Journal of Vertebrate Paleontology 20(4):645650, December 2000

q 2000 by the Society of Vertebrate Paleontology

A MIOCENE CUSK-EEL (OPHIDIIFORMES: OPHIDIIDAE) FROM

S, ARGENTINA
PENINSULA VALDE
CARLA M. RIVA ROSSI1, ATILA E. GOSZTONYI1, and MARIO A. COZZUOL2
Centro Nacional Patagonico (CONICET), Blvd. Brown s\n, Puerto Madryn, 9120, Chubut, Argentina;
2
Departamento de Ciencias Biomedicas, Universidade Federal de Rondonia, Br. 364Km 9.5, 78900700, Brasil
1

ABSTRACTGenypterus valdesensis, sp. nov. from the Puerto Madryn Formation in Pennsula Valdes, Argentina is
the first fossil ophidiid reported from South America and represents the first well preserved record of the genus
worldwide. The specimen consists of a nearly complete articulated skull and some disarticulated postcranial elements.
A thorough description is given of the osteological features of the species, which can be used to recognize it as a
distinct taxon. The fossil species is characterized by the presence of a foramen on infraorbital 3 and by the squareshape of infraorbital 4. An array of morphological features taken in combination contribute to further differentiate G.
valdesensis from the Recent species. This specimen also provides unequivocal evidence of the presence of the Ophidiiformes in the southwestern Atlantic Ocean 14 MA.

INTRODUCTION
The Tertiary marine deposits of Argentina are represented by
a single sedimentation cycle comprising two transgressions.
The first one principally affected Patagonia from the Late Eocene to the Early Miocene and is represented by the Julian and
Leonian Stages (Menda and Bayarsky, 1981; Scasso and Del
Ro, 1987; Cione and Cozzuol, in press). The second one
spanned the Middle to Late Miocene and occurred in northeastern Argentina and northeastern Patagonia. The most important outcrops in the latter region appear in the lower valley of
the Chubut River, in Pennsula Valdes (Fig. 1), and in northeastern Ro Negro Province; these are grouped in the Aonikan
Stage (Del Ro, 1988; but see also Frenguelli, 1926; Haller and
Menda, 1980; Haller, 1981; Menda and Bayarsky, 1981; Scasso y Del Ro, 1987; Cozzuol, 1993).
The Puerto Madryn Formation outcrops at Pennsula Valdes
and nearby Puerto Madryn city (Fig. 1) (Rovereto, 1913; Frenguelli, 1926; Haller, 1978; Scasso and Del Ro, 1987; Del Ro,
1988, 1990) and consists of approximately 100 m of nearshore
marine deposits, consisting of mudstones and yellowish and
brownish sandstones (Scasso and Del Ro, 1987).
Paleontological research in Pennsula Valdes has revealed a
rich coastal assemblage of vertebrates composed of marine
mammals (seals and cetaceans), penguins, marine birds, and
marine fishes (Cozzuol, 1991; Cozzuol et al., 1991a, b; Cozzuol, 1993).
The paleoichthyofauna of Pennsula Valdes has an important
fossil record. Sharks and rays are largely represented by isolated teeth or tooth plates and include Carcharodon megalodon,
Isurus hastalis, and Myliobatis sp. from the middle Miocene of
Patagonia (Cione, 1978), and an undetermined species of the
genus Squatina (Cozzuol et al., 1991b). Osteichthyans are represented by several taxa of Siluriformes, Ophidiiformes, Gadiformes, Perciformes and Scorpaeniformes (Cozzuol et al.,
1991b; Gosztonyi et al., 1993; Riva Rossi and Cozzuol, 1995).
One of the most important remains yielded by the lower beds
of the Puerto Madryn Formation in Pennsula Valdes consists
of an ophidiid fish (Ophidiidae: Ophidiiformes) attributed by
Gosztonyi to the genus Genypterus (Gosztonyi et al., 1993).
Modern fishes of the genus Genypterus, widely known as
cuskeels, include commercial species that inhabit the continental shelf and slope of the temperate Southern Oceans. Cohen
and Nielsen (1978) listed at least five species for this genus: G.
chilensis Guichenot 1848; G. capensis Smith 1847; G. blacodes

