THE WEST GONDWANAN OCCURRENCE OF

THE HYBODONTID SHARK PRIOHYBODUS,

AND THE LATE JURASSIC ± EARLY CRETACEOUS
AGE OF THE TACUAREMBOÂ FORMATION, URUGUAY

by D A N I E L P E R E A , M A R T ÂI N U B I L L A , A L E J A N D R A R O J A S and C EÂ S A R A. GOSO

ABSTRACT. Priohybodus cf. P. arambourgi is reported for the ®rst time from the Tacuarembo Formation of Uruguay.
This species is a hybodontid shark known previously only from Upper Jurassic and Lower Cretaceous deposits of
Saharan Africa and the Arabian Peninsula. The material (22 isolated teeth and a dorsal ®n spine) was found in a thin
bone bed, associated with abundant bone fragments, scales and teeth of semionotiform ®shes and theropod dinosaurs.
Until now, the age of the Tacuarembo Formation has been dif®cult to determine because its fossil content lacked useful
biostratigraphic indicators. The ®nding of Priohybodus cf. P. arambourgi in that unit greatly expands the
palaeobiogeographic range of the species, and allows us to propose a Late Jurassic±Early Cretaceous age for the
Tacuarembo Formation.

KEY WORDS: Priohybodus, Tacuarembo Formation, Jurassic±Cretaceous, Uruguay.

P R I O H Y B O D U S A R A M B O U R G I d'Erasmo, 1960 is a hybodontid shark only known, until now, from

Upper Jurassic and Lower Cretaceous deposits of Saharan Africa and the Arabian Peninsula (d'Erasmo
1960; Tabaste 1963; Goodwin et al. 1999; Duf®n in press). During recent ®eld work, 22 teeth and a dorsal
®n spine of Priohybodus cf. P. arambourgi were found in Mesozoic rocks in Uruguay (Text-®g. 1; Pl. 1).
This is the ®rst record of the taxon in West Gondwana and the ®rst Mesozoic shark to be discovered in
Uruguay. P. arambourgi was previously only found in the Lugh `Series' (Upper Jurassic), Somalia, the
Lower `Wealden' (Lower Cretaceous) of South Tunisia (d'Erasmo 1960; Tabaste 1963), the Tithonian
(Upper Jurassic) of Ethiopia (Goodwin et al. 1999), the Albian (Lower Cretaceous) of Libya, and the
Kimmeridgian±Tithonian of North Yemen (Duf®n in press). The Uruguayan specimens (Pl. 1) are almost
complete teeth collected from thin bone beds in the Tacuarembo Formation, Tacuarembo Department
(Text-®g. 1). They are associated with abundant bone fragments, scales and teeth of semionotiforms, and
theropod teeth.
This paper focuses on an analysis of the morphological variation in teeth of Priohybodus, its
chronostratigraphic and biogeographic value in the south-western Gondwana-Parana Basin, and on the
environmental conditions indicated by the associated fossil assemblage.

GEOLOGICAL SETTING
The Tacuarembo Formation is correlated with the Botucatu Formation of Brazil (FrancËa et al. 1995), both
of which crop out in the Parana Basin. Throughout the whole basin, Triassic ¯uvial±lacustrine deposits are
succeeded by sediments of a large desert that covered it and much of south-western Gondwana. The
Triassic rocks unconformably overlie continental Upper Permian (Paso Aguiar and YaguarõÂ formations).
By the end of the Jurassic, the break up of Gondwana was characterized in the Parana Basin by the
extrusion of tholeiitic basalts. These volcanic rocks constitute the Serra Geral Basalt (FrancËa et al. 1995),
which in Uruguay is called the Arapey Formation (Bossi and Navarro 1991).
In the North-Western Basin of Uruguay, the Tacuarembo Formation is characterized by ¯uvial systems
[Palaeontology, Vol. 44, Part 6, 2001, pp. 1227±1235, 1 pl.] q The Palaeontological Association
1228 PALAEONTOLOGY, VOLUME 44

TEXT-FIG. 1. Geographic location of the Uruguayan specimens of Priohybodus cf. P. arambourgi.

