Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The Lin-

nean Society of London, 2005? 2005

1442
145166
Original Article
EARLY OLIGOCENE CAINOTHERIIDSC. BLONDEL

Zoological Journal of the Linnean Society, 2005, 144, 145–166. With 5 figures

New data on the Cainotheriidae (Mammalia, Artiodactyla)

from the early Oligocene of south-western France
CECILE BLONDEL*
Laboratoire de Géobiologie, Biochronologie et Paléontologie Humaine, UMR 6046 CNRS, Faculté des
Sciences Fondamentales et Appliquées, Université de Poitiers, 40, avenue du Recteur Pineau F-86022
Poitiers Cedex, France

Received July 2003; accepted for publication March 2005

The Cainotheriidae are small artiodactyls that suddenly appeared in the late Eocene of western Europe. A revision
of early Oligocene cainotheriid lineages is proposed on the basis of newly dated material from the Quercy Phospho-
rites (south-western France). A significant diversification of the group occurred at the end of the Eocene. Few species
seem to have persisted through the Eocene/Oligocene boundary, but the Cainotheriidae subsequently diversified rap-
idly during the early Oligocene. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society,
2005, 144, 145-166.

ADDITIONAL KEYWORDS: Cainotheriinae – Oxacroninae – Palaeogene – Quercy Phosphorites – systematics.

INTRODUCTION owing to the uniformity of dental characters (Möltzer,

1975). The teeth of cainotheriids are brachydont; the
The Cainotheriidae are small artiodactyls, the largest
upper molars bear two anterior and three posterior
species reaching only the size of the extant Tragulus
cusps, a dental pattern that Stehlin (1910) used to
javanicus, the lesser Asian mouse-deer. They are
separate the Cainotheridia, the Hypoconifera and
known only from western Europe (Switzerland, Ger-
the Euartiodactyla among the artiodactyls, while the
many, Spain and southern England) and their
lower molars possess an additional mesiolingual cusp
remains are generally very abundant, particularly in
(= mesoconid; Sudre, 1977).
the Quercy Phosphorites, karstic localities in south-
Hürzeler (1936) provided the first systematic
western France (Schmidt-Kittler, 1987).
account of this family, based on material (the ‘Old
Although many ungulates endemic to Europe
Quercy collections’) which was largely undated. This
became extinct at the end of the Eocene (Sudre & Leg-
was later developed by Berger (1959), who proposed a
endre, 1992; Blondel, 1996, 2001), the Cainotheriidae
phylogenetic distribution of the group based on his
persisted beyond the Eocene/Oligocene boundary.
study of the Cainotheriidae from Gaimersheim, Ger-
They first appear in the fossil record at the end of
many (MP28) and gave details of the chronological
the Eocene (MP19, Fig. 1), flourished during the
ranges of the different cainotheriid genera.
Oligocene and appear to have become extinct in the
The Cainotheriidae contains two subfamilies: Oxa-
middle Miocene (Berger, 1959; Heizmann, 1983,
croninae Hürzeler, 1936 (genera Oxacron Filhol, 1884
1999). Distinctive features of the skeleton include the
and Paroxacron Hürzeler, 1936) and Cainotheriinae
morphology of the distal humerus, the well developed
Camp & VanderHoof, 1940 (genera Plesiomeryx Ger-
fibula diaphysis and the presence of huge auditory
vais, 1873, Caenomeryx Hürzeler, 1936 and Cainothe-
bullae.
rium Bravard, 1828). The Oxacroninae appear to
Few studies have been devoted to this family, prin-
occur only in the Eocene, the Cainotheriinae in the
cipally because identification of species is difficult,
Oligocene. Cainotherium is well represented by speci-
mens from late Oligocene localities in Auvergne, cen-
*E-mail: cecile.blondel@univ-poitiers.fr tral France. Our knowledge of Oxacron, Plesiomeryx

