Bartels A Transitional Paleocene-Eocene Reptile Fauna From The Bighorn Basin, Wyoming

Herpetologists' League
A Transitional Paleocene-Eocene Reptile Fauna from the Bighorn Basin, Wyoming

Author(s): William S. Bartels
Reviewed work(s):
Source: Herpetologica, Vol. 39, No. 4 (Dec., 1983), pp. 359-374
Published by: Herpetologists' League
Stable URL: http://www.jstor.org/stable/3892531 .
Accessed: 09/02/2013 20:35
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp
.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.
Herpetologists' League is collaborating with JSTOR to digitize, preserve and extend access to Herpetologica.
http://www.jstor.org
This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

All use subject to JSTOR Terms and Conditions
HERPETOLOGICA
December 19831
Arima, AMNH 87398-401, 87402 (skeleton); Mt.

Aripo, BMNH 1940.3.10.96-97; Mt. Tucuche,
AMNH 55760-63, BMNH 1934.12.2.14-18,
1935.1.5.6-11, 1965.635-36, 1947.2.22.4 1, MCZ
17894-97, 19895-98; Simla, Arima Valley, AMNH
62888, 79921-24, KU 192399-400; Spring Hill Estate, ArimaValley, USNM 166625 (1 + 2 tadpoles);
St. Pat's Estate, Arima Valley, BMNH 1956.1.4.63;
TamanaCaves, N slope Mt. Tamana,AMNH 87396,
87397 (tadpoles). VENEZUELA: Sucre: Mauraco, N
of Pilar, KU 181982-84.
Flectonotus pygmaeus. VENEZUELA: Aragua: Estaci6n Biol6gica Rancho Grande, AMNH 70434-47
(14 + 1 lot young, 3 lots tadpoles), KU 133346-87,
133463-64, 139518-19 (tadpoles), 166738-54,
166756-59, 167758-59 (skeletons), 167819-23 (tadpoles), 184957-59, 185719-31, 185777-78 (tadpoles), USNM 196335. Barinas: La Soledad,
UPR-M 5882-89, 5891-904. Carabobo: Puerto Cabello, SMF 2679. Merida: La Azulita, UPR-M429394. Tdchira: Mata Mula, UPR-M 6081, 6087-88,
6090.
Fritziana fissilis. BRASIL: Espirito Santo: Santa
Teresa, 700 m, KU 187470-74. Guanabara: Sumare
Herpetologica,
359
(Rio de Janeiro),AMNH 70260; Tijuca, KU 9223742. Rio dejaneiro: Angrados Reis, MNRJ1998 (tadpoles); Parati, MNRJ3119 (3); Teres6polis, AL (3),
BMNH 1949.1.1.61-62, El 1615. Sdo Paulo: Bocaina, WCAB 29253-442 (25 examined); Fazenda
da Posse, Sao Jose do Barreiro,WCAB 31362-75,
WCAB (tadpoles); Paranapiacaba,KU 71826; Saio
Jose do Barreiro,Serrada Bocaina, USNM 16402630; Serrada Bocaina, AL (25).
Fritziana goeldii. BRASIL: Guanabara: Realengo,
MNRJ2068; Represa dos Ciganos, MNRJ 1875 (3);
Rio de Janeiro,AL (5); Tijuca,AL (3), AMNH 7025758, 72324-25, El 1607, KU 84720, 84721 (skeleton),
92231-32 (skeletons), 92233-36; MNRJ 1846, 1862;
WCAB 18716 (tadpoles), 42496. Rio de Janeiro: CoIonia Alpina (Teres6polis), BMNH 1947.2.12.69-70;
Teres6polis, MNRJ267 (2), 269. Sdo Paulo: Campos
do Jordato,WCAB34669-98.
Fritziana ohausi. BRASIL: Rio de Janeiro: Teres6polis, AL (7 + 1 lot tadpoles), KU 92225, 92226
(skeleton), WCAB 1077, 7879, 7888, 7891, 7894,
7896, 8872, 9383-84, 19523-25, 36355-56. Sdo Paulo: Paranapiacaba, AMNH 69946-47, BMNH
1964.158, KU 71782, 92227-30.
39(4), 1983, 359-374
? 1983 by The Herpetologists' League, Inc.
A TRANSITIONAL PALEOCENE-EOCENE REPTILE FAUNA

FROM THE BIGHORN BASIN, WYOMING
WILLIAM
S. BARTELS
ABSTRACT: The Clarkforkian Land-Mammal Age is the transitional Paleocene-Eocene interval in the history of North American vertebrates. The Clarkforkian reptile fauna may be distinguished from earlier Paleocene faunas by the occurrence of the homed alligator Ceratosuchus
and the anguid lizard Melanosaurus. A more dramatic faunal change occurs at the ClarkforkianWasatchian boundary when champsosaurs and Ceratosuchus disappear, emydid turtles and glyptosaurine lizards undergo significant radiations, and several families of turtles presumably migrate into the region. The Bighorn Basin Clarkforkian reptile assemblage is dominated by large
aquatic taxa such as champsosaurs, alligators (Allognathosuchus in particular), the crocodylid
Leidyosuchus, and trionychid and emydid turtles. The terrestrial component of this fauna is far
less abundant and diverse, containing no snakes or distinctly terrestrial turtles and only two
abundant lizards, Melanosaurus and the small anguid Machaerosaurus.
Key words: Reptilia; Testudines;
Eocene; Wyoming
Choristodera; Lacertilia; Crocodilia; Fauna; Paleocene-
THE Clarkforkian Land-Mammal Age

has recently been redefined, based on a
thorough review of its mammalian taxa
(Rose, 1981). Clarkforkian deposits are
central to understanding the transition
from archaic Paleocene mammalian faunas to Eocene assemblages of modern as-
pect. The Clarkforkianis preceded in time

by the Tiffanian (late Paleocene) LandMammal Age, and followed by the Wasatchian (early Eocene) Land-Mammal
Age. The Clarkforkianrepresents an interval of two million years, from about 55
Ma to about 53 Ma (Gingerich, 1980).

