Journal of Vertebrate Paleontology 20(1):57–76, March 2000

q 2000 by the Society of Vertebrate Paleontology

A NEW SPECIES OF ARARIPESUCHUS (CROCODYLOMORPHA, MESOEUCROCODYLIA) FROM

THE LOWER CRETACEOUS OF PATAGONIA (ARGENTINA)

F. ORTEGA1, Z. GASPARINI2, A. D. BUSCALIONI1, and J. O. CALVO3

1
Departamento de Biologı́a, Unidad de Paleontologı́a, Universidad Autónoma de Madrid, 28049, Cantoblanco, Madrid, Spain;
2
Museo de La Plata, Dpto.Paleontologı́a de Vertebrados, Universidad Nacional de La Plata, Paseo del Bosque s/n. 1900,
La Plata, Argentina;
3
Museo de Geologı́a y Paleontologı́a, Universidad Nacional del Comahue, Buenos Aires 1400. 8300, Neuquén, Argentina

ABSTRACT—A new species of the genus Araripesuchus from the Albian–Cenomanian locality of the El Chocón
(Neuquén Province, Argentina) is described. The diagnosis of the genus is reviewed and its phylogenetic placement
within Crocodylomorpha discussed. Araripesuchus is proposed here as being the sister taxon of Neosuchia, corrobo-
rating previous phylogenetic analysis. The new species, Araripesuchus patagonicus, differs from the type species, A.
gomesii in the relationships of the prefronto-nasal and lachrymo-nasal sutures, the dermal placement of the postorbital
bar on the medial side of the jugal, and the greater extension of the squamosals in the skull dorsum. The African
species, ‘‘Araripesuchus’’ wegeneri, does not share the diagnostic traits of the genus, and its reassignment to a new
genus needs to be considered. The phylogenetic context of Araripesuchus and ‘‘A.’’ wegeneri permits a reanalysis of
the role played by their amphiatlantic distribution in the Aptian-Albian. ‘‘Araripesuchus’’ wegeneri and the South
American forms might share a pre-Aptian common ancestor, and have been already differentiated and isolated in the
African and South American continents by the time of the Aptian–Albian.

INTRODUCTION posed as a new Araripesuchus species; the westernmost and

The knowledge of basal mesoeucrocodylians (sensu Benton
and Clark, 1988) is mainly founded on Gondwanian evidence.
The Cretaceous beds of Africa and South America have pro-
vided the best record of notosuchians and araripesuchids. Up
until now the notosuchians described from South America have
been referred to the species Notosuchus terrestris Woodward,
1896 and Comahuesuchus brachybuccalis Bonaparte, 1991,
both from the Late Cretaceous of north-western Patagonia, Ar-
gentina (Bonaparte, 1996); and Candidodon itapecuruense Car-
valho and Campos, 1988 from the Albian of north-eastern Bra-
zil. The African genus Malawisuchus mwakayungutiensis
(Gomani, 1997) from the Aptian of Malawi has been related to
Notosuchus (Clark et al., 1989; Gomani, 1997). On the other
hand, the species Araripesuchus gomesii Price, 1959 from the
Middle Albian (Kellner, 1996) of north-eastern Brazil was com-
pared to the Aptian species Araripesuchus wegeneri Buffetaut,
1981 from the Tegama basin, Niger (Buffetaut and Taquet,
1979; Maisey, 1991; Hecht, 1991). Other taxa putatively related
to Araripesuchus come from the Mid-Late Cretaceous of Uru-
guay (Uruguaysuchus aznarezi Rusconi, 1933 and U. terrai
Rusconi, 1933), and from the Late Cretaceous of Madagascar
(Buckley and Brochu, 1996). Notosuchians and araripesuchids
provide clear evidence for the isolation and differentiation of
Gondwanic tetrapod faunas during the Cretaceous (Bonaparte,
1996), although a putative notosuchian (Chimaerasuchus par-
adoxus Wu et al, 1995) has been reported from the Lower Cre-
taceous of China. Moreover, the supposed close resemblance of
the African and Brazilian Araripesuchus species has played an
important role in supporting the hypothesis of a complete open-
ing of the South Atlantic Ocean later than the Aptian–Albian
(Buffetaut and Taquet, 1979; Buffetaut, 1981; Barron, 1987;
Bonaparte, 1991; Buffetaut and Rage, 1993).
A set of articulated crocodylomorph specimens found in
Neuquén, north-western Patagonia, provide an opportunity to
present a detailed description of specimens of distinct biolog-
ical ages, and to revise the interrelationships of the species of
the genus Araripesuchus. The Patagonian specimens are pro-
southernmost record of the genus.
Previous systematic considerations of Araripesuchus have
produced two major disagreements. First, Araripesuchus go-
mesii was originally proposed as a member of Notosuchidae
(Price, 1959; Hecht, 1991), and later assigned to Uruguaysuch-
idae (Gasparini, 1971; Buffetaut, 1982; Buffetaut and Rage,
1993). There is still no consensus regarding the phylogenetic
position of Araripesuchus within Mesoeucrocodylia. It is pro-
posed as being either the sister group of Notosuchus 1 Liby-
cosuchus (Wu et al., 1997), the sister taxon of Neosuchia
(Clark, 1994), the sister group of Sebecus 1 Neosuchia (Benton
and Clark, 1988), or a member of a clade including Libycosu-
chus, Sebecus, and Baurusuchus (Gomani, 1997). Secondly,
various authors (Michard et al. 1990; Kellner, 1994) have cast
doubt on the congeneric relationships of A. gomesii and A. we-
generi, and a revision of the African form is warranted. Other
material referred to aff. Araripesuchus and based on isolated
teeth from the Barremian–Aptian Koum Basin of Cameroon
and the Aptian of Malawi has been mentioned (Colins and Ja-
cobs 1990). The similarities of the teeth of the African speci-
mens to those from A. gomesii were discussed by Kellner
(1994).
Herein we describe in detail the specimens from Patagonia,
revising the diagnosis of the genus Araripesuchus and propos-
ing a new species. We also compare the putative closely related
forms of Araripesuchus within a phylogenetic framework in
which 27 crocodyliform taxa are considered.
MATERIAL EXAMINED
Araripesuchus gomesii: Holotype 423-R (a nearly complete
skull and lower jaw) housed in the Divisao de Mineralogia e
Geologia do Departamento Nacional da Produçao Mineral (Rio
de Janeiro D. F., Brazil). Locality: Unspecified location in the
Ladeira da Berlenga, western flank of the Araripe basin, Ceará
or Piauı́ State; Brazil. Horizon and age: Lower Cretaceous (Up-
per Aptian–Albian) of the Santana Formation (Hecht, 1991).
Other specimens attributed to the same species: specimen
AMNH # 24450 (an almost complete individual), housed in the

57
58 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 1, 2000
American Museum of Natural History (New York, USA). This
specimen came from the same basin as the holotype.
Araripesuchus wegeneri: Holotype GDF 700 (anterior por-
tion of a rostrum and lower jaw) housed in the Musée National
d’Histoire Naturelle de Paris. Locality: Gadoufaoua, Tegama
Basin, Niger (see Buffetaut, 1981, for details). Horizon and age:
Lower Cretaceous (Aptian) of the Elrhaz Formation (Buffetaut,
1981).
Other Taxa Studied from the Literature
Uruguaysuchus Rusconi, 1933: Several small crocodiles
found together during the construction of a hole in the Guichón
locality (Paysandú Department, Uruguay) with an imprecise age
ranging from the Senonian–Cenomanian to the Santonian–Cam-
panian (Price, 1959; Gasparini, 1996). Rusconi (1933) founded
two different species: U. aznarezi and U. terrai. Unfortunately,
the material is not easily available for revision. A part of the
holotype of U. terrai was destroyed before the description that
was supported solely by several photographs (Rusconi, 1933).
The remaining Uruguaysuchus specimens are currently housed
in a private collection formerly belonging to Jorge Aznarez.
GEOLOGICAL SETTING AND ASSOCIATED FAUNA
The specimens are contained in a ferruginous sandstone ma-
trix from the lower section of the Candeleros Member, Rio Li-
may Formation, Neuquén Group. The outcrop is located 5 km
south-east of El Chocón, Neuquén Province, Argentina. The
section is probably Albian in age, and the Candeleros Member
has been dated as Albian–Cenomanian (Calvo, 1991). The the-
ropod dinosaur Giganotosaurus carolinii (Coria and Salgado,
1995), the titanosaur Andesaurus delgadoi (Calvo and Bona-
parte, 1991), diplodocid-like dinosaurs (Calvo and Bonaparte,
1989), pipoid anurans (Báez and Calvo, 1990) and turtle re-
mains were all collected from the same formation and from
neighbouring localities (see also Calvo and Salgado, 1996, for
further information on the fossil association from this locality).
The geological sequence containing the crocodiles is thought
to be the product of flash flood conditions. The rock is a me-
dium- to fine-grained sandstone with pelitic intraclasts in tab-
ular bodies (Calvo and Gazzera, 1989). Rapid sedimentation is
indicated by the burial of this monospecific association of ar-
ticulated individuals. Sedimentological studies of Recent lake
Ezequiel Ramos Mexia show that the area is a depositional
fluvial environment crossed by an association of shallow water
bodies, some of them relatively wide, subjected to a regime of
sporadic seasonal flooding. The climate was temperate with al-
ternate rainy and dry periods (Calvo and Gazzera, 1989).
SYSTEMATIC PALEONTOLOGY
CROCODYLOMORPHA Walker, 1970 (sensu Benton and Clark,
1988)
CROCODYLIFORMES (sensu Benton and Clark, 1988)
MESOEUCROCODYLIA (sensu Benton and Clark, 1988)
Genus ARARIPESUCHUS Price, 1959
Type Species—Araripesuchus gomesii Price, 1959.
Revised Diagnosis—Short-snouted mesoeucrocodylians with
large orbits. Each orbit is dorsally covered by two palpebral
bones (primitive condition). Palpebrals do not contact each oth-
er. Rostrum with a trapezoidal section, slightly wider than high.
External surface of the rostrum lacking alveolar ornamentation
(primitive condition). Paired external nares (primitive condi-
tion), laterally exposed and anteroposteriorly expanded (auta-
pomorphy). Sharp anterior tip of the rostrum (autapomorphy),
due to the morphology of the internarial bar and the lateral
disposition of the external nares. Four premaxillary teeth. Tooth
row with different dental morphologies: conical anterior max-
illary teeth and low posterior teeth, anteroposteriorly expanded,
and without denticles at the anterior and posterior carinae. Hy-
pertrophied third maxillary tooth (derived condition shared by
basal neosuchians). Medium-sized antorbital fenestrae, slightly
larger than the foramen magnum. Flat skull table, with pitted
external surface of parietal, frontal and squamosals (derived
condition shared by Neosuchia). Posterior part of the squamosal
bends posteriorly to reach the paraoccipital process (primitive
condition). The supraoccipital lacks a sagittal crest. Occipital
region divided by a transverse crest extending between the par-
aoccipital processes. Pitted surface of osteoderms (derived con-
dition shared by Neosuchia). No multifenestrated quadrate (de-
rived with respect to Notosuchus). Infratemporal fenestra an-
teroposteriorly enlarged and laterodorsally oriented (derived
condition shared by Notosuchus). Short posterior branch of the
quadrate with posteroventrally directed condyles. Dentary ro-
bust, with a convex lateral surface. Glenoid mandibular fossa
posteriorly expanded but not as enlarged as in Notosuchus. The
palatal choana is located behind the nasopharyngeal duct and
divided by a pterygoidean bar. Depressed area behind the cho-
ana, narrower than the palatine tube (derived condition shared
by Neosuchia). Retroarticular process posteriorly expanded and
with its dorsal surface facing posterodorsally.
ARARIPESUCHUS PATAGONICUS, sp. nov.
(Figs. 2–6, 7B)
Holotype—Specimen MUCPV#269 (Figs. 2, 3, 4). Anterior
half of an articulated individual with skull and mandibles lack-
ing the most anterior portion of the rostrum. The specimen ex-
poses the precaudal dorsal armor, and bones of both shoulder
girdles (scapulae) and forelimbs (humeri, radius and ulnae).
Housed at the Museo de la Universidad Nacional del Comahue,
Paleontologı́a Vertebrados Neuquén, Argentina (MUCPV).
Diagnosis—Short prefronto-nasal suture. Lachrymal with ex-
panded dorsal surface, wider than that of A. gomesii and with
a long contact with the nasal. Postorbital bar flush with the
lateral surface of jugal, at least at its anterior border. Transverse
width of the parietal is one third of the posterior skull table
width, unlike A gomesii in which the parietal width is larger
than the squamosal.
Etymology—‘‘Patagonicus’’ refers to its occurrence from
Patagonia (Argentina).
Locality and Age—El Chocón (Embalse Ezequiel Ramos
Mexia; Neuquén Province, Argentina) from the Candeleros
Member, Rio Limay Formation, Neuquén Group (Fig. 1). The
Candeleros Member has been dated as Albian–Cenomanian, al-
though the section containing the crocodiles is likely to be Al-
bian in age (Calvo, 1991).
Referred Specimens—Specimen MUCPV#267 (Fig. 2), an-
terior half of an articulated individual preserving an almost
complete skull and attached mandibles. Specimen
MUCPV#268, postcranial articulated elements. Specimen
MUCPV#268b (Fig. 2), distal portion of tibia and fibula. Spec-
imen MUCPV#270 (Fig. 2), distal portion of a left femur in
articulation with the tibia and fibula. Specimen MUCPV#283
(Fig. 2), an isolated anterior portion of a rostrum comprising
the premaxillae and anterior tip of the maxillae, nasals and den-
taries. All the specimens considered were found in association
with the holotype.
DESCRIPTION
The sample is composed of at least four individuals, although
the presence of a fifth is very probable. The specimens have
been numbered on the basis of their relative position in the
block (Fig. 2). Beside these specimens, the same block contains
 ORTEGA ET AL.—NEW ARARIPESUCHUS FROM PATAGONIA 59