Schneider 1801; G. maculatus Tschudi 1846; and G. microstomus Regan 1903. Many authors have considered G. capensis
and G. microstomus as synonyms of G. blacodes (Barnard,
1972; Olivar and Sabates, 1989), and more recently Nakamura
(1986) and Cousseau and Perrota (1998) have also included G.
brasiliensis Regan 1903 as a valid species.
In this work we describe the skull of a new species of the
genus Genypterus from the middle Miocene of Pennsula Valdes, Argentina. It is a remarkable discovery because this specimen represents the first record of this group based on skeletal
remains (Gosztonyi et al., 1993). Hitherto the only fossil record
of Genypterus was otoliths from the early Oligocene of New
Zealand (Nolf and Steurbaut, 1989).
MATERIAL EXAMINED
The fossil cusk-eel skull is deposited in the Museo Paleontologico Egidio Feruglio, Departamento de Paleontologa de
Vertebrados (MPEFPV 614). In addition we examined Recent
crania of Genypterus blacodes, Genypterus brasiliensis, Genypterus capensis, Genypterus chilensis, and Genypterus maculatus. These skeletons are housed in the Centro Nacional Patagonico (CNP). Recent specimens included in this study are
listed in Appendix.
Institutional abbreviationsCNPICT, Centro Nacional
Patagonico-Ictiologa; MPEFPV, Museo Paleontologico Egidio
Feruglio, Departamento de Paleontologa de Vertebrados.
SYSTEMATIC PALEONTOLOGY
Order OPHIDIIFORMES, sensu Cohen and Nielsen 1978
Suborder OPHIDIOIDEI, sensu Cohen and Nielsen 1978
Family OPHIDIIDAE Rafinesque 1810
Subfamily OPHIDIINAE Rafinesque 1810
Tribe LEPOPHIDIINI Robins 1961
Genus GENYPTERUS Philippi 1857
Type SpeciesGenypterus nigricans Philippi 1857.
GENYPTERUS VALDESENSIS, sp. nov.
(Figs. 2, 3)
HolotypeMPEFPV 614, an almost complete skull articulated with the anterior precaudal vertebrae. The length of the
skull is 244 mm. In association, there are an isolated fragment
of the lachrymal, several precaudal vertebrae, and a few caudal

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species of Genypterus by the presence of a foramen on infraorbital 3 and by the square shape of infraorbital 4. Furthermore
an array of morphological features taken in combination differentiates G. valdesensis from the remaining species of the
genus; it differs from G. blacodes (5capensis), G. chilensis and
G. maculatus by the dorsal profile of the neurocranium which
is slightly slanted toward the supraoccipital, by the presence of
a projection on the infraorbital 4, the presence of a central projection on the extrascapula; it differs from G. blacodes, G. brasiliensis and G. chilensis by the posteriorly projected walls of
the foramen magnum. From G. chilensis and G. maculatus it
differs as follows: quadrate spine with a crest, an expanded
supracleithrum, and posterior lamina of the cleithrum backwardly directed. Finally, G. valdesensis differs from G. brasiliensis by the following: ventral fossa of the frontal expanded,
frontal and pterosphenotic bearing an oval depression; and the
straight posterior border of the opercle.
Description

FIGURE 1.

Map of Pennsula Valdes showing the fossil locality.

vertebrae. The jaws and the nasals are lost from the anterior
end of the neurocranium.
EtymologyThe specific epithet refers to the type locality
Pennsula Valdes, Chubut, Argentina.
Type Locality and HorizonPunta San Roman
(428149400S; 648119310W), on the northern shore of the Golfo
San Jose, Pennsula Valdes (Fig. 1). The fossil was collected
from the lower facies of the Puerto Madryn Formation (Middle
Miocene).
DiagnosisGenypterus valdesensis differs from all other

FIGURE 2.

