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Number 3342, 31 pp., 19 figures, 1 table June 22, 2001
A Partial Skeleton of Pseudaelurus (Carnivora:
Felidae) from the Nambé Member of the Tesuque
Formation, Espanola Basin, New Mexico
TOM ROTHWELL!
ABSTRACT
I describe a partial felid skeleton of Pseudaelurus validus, new species, from the early
Miocene, late Hemingfordian of New Mexico. This is the earliest known felid from North
America having cranial, upper, and lower dentition as well as postcranial information. A nearly
complete set of limb bones provides the first insight into the morphometry of this lynx to
puma-sized cat. The skull, similar to that of the earlier European Proailurus lemanensis, dis-
plays a high and thin sagittal crest with prominent nuchal crests. In the auditory region, the
tympanic bulla architecture resembles that of Proailurus and relates to modern felids. The
skeleton includes an articulated manus and pes with an apparently functional first metatarsal
bone, hitherto unknown in the genus Pseudaelurus. Log-ratio diagrams are used to compare
this skeleton with various extinct and modern felids. Dental morphology, as well as the new
postcranial information revealed by this skeleton, is contrasted with that of other felids.
INTRODUCTION ements. Only three skulls of Pseudaelurus
have been mentioned in the literature; all are
Pseudaelurus is a felid taxon that has been
from North America, and none have been as-
reported from the early Miocene of Europe,
sociated with postcranial material.
Asia, and North America. Most of the spec-
The relationship of modern cats to extinct
imens described in the literature consist of Felidae has been under study since the work
mandibular and maxillary fragments. Post- of Cope (1880). A common ancestry of the
crania, largely from Europe, are uncommon late Oligocene Proailurus lemanensis, the
and consist for the most part of isolated el- Miocene Pseudaelurus, and living felids has
' Graduate Student, Division of Paleontology, American Museum of Natural History and Columbia University.
Copyright © American Museum of Natural History 2001 ISSN 0003-0082 / Price $4.00
2 AMERICAN MUSEUM NOVITATES NO. 3342
been suspected since before the turn of the Pseudaelurus for the Blainville specimen. He
last century (Adams, 1897). Although a distinguished it from the genus Felis by a
small amount of cranial and postcranial ma- single character: “‘possessing an additional
terial has been described of the earliest genus inferior premolar (p2) in advance of the oth-
Proailurus (Filhol, 1888; Helbing, 1928), the ers’. This was the first recognition of an an-
postcrania of Pseudaelurus have been largely cestral state, or plesiomorphic anatomical
unknown. Considerable cranial and postcra- condition of a fossil specimen, with respect
nial material for Felis is found throughout to extant members of Felidae.
the fossil record from its earliest appearance In North America, Joseph Leidy (1858)
in the late Miocene of Europe (de Beaumont, described another new species of Miocene
1961b) and Asia (Heintz, 1981; Qi, 1985) cat, Felis intrepidus, from “‘the loose sands
(fig. 1). Information on the postcrania of of the Niobrara River’’ in western Nebraska.
Pseudaelurus, linked to a skull, is therefore This fossil also possessed evidence of the
needed to further the understanding of the lower second premolar, but Leidy apparently
relationships among these three felid genera. did not immediately recognize the generic
value of this character. By the time he had
ABBREVIATIONS published a drawing of the left ramus (Leidy,
1869) however, he had designated it as Pseu-
Anatomical daelurus intrepidus. He explained the signif-
icance of the additional tooth: “In both rami
a-p anterior to posterior
c lower canine tooth the premolar (p2), considered as the chief
C upper canine tooth character of the genus, is absent, but its al-
d-v dorsal to ventral veolus remains midway in the hiatus back of
il lower first incisor the canine tooth’’ (Leidy, 1869).
I3 upper third incisor Filhol (1879) proposed a new genus,
ml lower first molar Proailurus, to accommodate specimens from
mm millimeter the early Miocene (Aquitanian) of France
Mcl first metacarpal bone
with characters more plesiomorphic and fos-
Mtl first metatarsal bone
sils stratigraphically older than those already
M1 upper first molar
assigned to Pseudaelurus. Proailurus lema-
pl lower first premolar
phl first phalanx nensis mandibles, dating back to the late Ol-
P3 upper third premolar igocene Quercy fissures of France, possessed
four lower premolars where Pseudaelurus
Institutional
had three. When E.D. Cope (1880) published
AMNH American Museum of Natural His- his treatise: ‘““On the Extinct Cats of North
tory, New York America’’, he clearly suggested that Proail-
F:AM Frick Collection, American Museum urus and Pseudaelurus were ancestral felids.