360
HERPETOLOGICA
Prior to a preliminary version of this

study (Bartels, 1980), only a single Clarkforkianreptile (Simoedosaurus) had been
described from the Bighorn Basin (Sigogneau-Russell and Baird, 1978). The
only other published record of a Clarkforkian reptile was that of Ceratosuchus
(Schmidt, 1938) from the Plateau Valley
Beds of Colorado. Fortunately, late Paleocene Tiffanian (Estes, 1975; Gaffney,
1972; Gilmore, 1928, 1938, 1942) and early Eocene Wasatchian (Gilmore, 1942;
Hay, 1908; Hutchison, 1980; Mook, 1921,
1924; Sullivan, 1979) herpetofaunas are
more completely known and help to provide a frameworkfor elucidating the transitional Clarkforkianreptile fauna.
In the Bighorn Basin, Clarkforkianfossils are recovered from two formations of
Laramide-age basin fill, the Fort Union
Formation and the overlying Willwood
Formation. These elastic units are exposed in the northern Bighorn Basin
(Clark's Fork Basin-Polecat Bench area),
a structural and topographic depression
in northwestern Wyoming. The Fort
Union (Polecat Bench) Formation is
bounded below by Lance-equivalent Upper Cretaceous beds and above by the
Willwood Formation. The Fort Union
Formation reaches a maximum thickness
of nearly 3000 m in a foredeep along the
Beartooth Mountains to the west, but it
is much thinner throughout most of the
basin. It is comprised of drab fluvial gravels, sands, silts, muds, limestones and lignites, with sands and silts predominating.
The upper 100 m of the unit contain the
lower Clarkforkiandeposits of the Plesiadapis gingerichi Zone (Rose, 1981). This
interval is somewhat unfossiliferous and
is the most poorly understood portion of
the Clarkforkian.The remainder of the
Clarkforkianis contained in the lower 800
m of the Willwood Formation. Distinguished by the first appearance of red
banded beds representing well-drained
soils (see Bown, 1979), the Willwood Formation is comprised of variegated fluvial
sands, silts, muds, and limestones. Overbank silts and muds predominate and
lignites are locally absent. The Willwood
[Vol. 39, No. 4
contains the middle ClarkforkianPlesiadapis cookei Zone and the late Clarkforkian Phenacodus-Ectocion Zone (Gingerich, 1976).
SYSTEMATICPALEONTOLOGY
At least 21 species representing 10

families and four orders of reptiles are
now recognized from Clarkforkian Bighorn Basin deposits. Many of these taxa
need systematic revision beyond the
scope of this investigation. Some of the
following identifications should, therefore, be considered tentative.
Following Gingerich (1976) and Rose
(1981), stratigraphic occurrences of each
taxon are segregated into the Plesiadapis
gingerichi (early), Plesiadapis cookei
(middle), and Phenacodus-Ectocion (late)
ClarkforkianZones. The specimens listed below are in the University of Michigan Museum of Paleontology, except
where noted. Fossil localities listed for
the referred material are University of
Michigan Sand Coulee (SC) and Foster
Gulch (FG) sites in the northern Bighorn
Basin.
Class Reptilia
Order Testudines Linnaeus, 1758
Suborder Casichelydia Gaffney, 1975
InfraorderCryptodira(Cope, 1868)
Superfamily Baenoidea Williams, 1950
Family Baenidae Cope, 1862
Baenid (indeterminategenus and species)
Referred specimens.-UM
74606,
77682, carapace fragments that cannot be
identified below family.
Occurrence.-Phenacodus-Ectocion
Zone (SC-202, SC-234).
Discussion.-Baenids are among the
rarest Clarkforkianturtles. In addition to
these two specimens, there are a few carapace fragments from various horizons
that might be referable to this family.
Baenids are well-known from adjacent
older (Torrejonian and Tiffanian) and
younger (Wasatchian)horizons (Gaffney,
1972; Archibald and Hutchison, 1979), so
their scarcity during the Clarkforkianis
problematical.

HERPETOLOGICA
December 1983]
361
Superfamily Trionychoidea Gray, 1870

Family Trionychidae Bell, 1828
Plastomenus Cope, 1873
Zone (SC-156, SC-171). Phenacodus-Ectocion Zone (SC-71, SC-159).
Plastomenus sp.
are abundant throughout the Bighorn Basin Clarkforkian.Included here are those
specimens that would normally be assigned to Trionyx because of their greatly
reduced eighth costal and plastral elements. This genus is so poorly understood that such an assignment would only
further confuse the taxonomic status of
this form.
65240,
69680, 74596, 74610, portions of carapace
and plastron, none of which are complete.
Occurrence.-Plesiadapis cookei Zone
Discussion.-Scraps
of trionychid shell
(SC-62, SC-136). Phenacodus-Ectocion

Zone (SC-29, SC-163).
Description.-Each
contains one or
more of the following diagnostic characSuperfamily Testudinoidea Baur, 1893
teristics: hyo-, hypo-, and xiphiplastra
Family Emydidae Gray, 1825
massive and solidly united along midEmydid A
line; hyoplastron notched anteromedial(undescribed new genus and species)
ly; eighth costal not significantly reduced.
65492,
Discussion.-Plastomenus is the most
65493,65534,67562,67564,67565,71484,
completely understood trionychid genus;
74593, 74598, 74601, 74605, 74606, fragthe solid plastron is a clear, easily recogmentary
to nearly complete shells.
nized feature and is unique among triOccurrence.-Plesiadapis
gingerichi
onychiids. Specific relationships within
Zone
(SC-156,
SC-171),
Plesiadapis
cookthe genus are poorly known, however.
ei Zone (SC-120, SC-134, SC-135, SC-136,
Hay (1908:467) recognized species acSC-188), Phenacodus-Ectocion Zone (SCcording to the density and style of cara48, SC-50, SC-53, SC-234).
pace ornamentation. The systematic valDiscussion.-Included
here are speciue of these characters is questionable, and
mens
belonging
to
an
as
yet
unnamed gemost specimens do not fall clearly into
nus
and
species
of
macrocephalic
emyany of his categories. Specific assignment
did.
This
new
genus
is
characterized
by
should be delayed until a more satisfaca prominent anterior emargination in the
tory classification can be developed for
carapace and deep gutter-like epiplastra
the genus as well as for fossil trionychids
(both for the accommodation of the large
in general (see Gaffney, 1979).
This genus is the most common
Details of the carapace (lack of orna- skull).
Clarkforkian
turtle and is very similar to
mental ridges, expended seventh costal,
Wasatchian
forms
(Hutchison, personal
reduced eighth costal) and plastron (massive hypoplastron, strongly curved hyo- communication, 1981). It differs in detail
plastron) indicate that the Clarkforkian from a Tiffanian taxon referred to the European macrocephalic emydid PtychoPlastomenus is distinct from described
gaster
by Estes (1975:376). This TiffaniWasatchian forms (Hay, 1908). There is
an form is congeneric
with the
enough variation within Clarkforkian
Clarkforkian
and
genus
should
be considspecimens to suggest that two species of
ered
a
new,
evolved
convergently
taxon
Plastomenus may have lived in the Big(Hutchison,
personal
communication,
horn Basin at that time.
1980).
Trionychid
(indeterminate genus and species)
74597,
74600, 74608, 74609, shell fragments.
Occurrence.-Plesiadapis
gingerichi
The Clarkforkian macrocephalic genus

has previously been referred to as the
"Echmatemys-like emydid" by Bartels
(1980:77) and the "undescribed largeheaded emydid" by Hutchison (1980:
117). A full description of this unusual

HERPETOLOGICA
362
[Vol. 39, No. 4
as "unrelated to extant or other known

A
box turtles ....
This genus is the earliest
known example of a fully developed box

turtle morphology. There is a single
transverse plastral hinge separating fully
kinetic anterior and posterior lobes. The
genus is unique among emydid turtles in
having well developed elongate costiform processes on the nuchal bone."
This species is common throughoutthe
Bighorn Basin Clarkforkian.
B
~~c?-~
,>
C~~~~~
FIG. 1.-Champsosaurus
gigas UM 74562, anterior portion of right dentary. (A) Superior view. (B)
Internal view showing the relatively smooth symphyseal surface. (C) Lateral view. Scale = 2 cm.
turtle is currently being completed by J.

