 13. Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 187

The teeth of this species have a rather slender and elon­ Halaelurus burgeri (MuLLER & HENLE, 1838), Halaelurus gated principal cusp, which is more prominent in upper lineatus BAss, o' AuBREY & KISTNASAMY, 1975, Halaelu­ teeth, becoming lower toward the commissure. The princi­ rus natalensis (REGAN, 1904), Halaelurus quagga pal cusp is slightly inclined toward the commissure. It has (ALcOCK, 1899) with the type species Halaelurus burgeri. more or less sigmoidal mesial and distal cutting edges and Bythaelurus comprises Halaelurus canescens (GuNTHER, has commonly one (in upper and lower anterior teeth), 1878), Halaelurus dawsoni SPRINGER, 1971, Halaelurus two (in upper and lower lateral and upper posterior teeth) hispidus (ALcocK, 1891), Halaelurus immaculatus CHu & and three (in the lower commissural tooth). Also slender MENG, 1982, Halaelurus lutarius SPRINGER & o'AuBREY, and elongated cusplets are present at each side. The cus­ 1972 and Halaelurus alcocki GARMAN, 1913, which is of plets closest to the principal cusp reach half the height of uncertain status. The type species is Halaelurus canescens, the principal cusp or even more in lower lateral teeth. generotype Halaelurus biirgeri. The root is always secondary anaulacorhizid and shows two root lobes that are relatively long and narrow, forming an angle at the root base. The teeth are plurimillimetrical Halaelurus (Bythaelurus) canescens (GUNTHER, 1878) in range. (Plates 19, 20) The outer face of the principal cusp and cusplets is flat or very weakly convex, presenting well developed striae Scyllium canescens G NTHER, 1878. Annal and Magazine of Natu­ that run from just above the crown base to half the height ral History, (ser. 5), 2 (8): 18. of the principal cusp and apex of cusplets. Although these striae are rather regular in shape, they are sometimes The teeth of this species have a broad based, relatively slightly sigmoidal. They reach a maximum density of short principal cusp, which is more distinct in the lower seven or eight striae on the principal cusp on anterior and jaw. The principal cusp is more or less oblique toward the lateral teeth, decreasing to three or four on the lower commissure. Commonly, one also short and broad based posterior teeth. Striae on cusplets are always less distinct cuplet is present at each side. On the extreme edges, next than on the principal cusp. to each cusplet, an undeveloped second cusplet is present, Reticulated ornamentation is present on the crown base which is more distinct in upper lateral teeth. between the striae, which is more distinct on posterior The root varies between holaulacorhizid and secondary teeth. anaulacorhizid, showing two root lobes that are relatively The outer central crown base is depressed, caused by a long and narrow and form an angle at the root base. The deep depression of the root. teeth are plurimillimetrical in range. The inner face of the principal cusp and cusp lets is strongly The outer face of the principal cusp and cusplets is weakly convex, also presenting striae that are slightly less distinct convex, with well developed, short basal costules. The than the outer ones. crown tends to overhang a rather high, basal depression, They also run from the crown base toward the apex of which is present over the whole width of the crown base. the principal cusp and cusplets in a more or less regular Striae on the principal cusp and cusplets and basal orna­ way. Their density is rather steady, with eight to ten striae mentation are absent. on the principal cusp and three to four on the cusplets. The inner face of the principal cusp and cusplets is strongly The striae do not fully reach the apex of the principal cusp. convex, presenting striae that run from the crown base Basal ornamentation is absent. toward the apex of the principal cusp and cusplets in a The outer face of the root has a strong central depression, more or less regular way. Their density is rather high, with presenting a series of up to eight well developed foramina, about twelve to seventeen striae on the principal cusp and of which the central ones are often merged in anterior five to seven on the cusplets. Basal ornamentation is teeth. absent. The inner face of the root shows a ridge similar shaped The outer face of the root is rather high, presenting an as the angle of the root lobes. It divides the inner face of irregular series of three to five well developed foramina, the root into two sections. of which the central ones are often merged. Two to four foramina are present on each root lobe on The inner face of the root shows a ridge similar shaped the part toward the crown and are lined along the ridge. as the angle of the root lobes. It divides the inner face of Foramina are absent near the crown-root junction. A fora­ the root into two sections and its central part is more or men is always present just above the centre of the ridge. less protuberated. Some randomly scattered foramina are usually present on Two to three foramina are present on each root lobe on the basal face. the part toward the crown, in the depressions flanking the protuberation of the ridge. Foramina are absent near the crown-root junction. A foramen is always present in the Genus: Halaelu.rus GILL, 1862 centre of the ridge, which is sometimes more or less elon­ gated toward the root base, forming a partial groove or This genus comprises, after CoMPAGNO (1988), 2 subgene­ even sometimes a complete median one. ra : Halaelurus and Bythaelurus. Halaelurus comprises Some randomly scattered foramina are usually present on Halaelurus boesemani SPRINGER & o 'AUBREY, 1972, the basal face. I I