of Natural History Pseudaelurus specimens first appear in the
MHNL Muséum Guimet d’Histoire Natur-
early Miocene (MN3) of Europe (Dehm,
elle, Lyon
1950). The specimens are numerous in com-
USNM United States Natural History Mu-
parison to Proailurus. However, the over-
seum, Washington, DC
whelming majority of the material reported
in the literature consists of isolated lower
HISTORY OF THE GENUS PSEUDAELURUS
jaws. Four species have been named from the
In 1843, H. M. Ducrotay de Blainville de- Miocene of Europe: (1) Pseudaelurus quad-
scribed a maxillary fragment and a lower jaw ridentatus Blainville, 1843; (2) Pseudaelurus
fragment of a “‘highly specialized machai- turnauensis Hoernes, 1882; (3) Pseudaelurus
rodont”’ from the medial Miocene of Sansan, lorteti Gaillard, 1899; and (4) Pseudaelurus
France. The ramus contained three lower pre- romieviensis Roman-Viret, 1934.
molars (p2, p3, p4), whereas modern cats Charles Depéret (1892) described a
normally have only two (p3, p4). He gave small specimen, Pseudaelurus transitorius
this specimen the name Felis quadridentata. from the La Grive-Saint-Alban (Isere) lo-
In 1850, Paul Gervais established the genus cale of France. This species was placed in
ROTHWELL: PSEUDAELURUS FROM NEW MEXICO 3
NALMA EURO FAUNAL
PLEIST. IRVINGTONIAN MN21-
16
ba BLANCAN
MN15
14
HEMPHILLIAN OTnoT o—_— MN13
MN10
cunewowwn
= N
No| s©o o re
=/N=
BARSTOVIAN ~]
MN6
MN5
m2Zzmnmnoo0o-éez
MN4
|N eaibabaariay |
MN3
MN2
ARIKAREEAN MN1
OO-TO
Fig. 1. Stratigraphic ranges for Proailurus spp. (Pro), Pseudaelurus spp. in Europe (Eur), Pseudae-
lurus spp. in North America (NA) and Felis (Felis). There appears to be no overlap in the fossil record
among these three genera of Felidae. Proailurus is not found after MN2b. European Pseudaelurus range
is that of the smallest species, P. turnauensis: MN3 to MN9. Correlation references: McKenna and Bell
(1997), MacFadden and Hunt (1998).
synonymy with P. turnauensis Hoernes by sis) from the medial Miocene La Grive-
G. de Beaumont (1961). The synonymiza- Saint-Alban locality in France (Gaillard,
tion was confirmed by Heizmann (1973) 1899). G. de Beaumont (1961) described
and Wang (1998), but ignored by Ginsburg 19 separate postcranial bones from La Gri-
(1983). All four species was originally ve-Saint-Alban and the early Miocene Win-
described based on mandibular and maxil- tershof-West localities, but none was al-
lary fragments. There have been no Euro- leged to be from the same individual. L.
pean skulls yet described. The first post- Ginsburg (1961b) reported an even larger
cranial bone was described in 1899: a right assortment of isolated mandibular, maxil-
humerus of P. transitorius (= P. turnauen- lary and postcranial material from the me-
+ AMERICAN MUSEUM NOVITATES NO. 3342
dial Miocene of Sansan in France, again mandibles: Pseudaelurus pedionomus from
without a skull. the Minnechaduza Fauna of Nebraska and
Richard Dehm (1950), in his seminal re- Pseudaelurus thinobates from the Black
port on the carnivores from Wintershof-West Hawk Ranch Local Fauna of California. Da-
in Germany, described 18 lower jaws and 1 vid Kitts (1958) erected a new genus, Nim-
maxillary specimen. These are early Miocene ravides, for this material and made Pseudae-
specimens of Burdigalian (MN3) age. Dehm lurus thinobates its type species.