consisting of a mixture of wadi and terminal fan deposits (Preciozzi et al. 1985). The sandstone facies
association suggests a ¯uvial incision into an underlying aeolian sequence. Fine to medium-grained sands
with cross and ripple lamination dominate the sequence. The ¯uvial system supplied the bioclasts which
form the bone beds at the base of the section, and mollusc-rich levels (Text-®g. 2).
The Tacuarembo Formation crops out mainly in a discontinuous north-south trending belt in the
Uruguayan departments of Tacuarembo and Rivera. According to Sprechmann et al. (1981), two unnamed
members can be distinguished within the formation. In the lower member subhorizontal strati®cation
predominates, although there are also small ripples, ¯aser, and cross-bedded strati®cation (angles up to 30
degrees). There is an alternation of different lithologies, mainly ®ne- and medium-grained sandstones,
siltstones, shales, and mudstones. These were deposited in subaquatic continental conditions, and are
fossiliferous. The upper member is characterized by the presence of unfossiliferous sandstones produced
by the accumulation of sand dunes in arid conditions. The main sedimentary structures are tabular-planar
and cuneiform-planar, cross-bedded strati®cation. The average dip of these structures is 21´2 degrees.

MATERIAL AND METHODS

The material examined is deposited in the vertebrate fossil collection, Facultad de Ciencias, Montevideo,
Uruguay (FC-DPV). The teeth were isolated from the sediment in the laboratory using both thermal and
screen washing techniques. The rock fragments collected in the ®eld were dried for several days in an oven
at 808C and immediately submerged in water at room temperature. After one day, as a result of the sudden
change of temperature, the sandstone was weakened enough to be completely disaggregated gently with
the ®ngers on a sieve in a sink ®lled with water. The residue was picked and dried in small plastic boxes.
Institutional abbreviations. BMNH, British Museum of Natural History; FC-DPV, Departamento de PaleontologõÂa,
Facultad de Ciencias, Uruguay; IGF, Museo Geologico di Firenze; UCMP, University of California Museum of
Paleontology.
 PEREA ET AL.: MESOZOIC SHARK 1229

 Formation). The arrow shows the level at

TEXT-FIG. 2. Vertical stratigraphic section of the outcrop studied (Tacuarembo
which Priohybodus cf. P. arambourgi was found.
1230 PALAEONTOLOGY, VOLUME 44

SYSTEMATIC PALAEONTOLOGY
Class EUSELACHII Hay, 1902
Family HYBODONTIDAE Owen, 1846
Genus PRIOHYBODUS d'Erasmo, 1960

Type species by monotypy. Priohybodus arambourgi d'Erasmo, 1960 from the Lugh `Series', Upper Jurassic of
Somalia and the Lower Cretaceous (lower `Wealden') of southern Tunisia.

Priohybodus cf. P. arambourgi d'Erasmo, 1960

Plate 1, ®gures 1±9
*1960 Priohybodus arambourgi d'Erasmo, p. 11, pl. 1, ®gs 1±1a, 2±3.
1963 Priohybodus arambourgi; Tabaste, p. 459, pl. 7, ®g. 1.
1987 Priohybodus arambourgi; Cappetta, p. 32, ®g. 38F.
1999 Priohybodus arambourgi; Goodwin et al., ®g. 4I.
in press Priohybodus arambourgi; Duf®n.

Holotype. IGF 6600, a fragmentary, isolated crown (d'Erasmo 1960, pl. 1, ®gs 1±1a).

Type locality. 6±7 km south of El Mao, Somalia (d'Erasmo 1960, p.11).

Type horizon. Lugh `Series', Late Jurassic.

Other described and ®gured material. Original specimen of Tabaste (1963, pl. 7, ®g. 1); BMNH P44739, an isolated
fragmentary crown from the Lugh `Series' of Somalia (Cappetta 1987, ®g. 38F); JS-1/UCMP 170798, a partial crown
from the Mugher Mudstone (Tithonian) of Ethiopia (Goodwin et al. 1999). Nearly 200 isolated complete and
fragmentary crowns in the Lenuzza collection, including BMNH P65461, a complete tooth lacking only the tip of the
central cusp, and P65462 to P65467 inclusive; an incomplete dorsal ®n spine, P65468 (Duf®n in press).