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166 145
146 C. BLONDEL

and Caenomeryx is based on material collected from
the Quercy Phosphorites during the late nineteenth
century. Plesiomeryx, represented by P. cadurcensis
Gervais, 1873, and Caenomeryx, represented by
C. procommunis (Filhol, 1877), have been described
from undated specimens collected in karstic fissures in
Quercy.
Fieldwork in the Quercy Phosphorites, organized
by the Universities of Montpellier, Poitiers and Paris,
has been carried out since 1965. It has led to the dis-
covery of many new cainotheriid remains. Analysis of
the Quercy mammalian fauna, together with that of
southern Germany and some Palaeogene mammalian
lineages, shows that these lineages can be arranged
chronologically according to a reference level scale
(Mammalian Palaeogene, henceforth MP) defined at
the Mainz Symposium and revised at BiochroM in
1997 (Schmidt-Kittler, 1987; BiochroM, 1997; Fig. 1).
Using this newly dated material, we can now define
some episodes in the history of this family more pre-
cisely, in particular around the time of the Eocene/
Oligocene transition, which corresponds to the
Grande Coupure as originally defined by Stehlin
(1910).
Based on material collected in the Escamps locality
of Quercy (MP19), Legendre (1980) showed that the
Oxacroninae were more diverse than had been previ-
ously recognized in the late Eocene. A study in prep-
aration by Legendre & Sudre on the Cainotheriidae
from late Eocene localities in France supports the idea
of a rapid diversification of this group. At least five lin-
eages, including at least one member of the Cain-
otheriinae, were present in this period (Legendre &
Sudre, pers. comm., 1997).
In order to assess the relationships of late Eocene
and early Oligocene Cainotheriidae, it is necessary to
study the newly dated material from the Quercy Phos-
phorites, comparing the taxonomic status of these

Mya Marine Stages MP Reference Levels Quercy Localities

23 C 30 Coderet
O H 29 Rickenbach
A
L T 28 Pech du Fraysse
I T
27 Boningen
I
G A 26 Mas de Pauffié
28.5
O N
25 Le Garouillas
C R
U 24 Heimersheim
E P
N E 23 Itardies {Itardies - Roqueprune 2
Pech Crabit - Mounayne
STAMPIAN L Mas de Got A
E I 22 Villebramar {La Plante 2
A
N
21 Soumailles {Ravet
Aubrelong1
33.7 20 St-Capraise d'Eymet
19 Escamps Escamps
PRIABONIAN 18 LaDébruge
E 17b Perrière
O 17a Fons4
38 C 16 Robiac
E 15 La Livinière 2
N BARTONIAN
14 Egerkingen a et b
E
41
13 Geiseltal Obere Mittelkohle
LUTETIAN
12 Geiseltal Untere Mittelkohle

11 Geiseltal Unterkohle
48.5
10 Grauves
YPRESIAN
8-9 Avenay
7 Dormaal
55
P 6 Cernay
THANETIAN
A 1-5 Hainin
65 L

Figure 1. Subdivision of the European continental Palaeogene, based on reference levels proposed at the Mainz Sympo-
sium (Schmidt-Kittler, 1987) and revised at Biochrom ¢97 (BiochroM, 1997). The Quercy localities are correlated with the
reference levels. Numerical ages follow Odin (1994).