FIGURE 1. Geographic location of El Chocón, Neuquén Province, Argentina, A. The arrow indicates the outcrop where the associated specimens
of Araripesuchus patagonicus, sp. nov. were found. Simplified stratigraphic columns of the Neuquén Basin, the Neuquén Group, the local sedimen-
tology, and location where the individuals were found, B.
60 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 1, 2000
FIGURE 2. Monospecific association of Araripesuchus patagonicus, sp. nov. The sample (MUCPV#267-270) is composed of at least four
individuals; the presence of a fifth individual is very probable. Bold triangles indicate individuals, the arrow indicates the holotype. Scale bar
equals 45 mm.
more individuals probably belonging to the same taxon, but
which are yet to be examined.
Specimen MUCPV#267 consists of an almost completely
preserved skull and mandibles, with the exception of the ante-
rior-most portion of the rostrum. Dorsal and lateral views are
fully exposed, but the ventral surface is obscured by matrix.
The postcranial skeleton is in articulation, displaced a few mil-
limetres from the occipital condyle. The vertebral series is cov-
ered by the precaudal dorsal armor, with the exception of three
anterior cervical vertebrae. Both scapulae of the shoulder girdle
are present, the left being fully exposed in lateral view, and the
right being partially covered by the dorsal armor. Both humeri
are present, the right articulated with the scapula, and the left
slightly displaced from the shoulder girdle, but articulated with
the radius and ulna. Specimen MUCPV#268 exhibits part of its
caudal dorsal armor. The ilia and pubes of the pelvic girdle have
also been preserved, and some of the appendicular skeleton (left
femur, tibia and fibula). Specimen MUCPV#268b comprises a
distal portion of tibia and fibula. Their relative positions suggest
that they may belong to the left hind-limb of specimen
MUCPV#268. In the holotype specimen (MUCPV#269) the
skull has been completely removed from the matrix and
cleaned, providing a ventral view of skull and mandible. Spec-
imen MUCPV#270 is a distal portion of a left femur in artic-
ulation with the tibia and fibula. These bones might belong to
the holotype specimen (MUCPV#269), although the femoral
shaft does not fit with the direction and size of holotype. Spec-
imen MUCPV#283 is a badly crushed anterior portion of a
rostrum that does not belong either to specimen MUCPV#267
(because the hypertrophied third maxillary tooth is present in
both cases) or to specimen MUCPV#269 (because of their size).
Skull
General Description—The skull is ornamented by small pits
on the frontal and the parietal bones, there being no ornamen-
tation on the rostral bones. The length of the preserved portion
of the skull of the holotype is about 65 mm (with an estimated
total skull length of 80 mm) and about 90 mm for specimen
MUCPV#267 (estimated total length 100 mm). The known
range of the cranial length of Araripesuchus gomesii specimens
is 118 mm (holotype) and 83 mm (AMNH 24450) (Hecht,
1991). Araripesuchus wegeneri is only known from a rostral
fragment, but its total length might be as great as that of the
Brazilian type. The skull length in all the Araripesuchus is short
(80–130 mm). The largest specimen possesses a profuse orna-
mentation of wrinkles on the rostrum, unlike the Patagonian
individuals.
The skull is posteriorly broad and narrows in front of the
orbits, but does not widen at the same location as it does in
protosuchians and notosuchians. The skull is widest at the level
of the jugals, the bone bowing outward at its orbital border. The
estimated length of the rostrum is almost equivalent to the post-
rostral length (from the anterior border of the orbits to the rear
border of the skull table). The lateral contour of the rostrum in
front of the orbits is concave in dorsal view, becoming straight
 ORTEGA ET AL.—NEW ARARIPESUCHUS FROM PATAGONIA
anteriorly. The rostrum has no constriction at the premaxillo-
maxillary contact. The rostrum tapers to a narrow and acute tip.
The rostral section is slightly stretched in all the available spec-
imens although it is slightly wider than it is tall, with vertical
lateral walls. The orbits are long, constituting about one third
of the skull length and are laterally and dorsally directed.
The skull roof is typically crocodyliform, with the dorsal sur-
face of the postorbital and squamosal located at the same level
as the fronto-parietal surface. The dorsal surface of the frontal
is gently convex in its anterior portion. It descends from the
prefrontal suture, the dorsal profile of the rostrum being lower
than the skull roof in lateral view. The lateral sides of the skull
table are almost straight in dorsal view. The lateral margins of
the skull table widely overhang the otic aperture. A transverse
crest separating dorso-occipital and ventro-occipital surfaces di-
vides the occipital region. The dorso-occipital surface is a con-
tinuous shallow concavity, being bounded dorsally by a prom-
inent nuchal crest in the cranial roof and ventrally by the trans-
verse crest between the paraoccipital processes.
The major axes of the supratemporal fenestrae are antero-
posterior, and diverge anteriorly. The supratemporal fenestrae
are separated by a wide parietal surface. The fenestra is sur-
rounded by a supratemporal fossa that forms a smooth, shallow
depression, mainly comprised of the squamosal and the parietal.
A wide orbito-temporal foramen is dorsally visible, located at
the squamosal-parietal suture and the quadrate does not contact
it ventrally.
The infratemporal fenestrae face dorsally and laterally, and
are bordered by the jugal, postorbital and quadratojugal. The
infratemporal fenestra is subtriangular-shaped with its ventral
length equal to its height in the smaller specimen. In the largest
individual its length is about twice the height. In A. gomesii the
infratemporal fenestra is somewhat reduced, with subequal edg-
es, as occurs in the smallest Patagonian individual.
The antorbital fenestra is oval, laterally directed and com-
pletely surrounded by the lachrymal and the maxilla. The length
of the fenestra is one third of the orbital length.
Skull Elements
Rostral Region—The only available premaxilla (specimen
MUCPV#283) is badly crushed. The external nares open lat-
erally, and are ventrally bordered by the premaxilla and dorsally
by the nasals. The nasals probably reached the anterior pre-
maxillary symphysis, forming an internarial bar. The premaxilla
contacts the maxilla, forming an oblique sutural line, in which
the dorsal tip of the premaxilla is directed backwards. The pres-
ence of a foramen between the maxilla and the premaxilla on
the lateral sutural line is indicated by a depressed area filled
with matrix on both sides of the rostrum, although its actual
contour cannot be accurately distinguished. The external nares
are anteroposteriorly enlarged in the premaxilla and surrounded
by a smooth ventroposterior perinarial fossa. There is a foramen
between the nasal and the premaxilla of the posterior corner of
this perinarial fossa. Below this foramen, a vertical groove run-
ning towards the alveolar edge may correspond to the anterior
opening of the lachrymal duct. These two elements (foramen
and groove) are also evident in Baurusuchus and Notosuchus.
In palatal view, two fenestrae are parasagitally placed at the
contact between the premaxillae and maxillae. These fenestrae
probably connect the premaxillo-maxillary sinus (caviconchal
sinus) with the oral cavity. Their topographic position may cor-
respond to the cavitas neurovascularis, sensu Witmer (1997).
These fenestrae have also been observed in other basal me-
soeucrocodiles such as Notosuchus, the Fruita crocodile, Bau-
rusuchus and Lisboasaurus (Buscalioni et al., 1996).
The maxillary bones are vertical, although they diverge
slightly ventrally, to form a trapezoidal rostrum in which the
61
nasals form the dorsal roof. Maxillae are almost as wide as
long. Posteriorly, the maxillae end in a long narrow projection
beneath the jugal. Dorsally, the maxillojugal suture is located
in front of the orbital edge, and ventrally it extends beyond the
anterior orbital border. The ventral edge of the maxilla is slight-
ly festooned. It forms a convex wave, with its maximum cur-
vature at the level of the largest maxillary tooth (the third),
while the blunt posterior teeth are arranged along a shallow
concave wave. However, the Patagonian individuals do not have
such pronounced horizontal and vertical festooning as A. we-
generi and the type of A. gomesii. In A. patagonicus the last
maxillary teeth are caudally located on the anterior border of
the palatine fenestra.
The nasals prevent contact between the premaxillae behind
the external nares. Nasals widen backwards and meet the frontal
along a transverse suture. This suture is zigzagged as in A.
gomesii. The prefrontals contact the nasals in a short lateral
wedge at the level of the naso-frontal suture. Prefrontals and
frontal therefore meet at this suture. However, the lachrymals
contact the nasals laterally along a long suture. The relationship
between the prefrontal, lachrymal and nasal differs in A. go-
mesii. In the latter, the prefrontal touches the nasal laterally
along a long suture, while the lachrymo-nasal suture is much
shorter than in A. patagonicus.
The lachrymal is an L-shaped element with a wide dorsal
surface and a downwardly directed pillar. The dorsal portion
presents a subtriangular surface that contacts medially with the
prefrontal and the nasal. The lachrymal is narrower in A. go-
mesii than in A. patagonicus. The lachrymo-nasal suture is rel-
atively longer in the smallest individual. The orbital border of
the lachrymal is firmly attached to the anterior palpebral, while
its dorsal portion forms the roof of the antorbital cavity. The
descending pillar of the lachrymal is obliquely oriented, with
its dorsal tip anteriorly directed. At the orbital edge, the pillar
is robust and subcylindrical in section while it becomes a de-
pressed lamina at the antorbital fenestra. Ventrally, the pillar
touches the jugal posteriorly and the maxilla anteriorly. The
lachrymal expands posteriorly, forming the anterio-ventral edge
of the orbit. The lachrymal foramen is not visible because this
area is covered with sediment.
Only the dorsal surface of the prefrontal is available. This
element forms a narrow, bony band that runs parallel to the
frontal. The prefrontal does not expand laterally to any signif-
icant degree at the level of the anterior border of the orbits.
Posterolateral edges of the prefrontal contact the medial side of
the anterior palpebral.
The Temporal Region—The jugal is a triradiate bone. It is
folded so that its total height is not apparent in lateral view,
but is somewhat ventrally directed. It is outwardly bowed at
the level of the orbits. The jugal has an incipient alveolar or-
namentation. In lateral view, the jugal is as deep anteriorly as
it is posteriorly, as occurs in other oreinirostral (sensu Busbey,
1995) basal mesoeucrocodylians. The jugal almost contacts the
rear border of the antorbital fossa. The dorsal projection of the
jugal forms the ventral rod-like portion of the postorbital bar.
This projection is oblique rather than vertical, its dorsal tip be-
ing posteromedially directed. However, unlike the holotype of
Araripesuchus gomesii and specimen AMNH 24450, the inser-
tion of the postorbital bar in the main body of the jugal is
dermal, at least on its anterior edge. The posterior branch of
the jugal is a compressed bar that ends in a spine, overlapping
the lateral quadratojugal surface, and forming a V-shaped suture
with the caudally directed apex.
The quadrato-jugal forms the posteroventral border of the
infratemporal fenestra. The quadratojugal reaches the postor-
bital posterior lamina along much of its length. There is no
quadratojugal spine. The quadratojugal is narrow at the infra-
temporal fenestra, fully visible in dorsal view, and has a smooth
62 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 1, 2000