assigned them to P. transitorius Depéret (= Some North American cranial material has
P. turnauensis Hoernes). Dehm described been described. Chester Stock (1934) de-
these specimens as elements from a transi- scribed the first Pseudaelurus skull in a re-
tional species between Proailurus and the port on five individuals from the early Bar-
more derived specimens of Pseudaelurus re- stovian Tonopah, Nevada locality. He as-
ported from La Grive-Saint-Alban in France. signed all to P. intrepidus Leidy. MacDonald
What Dehm considered most interesting in (1948a) described seven assorted postcranial
this collection was the variable key lower bones referenced to P. pedionomus, but ma-
jaw characters: (1) presence or absence of
terial referred to this species has subsequent-
pl; (2) presence or absence of p2; (3) one or ly been shifted into the genus Nimravides (de
two roots of p2; (4) presence or absence of
Beaumont, 1990). Shotwell and Russel
m2; (5) presence or absence of ml metaco-
(1963) also described some assorted postcra-
nid; (6) length of m1.
nial fragments of Pseudaelurus, which they
Elmar Heizmann (1973) published a re-
thought “‘apparently represent a single indi-
view of the European Pseudaelurus radiation
vidual’’, but no species determination was
and recognized four species in order of in-
made.
creasing size: (1) Pseudaelurus turnauensis
In Asia, three species of Pseudaelurus
Hoernes; (2) Pseudaelurus lorteti Gaillard;
have been reported. All are described from
(3) Pseudaelurus romieviensis Roman and
mandibular, maxillary, or dental material:
Viret; (4) Pseudaelurus quadridentatus
Pseudaelurus guangheensis Cao, Du, Zhao,
Blainville. He suggested that Pseudaelurus
and Cheng, 1990, is a new species from the
evolved from Proailurus with the following
medial Miocene Guanghe District of Gansu,
character changes: (1) loss of pl; (2) loss of
China. Pseudaelurus cuspidatus Wang, 1998,
m2; (3) reduction of p2 to one root; (4)
is a new species from the early medial Mio-
lengthening of canines; (5) reductions in
cene Halamagai Formation in northern Jung-
height of lower premolars such that p3 and
gar Basin, in the Xinjiang Autonomous Re-
p4 became similar in height; (6) reduction of
gion, China. P. lorteti left ramus and dental
m1 protoconid height; (7) movement of ml
fragments were reported from the medial
metaconid from a position on the posterior
surface of the protoconid, lingual of median, Miocene of Xiacaowan, Sihong County in
Jiangsu Province, China (Qiu and Gu, 1986).
to a median position; (8) reduction of m1 tal-
From Africa, an m1 and tibia fragment
onid.
from the Al-Sarrar locality in what is now
In North America, three species have been
described from the medial Miocene. Again, Saudi Arabia were assigned to P. turnauensis
all species determinations were based on (Thomas et al., 1982). This formation is early
mandibular and maxillary fragments. Pseu- Miocene (MN4a) in age. There are no other
reports of any nonmachairodont members of
daelurus intrepidus Leidy, 1858, and Pseu-
Felidae from the African Miocene.
daelurus marshi Thorpe, 1922, are both late
Barstovian specimens from the Valentine The Nambé skeleton (F:AM 62128) de-
Formation in western Nebraska. Pseudaelu- scribed in this paper was collected in 1939
rus aeluroides Macdonald, 1954, is from the by J. C. Blick of the Frick Laboratory. The
early Barstovian Olcott Formation in Sioux specimen was recovered from the late Hem-
County, Nebraska. J. R. Macdonald (1948a, ingfordian Nambé Member locality (Galusha
1948b) described two large Clarendonian and Blick, 1971) in the Tesuque Formation,
Pseudaelurus species that have enlarged Espanola Basin, near East Cuyamunque,
mental ridges on the ventral margins of the New Mexico.
2001 ROTHWELL: PSEUDAELURUS FROM NEW MEXICO Gy)
SYSTEMATIC PALEONTOLOGY right and left ulnae, left manus, articulated
ORDER CARNIVORA BOWDICH, 1821 right manus, distal fragment of left femur
with patella, left tibia, right tibial fragment,
SUBORDER FELIFORMIA KRETZOI, 1945
left and right fibular fragments, right pes, ar-
FAMILY FELIDAE FISCHER DE WALDHEIM, 1817
ticulated left pes, partially prepared elements
in plaster jacket, including exposed scapulae,
Genus Pseudaelurus Gervais, 1850
ribs, vertebrae.
DISTRIBUTION: Early to medial (MN3 to
TYPE LOCALITY: Nambé Member, Tesuque
MNQY) Miocene of Europe; early to medial
Formation, Espanola Basin near East Cuy-
(late Hemingfordian to late Barstovian) Mio-
amunque, New Mexico.
cene of North America; early medial Mio- AGE: Late Hemingfordian (Galusha and
cene of Asia; early (MN4a) Miocene of Af- Blick, 1971).
rica.