188 J. HERMAN, M. HOVESTADT-EULER & D.C. HOVESTADT

Halaelurus (Halaelurus) burgeri (MOLLER & HENLE, 1838) The root is secondary anaulacorhizid and shows two root (Plate 25) lobes that are relatively long and narrow forming an angle at the root base. The teeth are plurimillimetrical in range. Scyllium biirgeri M OLLER & HENLE, 1838. Systematische Beschrei­ The outer face of the principal cusp and cusplets is flat hung der Plagiostomen. Berlin. Part 1 : 8. or very weakly convex, presenting well developed, elon­ gated basal costules. A reticulated ornamentation is present The teeth of this species have a broad based, relatively on the crown base between the costules, which is more short principal cusp, which is poorly developed in lateral distinct on posterior teeth. teeth. The mesial and distal cutting edges are extended. The inner face of the principal cusp and cusp lets is strongly Cusplets are absent. convex, presenting striae that run from the crown base to The secondary anaulacorhizid root shows two root lobes 3/4 of the height of the principal cusp and reach the apex that are relatively long and narrow and form a very obtuse on cusplets in a more or less regular way. Their density angle in anterior teeth and one line at the root base in varies between .seven to twelve on the principal cusp but lateral teeth. The teeth are plurimillimetrical in range. is less on cusplets. The outer face of the principal cusp is weakly convex, Basal ornamentation is absent. presenting well developed, short basal costules. The crown The outer face of the root has a strong central depression, more or less overhangs the root. presenting a series of four to ten well developed foramina, A basal reticulated ornamentation is present between the of which the central ones are often merged. costules. The inner face of the root shows a ridge that is angled as The inner face of the principal cusp and cusp lets is convex, the root lobes. It divides the inner face of the root into and some poorly developed striae are present. two sections. A basal reticulated ornamentation is present on lateral Two to four foramina are present on each root lobe on teeth. the part toward the crown, lined along the ridge. Foramina The outer face of the root is rather high, presenting an are absent near the crown-root junction. A foramen is irregular series of three of five well developed foramina. sometimes present in the centre of the ridge. The inner face of the root shows a ridge, similar shaped Some randomly scattered foramina are usually present on as the angle of the root lobes. It divides the inner face of the basal face. the root into two sections and its central part is, in anterior teeth, more or less protuberated. Two to three foramina are present on each root lobe on the part toward the crown on the mesial and distal parts. Genus: Holohalaelurus FowLER, 1934 Foramina are absent on the crown-root junction. A fora­ men is always present in the centre of the ridge. This genus comprises, after CoMPAGNO (1988), two spe­ Some randomly scattered foramina are usually present on cies: Holohalaelurus punctatus (GILCHRIST, 1914) and the the basal face. type species Holohalaelurus regani (GILCHRIST, 1922).

Genus: Haploblepharus GARMAN, 1913 Holohalaelurus regani (GILCHRIST, 1922) This genus comprises, after CoMPAGNO (1988), three spe­ (Plates 23, 24) cies: Haploblepharus fuscus SMITH, 1950, Haploblepha­ rus pictus (MOLLER & HENLE, 1838) and the type species Scyliorhinus regani GILCHRIST, 1922. Special report no. 3. Report Haploblepharus edwardsii (VOIGT, 1832). of the Fishery and Marine Biological Survey. Union of South Africa, Part 1, 2 : 45.

Haploblepharus edwardsii (VoiGT, 1832) The teeth of this species have a broad based, but elongated (Plates 21, 22) principal cusp. The principal cusp is more or less inclined toward the commissure in lateral and commisural teeth, Scyllium edwardsii VoiGT, 1832. In Cuvi ER, 1832, Tierreich, 2 : and two also short and broad based cusplets are present 504. at each side. The root is always secondary anaulacorhizid and shows The teeth of this species have a rather slender and elon­ two root lobes that are relatively long and narrow and gated principal cusp. form an angle at the root base. The teeth are plurimillime­ The principal cusp is slightly inclined toward the commis­ trical in range. sure and two also slender and elongated cusplets are pre­ The outer face of the principal cusp and cusplets is almost sent at each side. flat, presenting well developed, relatively long basal costu­ The cusplets closest to the principal cusp reach half the les. The crown overhangs the root. height of the principal cusp or even more in lower lateral A weak reticulated basal ornamentation is present between teeth. the costules. Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 189