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166

early Oligocene forms with those described from the
undated old collections.
MATERIAL AND METHODS
This study is based on dental material from the fol-
lowing karstic localities of the Quercy Phosphorites:
Aubrelong, Ravet (MP21); Mas de Got, La Plante 2
(MP22); Pech Crabit, Mounayne, Roqueprune 2,
Itardies (MP23). Only specimens with associated
molars and premolars were considered. The difficulty
of distinguishing the rank of several isolated molars
makes their attribution to a particular species uncer-
tain or doubtful. Recognition of different species is
based on variation in size and on morphological fea-
tures of the second and third premolars in relation to
types defined by Hürzeler (1936) and Berger (1959).
This is because the characters of canines, fourth pre-
molars, molars are not diagnostic to define species
within this family.
Measurements (in millimetres) were compiled using
a digital table (Jandel Scientific) and Sigma Scan v.
3.9. Length L is the largest mesiodistal dimension,
width l the largest transverse dimension. All speci-
mens are housed in the collections of Montpellier II
University.
The normality of the distribution of dental measure-
ments (length of lower premolars) was checked by the
non-parametric Kolmogorov-Smirnov (K-S) and Sha-
piro-Wilk (W) tests, when the number of specimens
was adequate. Variability was estimated by the vari-
ation (V) or Pearson coefficient (see tables). Dental
measurements of mammalian populations show a
variation coefficient between 4 and 10; this can, how-
ever, vary according to the mammalian order or type of
tooth (Simpson, Roe & Lewontin, 1960; Vianey-Liaud
& Legendre, 1986).
Institutional and karstic locality abbreviations
ABL1 Aubrelong 1
BMNH British Natural History Museum, London,
UK
BSP Bayerische Staatssammlung für Paläont-
ologie und historische Geologie, München,
Germany
ITD Itardies
MGT Mas de Got
MNHNP Museum National d’Histoire Naturelle,
Paris
MOU Mounayne
NMB Naturhistorisches Museum Basel,
Switzerland
PCT Pech Crabit
PL2 La Plante 2
RAV Ravet
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
EARLY OLIGOCENE CAINOTHERIIDS 147
ROQ2 Roqueprune 2
UMII Montpellier II University, France
Anatomical abbreviations
MX/ upper molar
M/X lower molar
PX/ upper premolar
P/X lower premolar
SYSTEMATIC PALAEONTOLOGY
FAMILY CAINOTHERIIDAE CAMP &
VANDERHOOF, 1940
SUBFAMILY OXACRONINAE HÜRZELER, 1936
GENUS PAROXACRON HÜRZELER 1936
Type species: Paroxacron valdense (Stehlin, 1906)
from Mormont Entreroches, Switzerland (MP19). A
left mandibular fragment with I/1or I/2 and P/2-M/3
(NMB Mt. 232; Stehlin, 1906).
Diagnosis: P1/, P/1, P2/ and P/2 are elongate and
premolariform; no or only short diastema between P1/
-P2/, P/1-P/2 and P/2-P/3; the anterior lobe of P3/ is
elongate, upper molars with mesostyle notch (Hür-
zeler, 1936).
PAROXACRON SP. (FIG. 2A-D)
Material: Escamps (Legendre, 1980). Aubrelong 1: a
right maxillary fragment with P2/-P4/ (ABL1 1455), a
left maxillary fragment with P3/-M1/ (ABL1 1454);
two isolated right P3/ (ABL1 1558, ABL1 1565), an
isolated left P3/ (ABL1 1557), two left mandibular
fragments with P/2-P/3 (ABL1 1479, ABL1 1485); an
isolated left P/3 (ABL1 1792). Ravet: a left maxillary
fragment with P4/-M3/ (RAV 1055); five isolated upper
teeth, a right P2/ (RAV 595), three left P3/ (RAV 599,
RAV 602, RAV 604), a right P3/ (RAV 771); three left
mandibular fragments, one with P/2-M/1 (RAV 1022),
one with P/1-P/3 (RAV 1019), and one with P/3-P/4
(RAV 1023); an isolated left P/2 (RAV 595), and an iso-
lated right P/3 (RAV 902). La Plante 2: two left max-
illary fragments, one with P2/-M1/ (PL2 1202) and one
with P2/-P3/ (PL2 1203).
Measurements: See Table 1.
Description
Upper dentition: The premolariform P2/ (Fig. 2A) is
more elongate than that of P. cadurcensis. A cingulum
occurs on the lingual side of the crown. The triangular
P3/ (Fig. 2B) bears a very elongate anterior lobe sim-
ilar to that of the P3/ of P. cadurcensis, and the para-
style is well pronounced. The lingual cusp is developed
distally. M1/ possesses a lingual paraconule which
gives this molar a quadrangular lingual outline. Hür-
148 C. BLONDEL