H. Hutchison and R. C. Wood.
Emydid B
(unnamed new genus and species)
65547,
74595, 74602, 77666-77678, carapace and
plastron fragments.
gingerichi
Zone (SC-83, SC-250), Plesiadapis cookei
Zone (SC-62, SC-84, SC-127, SC-136, SC197), Phenacodus-Ectocion Zone (SC-50,
SC-72, SC-158, SC-159, SC-162).
Discussion.-Included here are specimens referable to an as yet unnamed new
genus and species of emydid box turtle.
Hutchison (1980:117) described this form
Family Chelydridae Gray, 1870

Protochelydra cf. P. zangerli
Erickson, 1973
Referred specimen.-UM 71483.
Occurrence.-Phenacodus-Ectocion
Zone (SC-234).
fairly complete but
Discussion.-A
poorly preserved shell referable to the
Chelydridae, and probably P. zangerli. A
single incomplete shell of this form is also
known from the latest Tiffanian of the
Bighom Basin (UM 77658, SC-178).
Family Chelydridae? Gray, 1870
Clemmys cf. C. backmani Russell, 1934
69689,
74592.
(SC- 127), Phenacodus-Ectocion Zone
(SC-202).
Discussion.-Two partial shells representing a second, much largertaxon probably referable to the Chelydridae. This
form is closely related to or conspecific
with the problematical "Clemmys" backmani, which may be a chelydrid (Hutchison, personal communication, 1982).
Order Eosuchia
Suborder Choristodera Cope, 1884
Family Champsosauridae Cope, 1876
Champsosaurus Cope, 1876
Champsosaurus gigas Erickson, 1972
Referred specimens.-Jaw fragments
(UM 74562, Fig. 1), and a fragmentary
skeleton (UM 71808) referable to C. gigas.
Occurrence.-Plesiadapis gin gerichi
Zone (SC-250), Plesiadapis cookei Zone
(SC-176).

December 19831
HERPETOLOGICA
Discussion.-Most elements of typical

Champsosaurus construction. Snout long
and very slender, mandibular symphysis
very long and shallow. Clavicle and
quadrateexceptionally massive. The most
complete specimen (UM 71808) possesses the diagnostic distally flattened ribs
and long femora of C. gigas (Erickson,
1972).
Simoedosaurus Gervais, 1877
Simoedosaurus sp.
Referred specimens.-Princeton University Nos. 22199, 22334, posterior portions of mandibles and some fragmentary
postcrania.
gingerichi
Zone (Burns Mine, Bear Creek, Montana).
Discussion.-Included
here are two
previously described (Sigogneau-Russell
and Baird, 1978) specimens of a shortsnouted champsosaur characterized by
palatal elements covered with broad
patches of teeth and anterior marginal
teeth developed into large canines. The
mandible is robust and the splenial is excluded from the short, rugose symphysis.
First recognized and extensively studied
in Europe (Sigogneau-Russell and Russell, 1978), Simoedosaurus has only recently been reported from western North
America. Sigogneau-Russell and Baird
(1978) assigned two additional specimens to Clarkforkianhorizons, but these
are probably latest Tiffanian in age (Rose,
personal communication, 1979). Comparative work is still in progress, but the unnamed American species appears to be
very similar to the European material (D.
Baird, personal communication, 1979).
Numerous champsosaur remains have
been recovered from each of the three
Clarkforkianzones. Most of these are too
incomplete to assign to either of the above
taxa. Champsosaurpostcranial material is
very abundant in Tiffanian and early
Clarkforkianhorizons in the Fort Union
Formation but is quite rare in Willwood
Formation deposits. This apparent drop
in abundance is probably related to the
fact that champsosaurs, being highly
363
aquatic and fairly large (body length to 4

m or more), were piscivorous dwellers of
large streams and rivers, reminiscent of
the modem gharial (Gavialis gangeticus)
of India and Burma. The shift from Fort
Union to Willwood sedimentation is
characterized by a reduction in the number and size of streams, thereby discouraging habitation by and/or preservation
of the champsosaurs.
Order Squamata
Suborder Lacertilia
Family Anguidae Cope, 1864
Subfamily Glyptosaurinae
McDowell & Bogert, 1954
cf. Odaxosaurus piger (Gilmore, 1928)
Referred specimen.-UM 77641, posterior half of right frontal.
(FG-6).
Discussion.-This
specimen represents a small, primitive glyptosaurinevery
similar to Odaxosaurus (see Gauthier,
1982; Meszoely, 1970). The width of the
armored portion of the frontal along the
posterior margin is 3.7 mm, and the pattern of its osteodermal ornamentation is
generally vermiculate.
Gauthier (1982) erected Proxestops
jepseni to include Peltosaurusjepseni and
some specimens referred to Odaxosaurus
(Pancelosaurus) piger by Meszoely
(1970). Proxestops is intermediate between Odaxosaurus (small, vermiculate
armor)and Xestops (relatively large, pustulate armor).Although this Clarkforkian
specimen is most similar to Odaxosaurus, it is slightly larger than described
Odaxosaurus material, and shows incipient development of pustules along the
vermiculate ridges of the osteodermal
roofing. This assignment to Odaxosaurus
should, therefore, be considered tentative.
Xestops Cope, 1873
Xestops vagans (Marsh, 1872)
Referred specimens.-Partial
right
mandible (UM 67188) and left frontal(UM
74616).