NeurocraniumThe braincase is almost intact, except for

the loss of the vomer, the ethmoid, the anterior region of the
parasphenoid and both lateral ethmoids. The general shape of
the neurocranium is rectangular, elongate and slightly flattened.
The upper profile slants gently posterodorsally as in G. brasiliensis. The frontal (Figs. 2, 3), lacking the anterior end, is
almost rectangular in shape, has a weakly developed supraorbital process above the orbit, and a deep groove in its upper
surface containing the lateral line system. Ventrally it has a
deep and expanded fossa as in G. blacodes and G. maculatus.
In the postfrontal the outer margin of its anterior region is
strongly bent outward (Fig. 2A). Between the frontal and pterosphenotic (Figs. 2A, 3A) there is a marked oval depression as
in G. blacodes, G. chilensis and G. maculatus. The sphenotic
(Figs. 2, 3) has a posterodorsally directed, lateral, short and

Genypterus valdesensis, sp. nov., holotype (MPEFPV 614). Skull in right (A) and (B) left views.

RIVA ROSSI ET AL.A MIOCENE CUSK-EEL

647

FIGURE 3. Drawing of the skull of Genypterus valdesensis, sp. nov., holotype (MPEFPV 614). (A) in right and (B) left views. Abbreviations:
aa, anguloarticular; brs, branchiostegal rays; cl, cleithrum; d, dentary; ecp, ectopterygoid; enp, entopterygoid; eo, exoccipital; epo, epiotic; es,
extrascapula; f, frontal; hm, hyomandibular; i 34, infraorbitals; iop, interopercle; mt, metapterygoid; mx, maxilla; op, opercle; p, parietal; pal,
palatine; pch, posterior ceratohyal; pop, preopercle; ps, parasphenoid; pt, posttemporal; pts, pterosphenoid; q, quadrate; scl, supracleithrum; so,
supraoccipital; sop, subopercle; spo, sphenotic; sy, symplectic; vertebrae. Scale bar equals 1 cm.

broad projection that also occurs in the Recent species. In the

parasphenoid (Figs. 2A, 3A) the anterior region is lost and the
middle part is exposed under the orbit, with a prominent keel
on the ventral edge. In the rear part the winged process is upraised. Posterodorsally, the pterotic (Figs. 2, 3) has a strongly
posteriorly tapered processthe pterotic wing. In the dorsal
surface it has a deep and wide groove for the lateral line system.
In the posterior region of the skull roof the limits of the bones
are not clear. The parietals are poorly preserved (Figs. 2B, 3B).
They are barely separated from each other by the anterior limb
of the supraoccipital, their dorsal surface is slightly upraised
and at the rear there is a well-marked transverse crest abutting
posteriorly with the temporal fossa. The epiotics (Figs. 2B, 3B)
are not well preserved. At their posterior end the crest that
occurs in extant species is not obvious. The supraocciptal (Figs.
2B, 3B) is cross-like, with its anterior limb markedly narrow
as in G. brasiliensis. Its posterior limb is badly damaged and
projected ventrally toward the exoccipital through the laminae

FIGURE 4. Right infraorbital 3 (right) and lachrymal (left) of Genypterus valdesensis, sp. nov.