ETYMOLoGy: validus, Latin for strong, ro-
GENERIC DIAGNOSIS: Members of Felidae
bust, able.
with the unique combination of the following
REFERRED SPECIMENS: Sheep Creek Fauna,
derived and primitive characters: presence of
Sheep Creek Formation (late Hemingfordi-
p2 with the usual absence of pl and m2, ml
an), Sioux County, Nebraska: F:AM 61827,
with reduced metaconid and talonid, P2 with
right ramus with c, alveolus p2, p3, p4, bro-
one root, alisphenoid canal present, paroccip-
ken ml, Greenside Quarry; F:AM 61837, left
ital process cupped about the posterior sur-
maxillary fragment with I3, C-M1, right tibia
face of an enlarged caudal entotympanic.
and fibula, partial right radius, partial right
Differing from extant felid genera by c cross
ulna, astragalus, vertebrae and right fourth
section showing flattened inner surface and
metatarsal, Head of Pliohippus Draw; Lower
posterior trenchant edge, presence of p2, m1
Snake Creek Fauna, Olcott Formation (early
with variable metaconid and reduced talonid,
Barstovian), Sioux County, Nebraska: F:AM
presence of alisphenoid canal. Differing from
61834, complete skull with upper dentition,
Metailurus by presence of alisphenoid canal
zygomatic arches, intact basicranium and as-
and p2, absence of enlargement of mental
sociated lower jaws (F:AM 61829), Humbug
ridge. Differing from Nimravides by smaller
Quarry; F:AM 61803, left maxilla with [3
size and absence of any ventral mandibular
alveolus, C, alveolus Pl and P2; P3-M1,
mental ridge enlargement. Differing from
Humbug Quarry; F:AM 61833, partial skull
Proailurus by absence of pl and m2.
with upper dentition and partial right zygo-
TYPE SPECIES: Pseudaelurus quadridenta-
matic arch, Quarry 3 (Far Surface Quarry),
tus, (Blainville, 1843) (=Felis quadridenta
F:AM 61832, left ramus with 13, c, p2, bro-
Blainville, 1843).
ken p3, p4 and broken m1, East Wall Quarry;
INCLUDED SPECIES: Pseudaelurus intrepi-
F:AM 61830, partial left ramus with p4, m1,
dus Leidy, 1858; Pseudaelurus turnauensis Quarry 3 (Far Surface Quarry); F:AM
Hoernes, 1882; Pseudaelurus lorteti Gail-
61828, left and right rami with left and right
lard, 1899; Pseudaelurus marshi Thorpe,
c, p2 alveoli, p3-m1, right humerus, left ra-
1922; Pseudaelurus romieviensis Roman-V1- dius, left and right tibiae, Echo Quarry; F:
ret, 1934; Pseudaelurus aeluroides Macdon-
AM 61835, a nearly complete skull with up-
ald, 1954; Pseudaelurus guangheensis Cao,
per dentition and intact basicranium, Echo
Du, Zhao, and Cheng, 1990; Pseudaelurus
Quarry; F:AM 61836, skull fragment with
cuspidatus Wang, 1998.
left P4, left and right partial rami with partial
dentitions, maxillary fragment with canine
Pseudaelurus validus, new species
tooth, left radius, Echo Quarry.
HoLotyPe: F:AM 62128, skull with left DISTRIBUTION: Late Hemingfordian of New
and right I3, left C, broken right C, single- Mexico, late Hemingfordian and early Bar-
rooted alveoli for Pl and P2, left and right stovian of Nebraska.
P3-M1. Articulated lower jaws with left 13, DIAGNosis: Differing from other species
left and right c, left and right single-rooted by the combination of large size, long dis-
alveoli for p2, left and right p3-m1. Postcran- tance between c and p3, extremely reduced
ia: right and left humeri, right and left radii, or absent metaconid on m1, and robust den-
6 AMERICAN MUSEUM NOVITATES NO. 3342
tary with large and erect rectangular-shaped is at a slightly more obtuse angle to the lin-
coronoid process. P. validus overlaps in size gual surface of P4. In modern felids, the pro-
with P. intrepidus and P. marshi in North tocone is closer to the lingual surface of the
America and P. quadridentatus in Europe, tooth, resulting in a narrower, compressed
and can be differentiated from these species P4. The P4 of F:AM 62128 has a parastyle
by the longer c-p3 distance in P. validus, on the anterior surface which does not differ
which exceeds the length of its ml, and by from the condition in P. lemanensis, other
its large, erect, and rectangular coronoid pro- species of Pseudaelurus, or modern felids.