FIGURE 3. Araripesuchus patagonicus, sp. nov. (MUCP#269, holotype) from El Chocón, Neuquén Province, Argentina. A, dorsal view; B,
ventral view. Scale bar equals 10 mm.
 ORTEGA ET AL.—NEW ARARIPESUCHUS FROM PATAGONIA 63

FIGURE 5. Araripesuchus patagonicus, sp. nov. (MUCP#269, holotype) from El Chocón, Neuquén Province, Argentina. Dorsal (A) and ventral
(B) views. Abbreviations: an, angular; ar, articular; boc, basioccipital; bsp, basisphenoid; de, dentary; ect, ectopterygoid; exo, exoccipital; fr,
frontal; ju, jugal; la, lachrymal; mx, maxilla; na, nasal; pa, parietal; pal, palpebral; pfr, prefrontal; pl, palatine; po, postorbital; pt, pterygoid;
qj, quadratojugal; qu, quadrate; spl, splenial; soc, supraoccipital; sq, squamosal. Scale bar equals 10 mm.

surface. Caudally, the quadratojugal is laterally attached to the
quadrate, and does not reach the tip of the articular region.
Supratemporal Roofing Elements—The squamosal appears
triradiate in dorsal view. Its anterior projection is partially over-
lapped by the postorbital. In lateral view, it extends anteriorly
beneath the postorbital. The medial projection of the squamosal
contacts the dorsal surface of the parietal, comprising one third
of the cranial table width. Araripesuchus patagonicus differs
from A. gomesii in this trait in that the parietal is wider than
the squamosal in the latter species. The squamosal has a pos-
terior projection, forming a slender lobe that bends downward,
overlapping the paraoccipital process. In the figure of A. go-
mesii (Hecht, 1991), the squamosal overhangs the paraoccipital
process, but the description of this particular shape of the squa-
mosal is unclear. In A. patagonicus the suture between the squa-
mosal and the quadrate is visible laterally, and the squamosal
does not cover the dorsal ascending process of the quadrate at
the otic recess. Along the lateral edge of the skull roof, the
squamosal bears a tiny longitudinal groove, homologous to the
area for the insertion of the ear-flap muscles in modern croco-
diles.
The parietal is unpaired. There is no notable constriction be-
tween the supratemporal fossae, this area being as wide as that
of the interorbital. Its posterior margin cannot be clearly de-
fined, but it is likely that the dorsal surface of the supraoccipital
bone is barely exposed on the cranial table.
The frontal is an unpaired element. There is no trace of the
interfrontal suture even in the smallest specimen. The fronto-
parietal surface is flat and wide. The fronto-parietal suture runs
transversally, and is located on the anterior border of the su-

FIGURE 6. Araripesuchus patagonicus, sp. nov. (MUCP#269, holotype) from El Chocón, Neuquén Province, Argentina. Occipital (A) and
lateral (B) views. Abbreviations: additional to Figure 5; ot, otic notch. Scale bar equals 10 mm.

←
FIGURE 4. Araripesuchus patagonicus, sp. nov. (MUCP#269, holotype) from El Chocón, Neuquén Province, Argentina. A, lateral view; B,
occipital view. Scale bar equals 10 mm.
64 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 1, 2000
pratemporal fenestrae. The frontal forms part of the fossa, and
hence the parietal and the postorbital do not contact each other
on the skull table surface. A. gomesii (holotype and
AMNH#24450) shows the same disposition of the frontal, al-
though Hecht’s (1991) figure indicates a postorbital contacting
the parietal on the skull table. The frontal is not narrow between
the orbits.
The postorbitals present a narrow dorsal surface, with a
slightly concave anterior edge in dorsal view. At its anterolat-
eral corner, a lower shelf is differentiated from the dorsal post-
orbital surface, on which the posterior palpebral rests. The post-
orbital bar is inserted on a dorsally compressed and anteropos-
teriorly expanded lamina on the ventral surface of the main
body of the bone. The bar is robust and becomes rod-like ven-
trally. The postorbital bar contacts the jugal where it forms an
interdigitate suture. A vascular foramen can only be observed
in A gomesii, since this area is covered by matrix in A. pata-
gonicus.
Suspensorium—The cranial branch of the quadrate is more
laterodorsally inclined than in modern crocodiles. It has two
planes in lateral view, both of which are divided by a ridge
running parallel to the quadrate-quadratojugal suture. The lat-
eral plane is almost dorsally oriented, while the medial one is
more vertical. The dorsal quadrate surface is depressed around
the otic recess area. Medially, and anterior to this depressed
area, the quadrate has a wide preotic foramen. Posteriorly, the
quadrate forms the otic notch. Behind the otic notch the quad-
rate bears a dorsal sheet that forms the lateral surface of the
paraoccipital process. The posterior articular branch of the
quadrate is extremely short. The quadrate condyles do not ex-
tend beyond the base of the paraoccipital processes caudally,
and are as ventrally deflected as in Notosuchus. Both condyles
are equally developed. The medial branch of the quadrate ex-
tends beneath the exocipital, so that the quadrate is barely vis-
ible occipitally. The ventral surface of the quadrate is smooth,
without crests. The pterygoidal branch of the quadrate poste-
riorly contacts the exoccipital, but not the basioccipital, as is
the case in modern crocodiles. The quadrate has a relatively
short and narrow pterygoidean branch. The medial sutural line
of the ventral surface of the quadrate contacts the basisphenoid
along much of its length, avoiding the pterygoid and exoccipital
connection.
Braincase—The basisphenoid is broadly exposed in ventral
view. It broadly contacts posteriorly the exoccipital and, sagit-
tally, the basioccipital. Its anteroposterior length is similar to
that of the basioccipital. The ventral surface of the basisphenoid
is longitudinally crested by two prominent parasagittal ridges,
as in Protosuchus (Clark, 1986). These ridges occur in a similar
position to the distal ridges of the pterygoidal quadrate branch
in modern crocodylians.
The basioccipital is mainly ventro-occipitally oriented. The
occipital condyle is posteriorly oriented. The ventral tip of the
basioccipital forms the rear notch of the eustachian foramen.
The basioccipital plate has a sagittal crest and two shallower
parasagittal ones in front of the lateral eustachian foramina.
There are no basioccipital tubera present.
The exoccipitals form the main portion of the occipital view.
The surface of the exoccipitals is divided into two different
planes separated by a transverse crest that extends between the
paraoccipital processes. The two exoccipitals meet each other,
preventing the participation of the supraoccipital in the border
of the foramen magnum. The position of the foramina of the
cranial nerves and carotid artery cannot be accurately de-
scribed, since this area is poorly preserved. The exoccipitals
form the medial border of the cranioquadrate canal, which is
laterally enclosed by the quadrate and the squamosal.
The supraoccipital is barely exposed in dorsal view. Occip-
itally, the supraoccipital exposes a tall and transversely wide
polygonal flat surface. The postoccipital processes are poorly
developed, despite the presence of posterotemporal foramina.
Palate—The palate is partially visible in MUCPV#269, in
which the lower jaw is in articulation. The palatal surface (max-
illary and palatine secondary palate) is smooth and flat. The
palatines meet along the midline, enclosing the nasopharyngeal
duct and forming the rear extension of the secondary palate.
The caudal end of the palatine secondary palate is not continued
by the pterygoids, and the internal nares are formed by the
palatines.
The dorsal surface of the pterygoidal wings is flat rather than
concave, as seen in modern crocodiles. A sagittal sutural line
is evident along almost all of its length. The anterior extension
of the primary palate of the pterygoids forms a shallow con-
cavity and contacts the palatine primary palate. This concavity
forms the rear extension of the nasopharyngeal groove. The
internal nares are long and are longitudinally divided by a sep-
tum formed by a spine of the pterygoids. This spine has a T-
shaped transverse section. Anteriorly, this projection contacts
the palatal secondary palate, and slightly separates the rear ends
of both palatines. This condition could be preservational, but it
is also shared by the specimen of Araripesuchus gomesii drawn
by Hecht (1991). The ptegyoidean surface is rough. The de-
pressed area behind the internal nares is narrower than the pal-
atine tube. The posterior outline of the pterygoidean flanges
possesses a depressed, sagittal notch in which the anterior end
of the basisphenoid is lodged. The left ectopterygoid is slightly
exposed and attached to the lateral portion of the pterygoidean
flange, but no further details are available.
Palpebral—All the Araripesuchus have two palpebral ossi-
fications. The posterior palpebral is small, wider than it is long,
and restricted to the dorso-posterior area of the orbit. It is
lodged in a slight depression formed by the antero-lateral pro-
jection of the postorbital and probably does not contact either
the frontal or the anterior palpebral. The anterior palpebral is
hatchet-shaped, elongated anteroposteriorly, with a notch on its
posterior margin. Its antero-medial border is firmly sutured to
the lachrymal and prefrontal.
Dentition—There are four premaxillary teeth, the last being
the largest. The number of maxillary teeth cannot be directly
established in the specimens, but 11 to 13 would be a reason-
able estimate. Heterodonty is characterized by the change of
size and shape of the maxillary dental series. The anteriormost
teeth are subconical, sharp, and with the apex directed back-
wards. No denticles are present in the mesial and distal margins
(i.e., teeth not serrated). The third tooth is the largest (more
than twice the height of any other tooth). Behind the eighth
tooth, the maxillary teeth become blunt, with mesiodistally wid-
ened crowns. Their labial surfaces are hemispherical, and they
have a constriction at the base of the crowns, almost forming
a cingulum. Their apices are blunt without a clear acuminated
end. The worn facets are located at their distal side. The max-
illary teeth series occlude external to the dentary teeth. The
dentary teeth are more conical in shape in the caudal part of
the maxilla. Specimens can be considered to show marked het-
erodonty in shape, but size changes gradually along the series,
and unlike eusuchians, there are no undulations of size variation
in the posterior dentary and maxillary series.
Mandible—The lateral dentary surface has a lined set of
foramina running below the alveolar series over a smooth area.
The dentary ornamentation is composed of rare pits and an-
teroposterior grooves, with a smooth posterior area. The dentary
has a convex lateral profile that is dorsal and ventrally deflected.
In A. wegeneri the dentary is vertical, with a flat lateral surface.
The dentary is not as tall as in primitive crocodylomorphs. The
dentary rises slightly behind the dental series, but its relation-
ship with the surangular is hidden by the skull. Neither the
dentary teeth nor the dorsal edge of the dentary are fully visible
 ORTEGA ET AL.—NEW ARARIPESUCHUS FROM PATAGONIA 65