(Figs. 2B, 3B). The walls of the foramen magnum (Figs. 2B,
3B) are projected backwards meeting the exoccipital condyles
at the posterior end as in G. maculatus. On the outer face it has
a slightly downwardly bent heavy flange, along which runs a
laminar crest surrounding the glossopharyngeal nerve foramen
as in G. blacodes (Figs. 2B, 3B). Above this flange the vagus
nerve foramen opens.
InfraorbitalsOnly fragments of the lachrymal, infraorbital
3 and infraorbital 4 are preserved. The lachrymal (Figs. 4, 5A)
is an elongated laminar plate with the dorsal margin folded over
the outer face forming the infraorbital canal of the cephalic
lateral line system. Infraorbital 3 is L-shaped (Figs. 4, 5B). The
dorsal margin of the horizontal limb and the anterior of the
vertical limb is folded over the outer face forming the infraorbital canal. At the angle it has a foramen, absent in extant species. The posterior region is a rounded laminar, backward directed expansion. Infraorbital 4 has a square shape, whereas in
Recent species this bone is rectangular. Its anterior margin is
folded over the outer face.
JawsThe premaxilla (Figs. 6, 7) lacks the anterior end.
Although teeth are not preserved, the alveolar process reveals
that several rows of teeth were present. In the outer row the
alveoli are considerably larger than those of the inner rows and
the sizes of the alveoli decrease gradually toward the posterior
end. Posteriorly in the upper margin there is a triangular, low,
and slightly damaged postmaxillary process. The maxilla is
edentulous, its anterior end is lost. Anteriorly it is a thin bone,
oval in cross section and posteriorly flattened and dorsoventrally expanded. At the rear end the ventral margin is rounded
as in G. blacodes, G. chilensis and G. maculatus. The dorsal
margin has a well-developed triangular knob just in front of the

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 4, 2000

FIGURE 5. Drawing of Infraorbitals of Genypterus valdesensis, sp.

nov. A, right lachrymal and B, infraorbital 3. Scale bar equals 1 cm.

attachment to the supramaxilla. The inner face has a wide Vshaped fossa (Figs. 6B, 7B), the outer face has a shallow groove
(Figs. 6A, 7A). The supramaxilla (Fig. 6B) is elliptical. Its ventral margin is convex anteriorly and concave posteriorly; the
anterior end is acutely pointed as in G. blacodes, G. chilensis
and G. maculatus; the inner face has a shallow depression. The
lower jaw is long and relatively slender, and positioned at the
posterior level of the orbit. The dentary (Figs. 2A, 3A) is the
largest bone of the series, although its anterior part is lost. The
alveolar process bears numerous conical teeth arranged in a
single series. Since the medial face is not exposed, it is difficult
to determine whether inner rows of teeth are present. In the
outer face, under the alveolar process there is an elongated depression, whereas a deep and broad groove for the cephalic
lateral line system runs along the entire ventral surface. Anteriorly this groove terminates in numerous foramina. The articular (Figs. 2, 3) is wedge-shaped, strongly attached to the dentary, and posterodorsally thickened, forming a saddle-shaped
facet for the articulation with the quadrate; just in front of this
facet there is a marked flange. Above this flange the ascending
process of the articular is grooved and slightly slanted anterodorsally at the tip. The outer face has an enlarged fossa. The
angular (Figs. 2, 3) is strongly sutured to the articular. The
retroarticular (Figs. 2, 3) is a triangular, thickened bone.
Suspensorium and Opercular BonesThe palatine (Figs.
2A, 3A) is ventrally toothed. The conical, slightly curved teeth
are arranged in a single outer series. Since the medial surface
is not exposed it cannot be determined whether inner rows of
teeth are present as in Recent species. Anteriorly the maxillary
process is rod-like and has a groove in the outer face. At its
base it has a small facet for the articulation with the lateral
ethmoid. This process is at least three times shorter than the
total length of the bone. The quadrate (Figs. 2, 3) is triangular,
posteriorly thickening in a sturdy spine that gradually tapers at
the end. In the outer face of the spine runs a crest as in G.
blacodes and G. brasiliensis. Anteroventrally it bears the suspensorium condyle which has a small projection as in G. brasiliensis and G. maculatus. The ectopterygoid (Figs. 2A, 3A) is
thick and boomerang-shaped. Its suture with the quadrate is
dentate, and dorsally it is strongly attached to the mesopterygoid (Figs. 2A, 3A) which is a laminar bone. The metapterygoid (Figs. 2, 3) is sub-triangular with the anterior end rounded;
posteriorly it has a strong crest as in the adults of the Recent
species. The symplectics (Figs. 2, 3) are poorly preserved and
their limits are not clear. The hyomandibular (Figs. 2, 3) is a
heavy bone with a vertical shaft. It has three condyles, two of

