Hybodontidae teeth and spines from the Lourinha Formation, Sobral unit are described. These teeth and spines have been
ascribed to the genus Hybodus and regarded as Hybodus cf. reticulatus

Balbino, A. C.

English

New University of Lisbon's Repository

Upper Jurassic Hybodontidae (Selachii) from Lourinha, Portugal
A. c. Balbino
Projecto "POCTI/3653 I/PAL/2000 - Studies on Portuguese Paleontology (Post - Paleozoic)". Fundacao para a Ciencia e a Tecnologial
Ministerio da Ciencia e do Ensino Superior.
Centro de Estudos Geologicos, Faculdade de Ciencias e Tecnologia (UNL)/Quinta da Torre 2829-516 Caparica, Portugal. Departamento de
Geociencias, Universidade de Evora, Apartado 94, 7002-554 Evora, Portugal. acaceres @ueYora.pt
Abstract
Key-words: Hybodontidae; Upper Jurassic; Lourinha.
Hybodontidae teeth and spines from the Lourinha Formation, Sobral unit are described. These teeth and spines have been
ascribed to the genus Hybodus and regarded as Hybodus cf. reticulatus.
Resumo
Palavras-chave: Hybodontidae; Jurassico superior; Lourinha,
Sao descritos dentes e espinhos de Hybodontidae colhidos em depositos da Formacao da Lourinha, unidade Sobral, do genero
Hyb odus , representado por Hybodus cf. reticulatus.
1. Introduction
Mesozoic shark remnants are known in Portugal since
long ago. However, jurassic shark material is scarce, and
have mostly been overlooked. As far as we can ascertain,
the first reference has been produced by Sauvage
(1897-1898: 10): a single tooth collected at Santa Cruz
"dans le Malm superieur" has been briefly referred (without
any figure) by him to "Hybodus aff. polyprion Ag.".
Another, isolated Asteracanthus tooth was collected
in the older beds at the Fonte Quente limestone quarry
close by the Tomar to Pedreira road, ca 2 Km from this
last locality. Limestones have been ascribed to the
Hammatoceras insigne main level , upper Toarcian
(Antunes , 1967: 13).
Later on, an isolated Orthacodus (Sphenodus) sp. tooth
was collected by G.Manuppella at Ribeira do Barranco in
Algarve, upper Jurassic (determ. by M. T. Antunes;
unpublished so far).
On the other hand, a lot ofexcavation and other work
has been carried on upper Jurassic near Lourinha. The most
remarkabl e results concern the discovery and collecting
of dinosaur material, most of which is being kept at the
GEAL/Lourinha Museum (see Antunes & Mateus, 2003;
Manuppella, 1996 and Mateus, 1998). An account was
presented in a Colloquium held at theAcademy ofSciences
from Lisbon - "Upper Jurassic Palaeoenvironments in
Portugal- Geology,plants, eggs & Dinosaurs, mammals",
April 1998. Most of dinosaur the specimens have been
found in the Sobra1 Member of Lourinha Formation,
45
Ciencias da Terra (UNL) , 15
Genus Hybodus Agassiz, 1837
2. Systematics
Hybodus cf. reticulatus Agassiz, 1837
(Text-fig.1 ;PI.1, fig. 1-2; Pl. 2, fig. 1-4)
corresponding to a lower deltaic plain dated from early
Tithonian. However, other fossils were collected there,
among them the shark remnants under study.
Quite large-sized, upper Jurassic sharks ascribed to
Hybodus cf. reticulatus Agassiz, 1837 are represented at
Peralta and Porto das Barcas (near Lourinha) by mostly