The inner face of the principal cusp and cusp lets is strongly varies between six to ten on the principal cusp and between convex, presenting well developed striae. These run from two to five on cusplets. 1/4 to 3/4 of the height of the principal cusp and cusplets Basal ornamentation is absent. in a rather irregular way. Their density varies between five The outer face of the root represents an irregular series of and nine on the principal cusp and two to three on cusplets. five to eight well developed foramina, of which the central Basal ornamentation is absent. ones are often merged. The outer face of the root is rather high, presenting an The inner face of the root shows a ridge, which is angled irregular series of five to eight well developed foramina, as the root lobes. It divides the inner face of the root into of which the central ones are often merged. two sections and is more or less protuberated in the centre. The inner face of the root shows a ridge that is angled as Two to three foramina are present on each root lobe on the root lobes. It divides the inner face of the root into the part toward the crown in the depressions, flanking the two sections, and its central part is more or less protube­ protuberation of the ridge. Foramina are absent near the rated. crown-root junction. A foramen is always present in the Two to three foramina are present on each root lobe, on centre of the ridge. the part toward the crown in the depressions flanking the Some randomly scattered foramina are usually present on protuberation of the ridge. Foramina are absent near the the basal face. crown-root junction. A foramen is always present just below the centre of the ridge. Some randomly scattered foramina are usually present on the basal face. Genus : Poroderma SMITH, 1837

This genus comprises two species after CoMPAGNO (1988) : the type species Poroderma africanum (GMELIN, 1789) and Genus: Parmaturus GARMAN, 1906 Poroderma pantherinum (SMITH, 1837). This latter only was available for examination. The genus comprises, after CoMPAGNo (1988), 5 species : Parmaturus campechiensis SPRI NGER, 1979, Parmaturus macmillani H ARDY, 1985, Parmaturus melanobranchius (CHAN, 1966), Parmaturus xaniurus (GILBERT, 1892) and Poroderma pantherinum (SMITH, 1837) the type species Parmaturus pilosus GARMAN , 1906). (Plates 27, 28)

Scyllium pantherinum SMITH, 1837, in M OLLER & H ENLE, 1838c. Systema­ tische Beschreibung der Plagiostomen, Berlin (pt. I) : 13. Parmaturus pilosus GARMAN, 1906 (Plate 26, Plate 35, figs. 3 & 4) The teeth of this species have a broad based, triangularly shaped, but elongated principal cusp in both upper and

Parmaturus pilosus G A RMAN, 1906. New Plagiostoma. Bulletin lower jaws. The principal cusp is erect on anterior teeth, of the Museum of Comparative Zoology. Harvard College, becoming almost lightly oblique on lateral and commissu­ 46 (II) : 204. ral teeth. One cusplet is always present at each side. They are relatively small, weakly developed, and tend to curve The teeth of this species have a broad based but elongated to the principal cusp. The root is always secondary hemiau­ principal cusp, which in lateral teeth, is more or less obli­ lacorhizid and shows two root lobes, which are relatively que toward the commissure. The principal cusp commonly elongated and form an angle at the root base. has one also short and broad based cusplet at each side. The teeth are plurimillimetrical in range. On the most extreme mesial and distal part, next to each The outer face of the principal cusp and cusplets is rather cusplet, a vaguely developed second cusplet is present. convex, presenting well developed rather regular shaped The crown base overhangs a rather high basal depression fine costules at crown base. The outer surfaces of the that is present over the whole crown base. principal cusp and cusplets are smooth. The root is secondary anaulacorhizid and shows two root Ornamentation is absent. lobes that are relatively long and narrow, forming an angle The inner face of the principal cusp and cusp lets is strongly at the root base. The teeth are plurimillimetrical in range. convex. The surfaces of the principal cusp and cusplets The outer face of the principal cusp and cusplets is weakly are smooth. convex, presenting well developed, short basal costules. Basal ornamentation is absent. The crown is basally depressed. The outer face of the principal cusp and cuspl ets is strongly Striae may be present on the principal cusp and cusplets. convex. The surfaces of principal cusp and cusplets are Basal ornamentation is absent. smooth. The inner face of the principal cusp and cusplets is strongly Basal ornamentation is absent. convex, presenting well developed striae. These run from The outer face of the root is rather high, presenting up to a quarter of the height of the principal cusp and cusplets twenty foramina along the root base, of which the central to their apex in a more or less sigmoidal way. Their density ones are often merged. 190 J. HERMAN, M. HOVESTADT-EULER & D.C. HOYESTADT