FIGURE 7. Schematic drawing not to scale of the cranio-mandibular articulation of (A) Araripesuchus gomesii (AMNH#24450) in dorsolateral
view; grey line represents the reconstructed articular. (B), A. patagonicus in dorsolateral view. and (C), Notosuchus terrestris (specimens from
MUCP) in dorsal view. Quadrate articulation shadowed. Abbreviations: additional to Figure 5; c, crest along the mandibular glenoid fossa; ef;
external fossa; if, inner fossa; gl, glenoid fossa; rar, retroarticular process. The arrows delimit the cranio-mandibular articulation in Araripesuchus.

in the specimens, since the maxillary teeth occlude externally.
The dentary symphysis is not completely preserved. The splen-
ials form part of the mandibular symphysis, forming a stout
sutural join, and probably constitute half of the symphysial
length. The splenial deflects its lamina ventrally and is exposed
in this view. However, in A. wegeneri the splenial constitutes
more than half the width of the ventral surface of the mandible
in the postsymphysial region. An external mandibular fenestra
is displayed, bordered by dentary, surangular and angular. The
angular has a very restricted alveolar ornamentation below the
external mandibular fenestra. The angular borders the retroar-
ticular process laterally, and its internal lamina is low. In ventral
view, the articular is anteroposteriorly enlarged and possesses
a short horizontal descending process. It expands medially with
a convex medial contour. The ventro-lateral edge of the retroar-
ticular is covered by the spine of the angular. The dorsal surface
of the retroarticular process is notably wide and flat. There is
no evidence of the foramen aërum on the dorsal surface. The
retroarticular process does not curve dorsally, as in neosuchi-
ans, although it faces dorsally and has a slight medial deflec-
tion. The mandibular articular surface extends backwards and
medially to the quadrate condyles, and does not have a posterior
buttress of the articular, as occurs in modern species. This is
not apparent in A. gomesii because of its state of preservation.
However, in lateral view, the articular area of AMNH 24450 is
enlarged, as is the case in the Patagonian specimens (Fig. 7A).
This trait has been previously considered exclusive to notosu-
chians (Clark et al., 1989; Gomani, 1997), and, in fact, the
retroarticular process of Notosuchus is much more extensive
than that of Araripesuchus (Fig. 7C).
Postcranial
Osteoderms or matrix mostly hides the axial skeleton. The
few vertebrae that are exposed are amphicoelous. The neural
spine of the atlas is ring-shaped. An accurate description of the
axis of the neural spine is difficult since it is broken away at
the base. Cervical diapophyses and long, stout ribs are exposed.
Scapulae of two individuals are the only preserved elements
of the pectoral girdle. The scapular lamina is dorsally broad
(three times its ventral anteroposterior width), and expanded
particularly in the anterior part. The scapula has a concave an-
terior border and a straight, or slightly convex, posterior one.
A weak prominence is anteriorly developed at the level of the
acromial area. Laterally and medially a keel-like prominence
forms the posterior edge above the level of the glenoid up to
66 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 1, 2000

FIGURE 8. Cladogram summarising the definition of Mesoeucrocodylia. The clade Ziphosuchia is here exemplified by Notosuchus, and Neo-
suchia by Theriosuchus (see Appendix 4 for the extended phylogenetic analysis). Araripesuchus is proposed as the closest sister taxon of Neosuchia.
The genus Araripesuchus is composed of two species: A. gomesii and A. patagonicus. ‘‘Araripesuchus’’ wegeneri should be reassigned to a new
genus. It is discussed either as a sister taxon of Araripesuchus plus Neosuchia, or as a sister taxon of Araripesuchus. Unambiguous diagnostic
characters for each node are discussed in the text. Node 1, Mesoeucrocodylia; node 2, unnamed node; node 3, Neosuchia.

the mid-constriction of the scapular blade. This ridge may cor-
respond to the site of attachment of a head of the triceps mus-
culature.
The humerus has a straight, cylindrical shaft, without the
pronounced posterior curvature of the shaft of the recent spe-
cies. The proximal lateromedial width is wider than the distal
articulation. The humeral head is reflected posteriorly, below
which there is a pit or depressed area on the posterior surface.
The deltopectoral crest is well-developed and it has a convex
lateral profile. It extends along the first third of the shaft length.
The inner tuberosity bends ventrally. There is a prominent,
well-developed proximodistal ridge on the posterior surface, be-
low the lateral tuberosity, with a pit at the distal end of its ridge.
The distal condyles are anteroposteriorly enlarged. In lateral
view, the development of the lateral condyle modifies the distal
end of the shaft, forming a flat, expanded surface. No anterior
view is available in the specimens.
The radius and the ulna are shorter than the humerus. Few
fine details are preserved. Both shafts are cylindrical. The ulna
is a slender bone of about the same length as the radius. Un-
fortunately, distal forelimb elements are not exposed.
The pubis and ilium constitute the two preserved elements
of the pelvic girdle. Iliac blades have long posterior, laterally
curving processes. The pelvic girdle is notable for its deep ac-
etabular area and the dorsal margin of the iliac blades. Ante-
riorly, the ilium is poorly preserved; although it probably pos-
sesses a short anterior process. The pubis has a proximal rod-
like shaft and is flat and distally expanded.
Two femora are exposed. The femoral shaft is twisted so that
the proximal head and the distal condyles form an angle of
about 608, as in extant species. The femoral head is short with
respect to the shaft length. Distal condyles are hidden by other
bones or by the matrix. However, the lateral condyle is antero-
posteriorly wide.
The tibia is as long as the femur. It has a proximal head with
two articulating surfaces separated by a crest. Proximally, its
lateral border is taller than the medial one. A posterior view of
the tibia is exposed in one specimen, showing the distal artic-
ulating surface. The proximal fibular end is medially curved.
Dermal Armor
Two paramedian rows comprise the dorsal armour covering
the entire animal body, and range continuously from the occip-
ital area (except for the first three cervical vertebra, which are
uncovered). There is no evidence of ventral scutes. The osteo-
 ORTEGA ET AL.—NEW ARARIPESUCHUS FROM PATAGONIA
FIGURE 9. Holotype of ‘‘Araripesuchus’’ wegeneri from the Aptian
of Gadoufaoua (Tegama Basin, Niger). A, ventral view; B, lateral view;
C, detail of the fourth to sixth maxillary teeth. Abbreviations: addi-
tional to Figure 5; 3 mx, third maxillary tooth; 5 pmx, fifth premaxillary
tooth; orb, orbit. Scale bar equals 2 cm.
derms are thin and smoothly sculptured with infrequent and
shallow radiating pits. Osteoderms are wider than they are long,
with an unsculptured anterior articulating area. No anterolateral
peg is apparent, probably due to the state of preservation. Os-
teoderms have a dorsal keel, dividing each into two parts. The
external part of the osteoderm bends ventrally and is slightly
exposed laterally. The osteoderm shape changes along the dor-
sal armour; the anterior ones are rectangular while the caudal
ones are square and more profusely sculptured. The distal edges
become rounded, as osteoderms become more caudal.
DISCUSSION
The Phylogenetic Context of Araripesuchus
To place Araripesuchus in a phylogenetic context, we discuss
below the apomorphic traits that it shares with Notosuchus, and
Neosuchia, based on a phylogenetic analysis of Crocodylifor-
mes detailed in the Appendices 1–4. The synapomorphies ex-
plained below unambiguously diagnose the internal nodes of
the cladistic analysis discussed in Appendix 4 and summarised
in Figure 8. We define Mesoeucrocodylia comprising Zipho-
suchia 1 Araripesuchus 1 Neosuchia. Ziphosuchia is a new
proposed taxon that involves Notosuchus 1 Libycosuchus 1
Sebecosuchia. This new taxon will be elsewhere discussed in
67
detail. According the present analysis the Fruita Formation
crocodile is not placed within Mesoeucrocodylia, unlike the re-
sult shown in previous phylogenetic analysis (Benton and
Clark, 1988; Clark, 1994).
Members of Mesoeucrocodylia (Notosuchus and Araripesu-
chus 1 Neosuchia in Fig. 8) share the differentiation of the
posterior region of the skull. The infratemporal fenestrae are
dorsolaterally oriented (see character 46 in Appendix 1). The
plesiomorphic condition shared by basal crocodyliforms and the
Fruita Formation crocodile is interpreted as a vertical infratem-
poral fenestrae that is also dorsally covered by the lateral ex-
tension of the skull roof. At this node, the anteroventral border
of the quadratojugal extends forward to form part of the ventral
margin of the infratemporal fenestra (character 39). The closure
or disappearance of a depression on the dorsal surface of the
posterior branch of the quadrate delimiting the attachment area
of the tympanic membrane (Walker, 1990) is another diagnostic
trait (character 154) of node 1 (Fig. 8). In the palatal region of
Notosuchus 1 (Araripesuchus 1 Neosuchia) the ectopterygoid
contacts the maxilla preventing the participation of the jugal in
the suborbital fenestrae (character 61) and pterygoids are cau-
dally fused (character 58). Members of this node bear a sepa-
rated foramen for the cranial nerve IX (character 64). The ab-
sence of a premaxillo-maxillary notch (character 14) also di-
agnoses node 1 (Fig. 8). In these taxa the symphysis of the
dentary lengthens posteroventrally, enlarging the sutural area
(character 151), while in the primitive condition the symphysis
is short and vertically oriented. All Mesoeucrocodylia (as di-
agnosed in the Appendix 4) share the posterior extension of the
secondary palate involving the palatines and the position of the
internal choana, which is located on the rear border of these
elements. This condition is not an unambiguous trait of node 1
since the Fruita Formation crocodile also shares it.
Araripesuchus definitely does not constitute a clade with No-
tosuchus, but instead must be considered as the sister group of
Neosuchia (node 2, Fig. 8). This node is diagnosed by the ap-
pearance of a pitting pattern of ornamentation on the dermal
bones (characters 1 and 111). Initially, in the basal members of
this node and basal neosuchians (e.g., Theriosuchus, Pelago-
saurus) this pattern mainly involves osteoderms and dermal
bones of the cranial roof (parietal, frontal and squamosal). The
rostrum is differentiated in a number of ways related to the
trend toward the platyrostral condition (sensu Busbey, 1995),
and the rise of heterodonty (characters 20 and 132). The pla-
tyrostral skull is defined by a set of characters shared by Neo-
suchia and which are exaggerated in modern crocodiles. Al-
though Araripesuchus does not have a platyrostral skull, it nev-
ertheless follows the trend towards it, with a rostrum that is
slightly wider than it is tall. Heterodonty is expressed in the
maxilla in which the dental series is variable in size and shape.
In the basal members of the Neosuchia clade (except in the
marine crocodiles) the third maxillary tooth is the largest, as in
Araripesuchus, whereas in more advanced members of Neo-
suchia the fourth and fifth maxillary teeth become enlarged.
Furthermore, the teeth of the members of this node are not
disposed in a groove (character 19). In Araripesuchus and other
basal neosuchians (i.e., Pelagosaurus, Theriosuchus), the pre-
frontal pillars are transversely broad at their roof but more rod-
like at their base (character 30). On the other hand, the pillars
form a near-complete transverse wall where it makes contact
with the palatines in the plesiomorphic condition (shown in
Dibothrosuchus and Notosuchus). The palatal construction of
Araripesuchus and Neosuchia exhibits a narrowing of the post-
choanal depression behind the nasopharyngeal duct (character
139), unlike Notosuchus, in which it is expanded transversely.
In the members of node 2 the palatal surface is flat or even
convex (character 175) instead of the primitive, concave con-
dition. In Araripesuchus 1 Neosuchia, the retroarticular process
68 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 1, 2000