New material. 22 tooth crowns, some fragmentary: FC-DPV-903±909 and FC-DPV-1002±1016. The greater part of
the distal portion of a dorsal ®n spine: FC-DPV-1049.

Locality and horizon. Tacuarembo Department, near Martinote, 10 km west of Batovõ hills (Text-®g. 1), in a thin bone
bed within yellowish-white and pink sandstones of the Tacuarembo Formation (Text-®g. 2).

Description. The teeth range from 5´9 mm to 18 mm wide (see Pl. 1 and Table 1). They are symmetrical, with a broad,
triangular, smooth and not very thick cusp, on each side of which there are two (rarely three) pairs of lateral cusplets
that are very divergent and asymmetrical. The cutting edges are regularly indented by ®ne rounded serrations (c. 4´0
per mm). The roots are not preserved. The central cusp cross-section is compressed, with the labial face being much
¯atter than the relatively more convex lingual face. The central cusp and lateral cusplet bases are con¯uent. The ®rst
lateral cusplet pair is approximately one-third of the height of the central cusp. The cusplets are slightly asymmetrical,
with a ¯attened labial face and a rather more convex lingual face. The central cusp is upright to only slightly distally
inclined and is slightly more slender in some cases (cusp height: cusp base ratio 1´28 compared to 1´01 in more obtuse
cusps).
The spine fragment is 56´2 mm in length (Pl. 1). The tip of the spine is missing, but the overall pro®le shows a

EXPLANATION OF PLATE 1
Figs 1±9. Priohybodus cf. P. arambourgi. 1±8, teeth. 1±2, FC-DPV-903; 1, labial, and 2, lingual views; ´ 10. 3±4,
FC-DPV-907; 3, labial, and 4, lingual views; ´ 9. 5±6, FC-DPV- 909; 5, labial, and 6, lingual views; ´ 15. 7±8,
FC-DPV-1008; 7, labial, and 8, lingual views; ´ 12. 9, FC-DPV-1049, dorsal ®n spine, lateral view; ´ 4.
 PLATE 1

PEREA et al., Priohybodus

1232 PALAEONTOLOGY, VOLUME 44

TABLE 1. Tooth Measurements in mm of the Uruguayan specimens of Priohybodus cf. P. arambourgi. Dent. per mm,
denticles per mm; the value on the left corresponds to the anterior edge of the tooth and the value on the right
corresponds to the posterior edge of the tooth. h c c, height of the central cusp; h 1st c, height of the ®rst cusplet; w c c,
width of the central cusp; % 1st c c c, percentage of the ®rst cusplet in relation to the central cusp; R (h/w c c), relation
between the height and the width of the central cusp.

FC-DPV Dent. per mm hcc h 1st c wcc % 1st ccc R (h/w c c)

903 3´5/3´3 ± ± ± ± ±
905 3´49/± ± ± ± ± ±
906 ±/2´7 ± ± ± ± ±
907 2´86/2´84 7´7 2´15 7´2 27´9 1´07
908 3´8/± ± ± ± ± ±
909 ± 4´75 ± 3´7 ± 1´28
1002 ± ± ± ± ± ±
1003 ± 4´5 1´3 3´7 28´9 1´22
1004 ±/3´67 ± ± ± ± ±
1005 4´28/4´37 ± ± ± ± ±
1006 5´22/4´16 5´9 1´4 5´6 23´7 1´05
1007 ±/3´75 ± ± ± ± ±
1008 3´5/± ± ± ± ± ±
1009 ± 4´4 1´5 3´9 34´1 1´13
1010 ±/4´05 ± ± ± ± ±
1011 ± ± ± ± ± ±
1012 4´45/5´06 ± ± ± ± ±
1013 ± 3´9 1´3 3´5 33´3 1´11
1014 ± ± ± ± ± ±
1015 ±/5´08 ± ± ± ± ±
1016 5´35/4´63 4´15 ± 4´4 ± 1´01

TABLE 2. Measurements in mm of Libyan (from Duf®n, unpublished information) and Uruguayan specimens of
Priohybodus arambourgi and Priohybodus cf. P. arambourgi. HCC, central cusp height; CBL, central cusp base
length.