FIGURE 6. Genypterus valdesensis, sp. nov. A, lateral view of the

right maxilla and premaxilla and B, inner view of the right maxilla,
premaxilla and supramaxilla.

which articule with the neurocranium, and the third one with
the opercle. Between the posterior neurocranial condyle and the
opercular condyle an upright lamina extends. The facial foramen opens in the lateral face and not covered by a lamina. From
the mesial rim of the foramen a strong crest is developed. The
preopercle (Figs. 2, 3) is crescent-shaped, with the ventral and
dorsal limbs almost equal in length. The dorsal margin is not
preserved. A broad slightly forward curved fold, which contains the preoperculo-mandibular canal of the lateral line system
runs along the entire length. At the level of the angle, a single
blunt projection develops from the fold as in Recent species.
The opercle (Figs. 2, 3) is triangular with the dorsal margin
thickened and projecting backwards in a strong spine, above

FIGURE 7. Reconstruction of the right maxilla and premaxilla of

Genypterus valdesensis, sp. nov. A, lateral view and B, inner view.
Abbreviations: mx, maxilla; pmx, premaxilla. Scale bar equals 1 cm.

RIVA ROSSI ET AL.A MIOCENE CUSK-EEL

which is a laminar projection. The posterior margin is straight
as in G. blacodes, G. chilensis and G. maculatus. The interopercle (Figs. 2, 3) is exposed only at the rear end, the rest being
covered by the preopercle. The subopercle (Figs. 2, 3) is rounded, and partially covered by the opercle.
Hyoid ArchOnly the ventral margin of the branchiostegal
rays (Figs. 2, 3) are exposed and only a few of them along the
entire length. The remaining bones are unexposed.
Pectoral GirdleThe extrascapula (Figs. 2B, 3B) is a thin
bone with the lateral margin folded over the dorsal surface. In
its mid section this flange abruptly broadens as in G. brasiliensis. Along the dorsal surface runs a groove that branches off
anteriorly. The posttemporal (Figs. 2A, 3A) is V-shaped with
two limbs. The dorsal limb is flattened, tapering toward the end,
with an upright groove in the ventral region as in G. blacodes,
G. brasiliensis and G. maculatus, and without the ventral projection that is present in G. blacodes. The ventral limb is rodlike and shorter than the dorsal one. It is broken in the middle
and the dorsal portion is displaced. The supracleithrum (Figs.
2, 3) is rod-like as in G. blacodes and G. brasiliensis with a
well developed articular facet for the posttemporal. The beveled
edge is not preserved. In the cleithrum (Figs. 2B, 3B) the ventral limb is projected forward as a filamentous process. The
dorsal limb has a posterior lamina projected posteriorly as in
G. brasiliensis and G. blacodes.
VertebraeThe first precaudal vertebrae (Figs. 2, 3) are still
articulated with the neurocranium. In the third precaudal vertebra the parapophyses are greatly enlarged as in Recent ophidiids. The rest of the precaudal vertebrae and a few caudal
vertebrae are preserved as isolated elements.
DISCUSSION
Genypterus valdesensis is attributable to the order Ophidiiformes based on the position of the pelvic fins, which, when
present, are inserted at the level of preopercle or farther anterior
(mental or jugular) (Cohen and Nielsen, 1978; Nelson, 1994).
Its classification in the family Ophidiidae is supported by the
presence of: (1) at least one opercular spine; (2) supramaxilla
present; and (3) pelvics rarely absent (Cohen and Nielsen, 1978;
Nelson, 1994). Moreover, its status as an ophidiine is supported
by the presence of the ventral arms of the cleithra approximating each other anteriorly under the level of the preopercle from
which a slender element extends forward below the orbital region and by the position of the pelvic girdle between these
elements (Cohen and Nielsen, 1978).
The assignment of Genypterus valdesensis to the genus Genypterus is based upon the occurrence of several diagnostic features that are distinctively shared by the Recent species of the
genus. These characters are: (1) neurocranium flattened and
elongated; (2) sphenotic with a short and blunt lateral projection; (3) posterior margin of the infraorbital 3 a laminous projection; (4) supramaxilla elliptical; (5) dentary and palatine
bearing canine-like teeth arranged in a single series; (6) length
of the maxillary process one third the total length of the palatine; (7) posterior margin of the preopercular flange with an
unique blunt projection; and (8) post-temporal with an upright
groove.
Finally, Genypterus valdesensis has at least two derived features and a combination of characters which taken together support its designation as a new species, that inhabited the South
American seas in the upper Tertiary.
The order Ophidiiformes has an extensive fossil record based
on otoliths. They abound in the Tertiary of the North Sea Basin,
Paleogene of Europe, EoceneMiocene from Aquitaine, MiocenePliocene from the Dominican Republic, early Eocene to
present from New Zealand, and PleistoceneHolocene in the
southern USA (Nolf and Steurbaut, 1989; Fitch, 1964; Stringer,