incomplete teeth and dorsal fin spines.
Localities: Peralta and Porto das Barcas.
Material: 33 (some incomplete) teeth and spines.
Description: The best preserved teeth are small, with a
straight main cusp. Enamel presents several parallel, well
marked and straight folds, from the crown's base until nearly
half of the height of the main cusp.
The cusps has well defined cutting edges. The lateral
cusplets are small, pointed and folded on both faces .
The crown's labial face does not overhang the root.
The not very thick root is nearly perpendicular to the crown.
The basal face is flat and slightly depressed in the
labial face, which bears many foramina ; the lingual face is
rather convex .
Dorsal fin spines show several longitudinal crests
presenting small nodules (PI. 2, fig. 3). Some smaller spines
also have longitudinal crests; their posterior edges bear a
double row of denticles (PI. 2, fig. 2). Spines are roughly
triangular in dorsal section .
3. Conclusions
Discussion: The fossil teeth under study have been
compared to those from Hybodus plicatilis, H. sublaevis,
H. cuspidatus andH. marginalis figured by Agassiz (1836).
There are some differences as far as the crown, the crests
and the number of cusplets are concerned.
In H. grossiconus Delsate, 1995 teeth have a thicker
and broader crown, with thicker and more numerous cusps
than in our specimens. There are also significant differences
between Lourinha's specimens and the H. obtusus and H.
grossiconus teeth shown by Priem (1908) .
The abo ve proposed classification is a result of
comparison with teeth from Hybodu s reticulatus Agassiz,
1837 kept at the Laboratoire de Paleontologie collections
(Museum National d'Histoire Naturelle de Paris).
This work concerns the N° 1 Line of the Centro de
Estudos Geologicos, Universidade Nova de Lisboa, as
well as the POCTI/36531/PAL/2000 Project "Studies on
Portuguese Paleontology (Post-Paleozoic)".
Observation ofJurassic selachian material was possible
owing to the courtesy of P. Taquet and D. Goujet at the
Museum National d'Histoire Naturelle de Paris .
At the GEAL/ "Museu da Lourinha", H. Mateus
(Curator) allowed the study of specimens from this
Institution, and O. Mateus contributed with valuable data.
The author thanks Prof. M. T. Antunes for his critical
reading of this article .
Acknowledgements
CHONDRICHTYES Huxley, 1880
ELASMOBRANCHII Bonaparte , 1838
Euselachii Hay, 1902
Hybodontoidea Zangerl, 1981
Hybodontidae Owen, 1846
This genus is known from complete skeletons and
skin impressions as well as by a large number of isolated
teeth. The body is long and massive, with a subterminal
mouth (fig. 1).
The two dorsals arc large and supported by a powerful
anterior fin spine. In certain species, the fin spines are
ornamented with rounded tubercles.
The teeth are rather mesio-distally expanded
and have a straight main cusp. The lateral cusp lets
are generally numerous, straight, pointed, broadly
united at their base and decreasing in size toward the
margins ofthe tooth. In certain species they are very short.
The cusp has more or less convex labial and lingual
faces and well defined cutting edges; it shows strong and