Variable Valid N Mean Minimum Maximum Std. Dev.

Teeth from Libya HCC 28 9´68 5 15 2´55

CBL 28 8´61 4 15 2´44
Teeth from Uruguay HCC 7 5´06 3´9 7´7 1´33
CBL 7 4´57 3´5 7´2 1´36

pronounced distal curvature in lateral view and a widening toward the base. The spine is a laterally compressed oval in
cross-section. The lateral walls are costate, ornamented by a series of longitudinal ridges with intervening grooves. A
distally single but proximally double row of alternating downturned denticles extends for the greater length of the
central part of the posterior spine wall. Nineteen denticles, with a base length ranging from 2´9 mm distally to 2´2 mm
proximally, are represented.

DISCUSSION
Systematics
The morphology of Priohybodus, with its symmetrical, smooth, compressed and serrated teeth, is easily
distinguished from other hybodontoid sharks (Cappetta 1987; Brito and Ferreira 1989; Brito 1991, 1992;
Duf®n in press).
 PEREA ET AL.: MESOZOIC SHARK 1233

We consider that the number of teeth available in the sample is inadequate for constructing an arti®cial
odontographic series (Applegate 1965; Cappetta 1986). We can only propose that the more symmetrical
the teeth, the more proximal their position in the dentition. The serrated cutting edges of the cusps are a
distinctive feature of Priohybodus among the hybodonts, and the character has been developed
independently in other selachian lineages in which there is a tendency towards relatively homodont
dentitions with upright teeth (Duf®n in press).
Compared with the Libyan specimens studied by Duf®n (op. cit.), our sample contains smaller teeth,
each with more serrations per mm and a proportionally lower central cusp (Pl. 1; Tables 1±2). Also, the
Uruguayan dorsal ®n spine fragment is more curved and broader at the base than the only African
specimen described by Duf®n. These differences are insuf®ciently diagnostic for distinction between the
Uruguayan and African specimens, being much more likely to be the product of geographical variation in a
single widespread species.

Biostratigraphy and geochronology

The previously known fauna of the Tacuarembo Formation provides questionable evidence for its age. The
fauna consists of bivalves (MartõÂnez et al. 1993), gastropods, ostracods, conchostracans, semionotiform
and dipnoan ®shes, a crocodile, and ichnofossils (Mones 1980; Mones and Figueiras 1980; Da Silva 1990).
The age of the formation has been estimated, until now, on the basis of purely stratigraphic criteria; there is
no general agreement in the published literature, with estimates ranging from Late Triassic to Early
Cretaceous (MartõÂnez et al. 1993).
The discovery of Priohybodus cf. P. arambourgi in the Tacuarembo Formation allows us to place that
unit in the Late Jurassic±Early Cretaceous interval. The Arapey Formation has been radiometrically dated
by K/Ar, providing 126´8 6 3´1 Ma for the younger volcanic events to 152´4 6 8´2 Ma for the older ones
which lie on, or are interbedded with, the upper section of Tacuarembo Formation (Bossi and Navarro
1991). Taking into account the radiometric dates, the referred stratigraphic relationships and the known
fossil record of Priohybodus arambourgi, the Tacuarembo Formation is considered to be not older than
Kimmeridgian or younger than Hauterivian in age. Mones and Figueiras (1980) proposed a similar
chronological hypothesis (Late Jurassic±Early Cretaceous) based exclusively on absolute dates of the
overlying Arapey Formation.