HERPETOLOGICA
364
[Vol. 39, No. 4
FIG. 2.-(A) Xestops vagans UM 67188, posterior portion of right dentary. (B) Machaerosaurus sp. UM
73570, partial right dentary. Scale = 5 mm.
Zone
(SC-136), Phenacodus-Ectocion
(SC-81).
characteristics
Discussion.-Xestops
include: frontals strongly sutured but not
fused; frontoparietal scutes separated at
the midline by interparietal scutes; and a
lateral depression demarking the junction of the prefrontal and frontal. The
teeth of Xestops vagans are relatively robust and are almost uniform in size except for the very reduced posteriormost
teeth (Fig. 2), the crowns are blunt with
distinct carinae and striations radiating
perpendicular to the weakly developed
cutting edge. The intramandibular septum has a free ventral border posteriorly.
Xestops is a small, primitive glyptosaur
characterized by unfused frontals with
nearly parallel sides (Meszoely et al.,
1978). It may be distinguished from Melanosaurus (to which it is very similar and
closely related) by its less bulbous and
more isodont dentition, more gracile
mandible, more concave internal dental
border, smaller size, and somewhat less
distinct osteodermal ornamentation.
Originally described from the middle
Eocene Bridger Formation(Marsh, 1872),
Xestops vagans is now known throughout at least half of the Eocene, making it
one of the longest ranging fossil lizard
species.
Melanosaurus Gilmore, 1928

Melanosaurus maximus Gilmore, 1928
Referred specimens.-Partial
skulls
(UM 65550, 66709, 68044, 69258, 71638),
frontals (UM 73787, 74618), parietals

365
HERPETOLOGICA
December 1983]
'__
portionof rightdentary
FIG. 3.-Melanosaurus maximus. (A) Partialleft maxillaUM 66709. (B) Posterior
UM 73408. Scale= 1 cm.
(74615, 74617), and mandibles (UM

65494,65545,67246, 73408,74614, 76860,
77564, 77692), often associated with a variety of postcrania.
(SC-116, SC-117, SC-127, SC-143, SC-197,
SC-238, FG-6), Phenacodus-Ectocion
Zone (SC-23, SC-24, SC-50, SC-70, SC159, SC-164).
Discussion.-Included here are only
those specimens clearly referable to Melanosaurus maximus. Melanosaurus is a
large, heavily armored glyptosaurine
characterized by divided external nares,
fused parietals with an open parietal foramen, and massive, completely fused
frontals that taper anteriorly. Denticles
occur in large patches on the pterygoids,

but there are few on the vomers (Gilmore, 1928). The teeth have large, bulbous crowns with well-developed carinae and radiatingstriations.The shafts are
very inflated and possess broad, wrinkled
bases containing large basal foraminae
(Fig. 3). The posterior teeth are very reduced in size, and the anterior teeth are
slightly less robust, but never acute. Postcranially, Melanosaurus possesses large,
thick osteoderms in a variety of shapes
and sizes, stout limb elements, and very
massive vertebrae.
Melanosaurus is by far the most common Clarkforkianlizard, especially in the
more well-drained deposits of the Will-

366
HERPETOLOGICA
[Vol. 39, No. 4
wood Formation. It has not been recovered from the early Clarkforkian,
however; thus Melanosaurus, as presently known, must be considered an exclusively Eocene taxon.
the maxillae, very gracile mandibles, and

isodont dentition with dagger-like crowns
bearing sharp cutting edges that occasionally develop a "shouldered" appearance (Fig. 2). When fully developed, the
anterior shoulder forms a very small secondary cusp. The teeth are slightly recurved posteriorly and possess slender
shafts.
is a
Discussion.-Machaerosaurus
small anguid almost certainly referable to
the Anguinae (Meszoely et al., 1978). Machaerosaurus torrejonensis was described from the middle Paleocene Torrejon Formationof New Mexico (Gilmore,
1928) and the Torrejonian Swain Quarry
of southeasternWyoming (Sullivan, 1982).
The Clarkforkian specimens described
here are very similar to the Paleocene
material and belong to the same species.
Machaerosaurus is an extremely rare
element in the Clarkforkianof the Bighorn Basin. Specimens clearly referable
to this genus have only been recovered
from washing operations at two very fossiliferous "microsites," where very small
vertebrates are concentrated. The relative abundance of Machaerosaurusat SC188 (a locality that has provided much of
our understanding of very small Clarkforkian lizards and mammals) suggests
that it may have been fairly common during the Clarkforkian,but has suffered from
preservational and collecting biases. The
discovery and washing of additional microsites in the Clarkforkianand Wasatchian should add greatly to our knowledge
of this and other small lizards.
Subfamily Anguidae
Machaerosaurus Gilmore, 1928
Machaerosaurus torrejonensis
Gilmore, 1928
73570,
73608, 73681, 74611, 77006, 77007, fragmentary mandibles and maxillae.
(SC-143, SC-188).
Description.-These
specimens possess fine vermiculate ornamentation on
Subfamily Gerrhonotinae
"cf. Gerrhonotus sp." sensu Estes, 1964
Referred specimen.-UM 74612.
(SC-116).
Description.-A
single fragmentary
dentary of a very small lizard characterized by laterally beveled crowns bearing
extremely fine striations (Fig. 4).
Discussion.-The specimen referred to
this "taxon" possesses the chisel-shaped
B
A
FIG. 4.-(A) "Cf. Gerrhonotus sp." UM 74614,

middle portion of right dentary. (B) Exostinus rugosus UM 73565, fragmentof right maxilla. Scale =
3 mm.

Decemitber 1983]
HERPETOLOGICA
367
crowns of the cf. Gerrhonotus material

previously described from the Late Cretaceous Lance Formation and the middle
Paleocene
Tongue River Formation
(Estes, 1964, 1976). Many additional
specimens of this form are needed before
m-ore can be said of its taxonomic position
and stratigraphic range.
Family Xenosauridae Cope, 1864
Restes Gauthier, 1982
Restes rugosus (Gilmore, 1942)
Referred specimen.-UM
73565, fragmentary maxilla.
cookei Zone
(SC-188).
Descriptiotn.-This
specimen possesses characteristic rugose dermal sculpturing on the maxilla (Fig. 4). The teeth have
nearly circular shafts and bilobate crowns.
Discussion.-Restes
rugosus was descril)ed as Exostinus by Gilmore (1942).
Gauthier (1982) noted that the skull roof
osteoderms are more flattened and closely
packed than in other xenosaurids (Exostinus and Xenosaurus) and erected Restes
for this late Paleocene and early Eocene
species. It is also characterized by very
high-crowned teeth (Estes, 1975), no intramandibular septum, and extreme anterior excursion of the coronoid laterally
along the dentary.
Restes rugosus, like many other Clarkforkian vertebrates, is known only from
the middle Clarkforkian SC-188 microsite. It is therefore difficult to speculate
on the abundance and stratigraphic distribution of this species, which is quite
from other small
easily distinguished
Early Tertiary lizards.
Superfamily Varanoidea Lydekker, 1888
Varanoid
(indeterminate family, genus, and species)
Referred speciient.-UM
74619, isolated parietal.
Occurrentce.-Plesiadapis
gin gerichi
Zone (SC-215).
Discussion.-This
parietal (Fig. 5) reseimlles that of the varanid Saniwa (R.
Estes, personal communication, 1981), but
FIG. 5.-Varanoid parietal UM 74619. Scale in mm.
it may be referable to the poorly known