high enamel folds; the cusplets are folded too. The root
is not very thick; it is nearly perpendicular to the
crown with a rather prominent labial face; the basal face
is flat, often slightly depressed in its labial part which
presents many foramina; the lingual face is generally
rather convex.
The type speci es is H. reticulatus Agassiz 1837.
Cappetta (1987) includes in the g~nus Hybodus the
following species: H. huangnidanensis Wang 1977; H.
multiconus Jaekel 1889; H. nevadensis Wemple 1906;
H. plicatilis Agassiz 1843; H. rapax Stensio 1921 and
H. shastensis Wemple 1906.
Class
Subclass
Cohort
Superfamily
Family
The family Hybodontidae, known as early as in late
Paleozoic is especially well represented in the Triassic
and in the Rhaetic. It persists as late as Maastrichtian
(Cappetta, 1987). Several genera are included in this
family. Dentitions are oftearing-type in some genera such
as Hybodus, while in other genera, as in Acrodus,
dentitions are ofgrinding-type.
Cappetta 1987 includes in this family the genera
Hybodus, Priohybodus, and Pororhiza .
The Hybodontidae are known in Europe, North
America, Northern and Western Africa, and Asia
(Cappetta, 1987).
46
Ciencias da Terra (UNL), 15
Fig. 1 - Restoration ofHybodus sp. after Maisey (1982), in Cappetta (1987).
References
Antunes , M. T. (1967) - Urn Mesosuquiano do Liasico de Tomar (Portugal) / Consideracoes sabre a origem dos Crocodilos. Mem .
ServoGeol. Portugal, Lisboa, N.S., 13,66 p., 6 est.
Antunes, M. T. & Mateus, o. (2003) - Dinosaurs of Portugal. Comptes Rendus Palevol, Paris, 2(1) : 77-95.
Agassiz, L. (1843) - Recherches sur les Poissons fossiles. Neuchatel. Tome III, 389 p., 44 pI.
Cappetta, H. (1987) - Chondrichthyes II. Mesozoic and Cenozoic Elasmobranchii. In Handbook ofPaleoichthyology, Gustav Fischer
Verlag, Stuttgart, III B, 193 p.
Delsate , D. (1995) - Chondrichthyens mesozoiques du Grand Duche de Luxembourg. Elasmobranches et Stratigraphie. Ministere
des Affaires Economiques. Profess ional paper 1995/3, 278, VIII, 278 p., 48 fig. , 74 pI.
Manupp ella, G.(Coord .) (1996) - Carta geologica de Portugall/50 000. Folha 30-A, Lourinha, Instituto Geologico Mineiro, Lisboa.
Mateus , O. (1998) - Lourinhanosaurus antunesi, a new Upper Jurassic allosauro id (Dinosauria:Theropoda) from Lourinha (Portugal) .
Memorias Academia Ciencias Lisboa, 37: 111-124.
Sauvage, H. E. (1897-1898) - Vertebres fossiles du Portugal- Contributions a l'etude des Poissons et des Reptiles du Jurassique
et du Cretacique, Direction Travaux Geologiques Portugal, Lisbonne, Imprimerie de I' Academic Royale des Sciences,
47 p., 10 pI.
47
Ciencias da Terra (UNL), 15
Plate 1
Fig. 1 - 2 - Hybodus cf. reticulatus Agassiz, 1837
Fig. 1 - Lateral teeth : a, labial view; b, lingual view; c, occlusal view (Porto das Barcas).
Fig . 2 - Lateral teeth: a, labial view ; b, lingual view (Peralta); c, occlusal view (Porto das Barcas).
48
Ciencias da Terra (UNL). 15
PLAT E J
49
Ciincias cia Terra (UNL). 15
Plate 2
Fig. 1 - 4 - Hybodus ef. reticulatus Agassiz, 1837
Fig. I - Lateral teeth: a, labial view; b, lingual view (Peralta).
Fig. 2 - 4 - Dorsal fin spines, profil view (Peralta).
50
Ciincias da Terra (UNL). 15
PLATE 2
51