649

1992). However, in constrast to the otolith-based records, skeletal remains of ophidiiforms are very scarce (Patterson and Rosen, 1989).
Hitherto, skeletal remains of the family Ophidiidae were exclusively known from the Palearctic Region. The ophidiid
Eolamprogrammus occurs in the Paleocene of Turkmenistan
(Cohen and Nielsen, 1978). An ophidiid fish is represented in
the early Eocene deposits from Monte Bolca, Italy; also in this
period occur Ampheristus and Hoplobrotula (Patterson and Rosen, 1989). Finally, Glyptophidion is known from the Miocene
of Japan (Cohen and Nielsen, 1978).
Genypterus valdesensis represents the first fossil record of an
ophidiid in South America. The occurrence of this species in
the southwest Atlantic seashore is not unexpected, since the
fossil was found within the present-day distribution area of the
genus. However, it should be noted that the fossil species must
have been adapted to warm seas, since at that period the Patagonian marine environments were characterized by high temperatures, ranging from 188C to 248C (Del Ro, 1990).
It was not until the Late Miocene that the influence of cold
circumpolar currents caused by the development of the Antarctic ice cap, produced a decrease in the temperatures of South
American coastal waters (Marshall and Sempere, 1993). Consequently, most of the marine ichthyofauna inhabiting these latitudes endured a gradual northward migration toward more favourable conditions (Cione, 1978). Thus, the only original
groups that could survive in the cooler environment and not
migrate, were those fishes that were able to adjust to the decreased temperature of their new habitat.
Although most Recent species of Genypterus are principally
distributed in cold-temperate seas of the Southern Hemisphere,
the presence of G. valdesensis in the exceptionally warm Entrerriense sea is not surprising. Nowadays, the distribution of
the extant species G. brasiliensis ranges from the southern tip
of South America to the warmer waters off southern Brazil.
Thus it could be thought that the climatic conditions in the
present Brazilian sea resemble to some extent the Tertiary marine environment in which G. valdesensis occurred 14 Ma.
At present there is barely enough information to hypothesize
any explanation about the historical biogeography or phylogeny
of this group. This is due, in part to the scarcity of fossil material, since apart from the present record, there is only one
other based on otoliths from the Lower Oligocene of New Zealand (Nolf and Steurbaut, 1989). These two discoveries provide
enough evidence to indicate that Genypterus was already well
differentiated and widespread in Southern Oceans as early as
that epoch. However, additional fieldwork and analysis are
needed in order to remedy the lack of information about the
evolutionary history of this taxon.
ACKNOWLEDGMENTS
We thank the Museo Paleontologico Egidio Feruglio and
the Laboratory of Paleontology of the Centro Nacional Patagonico for its collaboration. We also are indebted to N. Garca,
M. Coscarella and A. Rubilar for providing the necessary material for this work. We gratefully acknowledge the thorough
work done by L. Reiner for the preparation of the fossil specimen. We also appreciate the support of M. T. Dozo during the
preparation of the manuscript and finally we thank M. M. Azpelicueta and M. Pascual for reviewing the drafts of the manuscript.
LITERATURE CITED
Barnard, K. H. 1972. A monograph of the marine fishes. Annals of
South African Museum 21:4191065.
Cione, A. 1978. Aportes paleoictiologicos al conocimiento de la evol-