varanid Paleosaniwa or parasaniwid Provaranosaurus. This specimen is the only
certain record of a Clarkforkian varanoid.
Subclass Archosauria
Order Crocodilia Gmelin, 1799
Family Alligatoridae Gray, 1844
Allognathosuchus Mook, 1921
Allognathosuchus wartheni Case, 1925
Referred specimens. -Partial skull and
mandibles (UM 65778) and fragmentary
mandibles (UM 65649, 71487, 74541).
cookei Zone
(SC-62), Phenacodus-Ectocion Zone (SC57, SC-235, SC-289).
Description.-All
specimens include
robust mandibles lacking the strong surangular rise posterior to the tooth row and
highly differentiated dentition in the
middle portion of the dentary characteristic of other described Eocene Allognathosuchus species.

368
HERPETOLOGICA
M.~
I~~
[Vol. 39, No. 4
~ ~ ~~.
6.
FIG. Allognathosuchus wartheni UM 65778. (A) Superior and (B) lateral views of partial right and
UM 72573. (C) Superior and internal views of anterior fragment
left mandibles. Leidyosuchusforinidabilis
of right dentary. (D) Lateral view of posterior fragment of right mandible. Scale = 2 cm.
here are only

Discussioni.-Included
those specimens clearly referable to A.
wartheni (Case, 1925). Many of the undiagnostic specimens listed below probably belong to this species.
is a primitive alliAllognathosuchus
gator of small to medium size characterized by a broad, highly vaulted snout with
moderately large external nares divided
by a poorly ossified nasal septum formed

December 1983]
HERPETOLOGICA
by the premaxillae and nasals. The supraoccipital is included in the strongly

ornamented cranial table. The choanae
are placed posterior to the middle of the
robust pterygoids. The mandible contains prominentvertical undulations along
the very short tooth row, a small external
fenestra in the deep postdentary region,
a wide and open adductor fossa, a leaflike splenial covering of the internal surface of the dentary, and a large symphysis
that includes the splenial. The dentition
is heterodont, containing enlarged anterior canines, medial spatulate teeth, and
bulbous posterior teeth that are often
heavily worn (Fig. 6). All the teeth are
set close together and bear fine striations
and strong carinae.
369
standing of this ubiquitous Early Tertiary

crocodilian.
Ceratosuchus Schmidt, 1938

Ceratosuchus burdoshi Schmidt, 1938
Referred specimens.-Isolated
squamosals (UM 72560), partial skull and
mandibles (UM 68238, 71489), fragmentary skulls and skeletons of several individuals (UM 71490), and type skull (Field
Museum of Natural History P15576).
Occurrence.-Plesiadapis gingerichi
Zone(?) Plateau Valley Beds, Piceance
Basin, Colorado, Phenacodus-Ectocion
Zone (SC-163, SC-235).
Discussion.-The Bighorn Basin material agrees closely with the type skull
(FMNH P15576) described by Schmidt
(1938) from the Plateau Valley Beds of
Allognathosuchus
western Colorado. These horizons are
(indeterminate species)
probably early Clarkforkianin age (Rose,
64706,
1981).
65061, 65102,65551, 65659,66291,66292,
Ceratosuchus burdoshi is a broadsnouted alligator closely related to the
66538, 66546,66583,68423,69681,69914,
more common genus Allognathosuchus.
71459, 72551-72555,
72557-72559,
72561-72566, 72568-72572, 74542,74546, Ceratosuchus may be distinguished from
74550, 74623, 74624, fragmentaryskulls, the latter (which it most closely resemmandibles, and postcrania not certainly bles) by its larger and more rounded exassignable to Allognathosuchus war- ternal mandibular fenestra, more spatutheni.
late and procumbent anterior dentition,
Occurrence.-Several localities in each deeper anteriorsplenial covering over the
ClarkforkianZone.
primordial canal, less highly vaulted
re- snout, reduced dermal ornamentation of
Discussion.-Allognathosuchus
mains are common throughout the Big- the skull, and the development of large,
horn Basin Clarkforkian. Most of these horn-like squamosals (Fig. 7). Ceratosuspecimens probably belong to a species chus is much rarer than Allognathosuof the ill-defined Allognathosuchus het- chus and is thus far known only from
erodonlpolyodon morphological group Clarkforkianhorizons.
loosely characterized by their small size,
Diplocynodon Pommel, 1847
gracile skull and mandible, poorly develDiplocynodon sp.
oped anterior vertical undulation on the
dentary, high surangular rise, and deep
Referred specimen.-UM 69867, antepostdentary region. The inadequate de- rior portion of a left dentary.
scriptions and illustrations of A. heteroOccurrence .-Phenacodus -Ectoci on
don and A. polyodon (Cope, 1872; Mook, Zone (SC-289).
1921, 1961) prohibit specific assignment
Discussion.-This specimen belongs to
of small, generalized early Eocene Allo- a form very similar to the narrow-snouted
gnathosuchus specimens. A thorough re- Diplocynodon stuckeri from the middle
view of these forms combined with the Eocene Bridger Formation (Mook, 1960).
description of additional early Eocene Diplocynodon is a large primitive caimaterial will add greatly to our under- man. It is the most common Tertiarycroc-

370
,
i
|--l
lg
:t
8
sE_q
,-
#:-
Q
X
y .t. ....
...:+
__
!..
_L_W
__
_l
*^
5;il_
ff
>
_wa
rj
,$S;_
o
111
IlIwofs
S1s_I
(_ lb1_1
w
4 ..,,;;! r isWSew
r
s
[Vol. 39, No. 4
HERPETOLOGICA
t
..
.i
fl
_
_
S
4
',
X#
__
:.
.........e.
>
{' is:
:
_
.
1
.L
^.
:,
' ;^;
'
,s
i-
i_
..
bi-S
>
<R9v_
..
uu
*r
.5
-SEis
_7sH-
J{7
*'
to
;3
>
b*v
{*
* to ww6s
o.v
>t
* x,
; os
'
>;'st"'j.tJ'9;.+<t
.Xv.-e
''.
'J
%,
.'v.#.
'-
. _
<>,sr.
v
'
;4_
;,.
*,
a5,
{_
'W'
3Ffd f
;
_ffi>>
--*
?A
^X_
_.,>
3SK
't
o-
<'Be3* *
tZ'
%.
s_
g.F
is95s
_.
?
w
__
x-
.,^\
M
-
Y
,
1Y
t
Z f
tst@
* 4h01
&
w
_
-?-*-
w-s
w
i
-- flh;_;
oW.o_Rj
-X#LB[=
>_
^XDg_:-
S.-o
s-^-.
Un. . *xxj
_
ht
._
F
.
_
FIG. 7.eratosuchus burdoshi.(A)Superiorandlateralviews of partialrightmandibleUM 71490.(B)
Posteriorand superiorviews of the horn-likesquamosalsUM 72560.Scale = 2 cm.