Taphonomy
As noted above, the fossiliferous level is a patchy bone-bed 30±60 mm thick within a well-sorted sandy
matrix toward the base of a sandy horizon (Text-®g. 2). The vertebrate assemblage is dominated by
ganoid semionotiform scales and subsidiary teeth, including those of Priohybodus, theropod dinosaurs,
semionotiform ®shes, and several fragmented bones and spines.
The bone concentration is bioclast- to matrix-supported with loosely packed to dispersed bone-units
depending on the thickness of the patch. In bedding plane and cross-section views the bone assemblage
has a random and disarticulated pattern. The scales and teeth are complete and unabraded; the ganoid
scales have cracked ganoin layers. The bone assemblage is parautochthonous to allochthonous; the
semionotiform scales and teeth are parautochthonous and the dinosaur teeth are the allochthonous
elements of the assemblage. Judging from the taphonomic features and the sedimentological frame-
work, a shallow subaqueous depositional environment produced by unidirectional ¯uvial ¯ow can be
inferred.

Biogeography and palaeoecology

The family Hybodontidae comprises marine and freshwater sharks represented in Middle Triassic±Upper
Cretaceous rocks in Europe, North America, North and West Africa, and Asia (Maisey 1987; Cappetta
1987; Cappetta et al. 1993).
The separation of South America from Africa was a heterochronous event occupying much of the Lower
1234 PALAEONTOLOGY, VOLUME 44

Cretaceous (Maisey 1991, 2000). The ®nding of Priohybodus in Uruguay demonstrates that it was a
widespread and conspicuous representative of the family. The South American and African populations of
the genus probably had a common evolutionary history in Gondwana before its break up during the
Jurassic and Cretaceous. The Afro-Brazilian rifting led to the development of rift valleys containing lakes
(Maisey 1991). The occurrence of very closely related species of vertebrates in the Jurassic±Cretaceous of
South America and Africa provides evidence for the existence of a continental fauna whose composition
diverged little between both continents until late in the Early Cretaceous (Buffetaut and Taquet 1979;
Gasparini 1996). According to Goodwin et al. (1999) Africa and South America retained the connection
through desert ecosystems until the mid Cretaceous. The separation between the southern parts of South
America and Africa started at the end of the Jurassic and rifting of the more northerly parts commenced
during Kimmeridgian±Early Cretaceous times.
Priohybodus provides additional support for this. It is associated, on both continents, with semionotid
and dipnoan remains. The former inhabited marine and freshwater environments (Gardiner 1993). The
ceratodontid and neoceratodontid dipnoan ®shes are largely represented in freshwater environments from
Triassic to Recent (Schultze 1993). The invertebrate fossils of the Tacuarembo Formation, particularly the
bivalves and conchostracans, indicate freshwater environments (Da Silva 1990; MartõÂnez and Figueiras
1991; Briggs et al. 1993).
A fauna that includes freshwater unionid bivalves has evolved in long-lived, large river systems
(Seilacher 1998). The conspicuous and bizarre freshwater fauna of the Tacuarembo Formation includes the
putative unionid bivalve Tacuaremboia caorsii MartõÂnez, Figueiras and Da Silva, 1993. This giant clam
suggests not only an `ancient' river system but high water energy inferred by its large size and thick shell,
which were presumably useful in giving stability in high-energy ¯ooded environments (MartõÂnez et al.
1993). In addition, judging from the large size of some ganoid scales (40 ´ 30 mm) big semionotiform
®shes measuring up to 1´5±2 m in length were present; Priohybodus itself could not have been less than
1´5 m long. The prey species of Priohybodus are not clear; it is hard to believe that semionotiforms were
included in its diet because of their heavy body scales.

Acknowledgements. We sincerely thank C. J. Duf®n for giving us much unpublished information and for his very
useful comments and improvement of the manuscript; the referees J. Maisey and C. Underwood, for their valuable
suggestions; M. Verde, who discovered the bone-bed; N. Lorenzo, for her collaboration in the ®eld; and R. PeÂrez, for
his help with the photographs.

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DANIEL PEREA
MARTIÂN UBILLA
ALEJANDRA ROJAS
CEÂSAR A. GOSO
INGEPA-Facultad de Ciencias
Igua 4225,
Typescript received 13 September 2000 11400 Montevideo, Uruguay
Revised typescript received 9 January 2001 e-mail perea@fcien.edu.uy