Upper Jurassic Hybodontidae (Selachii) from Lourinhã, Portugal

Repositório da Universidade Nova de Lisboa

Upper Jurassic Hybodontidae (Selachii) from Lourinha, PortugalA. c. BalbinoProjecto "POCTI/3653 I/PAL/2000 - Studies on Portuguese Paleontology (Post - Paleozoic)". Fundacao para a Ciencia e a TecnologialMinisterio da Ciencia e do Ensino Superior.Centro de Estudos Geologicos, Faculdade de Ciencias e Tecnologia (UNL)/Quinta da Torre 2829-516 Caparica, Portugal. Departamento deGeociencias, Universidade de Evora, Apartado 94, 7002-554 Evora, Portugal. acaceres @ueYora.ptAbstractKey-words: Hybodontidae; Upper Jurassic; Lourinha.Hybodontidae teeth and spines from the Lourinha Formation, Sobral unit are described. These teeth and spines have beenascribed to the genus Hybodus and regarded as Hybodus cf. reticulatus.ResumoPalavras-chave: Hybodontidae; Jurassico superior; Lourinha,Sao descritos dentes e espinhos de Hybodontidae colhidos em depositos da Formacao da Lourinha, unidade Sobral, do generoHyb odus , representado por Hybodus cf. reticulatus.1. IntroductionMesozoic shark remnants are known in Portugal sincelong ago. However, jurassic shark material is scarce, andhave mostly been overlooked. As far as we can ascertain,the first reference has been produced by Sauvage(1897-1898: 10): a single tooth collected at Santa Cruz"dans le Malm superieur" has been briefly referred (withoutany figure) by him to "Hybodus aff. polyprion Ag.".Another, isolated Asteracanthus tooth was collectedin the older beds at the Fonte Quente limestone quarryclose by the Tomar to Pedreira road, ca 2 Km from thislast locality. Limestones have been ascribed to theHammatoceras insigne main level , upper Toarcian(Antunes , 1967: 13).Later on, an isolated Orthacodus (Sphenodus) sp. toothwas collected by G.Manuppella at Ribeira do Barranco inAlgarve, upper Jurassic (determ. by M. T. Antunes;unpublished so far).On the other hand, a lot ofexcavation and other workhas been carried on upper Jurassic near Lourinha. The mostremarkabl e results concern the discovery and collectingof dinosaur material, most of which is being kept at theGEAL/Lourinha Museum (see Antunes & Mateus, 2003;Manuppella, 1996 and Mateus, 1998). An account waspresented in a Colloquium held at theAcademy ofSciencesfrom Lisbon - "Upper Jurassic Palaeoenvironments inPortugal- Geology,plants, eggs & Dinosaurs, mammals",April 1998. Most of dinosaur the specimens have beenfound in the Sobra1 Member of Lourinha Formation,45Ciencias da Terra (UNL) , 15Genus Hybodus Agassiz, 18372. SystematicsHybodus cf. reticulatus Agassiz, 1837(Text-fig.1 ;PI.1, fig. 1-2; Pl. 2, fig. 1-4)corresponding to a lower deltaic plain dated from earlyTithonian. However, other fossils were collected there,among them the shark remnants under study.Quite large-sized, upper Jurassic sharks ascribed toHybodus cf. reticulatus Agassiz, 1837 are represented atPeralta and Porto das Barcas (near Lourinha) by mostlyincomplete teeth and dorsal fin spines.Localities: Peralta and Porto das Barcas.Material: 33 (some incomplete) teeth and spines.Description: The best preserved teeth are small, with astraight main cusp. Enamel presents several parallel, wellmarked and straight folds, from the crown's base until nearlyhalf of the height of the main cusp.The cusps has well defined cutting edges. The lateralcusplets are small, pointed and folded on both faces .The crown's labial face does not overhang the root.The not very thick root is nearly perpendicular to the crown.The basal face is flat and slightly depressed in thelabial face, which bears many foramina ; the lingual face israther convex .Dorsal fin spines show several longitudinal crestspresenting small nodules (PI. 2, fig. 3). Some smaller spinesalso have longitudinal crests; their posterior edges bear adouble row of denticles (PI. 2, fig. 2). Spines are roughlytriangular in dorsal section .3. ConclusionsDiscussion: The fossil teeth under study have beencompared to those from Hybodus plicatilis, H. sublaevis,H. cuspidatus andH. marginalis figured by Agassiz (1836).There are some differences as far as the crown, the crestsand the number of cusplets are concerned.In H. grossiconus Delsate, 1995 teeth have a thickerand broader crown, with thicker and more numerous cuspsthan in our specimens. There are also significant differencesbetween Lourinha's specimens and the H. obtusus and H.grossiconus teeth shown by Priem (1908) .The abo ve proposed classification is a result ofcomparison with teeth from Hybodu s reticulatus Agassiz,1837 kept at the Laboratoire de Paleontologie collections(Museum National d'Histoire Naturelle de Paris).This work concerns the N° 1 