The inner face of the root shows a ridge that is strongly Genus : Scyliorhinus BLAINVILLE, 1816 protuberated in the central part. It divides the inner face of the root into two sections. This genus comprises, after CoMPAG NO (1988), 13 species: A first aperture is present in the centre of the protuberated Scyliorhinus besnardi SPRINGER & SADOWSKY, 1970, Scy­ part of the ridge. Another one appears on the basal root liorhinus boa GooDE & BEAN, 1896, Scyliorhinus capensis face, as the relict of a median groove. Numerous randomly (SMITH, 1838), Scyliorhinus cervignoni MAUR I & BoN­ scattered foramina are present on the basal faces of both NET, 1970, Scyliorhinus garmani (FowLER, 1934 ), Scylio­ root lobes. rhinus haeckeli (RIBEIRO, 1907), Scyliorhinus hesperius SPRI NGER 1966, Scyliorhinus meadi SPRINGER, 1966, Scy­ liorhinus retifer (GARMAN, 1881 ), Scyliorhinus stellaris Genus: Schroederichthys SPRINGER , 1966 (LINNAEUS, 1758), Scyliorhinus torazame (TANAKA, 1908), Scyliorhinus canicula (LINNAEUS, 1758). This genus comprises, after CoMPAGNO (1988), 4 species : CoMPAGNO ( 1988.) proposed to divide Scyliorhinus into Schroederichthys bivius (MOLLER & HENLE, 1838), Schroe­ two groups. The odontological differences are not drama­ derichthys chilensis (GuiCJ-JENOT, 1848), Schroederichthys tic, but nevertheless, Scyliorhinus torazame is illustrated tenuis SPRINGER, 1966 and the type species Schroederich­ for comparison (Plates 33, 34). thys maculatus SPRINGER, 1966.

Schroederichthys maculatus SPRINGER, 1966 Scyliorhinus canicula (LINNAEUS, 1758) (Plates 29, 30) (Plates 31, 32)

Fishery Bulletin of the United States. Fish and Wildlife Service, Squalus canicula LI NNAEUS , 1758. Systema naturae, Ed. X : 234. 65 (3) : 605. The teeth of this species have a rather broad based but The teeth of this species have a broad based, relatively elongated principal cusp, becoming lower toward the com­ short principal cusp. The principal cusp is more or less missure. The principal cusp is slightly oblique toward the oblique toward the commissure in lateral teeth. One short commissure and commonly one cusplet is present at each and poorly developed cusplet is present at each side. side in upper lateral and posterior teeth and in lower ones. The root is secondary anaulacorhizid and shows two root The poorly developed cusplets are absent in upper anterior lobes that are relatively long and narrow and form an teeth. obtuse angle or an almost straight line at the root base. The root is always secondary anaulacorhizid and shows The teeth are plurimillimetrical in range. two root lobes that are relatively long and narrow and The outer face of the principal cusp and cusplets is weakly form an angle at the root base. The teeth are plurimillime­ convex, presenting well developed, very short basal costu­ trical in range. les. The crown overhangs the root. The outer part of the principal cusp and cusplets is flat or Striae are present on the principal cusp and cusplets in very weakly convex and presents poorly developed cus­ upper teeth and in lower posterior ones as extensions of plets, which are even absent on lower anterior teeth. A the costules. A reticulated basal ornamentation is present reticulated basal ornamentation is present in upper poste­ on the lower part of the costules. rior teeth only. The inner face of the principal cusp and cusplets is strongly The inner part of the principal cusp and cusp lets is strongly convex, presenting well developed striae. They run from convex and presents striae that are slightly less distinct the costules to the apex of the principal cusp and cusplets than the outer ones. They run from the base to the apex in a more or less regular way. Their density is rather of the crown of the principal cusp and cusplets in a more steady, of about seven striae on the principal cusp and of or less regular way. Their density varies between eleven one or two on cusplets. to sixteen on the principal cusp and from three to four on Basal ornamentation is absent. cusplets. The striae do not fully reach the apex of the The outer face of the root presents a regular series of five principal cusp. to eight well developed foramina, of which the central Basal ornamentation is absent. ones are sometimes merged. The outer part of the root presents a series of up to eight The inner face of the root shows a ridge that is obtusely well developed foramina, of which the central ones are angled as the root lobes. It divides the inner face of the often merged. root into two sections, and is more or less protuberated in The inner part of the root shows a ridge that is angled as the central part. the root lobes. It divides the inner face of the root into Foramina are absent on the root section toward the crown two sections. in the depressions flanking the protuberation of the ridge. Foramina are absent on the root part toward the crown as Nor there are foramina near the crown-root junction. A well as near the crown-root junction. A foramen is always foramen is always present in the centre of the ridge. present in the centre of the ridge. Some randomly scattered foramina are usually present on Some randomly scattered foran1ina are usually present on the basal face. the basal face. '' Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 191