650

JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 4, 2000

ucion de las paleotemperaturas en el area austral de America de

Sur durante el Cenozoico. Ameghiniana 15:183208.
Cohen, D., and J. Nielsen. 1978. Guide to the Identification of Genera
of the Fish Order Ophidiiformes with a Tentative Classification of
the Order. NOAA Technical report NMFS. Washington, D.C. Circular 417, 72 pp.
Cousseau, M. B., and R. G. Perrota. 1998. Peces marinos de Argentina.
Biologa, distribucion, pesca. INIDEP (Instituto Nacional de Investigacion y Desarrollo Pesquero). Mar del Plata, Argentina, 163 pp.
Cozzuol, M. A. 1991. Primer registro de un pinnpedo terciario en Patagonia. VIII Jornadas de Paleontologa de Vertebrados, La Rioja,
Resumenes. Ameghiniana 28(34):406.
1993. Los mamferos marinos fosiles del Terciario de America
del Sur. Ph.D. dissertation. Facultad de Ciencias Naturales y Exactas, Universidad Nacional y Museo de La Plata, La Plata, 148
pp.
, R. E. Fordyce, and C. Jones. 1991a. La presencia de Eretiscus
tonnii en el Mioceno temprano de Nueva Zelandia y Patagonia.
VIII Jornadas de Paleontologa de Vertebrados, La Rioja, Resumenes. Ameghiniana 28(34):406.
, A. E. Gosztonyi, and L. Kuba. 1991b. Una asociacion de vertebrados marinos de la Formacion Puerto Madryn (Mioceno medio)
en Pennsula Valdes, Chubut. Resumenes Jornadas Nacionales de
Ciencias del Mar 1991:77.
Del Ro, C. J. 1988. Bioestratigrafa y Cronoestratigrafa de la Formacion Puerto Madryn (Mioceno Medio)Provincia del Chubut
Argentina. Anales de la Academia Nacional de Ciencias Exactas,
Fsicas y Naturales 40:231254.
1990. Composicion, Origen y Significado Paleoclimatico de la
Malacofauna Entrerriense (Mioceno Medio) de la Argentina.
Anales de la Academia Nacional de Ciencias Exactas, Fsicas y
Naturales 42:205224.
Fitch, J. E. 1964. The fish fauna of the Playa del Rey locality, a Southern California marine Pleistocene deposit. Contributions in Science.
35 pp.
Frenguelli, J. 1926. El entrerriense en el Golfo Nuevo del Chubut. Academia Nacional de Ciencias, Boletn 29:191270.
Gosztonyi, A., M. A. Cozzuol, and L. Kuba. 1993. Primer registro de
un abadejo fosil, Genypterus sp. (Osteichthyes, Ophidiidae) de la
Formacion Puerto Madryn (Mioceno Medio) en Pennsula Valdes,
Argentina. Resumenes Jornadas Nacionales de Ciencias del Mar,
114 pp.
Haller, M. 1978. Estratigrafa de la region al poniente de Puerto Madryn, provincia del Chubut, Republica Argentina. Actas VII8 Congreso Geologico Argentino 1:285297.
1981a. Descripcion geologica de la hoja 43H, Puerto Madryn,
Chubut. Servicio Geologico Nacional, Boletn 184.
1981b. Las sedimentitas plegadas de la margen oriental de Pennsula Valdes. Actas VIIIO Congreso Geologico Argentino 3:25
32.
, and J. E. Mendia. 1980. Las sedimentitas del Ciclo Patagoniano
en el litoral atlantico nordpatagonico. Asociacion Geologica Argentina, Buenos Aires, 11\ 28\80; pp. 593606 in J. E. Menda, and A.
Bayarsky (eds.), Estratigrafia del Terciario en el valle inferior del ro
Chubut. VIII8 Congreso Geologico Argentino, Actas 3.
Marshall, L. G., and T. Sempere. 1993. Evolution of the Neotropical
Cenozoic land mammal fauna in its geochronologic, stratigraphic,
and tectonic context; pp. 329392 in P. Goldblatt (ed.), Biological
Relationships between Africa and South America. Yale University
Press, New Haven, Connecticut.