Line of the Centro deEstudos Geologicos, Universidade Nova de Lisboa, aswell as the POCTI/36531/PAL/2000 Project "Studies onPortuguese Paleontology (Post-Paleozoic)".Observation ofJurassic selachian material was possibleowing to the courtesy of P. Taquet and D. Goujet at theMuseum National d'Histoire Naturelle de Paris .At the GEAL/ "Museu da Lourinha", H. Mateus(Curator) allowed the study of specimens from thisInstitution, and O. Mateus contributed with valuable data.The author thanks Prof. M. T. Antunes for his criticalreading of this article .AcknowledgementsCHONDRICHTYES Huxley, 1880ELASMOBRANCHII Bonaparte , 1838Euselachii Hay, 1902Hybodontoidea Zangerl, 1981Hybodontidae Owen, 1846This genus is known from complete skeletons andskin impressions as well as by a large number of isolatedteeth. The body is long and massive, with a subterminalmouth (fig. 1).The two dorsals arc large and supported by a powerfulanterior fin spine. In certain species, the fin spines areornamented with rounded tubercles.The teeth are rather mesio-distally expandedand have a straight main cusp. The lateral cusp letsare generally numerous, straight, pointed, broadlyunited at their base and decreasing in size toward themargins ofthe tooth. In certain species they are very short.The cusp has more or less convex labial and lingualfaces and well defined cutting edges; it shows strong andhigh enamel folds; the cusplets are folded too. The rootis not very thick; it is nearly perpendicular to thecrown with a rather prominent labial face; the basal faceis flat, often slightly depressed in its labial part whichpresents many foramina; the lingual face is generallyrather convex.The type speci es is H. reticulatus Agassiz 1837.Cappetta (1987) includes in the g~nus Hybodus thefollowing species: H. huangnidanensis Wang 1977; H.multiconus Jaekel 1889; H. nevadensis Wemple 1906;H. plicatilis Agassiz 1843; H. rapax Stensio 1921 andH. shastensis Wemple 1906.ClassSubclassCohortSuperfamilyFamilyThe family Hybodontidae, known as early as in latePaleozoic is especially well represented in the Triassicand in the Rhaetic. It persists as late as Maastrichtian(Cappetta, 1987). Several genera are included in thisfamily. Dentitions are oftearing-type in some genera suchas Hybodus, while in other genera, as in Acrodus,dentitions are ofgrinding-type.Cappetta 1987 includes in this family the generaHybodus, Priohybodus, and Pororhiza .The Hybodontidae are known in Europe, NorthAmerica, Northern and Western Africa, and Asia(Cappetta, 1987).46Ciencias da Terra (UNL), 15Fig. 1 - Restoration ofHybodus sp. after Maisey (1982), in Cappetta (1987).ReferencesAntunes , M. T. (1967) - Urn Mesosuquiano do Liasico de Tomar (Portugal) / Consideracoes sabre a origem dos Crocodilos. Mem .ServoGeol. Portugal, Lisboa, N.S., 13,66 p., 6 est.Antunes, M. T. & Mateus, o. (2003) - Dinosaurs of Portugal. Comptes Rendus Palevol, Paris, 2(1) : 77-95.Agassiz, L. (1843) - Recherches sur les Poissons fossiles. Neuchatel. Tome III, 389 p., 44 pI.Cappetta, H. (1987) - Chondrichthyes II. Mesozoic and Cenozoic Elasmobranchii. In Handbook ofPaleoichthyology, Gustav FischerVerlag, Stuttgart, III B, 193 p.Delsate , D. (1995) - Chondrichthyens mesozoiques du Grand Duche de Luxembourg. Elasmobranches et Stratigraphie. Ministeredes Affaires Economiques. Profess ional paper 1995/3, 278, VIII, 278 p., 48 fig. , 74 pI.Manupp ella, G.(Coord .) (1996) - Carta geologica de Portugall/50 000. Folha 30-A, Lourinha, Instituto Geologico Mineiro, Lisboa.Mateus , O. (1998) - Lourinhanosaurus antunesi, a new Upper Jurassic allosauro id (Dinosauria:Theropoda) from Lourinha (Portugal) .Memorias Academia Ciencias Lisboa, 37: 111-124.Sauvage, H. E. (1897-1898) - Vertebres fossiles du Portugal- Contributions a l'etude des Poissons et des Reptiles du Jurassiqueet du Cretacique, Direction Travaux Geologiques Portugal, Lisbonne, Imprimerie de I' Academic Royale des Sciences,47 p., 10 pI.47Ciencias da Terra (UNL), 15Plate 1Fig. 1 - 2 - Hybodus cf. reticulatus Agassiz, 1837Fig. 1 - Lateral teeth : a, labial view; b, lingual view; c, occlusal view (Porto das Barcas).Fig . 2 - Lateral teeth: a, labial view ; b, lingual view (Peralta); c, occlusal view (Porto das Barcas).48Ciencias da Terra (UNL). 15PLAT E J49Ciincias cia Terra (UNL). 15Plate 2Fig. 1 - 4 - Hybodus ef. reticulatus Agassiz, 1837Fig. I - Lateral teeth: a, labial view; b, lingual view (Peralta).Fig. 2 - 4 - Dorsal fin spines, profil view (Peralta).50Ciincias da Terra (UNL). 15PLATE 251

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Upper Jurassic Hybodontidae (Selachii) from Lourinhã, Portugal

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