FIGURE 10. Reconstruction of the skull of Araripesuchus patagonicus, sp. nov. (A) in comparison with A. gomesii (B) in dorsal view. The
Patagonian species have a set of juvenile traits and proportions: large orbits; bowed jugals; and large orbital/rostrum length. The species differ
in: (1) the dorsal extension of the prefrontal and lachrymal bones; (2) the enlargement of the squamosal; and (3) the inset/outset placement of
the ascending process of the jugal to form the postorbital bar. The skull length of the holotype of A. gomesii is 118 mm, and A. patagonicus is
about 80 mm. Although the specimens used for the reconstruction of the skull have unequal skull length the differences reported are also evident
in the smallest specimen of A. gomesii (Hecht, 1991).

faces dorsally, instead of medially, as is the case in Notosuchus.
Finally, the lateral wall of the dentary is convex (character 81)
and the inner shelf of the retroarticular process is dorsally ex-
posed (character 147) in members of node 2 (Fig. 8), differing
from the flat vertical wall of Notosuchus.
The distribution of unambiguous characters discussed here
places Araripesuchus in a phylogenetic position that is taxo-
nomically consistent with that proposed by Clark (l994), i.e.,
as the sister group of Neosuchia. In the cladogram depicted by
Clark (1994) Baurusuchus, Notosuchus, Sebecus and Libyco-
suchus appear as the basal mesoeucrocodylians, forming suc-
cessive sister groups of (Araripesuchus 1 Neosuchia). The de-
rived conditions of the characters that diagnose Araripesuchus
1 Neosuchia mentioned above are not shared by any of the
other basal mesoeucrocodylians (i.e., Baurusuchus, Sebecus,
Bretesuchus, Libycosuchus, and Iberosuchus, which, in turn, are
members of Ziphosuchia) considered in Figure 11.
Two autapomorphic traits (not included in the character list
of Appendix 1) defining Araripesuchus have been remarked
upon in the systematic paleontology section: an acuminate an-
terior rostral contour, due to the projection of the nasals be-
tween the external nares, forming an internarial bar, and the
lateral orientation of the external nares, which are extended an-
teroposteriorly. The two species of the genus Araripesuchus (A.
patagonicus 1 A. gomesii) share a dental morphology (char-
acter 131) whereby the teeth are mesiodistally enlarged (con-
vergent in Theriosuchus and Bernissartia). They also share the
posterior extension of the dorsal surface of the glenoid articular
fossa (character 105) without a posterior buttress (as occurs in
Notosuchus and Malawisuchus) (Gomani, 1997). The develop-
ment of this surface is much larger in Notosuchus and Mala-
wisuchus. In Notosuchus, the articular fossa has a bilobate
asymmetrical surface, divided by a medium-shallow crest. The
lobes delimit two trails for the anteroposterior movements of
the quadrate condyles. The external fossa is longer than the
medial one. In Araripesuchus the dorsal surface of the articular
fossa is small and rather flat (Fig. 7).
Other Species Assigned to the Genus Araripesuchus
Buffetaut (1981) described Araripesuchus wegeneri, based
on a rostral fragment with joint mandibles (Fig. 9) from the
Aptian of the Elrhaz Formation (Niger). The anterior tip of the
rostrum and the most anterior part of the dentary are eroded.
Araripesuchus wegeneri may well belong to Mesoeucrocodylia,
but there is a lack of information concerning the nature of the
palate. Nonetheless, the specimen has a secondary palate
formed by the palatines, and although the palatine nasopharyn-
geal duct has broken off, the choana should be located, at least,
at the caudal end of the palatines. Araripesuchus wegeneri
shares the pattern of maxillary heterodonty and the flat palatal
surface with Neosuchia. As in the South American Araripesu-
chus, A. wegeneri has a trapezoidal section of rostrum. Arari-
pesuchus wegeneri differs from the South American Araripe-
suchus in a number of features. It has five premaxillary teeth,
a widespread condition in Crocodylomorpha, but absent in Ar-
 ORTEGA ET AL.—NEW ARARIPESUCHUS FROM PATAGONIA 69

FIGURE 11. Concensus tree of two most parsimonious solutions resulting form a strict parsimony analysis (see Appendices 1–4). Data matrix
processed using PAUP 3.1.1 (Swofford, 1993). The tree has a length of 388 steps. Consistancy Index is low (0.469), but Retention Index is high
(0.745). The two most parsimonious trees differ only in the placement of Thalattosuchia within Neosuchia. Either as the sister group of Itasuchus
1 all remaining neosuchians, or as the sister group of Goniopholis 1 Bernissartia 1 Las Hoyas Neosuchia 1 Eusuchia. (See Appendix 4 for
information on nodes.)