DIFFERENTIAL DIAGNOSIS deeply concave and longitudinally hollowed crown base. Asymbolus and Haplob/epharus share a rather elongated Scyliorhinid teeth have a root type, which is in an evolving principal cusp. stage. Their original holaulacorhizid stage is adapting via Schroederichthys and Ha/aelurus are rather different from a secondary hemiaulacorhizid stage into a secondary anau­ the other Scyliorhinidae by their broad root and crown lacorhizid stage. Usually two of the three stages are present base and by their relatively low, triangularly shaped crown. in one species. However, all the three stages can sometimes Furthermore, a strong monognathic heterodonty is present. be observed. Though a clear evolutionary trend in root Schroederichthys is separable from Halaelurus by the pre­ stage developments could not be found, some remarkable sence of poorly developed cusplets, which are absent in features could be noted in root development. Ha/aelurus. The crown development also presents a remarkable diffe­ Cephalurus and the Cephalurus-like SE-Pacific specimen rentiation, which was very useful but the degree of reticu­ have an outstanding root formed by a very flat root, which lated ornamentation and the amount of striae were not has merged root lobes and a concave basal surface. it used. Both are variable features depending on sexual and makes this genus very remote from all the other scyliorhi­ ontogenitic heterodonty. nid genera. Both are separable from each other. Cephalu­ Atelomycterus and Aulohalaelurus have the same extraor­ rus is characterised by the erect or slightly oblique, elonga­ dinary root type and the same shape of principal cusp. The ted and slender cusp, flanked by a poorly developed short and broad root lobes have a basal subquadrangular cusplet, while Cephalurus-Iike SE-Pacific specimens are shape and run almost parallel to each other. However, both characterised by a short inclined principal cusp, up to three genera differ in the shape of the principal cusp and in the mesial and one distal cusplets. amount and development of cusplets flanking the principal cusp : Atelomycterus has one well developed cusplet on Key to the scyliorhinid genera based on odontological each side of the elongated but broad based principal cusp, characters : while Aulohalaelurus has two rather poorly developed cus­ plets on each side of the slender and elongated principal la - Root lobes, in basal view, relatively short and cusp. subquadrate, forming a straight line at root base : 2 Scyliorhinus, Poroderma and Cephaloscyllium share the 2a - Principal cusp elongated but broad based and same massive, short principal cusp, which is relatively flanked by one cusplet. convex on the inner face. - Outer and inner striae well developed : . . . Poroderma is distinguished from Scyliorhinus and Cepha­ ...... Atelomycterus /oscyllium by the vertically high inner part of the root. Plates 7 & 8 Cepha/oscyllium differs from Scyliorhinus by well deve­ 2b - Principal cusp elongated and slender. loped cusplets and short outer costules. - Poorly developed second cusplet present. Apristurus, Galeus and Holohalaelurus have a very similar - Outer and inner striae poorly developed or tooth morphology, which differs from the other genera by absent : ...... Aulohalaelurus the inclination of the principal cusp toward the commis­ Plates 9 & 10 sure. This implies an arched mesial cutting edge of the I b - Root lobes, in basal view, long and nanow : . . 3 principal cusp. Holohalaelurus can be separated from 3a - Outer view : root very low ; root lobes merged, Apristurus and Galeus by less developed striae on inner concave at basal view : ...... 4 and outer faces, which do not reach the apex in Holohalae­ 4a - Principal cusp slender and elongated. lurus. - Cusplets poorly developed, flanking the princi- Apristurus is characterized by two or three cusplets flank­ pal cusp : ...... Cephalurus ing the principal cusp, as compared with one or two in Plates 13 & 14 Gale us. 4b - Principal cusp short, inclined toward commis­ Apristurus, unlike Galeus, has a second well developed sure. cusplet. - Up to three mesial and one distal cusplets : The subgenera Halaelurus and Bythae/urus have such a . . . . . Cepha/urus-like SE-Pacific specimen different tooth morphology that odontologically they could Plates 15 & 16 be considered as separate genera, or even as subfamilies 3b - Root lobes rather plain in basal view : . . . 5 (they will be treated so below). Sa - Toot lobes very long and nanow, forming an Bythae/urus, Hap/oblepharus, Parmaturus and Asymbolus obtuse angle : ...... 6 all share a very strong similarity in tooth morphology. The 6a - Principal cusp massive, with outer face triangu- principal cusp of their teeth do not incline toward the larly and strongly convex : ...... 7 commissure, but are erect. In some cases, they become 7a - Outer face of root very high : Poroderma slightly oblique toward the commissure, which implies that Plates 27 & 28 the mesial cutting edge is not arched nor bended. 7b - Outer face of root low : ...... 8 Minor but stable characteristic features make possible the 8a - Cusplets poorly developed : . . Scyliorhinus separation into subgroups. Plates 31 & 32 Bythae/urus and Parmaturus share a high, transversally 8b - Cusplets well developed. 192 J. HERMAN, M. HOVESTADT-EULER & D.C. HOYESTADT