December 1983]
371
HERPETOLOGICA
odilian in Europe (Berg, 1966) but is quite

rare in North American deposits of simi- A
lar age. The North American specimens
differ from their European counterparts
in the possession of more pronounced
vertical undulations along the tooth row,
somewhat more robust mandible, larger
symphysis, and a stronger confluence of
the third and fourth dentary teeth (Fig. B
8).
The referred Clarkforkian specimen
differs in detail from D. stuckeri and
probably represents a new species.
Family Crocodylidae Cuvier, 1807
Subfamily Leidyosuchinae Nopsca, 1928
Leidyosuchus Lambe, 1907
Leidyosuchusformidabilis Erickson, 1976
Referred specimens.-Fragmentary
skulls and postcrania (UM 72573, 74552)
and isolated teeth (UM 74553, 74554,
77001, 77003).
gingerichi
Zone (SC-171, SC-249), Plesiadapis cookei Zone (SC-108, SC-117, SC-120, SC-134,
SC-136, SC-200), Phenacodus-Ectocion
Zone (SC-il, SC-49, SC-81, SC-163, SC164, SC-183, SC-202, SC-234).
Discussion.-These specimens are referable to the large crocodylid Leidyosuchus formidabilis (Fig. 6). Leidyosuchus is a common element of Late
Cretaceous and Early Tertiary faunas of
western North America. It is a primitive
crocodylid characterized by frontal participation in the anteromedial border of
the supratemporalfenestrae, a nasal-frontal contact, and a gently sloping occiput.
The posterior portion of the skull is very
flat and broad, and there is a pronounced
posteriorexcursion of the squamosalsonto
the dorsal surface of the quadrates. The
mandible is very deep posteriorly and
possesses a large external fenestra and
extremely long, shallow symphysis. Leidyosuchusformidabilis is a larger, narrowsnouted species, previously reported from
the Paleocene (Tiffanian) of North Dakota (Erickson, 1976).
Leidyosuchus teeth occur through the
Tiffanian, Clarkforkianand Wasatchianof
FIG. 8.-Diplocynodon sp. UM 69867. (A) Superior view showing cross sections of broken first and
third teeth, (B) lateral view, and (C) internal view
showing the rugose area of splenial attachment behind the large, curved symphysis. Scale = 1 cm.
the Bighom Basin. These teeth (and more
complete specimens) are common in Fort

Union horizons but become quite rare
with the onset of Wiliwood sedimentation. Habitational and preservational
biases (similar to those postulated for the
champsosaurs) may have diminished the
fossil record of Leidyosuchus during the
early Eocene. This change from large
stream and swamp environments (Fort
Union) to smaller stream and broad floodplain conditions (Willwood) seems to
have greatly benefited the small alligators, however, and they become very
abundant in Willwood sediments (perhaps at the expense of the larger aquatic
predators).
FAUNAL
COMPARISONS
Clarkforkiandeposits record the transition from relatively archaic Paleocene

herpetofaunas to Eocene assemblages of

372
HERPETOLOGICA
a more modern aspect. North American

Paleocene reptile communities are, in
general, little changed from Late Cretaceous assemblages, and comparisons with
Lancian reptile faunas are also pertinent
(Estes, 1970; Estes et al., 1969).
Estes (1964) described extensive University of California collections from the
type area of the Lance Formation in eastern Wyoming. Except for the lizards (and
dinosaurs), the Lancian fauna is quite
similar to that found in the Clarkforkian.
The aquatic component of the Cretafauna contains
ceous
of
species
Clarkforkian genera ("Baena," Champ-
sosaurus, and Leidyosuchus), trionychids

similar to later forms, and the primitive
alligator Brachychampsa (Gilmore, 1911).
The major aquatic element lacking from
the Lance beds are the emydids (J. H.
Hutchison, personal communication,
1980). The Late Cretaceous lizards are
very different from those of the Clarkforkian. Of the fourteen Lancian lizards, only
Odaxosaurus and the problematical "cf.
Gerrhonotus" are known from Clarkforkian horizons.
A few undescribed specimens of large
reptiles are known from the basal Paleocene Mantua Quarry (Polecat Bench, Bighorn Basin, Wyoming). This small assemblage contains a baenid, a trionychid,
indeterminate species of Champsosau-
rus and Leidyosuchus, and Diplocynodon. Allognathosuchus mooki has not

been recovered from the quarry, but it is
known from the very early Paleocene of
New Mexico (Simpson, 1930). Except for
the absence of emydids, the early Paleocene reptile fauna differs only slightly
from that of the Clarkforkian. No lizards
are present in the Mantua sample.
Middle Paleocene reptile assemblages
are slightly more advanced than those of
the early Paleocene. Estes (1976) described the lower vertebrates from the late
Torrejonian Tongue River Formation of
Montana. This fauna is a mixture of Late
Cretaceous holdovers and new Paleocene taxa. Clarkforkian taxa present in the
[Vol. 39, No. 4
Bighorn Basin by this time include

Champsosaurus gigas, Plastomenus, Machaerosaurus, Odaxosaurus and Allognathosuchus. The middle Paleocene
fauna is still somewhat primitive in that
it lacks a wide assortment of anguid lizards and alligators.No emydids are known
from the middle Paleocene of the Bighorn Basin, although a macrocephalic
form is present in New Mexico at this time
(J. H. Hutchison, personal communication, 1982).
The late Paleocene (late Tiffanian) reptilian fauna of the Bighorn Basin is much
more advanced than earlier Tertiary assemblages. Clarkforkiantaxa present by
this time include Restes rugosus, Leidyosuchus formidabilis, Allognathosuchus heterodon, and a variety of turtles,
including the macrocephalic emydid and
two chelydrids (J. H. Hutchison, personal
communication, 1981). The only Clarkforkian elements not present by the late
Tiffanian are Machaerosaurus, Ceratosuchus, Allognathosuchus wartheni and
the emydid box turtle.
In contrast to the gradual transformation between older Paleocene and Clarkforkian faunas, there is a fairly dramatic
faunal break at the Clarkforkian-Wasatchian boundary. Extinctions at this time
include Ceratosuchus and the champsosaurs. Wasatchian faunas are also characterized by a diversification of emydids
and glyptosaurinids, the first appearance
of kinosternid, dermatemydid and testudinid turtles (Hutchison, 1980), and the
occurrence of modern rhineurid amphisbaenians and the problematical crocodilian Orthogenysuchus (Mook, 1924).
The Clarkforkiantaxa that continue into
the Wasatchianwith little change include
Allognathosuchus, Leidyosuchus, Melanosaurus and some of the emydid and
trionychid turtles.
In summary, the Clarkforkian reptile
fauna represents an assemblage that
evolved slowly throughoutthe Paleocene
from the typical Late Cretaceous non-dinosaurian herpetofauna. The Wasatchian