Menda, J., and A. Bayarsky. 1981. Estratigrafa del Terciario del Valle
inferior del ro Chubut. Actas VIII8 Congreso Geologico Argentino
3:593606.
Nakamura, I. 1986. Important Fishes trawled off Patagonia. Japan Marine Fishery Resource Research Center, 369 pp.
Nelson, J. S. 1994. Fishes of the World. John Wiley and Sons Inc. New
York, 600 pp.
Nolf, D., and E. Steurbaut. 1989. Importance and restrictions of the
otolith-based fossil record of gadiform and ophidiform fishes; pp.
3745 in D. M. Cohen (ed.), Papers on the Systematics of the
Gadiform Fishes. Natural History Museum of Los Angeles County,
Science Series No. 32.
Olivar, M. P., and A. Sabates. 1989. Early life history and spawning of
Genypterus capensis in the Southern Benguella System. South African Journal of Marine Sciences 8:173181.
Patterson, C. 1993. An overview of the early fossil record of Acanthomorphs. Bulletin of Marine Science, 52:2959.
, and D. Rosen. 1989. The Paracanthopterygii revisted: Order
and disorder; pp. 536 in D. M. Cohen (ed.), Papers on the Systematics of the Gadiform Fishes. Natural History Museum of Los
Angeles County, Science Series No. 32.
Riva Rossi, C. M., and M. A. Cozzuol. 1995. Lista preliminar de los
peces oseos de la Formacion Puerto Madryn (Mioceno medio) en
Pennsula Valdes. Resumenes XI Jornadas Argentinas de Paleontologa de Vertebrados:17.
1996. Una nueva especie del genero Genypterus (Pisces, Ophidiiformes) del Mioceno Medio de Pennsula Valdes (Chubut) y sus
relaciones filogeneticas con los abadejos actuales. Ameghiniana 33:
471.
Rovereto, G. 1913. La Pensola Valdez, e la forme costiere della Patagonia settentrionale. Reale Academia dei Lincei. Estratto del Col.
Col. XXIII 5:103105.
Scasso, R. A., and C. Del Ro. 1987. Ambientes de sedimentacion,
estratigrafa y proveniencia de la secuencia marina del Terciario
superior de la region de Pennsula Valdes, Chubut. Asociacion Geologica Argentina 42:291321.
Stringer, G. 1992. Late Pleistoceneearly Holocene teleostean otoliths
from a Mississippi River mudlump. Journal of Vertebrate Paleontology 12:3341.
Received 5 July 1999; accepted 20 May 2000.

APPENDIX 1
Recent skeletons of Genypterus included in this study (all CNPICT
specimens).
Genypterus blacodesCNPICT1995/1, CNPICT1995/2, CNPICT1995/3,
CNPICT1995/4, CNPICT1995/5, CNPICT1995/6, CNPICT1995/7,
CNPICT1995/8, CNPICT1995/9, CNPICT1995/10, CNPICT1995/11,
CNPICT1995/12, CNPICT1995/13.
Genypterus
brasiliensisCNPICT1995/14,
CNPICT1995/16, CNPICT1996/1.

CNPICT1995/15,

Genypterus chilensisCNPICT1996/2.1, CNPICT1996/2.2.

Genypterus maculatusCNPICT1996/3.
Genypterus capensisCNPICT1997/2, CNPICT1997/3, CNPICT1997/
4, CNPICT1997/5.