- Outer basal costules on crown well developed : Despite some clear distinctions, odontology of several ...... Cephaloscyllium groups of the family Scyliorhinidae shows their close rela­ Plates 11 & 12 tionship. The genera of this family are much closer related 6b - Principal cusp slender, with the outer face to each other than for example the Squalid genera. This weakly convex : ...... 9 is obvious through the rather minor differences within the 9a - Principal cusp inclining toward commissure Squalidae. The present example of the Scyliorhinidae indi­ (more clearly vi sable in lateral teeth) : . . . 10 cates that the significance of characters, diagnosing supras­ lOa - Principal cusp strongly inclined toward com­ pecific taxa, should be evaluated more carefully. missure. Two or three elongated, narrow mesial The instability of holaulacorhizy, secondary hemiaulacor­ cusplets : ...... Apristurus hizy and secondary anaulacorhizy, not only on generic or Plates 1 to 4 specific levels but also intraspecifically, indicates the inter­ lOb - Principal cusp weakly inclined toward commis- mediate evolutionary stage of Scyliorhinids. sure : ...... 11 From the odontological point of view, 5 groups could very 11 a - One or two elongated, narrow mesial cusp lets well be seen as subfamilies. (second one less developed) : . . . . . Galeus The first group with Atelomycterus and Aulohalaelurus is Plates 17 & 18 well distinguished from the other genera by the shape of 11 b - Two or three short mesial cusp lets : . . . . the root and the root lobes...... Holohalaelurus The second group with Scyliorhinus, Poroderma and Plates 23 & 24 Cephaloscyllium can be separated by their short massive 9b - Principal cusp always erect or slightly oblique: 12 principal cusp. 12a - High outer depression at crown base : 13 The odontological differentiation within the third group is 13a - Outer striae absent. Outer basal costules on not strongly developed, and tooth morphology is in some crown : ...... Bythaelurus cases rather uniform. However, Apristurus, Galeus and Plates 19 & 20 Holohalaelurus are clearly separable from Bythaelurus, 13b - Striae on inner face of crown, coarse and well Haploblepharus, Parmaturus and Asymbolus, by the way developed. Outer basal costules absent on their lateral teeth are directed toward the commissure. crown : ...... Parmaturus Apristurus and Galeus, as well as Bythaelurus and Parma­ Plate 26 12b - High outer depression at crown base absent: . 14 runts, have a very similar tooth morphology with rather 14a - Secondary cusplets poorly developed or minor differences, respectively, and lumping them into absent : ...... Asymbolus only two genera might be possible. Plates 5 & 6 Halaelurus differs so dramatically from Bythaelurus and 14b - Secondary cusplets well developed : . . . . the other genera of the group, that this genus should be ...... Haploblepharus reallocated to an other group. Plates 21 & 22 The fourth group with Schroederichthys and Halaelurus 5b - Root lobes at basal view very long and narrow, can be separated by extended root lobes, and by low to forming sometimes a nearly straight line instead extremely low triangularly shaped principal cusp both in of a very obtuse angle : ...... 15 lateral and in anterior teeth. Cusplets, if present, are small 15a - Crown low, more or less triangular. Cusplets and poorly developed. poorly developed. Lower teeth : strong outer The fifth group with Cephalurus and the Cephalurus-like costulation, striae absent. Upper teeth : strong SE-Pacific specimen can be separated by their peculiar outer costulation, striae present : ...... concave basal face of the root...... Schroederichthys Five major groups can odontologically be separated by Plates 29 & 30 root development : 15b - Crown low, triangular in anterior teeth. Princi- 1 2 pal cusp poorly developed with extended mesial Atelomycterus Scyliorhinus and distal cutting edges. Cusplets absent : . . . Aulohalaelurus Poroderma ...... Halaelurus Cephaloscyllium Plate 25 3 4 Apristurus Schroederichthys PRELIMJNARY CONCLUSIONS Galeus Halaelurus I-1 olohalaelurus It has always been problematic

Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 193

Ontogenitic heterodonty is strong in Scyliorhinidae. The Acknowledgements teeth of juveniles always possess crowns with a very flat outer face. Cusplets are always merged with the cutting We would like to thank Dr. M.L. BAUCHOT, Musee National edges of the principal cusp and are very flat on both inner d'Histoire naturelle, Paris ; Dr. M. BoESEMAN, Rijksmuseum van and outer faces. Natuurlijke Historie, Lei den ; Dr. G .. BuRG ESS, Florida State (See plate 35). Museum ; Dr. J.L.V. CoMPAGNO, formerly San Francisco State University, Cali fornia ; Dr. W.N. EscHMEYER, California Academy The squalid genera Centroscyllium and Aculeo/a have true of Sciences, San Francisco ; Dr. J.P. GossE, Institut Royal de but modified anaulacorhizid root types, as it is usual for Sciences naturelles de Belgique, Brussels; Mrs. S.L. JEWETI, the Squalifonnes, and far remote from Scyliorhinidae. Unites State National Museum of Natural History ; B. SERET, However both have more or less scyliorhinid-like crowns, formerly 0RSTOM, Dakar; Dr. V.G. SPRI NGER, United States which may indicate a presently unexplained similarity with National Museum, New York and A. WHEELER, British Museum the Scyliorhinidae (HERMAN et al. , 1990). of Natural History, London, for the permission to examine speci­ mens at their disposal. Dr. M. STEHMANN provided specimens of I.S.H., Hamburg. We also would like to thank Captain P. GuEGUEN (Lorient) for allowing us to collect specimens during his cruises. We thank particularly Miss F. LADEUZE, F.N.R.S., Brussels, for her technical assistance and careful corrections of the proofs. The SEM-photographs were take by Mr. J. CiLus, Institut Royal des Sciences naturelles de Belgique, Brussels, and were printed by Mr. H. STOUT, Brussels and our friends Mr. G. BROGNET, Mr. J. DEG REEF and Mr. M. VALLE, Brussels.

References

CAPPETIA, H., 1987. Chondrichtyes 2. Mesozoic and cenozoic STEHMANN, M., elasmobranchii. In Handbook of Paleoichthyology (ed. H.P. Aussenstelle lchthyologie ScHULZE). Vol. 3B : 193 pp. des Instit uts fUr Seefischerei, CAS lER, E., 1947. Constitution et evolution de Ia racine dentaire c/o Zoologisches Museum des Euselachii. Bulletin du Musee royal d' Histoire naturel/e de der Un iversitat Hamburg, Belgique, Bruxelles, 23, no. 13, 14, 15, 32 et 45 pp., 5 pl. Martin-Luther-King-Platz 3, D-2000 Hamburg 13, CoMPAGNO, L.J.V., 1984. FAO species catalogue Vol. 4. Sharks Fed. Rep!. of Germany of the world. An annotated and illustrated catalogue of shark species known to date. Part 2. Carcharhini fo rmes. FA O Fishery HERMAN, J., Synopsis, ( 125) Vol. 4. Part 2 : VIII + 655 pp. Service Geologique de Belgique, CoMPAGNO, L.J. V., 1988. Sharks of the Order Carcharhiniformes. Rue Jenner 13, Princeton University, Princeton, New Jersey, 486 pp., 35 pl. B-1 040 Brussels, Belgium HERMAN, J., HovESTADT-EULER, M. and HovESTADT, D.C., 1990. Order: Squaliformes. Families : Echinorhinidae, Oxynotidae and 1-IOVESTADT-EULER, M. Squalidae. In : Contributions to the study of the comparati ve and HovESTADT, D.C., morphology of teeth and other relati ve ichthyodorulites in li ving Merwedelaan 6, supraspecific taxa of Chondrychthyan fi shes. Ed. Stehmann, M. NL-4535ET Terneuzen, Bulletin de I' lnstitut Royal des Sciences Naturelles de Belgique, The Netherlands Biologie, 59: 101 - 157. · SPRINGER, S., 1979. A revision of the catsharks Family Scyliorhi ­ nidae. Nati onal Oceanic and Atmospheric Adminisn-ation Techni­ cal report NM FS, Circular 422. 194 J. HERMAN, M. HOVESTADT-EULER & D.C. HOVESTADT