373
HERPETOLOGICA
December 1983]
early Eocene reptile fauna is a diversified

Clarkforkian assemblage mixed with a
variety of immigrant taxa.
PALEOECOLOGY
Reptiles are environmentally sensitive

and provide some general information on
the climate and geography of the region
in which they live (or lived). Since some
Clarkforkianreptiles are closely related
to their Recent counterparts, some of
these ecological inferences can be applied to the latest Paleocene-earliest
Eocene environments of the Bighorn Basin.
The Clarkforkian trionychids are extremely similar to living taxa and provide
reliable comparisons. Living Trionyx
species inhabit larger water bodies (lakes,
large ponds and streams, rivers), indicating the presence of significant amounts
of permanent water during the Clarkforkian. Trionyx is quite temperature tolerant
(Recent species range from Florida to
Michigan), so little can be said regarding
paleotemperatures.
Few paleoecological inferences can be
drawn from the champsosaurs. They are
regarded as piscivorous and highly aquatic, and their large size probably restricted
them to major streams and ponds.
Clarkforkianlizards are all distantly removed morphologically and phylogenetically from their Recent relatives and
therefore provide little insight into Clarkforkian environments. The predominance of presumably terrestrial glyptosaurs may indicate a considerable amount
of dry ground between the streams and
ponds, particularly during Willwood deposition.
Snakes, amphisbaenians and amphibians are either extremely rare or absent
from most Bighorn Basin Tertiary horizons. This scarcity must be due in large
measure to taphonomic and collecting
biases. Each of these groups possess small
delicate skeletons that are not easily preserved, weathered to the surface, or seen
by the collector. It is unlikely that the
absence of these forms indicates any unusual paleoecologic condition during the
Clarkforkian.
The great abundance and diversity of
Clarkforkiancrocodilians suggest an equal
abundance and variety of aquatic habitats. Recent caimans and crocodylids do
not hibernate, and it is unlikely that their
primitive ancestors (Diplocynodon and
Leidyosuchus) were capable of this complex activity. If this is true, it indicates
that Clarkforkian temperatures were
equable and there was no hard winter
freeze. Allognathosuchus is closely related to the Recent Alligator and may have
shared its descendants' intoleranceto very
warm tropical climates (Alligator mississippiensis and A. sinensis are found only
in subtropical and warn temperate regions). Combined, the crocodilians may
bracket Clarkforkiantemperatures in the
subtropical range.
thank P. D. Gingerich for
Acknowledgments.-I
reviewing this manuscript and allowing me to study
the collections at the University of Michigan. Special thanks go to J. H. Hutchison for providing unpublished information on the evolution of Early
Tertiary turtles, and D. Baird for the loan of specimens at Princeton University. I also thank W.
Langston, Jr. and R. M. Sullivan for their helpful
insights on Tertiary reptiles. This research was supported in part by National Science Foundation grants
DEB 77-13465 and DEB 80-10846 to P. D. Gingerich.
LITERATURE CITED
J. D., AND J. H. HUTCHISON. 1979.

Revision of the genus Palatobaena (Testudines,
Baenidae), with description of a new species.
Postilla 177:1-19.
BARTELS, W. S. 1980. Early Cenozoic reptiles and
birds from the Bighorn Basin, Wyoming. In P. D.
Gingerich (Ed.), Early Cenozoic Paleontology and
Stratigraphy of the Bighorn Basin. Univ. Michigan Pap. Paleont. 24:73-79.
BERG, D. E. 1966. Die Krokoddile, insebesondere
Asiatosuchus und aff. Sebecus?, aus dem Eozan
von Messel bei Darmstadt/Hessen. Abh. Hess.
Landesamt. Bodenforsch. 52:1-105.
BowN, T. M. 1979. Geology and mammalian paleontology of the Sand Creek Facies, Lower Willwood Formation (Lower Eocene), Washakie
County, Wyoming. Geol. Surv. Wyoming Mem.
2:1-151.
ARCHIBALD,

374
1925. Note on a new species of the

Eocene crocodilian Allognathosuchus, A. wartheni. Contr. Mus. Geol. Univ. Michigan 2:93-97.
COPE, E. D. 1872. On the dentition of Metalophodon. Proc. Amer. Phil. Soc. 12:542-545.
ERICKSON, B. R. 1972. The lepidosaurian reptile
Champsosaurus in North America. Monogr. Sci.
Mus. Minnesota, Paleont. 1:1-91.
. 1976. Osteology of the early eusuchian
crocodile Leidyosuchus formidabilis, sp. nov.
Monogr. Sci. Mus. Minnesota, Paleont. 2:1-61.
ESTES, R. 1964. Fossil vertebrates from the Late
Cretaceous Lance Formation, eastern Wyoming.
Univ. California Publ. Geol. Sci. 49:1-180.
1970. Origin of the Recent North American lower vertebrate fauna: an inquiry into the
fossil record. Forma et Functio 3:139-163.
. 1975. Lower vertebrates from the Fort
Union Formation, late Paleocene, Big Horn Basin, Wyoming. Herpetologica 31:365-385.
1976. Middle Paleocene lower vertebrates
from the Tongue River Formation, southeastern
Montana. J. Paleont. 50:500-520.
ESTES, R., P. BERBERIAN, AND C. A. M. MESZOELY.
1969. Lower vertebrates from the late Cretaceous Hell Creek Formation, McCone County,
Montana. Breviora 337:1-53.
GAFFNEY, E. S. 1972. The systematics of the North
American family Baenidae (Reptilia, Cryptodira).
Am. Mus. Nat. Hist. Bull. 147:243-319.
. 1979. Description of a large trionychid
turtle shell from the Eocene Bridger Formation
of Wyoming. Contrib. Geol. Univ. Wyoming 17:
53-57.
GAUTHIER, J. A. 1982. Fossil xenosaurid and anguid lizards from the early Eocene Wasatch Formation, southeast Wyoming, and a revision of the
Anguioidea. Contrib. Geol. Univ. Wyoming 21:754.
GILMORE, C. W. 1911. A new fossil alligator from
the Hell Creek beds of Montana. Proc. U.S. Nat.
Mus. 41:297-302.
. 1928. Fossil lizards of North America.
Mem. Nat. Acad. Sci. 22:1-169.
1938. Descriptions of new and little-known
fossil lizards from North America. Proc. U.S. Nat.
Mus. 86:11-26.
. 1942. Paleocene faunas of the Polecat
Bench Formation, Park County, Wyoming. Part
II. Lizards. Proc. Amer. Phil. Soc. 85:159-167.
GINGERICH, P. D. 1976. Cranial anatomy and evolution of Early Tertiary Plesiadapidae (Mammalia, Primates). Univ. Michigan Pap. Paleont. 15:
1-141.
1980. Evolutionary patterns in Early Cenozoic mammals. Ann. Rev. Earth Planet. Sci. 8:
407-424.
HAY, 0. P. 1908. Fossil Turtles of North America.
Carnegie Institute, Washington.
CASE, E. C.
[Vol. 39, No. 4
HERPETOLOGICA
J. H. 1980. Turtle stratigraphyof the