aripesuchus gomesii and A. patagonicus. Furthermore, the last
premaxillary tooth is the largest in the South American species
whereas the fifth is very small in A. wegeneri. On the other
hand, the probably paired external nares in the latter species
are directed frontally rather than laterally, which is an autapo-
morphic trait of the Southamerican species. The jugal is deep
and flat instead of narrow, folded and bowed outwards. The
mandible of A. wegeneri has a dentary with a deep, vertical and
flat posterior lamina. Although the mandible of A. wegeneri is
incomplete, its lateral profile is primitive with respect to that of
Araripesuchus and Neosuchia. The splenial is widely exposed
ventrally in A. wegeneri, with the ventral mandibular width
formed by the equal participation of the splenial and dentary (a
condition shared by Baurusuchus and Notosuchus). In Arari-
pesuchus the splenial is less involved in the formation of the
ventral surface of the mandible. The crown morphology of the
posterior maxillary teeth is similar to that of A. patagonicus
and A. gomesii: low, blunt, mesiodistally enlarged teeth, with
tips that are not distally curved. However, there are denticles
on the margins of the teeth of A. wegeneri, with fine enamel
ridges on the labial side, and a constricted neck. Consequently,
taken together, these differences suggest that the African spe-
cies does not share the diagnostic traits proposed here. The
reassignment of Araripesuchus wegeneri to a new genus should
therefore be considered. We will refer it as ‘‘Araripesuchus’’
wegeneri until a description of new African material becomes
available.
The other crocodile that has been postulated as being a close
relative of Araripesuchus is Uruguaysuchus from the Upper
Cretaceous of Uruguay. It is heterodont, and it also has the
parietal, frontal, squamosals and osteoderms ornamented with
an alveolar sculpture. Both traits are derived and also shared
by Araripesuchus 1 Neosuchia. Uruguaysuchus has played a
key role in the revision of the taxonomic status of the Uru-
guaysuchidae family (Gasparini, 1971), in which Araripesuchus
gomesii was included as a member. However, its phylogenetic
relation to the latter genus have not yet been determined, and
unfortunately, the type species is not available for revision. It
is possible that, as in the case of the African species ‘‘A.’’
wegeneri, Uruguaysuchus was a member or the sister taxon of
the node defined by Araripesuchus and Neosuchia.
Biogeographic Implications of the ‘‘Araripesuchids’’
Araripesuchus gomesii and Notosuchus are the South Amer-
ican mesoeucrocodylians that have received most attention, not
only with regard to their phylogenetic relationships, but also to
their paleobiogeographic nexus with other related African spe-
cies (Gasparini, 1996; Gomani, 1997). The ‘‘araripesuchids’’
(Araripesuchus and ‘‘Araripesuchus’’ wegeneri) have been suc-
70 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 1, 2000
cessively used to date the African–South American splitting
(Buffetaut and Taquet, 1979). Based on the shared vertebrate
fauna of South America and Africa, some authors have postu-
lated that the two continents were connected by a land nexus
during the Aptian (Buffetaut and Rage, 1993). Others have sug-
gested that this land bridge was viable at least up to the Albian–
Cenomanian (Calvo and Salgado, 1996; Gasparini et al., 1998).
However, the faunal similarities between Africa and South
America involving phylogenetically related taxa, do not nec-
essarily presuppose the existence of a continental nexus by the
Aptian, and the archosaurian fauna could have been previously
isolated in each continent. In fact, Kellner (1994) has argued
that there is no coincidence at a specific or even generic level
in the amphiatlantic fish and reptiles.
In this study, we have stressed the differences between ‘‘Ar-
aripesuchus’’ wegeneri and South American species, arguing
that the former does not belong to the genus. Although the
phylogenetic context of ‘‘A.’’ wenegeri is not fully resolved,
the species shares some derived conditions (see above) with
node 2 (Fig. 8), so that it might therefore be considered either
as the sister taxon of Araripesuchus 1 Neosuchia, or as the
sister group of Araripesuchus. In the first case, it would rep-
resent a splitting from a common Jurassic pangeic ancestor
(since the Neosuchia fossil record is as early as Middle Juras-
sic), whereas in the second, it would imply the sharing of a
pre-Aptian Gondwanic common ancestor with Araripesuchus.
In this later case the distribution of ‘‘A.’’ wegeneri and the
South American Araripesuchus should be explained as a vicar-
iance event. Similar explanations of relationships and biogeo-
graphic distribution of the notosuchids Notosuchus and Mala-
wisuchus might be proposed. Our results disprove the use of
the distribution of the notosuchids and the ‘‘araripesuchids’’
crocodilian faunas to support hypotheses that propose a terres-
trial nexus between both Gondwanic continents (Africa and
South America) after the Albian.
ACKNOWLEDGMENTS
The specimens of A. patagonicus were found by the geologist
A. Gazzera and one of the authors (J. C. Calvo). Fieldwork was
carried out by the Museo Nacional ‘‘Bernardino Ribadiavia’’
and the Universidad Nacional del Comahue. Restoration of the
specimens was carried out in the Museo Nacional ‘‘Bernardino
Ribadiavia’’ and Museo de La Plata. Photographs of ‘‘A.’’ we-
generi by D. Serrette from the Museum national d’Histoire na-
turelle de Paris. We want to thank also France de Lapparent
who provided the ‘‘A.’’ wegeneri photographs. Photographs of
A. patagonicus by Gerardo Kurtz. Funds for the study were
provided by the Programa de Estancia de Investigadores Ex-
tranjeros en España en Regimen Sabático (to Z. Gasparini), the
Convenio UAM-UNLP, and the DGICYT Program from the
Ministerio de Educación y Cultura del Estado Español, Project
number PB97-0061. We also thank B. P. Pérez-Moreno for his
useful comments on the manuscript.
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APPENDIX 1
List of characters used in this analysis. Characters are either reela-
borated from different authors (Molnar, 1981; Clark, 1986, 1994; Clark
et al., 1989; Buscalioni and Sanz, 1990; Benton and Clark, 1988; Norell
and Clark, 1990; Parrish, 1991; Gasparini et al, 1991, 1993; Sereno and
Wild, 1992; Wu and Chatterjee, 1993; Wu et al., 1994, 1995; Wu and
Sues, 1996; Legasa et al 1994; Ortega and Buscalioni, 1995; Busbey,
1995; Ortega et al., 1996), or are newly developed based on the direct
study of most of the taxa included in the analysis.
1. Ornamentation of external surface of cranial dermal bones: smooth
or formed by grooves and ridges (0); with circular or subpolygonal
pits (1).
2. Sculpturing of palatal surface: maxillary palatal surface smooth (0);
maxillary palatal surface ornamented with ridges (1).
3. Rostral length: distance from anterior orbital edge to anterior con-
tour of rostrum equal or longer than distance from anterior orbital
edge to posterior parietal contour (0); distance from anterior orbital
edge to anterior contour of rostrum shorter than distance from an-
terior orbital edge to posterior parietal contour (1).
4. Rostral length: distance from anterior orbital edge to anterior con-
tour of rostrum shorter, equal or slightly longer than distance from
anterior orbital edge to posterior parietal contour (0); distance from
anterior orbital edge to anterior contour of rostrum at least twice
than distance from anterior orbital edge to posterior parietal contour
(1).
5. Rostral section: tubular, almost as deep as wide (0); wider than deep
(1).
6. Premaxillo-maxillary suture in lateral view: vertical (0); caudodor-
sally directed (1).
7. Premaxillo-maxillary joint: premaxilla overlapping maxilla (0); pre-
maxilla and maxilla sutured (1).
8. Premaxillo-maxillary suture in lateral view: straight (0); zigzag
shaped (1).
9. Direction of premaxillo-maxillary suture in palatal view: cranially
directed (0); caudal directed (1). (Direction of suture is evaluated
with respect to a theoretical line that passes between the lateral
contact of both bones).
10. Ventral edge of premaxilla with respect to ventral edge of maxilla
in lateral view: placed almost at same height (0); deeper, and an-
terior dorsal contour of dentary is also strongly concave (1).
11. Oral cavity communicates with nasal cavity through palatal perfo-
rations (naso-oral fenestrae), besides internal nares: no (0); yes (1).
12. Naso-oral fenestrae formed by: maxilla and premaxilla (0); pre-
maxilla (1).
13. Foramen at premaxillo-maxillary suture in lateral view: present (0);
absent (1).
14. Premaxillo-maxillary notch: absent (0); present (1).
15. Last premaxillary alveolous the largest of premaxillary tooth row:
no (0); yes (1).
16. External nares: facing frontal, laterofrontal or frontodorsally (0);
facing dorsally (1).
17. Position of external nares with respect to anterior rostral contour in
dorsal view: concealed (0); a premaxillary bar separates external
nares and anterior rostral contour (1).
18. Relative position of last maxillary tooth with anterior edge of pal-
atine fenestra: last maxillary tooth caudal to anterior edge of pal-
atine fenestra (0); last maxillary tooth cranial to anterior edge of
palatine fenestra (1).
19. Dental implantation: teeth set in isolated alveoli (0); teeth set dis-
posed in a groove (1).
20. Size of maxillary teeth: all maxillary teeth similar in size or with
the largest alveolous placed at middle of maxillary row (0); tooth
row with waves of size variation (1).
21. Ventral edge of maxilla in lateral view: straight or convex (0); si-
nusoidal (1).
22. Rostral tip of nasals: nasals reach the caudal edge of external nares
(0); nasals do not reach the caudal edge of external nares (1).
23. Rostral tip of nasals: nasals reach premaxillae (0); nasals do not
reach premaxillae (1).
24. Caudal tip of nasals: caudally nasals converge at sagittal plane (0);
nasals caudally separated by an anterior sagittal projection of frontal
(1).
25. Frontals: separated (0), fused (1).
26. Dorsal fronto-parietal surface: reduced to a narrow bar (0); wide
(1).
27. Parieto-postorbital suture: absent on dorsal surface of skull roof (0);
present on dorsal surface of skull roof (1).
28. Parietals: separated (0); fused (1).
29. Prefrontal pillars: do not reach palate (0); reach palate and solid
integrated (1).
30. Prefrontal pillars when integrated in palate: pillars transverselly ex-
panded (0); transversely expanded in their dorsal half and columnar
ventrally (1); pillars longitudinally expanded in their dorsal part and
columnar ventrally (2).
31. Postfrontals: present (0); absent (1).
32. Antero-lateral spine of postorbital: absent (0); postorbitals project
a short and sharp spine (1).
33. Relative length between postorbital and squamosal: squamosal is
longer (0); postorbital is longer (1).
34. Jugal portion of postorbital bar: flushes with lateral surface of jugal
(0); medially displaced and a ridge separates postorbital bar from
lateral surface of jugal (1).
35. Caudal tip of ectopterygoidean-jugal contact: placed cranial to post-
orbital bar (0); placed caudal and surpassing postorbital bar (1).
36. Ectopterygoid-postorbital suture: ectopterygoid does not contact
postorbital (0); ectopterygoids contacts postorbital on medial side
of postorbital bar (1).
37. Postorbital and jugal forming postorbital bar in lateral view: jugal
sets caudal to postorbital (0); postorbital is medial or caudal to jugal
(1).
38. Jugal and quadratojugal in lateral view: quadratojugal visible be-
neath jugal (0); quadratojugal is not exposed (1).
39. Corner of infratemporal fenestra in lateral view: jugal-quadratojugal
suture lies at posteroventral corner (0); quadratojugal extends an-
teriorly forming part of dorsal edge of infratemporal bar (1).
40. Skull roof: not developed, postorbito-squamosal dorsal surface on
a lower plane than fronto-parietal dorsal surface (0); developed,
postorbito-squamosal and fronto-parietal surfaces are on the same
plane (1).
41. Supratemporal fenestrae: relatively large, covering most of surface
of skull roof (0); relatively short, fenestrae surrounded by a flat and
extended skull roof (1).
42. Outer surface of squamosal: laterodorsally oriented (1); dorsally
oriented (1).
43. Contour of squamosal in dorsal view: curved (0); L-shaped (1).
44. Quadrate inclination with respect to a horizontal plane including
the cranial roof: craniocaudal axis of quadrate inclined more than
45 degrees (0); craniocaudal axis of quadrate inclined less than 45
degrees (1).
45. Longitudinal groove on dorsolateral surface of squamosal: absent
(0); present (1).
46. Infratemporal fenestrae: facing laterally (0); facing laterodorsally
(1).
47. Quadratojugal spine at caudal margin of infratemporal fenestrae:
absent (0); present (1).
48. Quadratojugal contacts postorbital: widely (0); punctually, quadra-
tojugal is dorsally acute, or there is no contact at all (1).
49. Quadratojugal contacts postorbital: yes (0); no (1).
50. Quadrate fenestrated: quadrate lacking fenestrae (0); quadrate fen-
estrated even it possesses one fenestra which in such case is placed
on dorsal surface of the cranial ascendent process of quadrate (1).
51. Quadrate fenestrated: with more than one fenestra (0); with just one
fenestra (1).
52. Relative position of cranio-mandibular joint: beneath basioccipital
ventral edge (0); aligned with basioccipital (1).
53. Quadrate condyles: almost aligned (0); medial condyle expands
ventrally (1).
54. Dorsocaudal edge of quadrate: straight or smoothly curved (0);
forming otic notch (1).
55. Dorsal surface of caudal branch of quadrate: concave or flat and
 ORTEGA ET AL.—NEW ARARIPESUCHUS FROM PATAGONIA
smooth (0); with a longitudinal ridge from base of paraoccipital
process to articular end (1).
56. Pterygoidean secondary palate: pterygoids of primary palate ex-
posed and they do not contact each other secondarily on midline
(0); pterygoids meet on midline forming a secondary palate (1).
57. Basipterygoid processes: expanded (0); reduced to a narrow arista
or absent (1).
58. Pterygoids: caudally separated (0); caudally fused (1).
59. Vomer: exposed on palate between premaxillae and maxillae (0);
hidden by palatal branch of maxillae (1).
60. Palatal secondary palate: palatines of primary palate exposed and
they do not contact each other secondarily on midline (0); palatines
meet on midline forming a secondary palate (1).
61. Ectopterygoid-maxilla contact: ectopterygoid does not connect to
palatal branch of maxilla (0); ectopterygoid makes contact with
maxillary palatal branch (1).
62. Supraoccipital exposure on cranial roof: no, parietals contact on
occiput avoiding dorsal exposition of supraoccipital (0); supraoc-
cipital connects parietal at posterior edge of skull roof or is clearly
exposed in dorsal surface of cranial roof (1).
63. Bones bounding foramen magnum: exoccipitals and supraoccipital
(0); exoccipitals (1).
64. Occipital foraminae for cranial nerves IX, X, and XI: all passing
through a common foramen (0); foramen metoptico (IX) in a sep-
arate passage (1).
65. Basioccipital surface under foramen magnum: caudoventrally ori-
ented (0); vertical and occipitally oriented (1).
66. Cranial projection of iliac blade: present and as long as the caudal
projection (0); reduced to a tuberosity (1); absent (2)
67. Basisphenoid in ventral view: widely exposed (0); almost excluded
from ventral view and hidden by pterygoid and basioccipital (1).
68. Relative length of basisphenoid and basioccipital: basisphenoid
shorter or equal than basioccipital (0); basisphenoid longer and
transversely wider than basioccipital (1).
69. Antorbital fenestra: present (0); absent or reduced to a tiny foramina
(1).
70. Nasal participation in antorbital fenestra: yes (0); no (1).
71. Jugal participation in antorbital fenestra: yes (0); no (1).
72. Supratemporal fenestra: present (0); absent (1).
73. Infratemporal fenestra: wide (0); forming a vertical slot (1).
74. Infatemporal fenestra: shorter than it is deep or at least as long as
deep (0); much longer than deep (1).
75. Anterior opening of temporo-orbital: exposed in dorsal view (0);
hidden in dorsal view, and overlapped by squamosal rim of supra-
temporal fossa (1).
76. Cranio-quadrate canal: laterally open (0); closed off by a thin lam-
ina formed by squamosal, quadrate and exoccipital (1) closed off
by a thick lamina formed by squamosal, quadrate and exoccipital
(2).
77. Iliac blade: with posterior lamina as high as anterior one (0); with
posterior lamina higher than anterior one (1).
78. Maxilla forms part of secondary palate: no (0); yes (1).
79. Palatines or pterygoids participating on caudal opening of naso-
pharyngeal duct (internal nares): no (0); yes (1).
80. External mandibular fenestra: present as a wide foramen (0); pres-
ent but very reduced (1); absent (2).
81. Mandibular compression: dentary compressed, formed by almost
vertical lateral and medial laminae (0); dentary transversely ex-
panded, almost as wide as high, and with convex lateroventral sur-
face (1).
82. Suture between dentaries: separated mandibular symphysis (0);
dentaries fused, at least in ventral mandibular symphysis (1).
83. Dorsocaudal branch of dentary: dentary extends caudally up to end
of tooth row (0); dentary extends beyond tooth row and with a
dorsal ascending projection (1).
84. Lateral contour of dentary in dorsal view: straight (0); sigmoidal
(1).
85. Post-caniniform dentary teeth: almost homodont in size (0); with
waves of size variation (1).
86. Outline of dentary tooth row in dorsal view: straight (0); sigmoidal
(1).
87. Proximal articular head of tibia: with two concavities separated by
a crest (0); with one concave articulating area (1).
88. Splenial symphysis: splenial not involved in mandibular symphysis,
73
splenials just touch or do not contact each other (0); splenials form
significant part of symphysis (1).
89. Splenials behind symphysis: thin and lateromedially compressed
(0); broad and robust (1).
90. Foramen intramandibularis oralis: small or absent (0); big slot-like
foramen (1).
91. Emergence of the mandibular branch of trigeminus nerve in the
medial side of mandible: caudal to symphysis (0); enclosed within
symphysis (1).
92. Transversal section of splenial: plane (0); convex (1).
93. Caudal edge of angular: does not ascend to the articular glenoid
cavity (0); ascend surpassing the articular glenoid cavity (1).
94. Caudal edge of surangular: slopes ventrally (0); slopes dorsally (1).
95. Rear portion of dentary in dorsal view: does not expand medial to
tooth row (0); medially expanded to tooth row (1).
96. Anterior projection of surangular in lateral view: unique and acute
(0); forked (1);
97. Prearticular: present (0); absent (1).
98. Coronoid size: short (0); long, anteriorly extended (1).
99. Surangular and quadratojugal taking part in craniomandibular joint:
no (0); yes (1).
100. Mesial and distal margin of tooth crowns: with denticulate carinae
(denticles are isolated units, distinctly shaped and sized) (0); with-
out carinae or with carinae smooth or crenulated (crenulation is
made of enamel wrinkles) (1).
101. Root of maxillary and dentary teeth: as wide or narrower than its
crowns (0); inflated roots, wider than is crowns (1).
102. Shape of tooth crowns in bucal view: triangular (0); trapezoidal
(1).
103. Lingual side of maxillary and dentary tooth: without a longitu-
dinal depression (0); with a longitudinal depression affecting root
and crown (1).
104. Maxillary and dentary teeth transverse section: labiolingually
compressed (0); subcircular (1).
105. Glenoid fossa of articular: craniocaudally similar to articular sur-
face of quadrate (0); craniocaudally longer than articular surface of
quadrate (1).
106. Tip of maxillary and dentary tooth crowns: caudally curved (0);
dorsal directed or lingually curved (1).
107. Number of longitudinal rows of osteoderms forming presacral dor-
sal armor: two (0); four (1).
108. Presence of accessory ranges of osteoderms (sensu Frey, 1988),
that is, more than four longitudinal rows of osteoderms forming
presacral dorsal armor or four longitudinal rows in which the par-
asagitals must show two longitudinal keels: no (0); yes (1).
109. Continuity of dorsal armor: dorsal armor continues from neck to
tail (0); dorsal armor shows a narrowing or gap at cervico-thoracic
juncture (1).
110. Presacral dorsal osteoderms: lateroventrally deflected (0); flat (1).
111. Ornamentation of presacral dorsal osteoderms; smooth or made of
ridges and grooves (0); with a pitting surface (1).
112. Dorsal paravertebral osteoderms constituting a hemiplastron: hem-
iband is narrower than its craniocaudal length (0); hemiband is
wider than its craniocaudal length (1).
113. Imbrication of osteoderms: osteoderms of presacral dorsal armor
with an anterior peg that articulates into a socket of precendent one
(0); osteoderms without anterior projection (1).
114. Imbrication of osteoderms: osteoderms of presacral dorsal armor
with a short caudal salient peg (0); osdeoderms with straight caudal
edge (1).
115. Number of keels on transverse bands of presacral dorsal armor:
two (0); more than two (1).
116. Coracoid shaft: short (0); long shaft extending ventrally (1).
117. Cervical and dorsal centra: amphicoelous (0); procoelous (1).
118. Styliform process of coracoid: present. (0); absent (1).
119. Centrum of first caudal vertebra: amphicoelous (0); biconvex (1).
120. Contour of scapular blade: dorsally broad and with concave cra-
nial and caudal edges (0); dorsally broad and with a concave cranial
edge but straight or convex caudal one (1); dorsally narrow, straight
cranial and caudal edges (2).
121. Relative length of coracoid and scapula: scapula is at least one
third longer than coracoid (0); scapula as long as coracoid (1).
122. Glenoid surface of coracoid: extended on a subhorizontal plane
(0); extended on a vertical plane (1); extended on a oblique plane,
and the glenoid lip facing outwards and posteroventrally (2).
74 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 1, 2000