Glossary (also applying to previous issues of thi s seri es, replacing the previous glossari es)

CONCERNJNG THE JAW Hemiaulacorhizid Vascularisation through a median and 1 or 2 lateral Anterior foramina on inner face (like Squatinidae and Orectolo­ Tooth position close to the junction of the left and bidae). right jaw parts. Holaulacorhizid Commissural Vascularisation through many small foramina concen­ Tooth position next to the end of jaw. trated in a median groove running from outer to inner face (like Rajidae). Dignathic Heterodont by having different tooth morphology in Polyaulacorhizid upper and lower jaws. Vascularisation through many small foramina concen­ trated in several grooves running parallel from the File outer to the inner face and next to the crown-root Tooth row from symphys is toward the end of jaw. junction, on both inner and outer faces (like Myliobati­ Heterodonty dae, etc.). Different tooth morphology within a set of teeth. There Apron are two types of heterodonty : dignathic and monogna­ Expansion of the central part of the outer crown base. thic. Basal Homodonty Bottom face concerned. Equal tooth morphology within a set of teeth. Costule Lateral Short, vertical ridge sometimes present on inner and/ Tooth position half way along the jaw. or outer crown base. Longitudinal Crown Symphysial/commissural direction of a file. Enamelated tooth part. Distal Monognathic Tooth edge or part toward the angle of jaw. Heterodonty within one jaw only (this can be gradient or di sjunct). Inner face Viewed from inside the mouth. Parasymphysial First anterior tooth row, if a symphysial tooth row is Longitudinally Apico/basally directed. absent. Median groove Posterior Groove running from the inner root base to the inner Tooth position toward the angle of jaw. crown-root junction, dividing anholaulacorhizid type Pseudosymphysial of root into two root lobes. It includes the main fora­ One of the parasymphysial tooth rows becomes the mina of the vascularisation system. character of a sy mphysial tooth row (symmetry). This Median keel phenomenon appears in some species. Transverse ridge dividing the crown into inner and Row outer faces. Tooth row from the inner to the outer face of jaw. Mesial Symphysial Tooth edge or part toward the junction (symphysis) Teeth at junction of both halves of a jaw. of left and right jaw halves. Outer face Transversal Viewed from outside the mouth. Outer/inner direction of a row. Pseudoapron CONCERNING THE TOOTH Apron-like vertical ridges that appear sometimes on lateral and posterior teeth. Considering their vascularisation, E. Casier ( 194 7) Root recogni sed and described 4 phylogenetically characte­ Non-enamelated tooth part that forms the junction with ri sti c root forms of elasmobranch teeth. the jaw and provides vascularisation of the tooth. Anaul acorhizid Striae Vascularisation through scattered fo ramina of equal Vertical ridge running from the base toward the apex size on both outer and inner faces (like Hexanchidae). of the crown. Contributions to the study of the comparative morphology of teeth and other relevant ichthyodorulites 195

Secondary anaulacorhizy Composition of the plates Syncreted median groove of holaulacorhizid type of root. As far as possible, one plate with SEM-photographs of isolated teeth is presented fo r each of the genera and subgenera. Secondary hemiaulacorhizy Upper teeth are pre ented wi th the principal cusp downwards. Beginning syncretion of a median groove of a holaula­ lower teeth with the principal cusp upwards. corhizid type of root. The choice of left or right jaw halves depends onl y on the quality of preservat ion of the specimens' tooth fi les. Sulcus The upper scale bar refers to the upper teeth, the lower scale bar Groove developed by the main vascularisation string to the lower teeth. Photographs of posterior and commissural teeth leading from root base to the main foramen on anala­ mostly are enlarged and have their own scale bar. corhizid root types. It differs from the median groove of the holaulacorhizid root type and from the parallel Legend grooves of the polyaulacorhizid root type, in which several foramina are concentrated, respectively. s Symphysial position a Anterior position Transversal al Antero-lateral position Mesio-/distally directed. Lateral position lp Latero-posterior position Uvula p Posterior position Lobate extension of the inner crown base. c Commissural position \0 0\

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Plate 2 1. - Haploblepharus edwardsii (VOIGT, 1 832), male 50 em (t.l.), Capetown, South Africa. Contributions to the study of the comparati ve morphology of teeth and other relevant ichthyodorulites 21 7

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