Willwood Formation, Wyoming: preliminary results. In P. D. Gingerich (Ed.), Early Cenozoic
Paleontology and Stratigraphyof the Bighorn Basin. Univ. Michigan Pap. Paleont. 24:115-118.
MARSH, 0. C. 1872. Preliminary description of
new Tertiaryreptiles, Part 1. Amer. J. Sci. 4:298309.
MESZOELY, C. A. M. 1970. North Americanfossil
anguid lizards. Bull. Mus. Comp. Zool. Harvard
University 139:87-150.
MESZOELY, C. A. M., R. ESTES, AND H. HAUBOLD.
1978. Eocene anguid lizards from Europe and a
revision of the genus Xestops. Herpetologica 34:
156-166.
MOOK,C. C. 1921. Allognathosuchus, a new genus of Eocene crocodilians.Amer.Mus. Nat. Hist.
HUTCHISON,
Bull. 44:105-110.
1924. A new crocodilianfromthe Wasatch

Beds. Amer. Mus. Novitates 137:1-4.
. 1960. Diplocynodon remains from the
Bridger Beds of Wyoming. Amer. Mus. Novitates
2007:1-4.
. 1961. Notes on the skull charactersof Allognathosuchus polyodon. Amer. Mus. Novitates
2072:1-5.
ROSE, K. D.
1981. The ClarkforkianLand-Mammal Age and mammalianfaunalcompositionacross

the Paleocene-Eocene boundary.Univ. Michigan
Pap. Paleont. 26:1-197.
SCHMIDT, K. P. 1938. New crocodilians from the
upper Paleocene of western Colorado.Field Mus.
Nat. Hist., Geol. Ser. 6:315-321.
SIGOGNEAU-RusSELL, D., AND D. BAIRD. 1978.
Presence du genre Simoedosaurus (Reptilia,
Choristodera)en Amerique du Nord. Geobios 11:
251-255.
D., AND D. E. RUSSELL.
1978. Etude osteologique du reptile Simoedosaurus (Choristodera).Ann. Paleontologie (Vert.)
64:1-84.
SIMPSON, G. G. 1930. Allognathosuchus mooki, a
new crocodile from the Pureco Formation.Amer.
Mus. Novitates 445:1-16.
SULLIVAN, R. M. 1979. Revision of the Paleogene
genus Glyptosaurus (Reptilia, Anguidae). Amer.
SIGOGNEAu-RusSELL,
Mus. Nat. Hist. Bull. 163:1-72.
. 1982. Fossil lizards from Swain Quarry

"Fort Union Formation,"middle Paleocene (Torrejonian), Carbon County, Wyoming. J. Paleont.
56:996-1010.
Accepted: 6 June 1983

Associate Editor: Stephen Tilley
University of Michigan, Museum of Paleontology, Ann Arbor, MI 48109, USA


Bartels A Transitional Paleocene-Eocene Reptile Fauna From The Bighorn Basin, Wyoming

Transféré par

Informations du document

Titre original

Copyright

Formats disponibles

Partager ce document

Partager ou intégrer le document

Options de partage

Avez-vous trouvé ce document utile ?

Ce contenu est-il inapproprié ?

Droits d'auteur :

Formats disponibles

Bartels A Transitional Paleocene-Eocene Reptile Fauna From The Bighorn Basin, Wyoming

Transféré par

Droits d'auteur :

Formats disponibles

Herpetologists' League

A Transitional Paleocene-Eocene Reptile Fauna from the Bighorn Basin, Wyoming

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

Arima, AMNH 87398-401, 87402 (skeleton); Mt.

39(4), 1983, 359-374

? 1983 by The Herpetologists' League, Inc.

A TRANSITIONAL PALEOCENE-EOCENE REPTILE FAUNA

Choristodera; Lacertilia; Crocodilia; Fauna; Paleocene-

THE Clarkforkian Land-Mammal Age

pect. The Clarkforkianis preceded in time

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

Prior to a preliminary version of this

[Vol. 39, No. 4

At least 21 species representing 10

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

Superfamily Trionychoidea Gray, 1870

Zone (SC-156, SC-171). Phenacodus-Ectocion Zone (SC-71, SC-159).

(SC-62, SC-136). Phenacodus-Ectocion

The Clarkforkian macrocephalic genus

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

[Vol. 39, No. 4

as "unrelated to extant or other known

box turtles ....

This genus is the earliest

known example of a fully developed box

turtle is currently being completed by J.

Family Chelydridae Gray, 1870

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

Discussion.-Most elements of typical

aquatic and fairly large (body length to 4

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

[Vol. 39, No. 4

Melanosaurus Gilmore, 1928

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

(74615, 74617), and mandibles (UM

occur in large patches on the pterygoids,

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

[Vol. 39, No. 4

the maxillae, very gracile mandibles, and

FIG. 4.-(A) "Cf. Gerrhonotus sp." UM 74614,

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

crowns of the cf. Gerrhonotus material

FIG. 5.-Varanoid parietal UM 74619. Scale in mm.

it may be referable to the poorly known

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

[Vol. 39, No. 4

here are only

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

by the premaxillae and nasals. The supraoccipital is included in the strongly

standing of this ubiquitous Early Tertiary

Ceratosuchus Schmidt, 1938

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

[Vol. 39, No. 4

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

odilian in Europe (Berg, 1966) but is quite

the Bighom Basin. These teeth (and more

complete specimens) are common in Fort

Clarkforkiandeposits record the transition from relatively archaic Paleocene

This content downloaded on Sat, 9 Feb 2013 20:35:40 PM

a more modern aspect. North American

sosaurus, and Leidyosuchus), trionychids

rus and Leidyosuchus, and Diplocynodon. Allognathosuchus mooki has not

[Vol. 39, No. 4