CeSS. Both M1 are present with two roots. The
M1 are compressed in the a-p direction, but
DESCRIPTION AND COMPARISON are elongate transversely. Individual cusps
can be seen on the left M1. The buccal end
SKULL
of this tooth is formed by the parastyle wing.
The skull (figs. 2a, 2b) has been crushed, The nearly conjoined paracone and metacone
giving it a low and wide appearance. This are midway along the occlusal surface. The
compromises the integrity of some skull lingual margin of M1 is formed by a distinct
measurements, especially those of height and protocone. The parastyle of F:AM 626128
width. However, the upper dentition §s ur- M1 is as tall as the caudal surface of the
vived, and can be associated with the lower metastyle blade of P4 and appears to extend
teeth for only the second time in this genus. the carnassial blade. The left M1 of F:AM
The premaxillary bones possess a pair of 62128 compares favorably with the M1 in
fragmented [3, the I1—2 having been lost. A the P. lemanensis skull (MNHN1903-—20),
short and smooth diastema separates I3 from but is in sharp contrast to the rudimentary
C. The left C is separated from the skull, M1 of modern cats.
revealing a large, cavernous alveolus. The The skull has a high, thin sagittal crest
right C root is in situ; however its crown is which is joined by equally large nuchal crests
fractured and missing. These two C are more that unite at the occiput to form a protuber-
elliptical than round in cross-section and lack ance that projects to a point caudal to the
any longitudinal grooves. Posterior to the al- level of the occipital condyles. This area of
veolus for the left C are a preserved P1 and the skull is very similar to the type skull of
an alveolus for P2, both single-rooted. This P. lemanensis (MNHN1903-—20) from the
agrees with the presence of these teeth in the early Miocene of France. The basicranial
skull described by Stock (1934) from Tono- anatomy of F:AM 62128 is somewhat dis-
pah, although the presence of Pl was inter- torted, but much information endures (figs.
preted by Stock as being ‘‘a rather unusual 2b, 3). On the right side of the skull, much
feature’’. of the ossified bulla is lost, but the close ap-
The lengths of P3 and P4 in P. validus proximation of the mastoid and paroccipital
agree with the Tonopah skull but are smaller process to the bulla can be appreciated. The
than the P. quadridentatus type. These two paroccipital process displays a concavity on
teeth are aligned parallel to the maxillary its anterior surface where it cradled the pos-
axis and do not exhibit any overlap, oblique- terior surface of the tympanic bulla in typical
ness, or crowding as seen in some extant spe- aeluroid fashion (Cope, 1880; Hunt, 1987,
cies (Salles, 1992). The left P3 is wider and 1989, 1998; Werdelin, 1996; Wyss and
more robust when compared to P3 in the P. Flynn, 1993). Medially, the ridge formed by
lemanensis type skull (MNHN1903-—20). On the caudal entotympanic indenting the basi-
the right P4 (fig. 2b) can be seen a prominent occipital bone can be seen (fig. 3). On the
protocone that projects in an anterolingual lateral margins of the right bulla, just proxi-
direction. The morphology of P4 in F:AM mal to the mastoid process, is the external
62128 resembles the P4 in the P. guadriden- acoustic meatus surrounded by most of the
tatus maxillary fragment from la Grive- ectotympanic. This evidence of the antero-
Saint-Alban (Isére) (cast = AMNH 103396) lateral compartment of the bulla gives a
described by Gaillard (1899). In the P. le- strong signal as to the inflated size of the
manensis type specimen, the protocone of P4 caudal entotympanic. This is further evi-
2001 ROTHWELL: PSEUDAELURUS FROM NEW MEXICO 7
Fig. 2a. Dorsal view of F:AM 62128 skull. The morphology of the cranium resembles that of the
P. lemanensis skull (MNHN S.G. 3509a) from Europe and the early North American Ginn Quarry skull
(F:AM 61847) from Nebraska. sc = sagittal crest, nc = nuchal crest.