123. Humeral proximal head: facing backwards (posterodorsally) (0);
facing dorsally, and with a lateromedial major axis (1).
124. Inner tuberosity of proximal head of humerus: developed and with
articular facet bending ventral or obliquely (0); not developed and
with articular surface facing dorsally (1).
125. Ligamental depression on anterior surface of humerus: inmediate-
ly lateral to inner tuberosity and below humeral head (0); displaced
laterally toward the border of the shaft and located lateral to hu-
meral head (1).
126. Lateral profile of deltopectoral crest: convex (0); concave (1).
127. Radial: longer than wide (0); as long as wide (considering its
proximal width as reference) (1).
128. Relative length of radial and ulnar: radial as long as ulnar (0);
radial longer, at least 1/3 of ulnar length.
129. Proximal carpals short, almost spherical (0); radial and ulnar elon-
gated (1).
130. Lateral contour of rostrum in dorsal view: straight (0); with con-
strictions and with a sinusoidal aspect (1).
131. Mesio-distal length of tooth crowns: decreasing towards the rear
of tooth row (0); increasing towards the rear of toothrow (1).
132. Heterondonty of maxilla and dentary teeth: homodonty (0); with
different dental morphologies (heterodonty) (1).
133. Number of premaxillary teeth: five (0); less than five (1).
134. Exposition of posterior part of angular in ventral view: angular
surface (where pterygoidean muscle attached) ventrally exposed
(0); angular laterally displaced and overlapping articular, with sur-
face for pterygoidean muscle attachment facing laterally (1).
135. Caudal branch of quadrate: at least as long as broad (0); shorter
than broad (1).
136. Palatal maxillary platform: absent (0); present, avoiding contact
between ectopterygoid and maxillary alveoli (1).
137. Septated internal nares: yes (0); no (1).
138. Pterygoidean flanges: laminar (0); bar-like (1).
139. Depression on primary pterygoidean palate posterior to internal
nares: depression wider than palatine bar (0); narrower than palatine
bar between palatal fenestrae (1).
140. Postmaxillary internal nares: present (0); absent (1).
141. Retroarticular process: without a medial shelf (0); with a medial
shelf (1).
142. Internal nares: far from caudal contour of skull (0); close to caudal
contour of skull (1).
143. Caudal edge of internal nares: cranial to rear edge of palatine
fenestra (0); caudal to de rear edge of palatine fenestra (1).
144. Ventral edge of external mandibular fenestra: smoth (0); with a
depression surrounding the edge (1).
145. Lateral surface of anterior branch of jugal: smooth, plane or con-
cave (0); with a pronounced triangular depression.
146. Ectopterygoidean medial descendent branch: single (0); distally
forked (1).
147. Medial shelf of retroarticular process: vertical and facing medially
(0); facing dorsally (1).
148. Coracoidal shaft: blade-like (0); rod-like (1).
149. Femoral shaft: slightly twisted, proximal and distal articular facets
are twisted each other about 30 degrees (0); strongly twisted, prox-
imal and distal articular facets are twisted each other about 60 de-
grees (1).
150. Proximal and distal ends of radial: almost equally expanded (0);
proximal head wider than distal one (1).
151. Mandibular symphysis: short (0); long, dentary symphysis pro-
longs caudal to 4th alveoli (1).
152. Craniocaudal length of retroarticular process: less than craniocau-
dal length of articular glenoid fossa (0); much longer than cranio-
caudal length of articular glenoid fossa (1).
continued next page

APPENDIX 2. Distribution of character states. ‘‘0’’ denotes the primitive condition; ‘‘-’’denotes missing or not applicable condition.

Character

1 2 3 4 5 6 7 8
Taxon 12345678901234567890123456789012345678901234567890123456789012345678901234567890
Postosuchus 0000000000--00000000000000000-0000000000000000000000000000000-00--0-00000000-000
Sphenosuchus 00000000000110000000000000010-1000-00000000000000000000000000000-00-011000000100
Dibothrosuchus 0000000000100000000000000001101000000100000000000000000000000000--0-01100000-100
Protosuchus 01100110001001100100000001010-10000011011011100001100100100000100001011010000101
Orthosuchus 00100110001001100100000001010-1000-011011011100001100-00100000100001011000000100
Sichuanosuchus 01100110-0---10001000-000111--10000011-110111000011----01--00---0-010110--00-100
Shantungosuchus 011000100010010001000-0--------0-00-1001---1-0-00110010011-00--10-01011-10-0-100
Gobiosuchus 0010001000011110001000001101101000001-011011100001100-0010000-100-0-010100-1-102
Las Hoyas Croc. 00100010-0---100001000001101--1000--1-01101110000------01-00----0-0-011100---102
Fruita Fm. Croc. 01100010001001000010000011010-100000110110111000011-0-10--01011-----0110000--112
Notosuchus 0010001000100010011000001111101000--10111010110001101100110111110100011000020110
Baurusuchus 00000011001-01000100000111111010000011111110110001001100111111110-0011100102-110
Itaborai Croc. 000000111001110000011001111110100001110111111100010-011011111-110-1011101-02-110
Bretesuchus -0000010-10111000100000-------1--001--------------0----111111--10-10111-01---110
Sebecus 0000001-011-11010100000-1101--10000111011111110-01000110-11111110-1011100002-110
Iberosuchus 0000001100--0100000000011101101100101011111111010100110-111--1110-1001000102-1-0
Libycosuchus 00100-1--00110100110000-11-1--1000-01111111011000100110-11-1111-0-1011100002-110
Ar. gomesi 10100010-0010000000110001101111001--1011101011000100-100111111110-0001100002-110
Ar. patagonicus 1010001--0--0000000100001101--1000--1011101011000100010011-1111-0-0001100002-110
Lomasuchus 1000101100--1-00000100001101--1000--1111101111-0010----011111-110-1001100-02-110
Itasuchus 10001-1-0011--01-00111011--------0001111----110-----------11--------011-01---110
Theriosuchus 101010100011110100011001110111100101111110111101--000110111111110100111000010112
Pelagosaurus 1001101000--1-010000011100010110110011100001100100-00000110111100-0011100101-110
Steneosaurus 0001101000101101000001110001--10110-11100001100100-000001-11-11-0100111001010110
Goniopholis 1000101000--11010001110111011210010111111111111101010110111111110110111000111112
Bernissartia 100010101011110110011-01110111100111111111111111010101101111111-0-1011100011-112
Las Hoyas Neos. 10001010-011110110011-0111-11110011111-1111111-1-10-010111111-1-1-1011100011-112
Hylaeochampsa 00---------------00----11101111001111--1111111-1-10101-1111111111-10---0--11-11-
Alligator 10001010101110011001100111111210011111011111110101010101111111111210111000121110
Crocodylus 10001010101111011001100111111210010111011111111111010101111111111210111000121110
 ORTEGA ET AL.—NEW ARARIPESUCHUS FROM PATAGONIA 75

153. Dorsal surface of retroarticular process: facing craniocaudally (0);
facing dorsal or craniodorsally (1).
154. Dorsal surface of caudal branch of quadrate: with a triangular
depression (0); without depression (1).
155. Dorsal contour of rostrum in lateral view: straight or convex (0);
concave (1).
156. Teeth at anterior part of maxilla: no prominent tooth (0): second
or third alveoli enlarged (1); fourth or fifth alveoli enlarged (2).
157. Skull roof: rectangular and with a major transverse axis (0); square
or rectangular and with a longitudinal dominant axis (1).
158. Caudolateral lobe of squamosal: not differentiated (0); squamosal
showing a smooth lobe differentiated from the skull by a caudo-
lateral groove (1).
159. Anterior opening of cranio-quadrate passage in otic area (when
cranio-quadrate canal is closed off): not expanded (0); opening ex-
panded forming a caudal notch (1).
160. Ventral border of exoccipital: straight, and leaving posterior open-
ing of cranio-quadrate passage visible in occipital view (0) convex
and ventrally overhanging and obscuring posterior opening of cran-
io-quadrate passage from occipital view (1).
161. Length from proximal articular facet of femur to distal end of
fourth trocanter: more than one third than total femoral length (0);
one third or less than total femoral length (1).
162. Largest mandibular teeth: unique (0); double, generally third and
fourth (1).
163. Premaxillo-maxillar suture in palatal view: acute (0); straight and
orthogonal with the sagittal plane (1).
164. Number of maxillary teeth: ten or more (0); less than ten (1).
165. Naso-lachrymal suture: nasal extensively contact lachrymal (0),
nasal and lachrymal do not contact or with a short contact (1).
166. Articulation of medial process of articular and otoccipital: absent
(0); present (1).
167. Internal edge of proximal ulna head: concave (0); straight (1).
168. Section of ulna shaft: circular (0); compressed at least its distal
end (1).
169. Prefronto-frontal joint: prefrontal overlaps frontal (0); frontal over-
laps prefrontal (1); frontal and prefrontal with a interdigitate suture
(2).
170. Distal lateral condyle of humerus in posterior view: projecting a
sharp ridge towards shaft and delimiting a flat lateral plane (0);
smooth (1).
171. Lachrymal descending lateral process: columnar (0); laminar (1).
172. Lachrymal orbital contour: facing laterally (0); facing laterodor-
sally.
173. Cranial jugal branch: as deep or slightly deeper than caudal branch
(0); much deeper than caudal one (1).
174. Lateral surface of jugal in ventral view: exposed lateral to maxilla,
jugal outwardly bowed (0); not visible in ventral view, jugal straight
(1).
175. Palatal surface: concave (0); plane (1).
176. Occipital condyle: caudally directed (0); ventrocaudally directed
(1).
177. Primary pterygoidean palate: without parasagittal depressions (0);
with deep parasagittal depressions (1).
178. Deltopectoral crest of humerus: partially visible in caudal view
(0); hidden in caudal view (1).
179. Ligamental pit below humeral head on posterior surface: delim-
iting a ridge on its medial border (0); pit displaced laterally between
humeral head and lateral tuberosity (1).
180. Humeral shaft: straight (0); sigmoidal, with a pronounced posterior
curvature of shaft on proximal area of humerus (1).
181. Proximal and distal head of humerus: each turned more than 30
degrees (0); each turned less than 30 degrees (1).
182. Inner distal condyle of humerus: with a vertical extension of troch-
lea (0); transervely expanded (1).