dence that this Nambé Pseudaelurus speci- region on the left side (figs. 2b, 3). Although
men possessed felid characters in the basi- the petrosal has been nudged in a caudal di-
cranial area (Hunt and Tedford, 1993). rection by the crushing, facets for its articu-
The petrosal bone and its promontorial lation with the ectotympanic can be seen on
process are preserved in the tympanic bulla its ventral surface. The grooving of the an-
8 AMERICAN MUSEUM NOVITATES NO. 3342
Fig. 2b. Ventral view of F:AM 62128 skull containing four upper premolars with P2 evidenced by
a single-rooted alveolus. PP = paroccipital process, P = petrosal part of the temporal bone, M = mastoid
bone.
terodorsal roof of the tympanic bulla, formed LOWER JAW
by the anterior invasion of the caudal ento-
tympanic to the level of the squamosal bone, The two rami of the lower jaw were ce-
can also be seen. The hypoglossal foramina mented together during early, cursory prep-
are visible bilaterally, within the depression aration (fig. 4). The size of this jaw is smaller
where the posterior lacerate foramina would than Leidy’s P. intrepidus type, and is more
be. similar in size to P. marshi (table 1). How-
2001 ROTHWELL: PSEUDAELURUS FROM NEW MEXICO 9
val Be
7
— + a! » inee ' “s OWew
» AS
+
=
Figs 3: Close view of basicranial area of skull. The x markers delineate the medial emargination of
the basioccipital caused by an enlarged caudal entotympanic. Also seen: the concavity of the paroccipital
process (PP) where it contacts the caudal surface of the expanded caudal entotympanic. T = ectotym-
panic, P = petrosal bone, PP = paroccipital process.
ever, it is obvious that this jaw was recon- (Matthew, 1918; Thorpe, 1922). Although
structed from multiple fragments, thus com- the number of roots of p2 varied in early Eu-
promising its measurements. The mandibles ropean specimens (Dehm, 1950), I am un-
are not slender, as described in Thorpe’s P. aware of any references, in North America,
marshi specimen, but rather display consid- to a double-rooted p2.
erable depth below the tooth row. The ventral The anterior mental foramina are located
surface of the lower jaw is slightly convex, beneath the posterior portion of the c-p3 di-
but the area immediately below the tooth row astema, and are best seen on the left ramus
is essentially straight. There is no ventral of this jaw (fig. 4). The left posterior mental
bulge in this location. foramen is underneath the posterior root of
Both of the large lower canines are pre- the third lower premolar. This location agrees
served. The inner surface of each is more well with Stock’s (1934) description of the
flattened than the outer surfaces. Their Tonopah specimens that he assigned to P. in-
crowns do not possess any grooves. In the trepidus and is in slight contrast to the more
space between c and p3 on both rami is an anterior location in the P. marshi specimen
alveolus for p2, located not at the midpoint (Thorpe, 1922). The coronoid process is wid-
in the diastema as in Leidy’s description er and more erect than in the Leidy or Marsh
(Leidy, 1869), but considerably closer to p3 types and is much taller than in P. lemanen-
(fig. 4). This single alveolus is also located sis. There is no terminal “‘hook”’ in the cor-
medial of the line of the tooth row, a char- onoid process as is seen in modern felids.
acter first described by Thorpe (1922) in the The deep masseteric fossa of F:AM 62128
P. marshi type specimen. This medial loca- extends anteriorly to a point just below the
tion of p2 is a consistent feature in the North talonid of ml. The F:AM 62128 condyle has
American specimens, and can be seen in the a mediolateral articular surface that is 20%
P. intrepidus and P. intrepidus sinclairi types longer than the Leidy type, and sits higher
10 AMERICAN MUSEUM NOVITATES NO. 3342
eatstanes!mieleclanliars
p2 alveolus
fami celi elalie
Fig. 4. Lateral (A) and oblique (B) views of lower jaw indicating dental characters of generic
significance: presence of p2, lack of pl and m2, and reduced metaconid and talonid on ml.
by an equal amount above the angular pro- and p4 is present in specimens of Proailurus
cess. lemanensis and persists, with little change, in
The p3 and p4 are damaged by crushing. modern felids. The left m1 is well preserved
However, their size is recoverable (table 1). and demonstrates a much-reduced metaconid
The posterior accessory cusps of the left p3 that blends smoothly into an abridged tal-
and both p4 can be seen on the posterior sur- onid. The protoconid is taller than the para-
face of the primary cusps. These posterior conid and the carnassial notch is open and
accessory cusps are separate from a well-de- deep. This description of the carnassial
fined posterior cingulum. This condition of a would fit the diagnosis of any of the previ-
conspicuous posterior accessory cusp on p3 ously described large Pseudaelurus species