APPENDIX 2. (Continued).

Character
1 1 1 1 1 1 1 1 1
9 0 1 2 3 4 5 6 7 8
123456789012345678901234567890123456789012345678901234567890123456789012345678901234567890123456789012
000000-0000001-100-00000000--------0--000000---00000--000000-0000------00000--00-00000--0-000---------
00000000000000000100000000000000000100000000000010000-00000000000--000000000--00-00010001000-0000000-0
000000-0000000-00000000000000000000100000000000010000-000000000000-000000000--00-00010--1-00----------
0010000000-000-00-000000-00000-100010001010000001000-0100000-00000-000-000000000-000-10-2-00000000-0-0
00000000000001--0-010000-00000-100010-011100-0001000--100000-00000-0-0-00-000000-00110-02-00---------0
00000--0---00------1---0-00000-1000--------------0011--00000-0000----0-00-0-0000-0001---2-0000--0-----
000000-0010000-0--011000-0----------0---11---------1-01000000000000----0-----000-00001--2-00-0--0-----
011000-100-000---0011010-11-0001-1--0-0-1-------10001-100000-00-00-----0-000000--00010--2-00-0-00-----
01-000-10--0---0---11010-10000010101010-110000001000---00-00-00-0---00-0--0000-0-0-000-020000----000-0
000000-000-000-0-0010100010---01010-1--11----0---000---00-00--0000----0000000000-10000--2-10000-0-----
001010010011000-10010100110---01-10101-111000010100010100000100001010-1001000000000100112-000001100000
001011-1110100001000000000----------0---------1-1000101010011010110--11101011000-00110--2-1011011-----
00100101010000001010000000-----1----0-----0------000110000111001-11---1111-10000-000-0--20--11-11----0
001001-11--1-0001--0000000-----------------------00010-0-011-10011----1---01-----000---------10-1-----
000000-1-00---001010000000-----------------------0001-00000110010-----111101-0---00000----10110-------
001000-11101-00010000000000--100110101-1110000---0000-0-01-1-1-01--10-1--1011010-00-00--20101101100000
001000-1-0--010-1-0-----0------------------------0--0010-0-110-0000---11010000-0-000-0--2-101001------
1010-0-1000000001001000111000011010101011100-0--111110100011100000101-1101010000-00000-1200000110000--
101--0-100-000-0--0100-111000011010101-1--0000--101110100011100000101-1101010000-0-000--2-00001100-0--
100110-----0-001---0-000-1------------------------01---0-011-00-0---------01-000--001---2-1011-0------
101111-100---1-1---1000101---011110--------------101-------1--0---1---1-1-1------0000---2-111-1-------
000110-10000010110010001-100-01101-10111-1-00-001111--000011-00000--101-011201-0100000-12-110-1-00-0--
100000-1000001010101000101000111010-0-0---------10001000101100000010--1101100010-00000--2-1001100-----
100000110010010--1010001010001110101010212111000100010001-1100000010101111100010100000012-100110001002
100110-110-101011001000101000111010101-1121111001100-000001111000010111101120111110000--21111110011111
100110-00010011-100100010101011111111111121111--11110100-011100000101-1111120011-11010--2-1111100-----
100110-000000---1001000101010111111----------1--1-----0010111000001-1-11-11200-1---0-0--2-11-110------
------------------------------------------------------001011-10-00-------11-0011----1---2-11-1000-----
10011110000001111001-001011111111111111212111111110001010011-10000101111111200101010101121111010011111
100110100000110110010001011111111111111212111111110001001011111000101111111200101000001121111110111111
76 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 20, NO. 1, 2000
APPENDIX 3
List of Taxa Used in this Analysis
Out-group is formed at: Postosuchus, Upper Triassic of North Amer-
ica, (Chatterjee, 1985); Sphenosuchus, Upper Triassic or Lower Jurassic
of South Africa, (Walker, 1990); and Dibothrosuchus, Lower Jurassic
of China (Wu and Chatterjee, 1993). In-group is composed of: Orthos-
uchus, Lower Jurassic of South Africa (Nash, 1975); Protosuchus*,
Lower Jurassic of North America (Colbert and Mook, 1951); Sichuano-
suchus, Upper Jurassic of China (Peng, 1995); Shantungosuchus, Upper
Jurassic/Lower Cretaceous of China (Wu et al., 1994); Fruita Fm. croc-
odile* (sensu Clark, 1985): Upper Jurassic of North America (Benton
and Clark, 1988), Gobiosuchus*, Upper Cretaceous of Mongolia (Os-
mólska, 1972; Osmólska et al., 1997); Las Hoyas crocodyliform*, Low-
er Cretaceous of Spain (Buscalioni et al., 1997); Notosuchus*, Upper
Cretaceous of Argentina (Woodward, 1896; Gasparini, 1971); Bauru-
suchus*, Upper Cretaceous of Brazil (Price, 1945); Iberosuchus*, Eo-
cene of southwestern Europe (Antunes, 1975; Ortega et al; 1996); Se-
becus*, Eocene of Argentina (Colbert, 1946; Gasparini, 1972); Libyco-
suchus*, Upper Cretaceous of Egypt (Stromer, 1914); Itaborai crocodile*
(sensu Buffetaut, 1982), Paleocene of Brazil; Bretesuchus*, Paleocene
of Argentina (Gasparini et al., 1993); Araripesuchus gomesii*, Lower
Cretaceous of Brazil (Price, 1959); Araripesuchus patagonicus*, Lower
Cretaceous of Argentina; Lomasuchus*, Upper Cretaceous of Argentina
(Gasparini et al., 1991); Pelagosaurus*, Lower Jurassic of Europe (Buf-
fetaut, 1980; Clark, 1986); Steneosaurus*, Jurassic (Buffetaut, 1982);
Itasuchus*, Upper Cretaceous of Brazil (Price, 1955); Theriosuchus*,
Upper Jurassic/Lower Cretaceous of Europe (Clark, 1986); Goniophol-
is*, Lower Cretaceous of Europe (Buffetaut, 1982; Clark, 1986); Ber-
nissartia*, Lower Cretaceous of Europe (Buscalioni and Sanz, 1990,
Norell and Clark, 1990); Las Hoyas Neosuchia*, Lower Cretaceous of
Spain (Ortega and Buscalioni, 1995); Hylaeochampsa*, Lower Creta-
ceous of England (Clark and Norell, 1992); Alligator mississippiensis*,
Recent; and Crocodylus niloticus*, Recent. Asterisks denote species di-
rectly reviewed by authors.
APPENDIX 4
Cladogram. Consensus tree of two most parsimonious solutions re-
sulting from a strict parsimony analysis of data matrix including 3 out-
groups and 27 crocodyliform taxa (see Appendix 3). Data matrix has
been processed using PAUP 3.1.1 (Swofford, 1993). They have a length
of 388 steps. Consistence index is low (0.469), but Retention index
(0.745) is rather high. The two most parsimonious trees just differ in
the placement of Thalattosuchia within Neosuchia. Either as the sister
group of Itasuchus plus the remainder neosuchians or as the sister group
of Goniopholis plus Bernissartia plus Las Hoyas Neosuchia plus Eu-
suchia. Unambiguous changes along branches on the strict consensus
and node names are below. (Abbreviations: ‘‘,’’ primitive condition,
‘‘.’’ derived condition).
Node 1. (Crocodyliformes): 3.; 7.; 14.; 26.; 37.; 40.; 41.; 43.;
44.; 45.; 50.; 51.; 54.; 57.; 63.; 100.; 112.; 120.; 121.;
122.; 133.; 135.; 169...
Node 2. 19.1; 25.; 62.; 106.; 114.; 165..
Node 3. 29.;83.; 88..
Node 4.(Mesoeucrocodylia): ,14; 39.; 46.; 58.; 61.; 64.; 154.;
168..
Node 5. 1.; ,19; 20.; 30.; 81.; 111.; 132.; 139.; 147.; 149.;
175..
Node 6. (Neosuchia) 5.; 13.; 96.; 171.; 174..
Node 7. 9.; 17.; 35.; ,88; 108.; 115.; 117.; 134.; 165..
Node 8. ,55; 56.; 65..
Node 9. (Eusuchia): 142.
Node 10. (Crocodylia): 27.; 76..; ,160.
Node 11. (Protosuchia): 6.; 18..
Node 12. 2.; 73.; 166..
Node 13. 132..
Node 14. (Gobiosuchidae): 72.; 82.; 101.; 103..
Node 15.(Ziphosuchia, new taxon): 18.; 53.; 92.; 127.; 148.;
177..
Node 16. 42.; 69.; 171.; 173..
Node 17. (Sebecosuchia): ,3; 14.; ,19; 74.; 90.; 145.; 156.;
174..
Node 18. 44.; ,135.
Node 19. (Sebecidae): 13.; 36.; ,39; ,53; 55.; 99..
Node 20. ,11; 86.; 139..
Node 21. (Araripesuchus): ,38; 105.; 131..
Autapomorphies:
Postosuchus: ,71.; ,94-.; ,96.; ,165.; ,169..
Sphenosuchus: 13.; 98..
Dibothrosuchus: 29..
Orthosuchus: 94.; 164..
Protosuchus: 80.; 83.; ,100; ,121.
Sichuanosuchus: no unambiguous changes
Shantungosuchus: ,6; ,38; ,165.
Fruita Fm. crocodile: 2.; 55.; 102.; 117.; 162.; 171..
Gobiosuchus: 15.; ,71; 107.; 165..
Las Hoyas crocodyliform: no unambiguous changes
Notosuchus: ,38; 91.; 102.; 105.; 164..
Libycosuchus: ,11; 13.; 94.; ,133.
Baurusuchus: 12...; 86.; 137.; 143.; 164.; 165..
Iberosuchus: ,18; 35.; ,38; 48.; ,69; ,71; ,112; ,133; 142.;
159..
Sebecus: 16.; ,74; ,83; ,90; ,145.
Itaborai crocodile: ,18; 20.; 21.; 73.; ,89; ,92; 134..
Bretesuchus: ,8; 56.; 142..
Araripesuchus gomesii: 21.; 34.; 130..
Araripesuchus patagonicus: no unambiguous changes
Lomasuchus: 8.; ,100, 165..
Pelagosaurus: ,59.
Steneosaurus: ,1; 91..
Goniopholis: ,,14; 30.; 89.; 92.; 142..
Bernissartia: 91.; 131.; 132..
Las Hoyas neosuchian: no unambiguous changes.
Hylaeochampsa: ,1; ,175.
Alligator: ,47; 86.; ,137; ,174.
Crocodylus: ,,14; ,35; 143.; ,165.