Table 1. Measurements and statistical data for teeth of Paroxacron sp. N, number of specimens; L, length; l, width; LV,
limit values; M ± SE, Mean ± standard error; s, standard deviation, V, variation coefficient

N L/l LV M ± SE s V

ABL1
P2/ 1 L 3.15
1 l 1.4
P3/ 5 L 2.99–3.36 3.19 ± 0.08 0.15 4.63
5 l 2.48–2.88 2.66 ± 0.08 0.15 5.65
P4/ 2 L 2.39–2.50 2.45
2 l 3.22–3.28 3.25
M1/ 1 L 3.23
1 l 3.95
P2/-P4/ 1 L 8.4
P3/-P4/ 2 L 5.5–5.8 5.7
P/2 2 L 2.74–3.08 2.91
2 l 0.97–1 0.99
P/3 3 L 3.28–3.42 3.37 ± 0.06 0.08 2.31
3 l 1.26–1.37 1.32 ± 0.04 0.06 4.29
RAV
P2/ 1 L 3.5
1 l 1.3
P3/ 4 L 2.74–3.16 2.97 ± 0.1 0.17 5.85
4 l 2.33–2.80 2.56 ± 0.12 0.2 7.94
P4/ 1 L 2.21
1 l 2.55
M1/ 1 L 2.8
1 l 3.56
M2/ 1 L 2.95
1 l 3.87
M3/ 1 L 2.84
1 l 3.75
M1/-M3/ 1 L 8.8
P/1 1 L 2.5
1 l 0.87
P/2 3 L 3.45–3.48 3.47 ± 0.01 0.02 0.44
3 l 0.95–1.20 1.05 ± 0.09 0.13 12.4
P/3 4 L 3.37–4 3.76 ± 0.16 0.28 7.36
4 l 1.31–1.73 1.52 ± 0.10 0.18 12
P/4 2 L 3.43 3.43
2 l 1.83–2.05 1.94
M/1 1 L 3.21
1 l 2.33
P/2-P/4 1 L 11
P/3-P/4 2 L 7.4–7.5 7.45
PL2
P2/ 2 L 2.79–3.19 2.99
2 l 1.22–1.32 1.27
P3/ 2 L 2.7–3.15 2.92
2 l 2.25–2.65 2.45
P4/ 1 L 2.38
1 l 3.07
M1/ 1 L 2.81
1 l 4.07

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
 EARLY OLIGOCENE CAINOTHERIIDS 149

A B

C D

Figure 2. Paroxacron sp. A-D, occlusal view. A, right P2/-P4/ (ABL1 1455). B, left P3/-M1/ (ABL1 1454). C, left P/1-P/3
(RAV 1019). D, left P/2-M/1 (RAV 1022). P. cf. bergeri. E, left P1/-M3/, MGT 3210, occlusal view. Scale bar = 1 mm.

zeler (1936) indicates that the mesostyle of Paroxacron
is split, although this feature, which only can be
observed on unworn teeth, is variable.
Lower dentition: The double-rooted P/1 is premolari-
form, elongate and narrow on the mandible from
Ravet (Fig. 2C). This premolar possesses a sharp
median cusp and a faint paraconid. The double-rooted
and elongate P/2 is separated from P/1 by a diastema
which varies from 0.8 to 1.4 mm in length. The para-
conid is lingual and more developed than on P/1. A
diastema varying from 0.5 to 1.1 mm long separates P/
2 from P/3. The elongate P/3 is wider distally than P/
2, its paraconid is more pronounced than that of P/2
and a crest extends from a median cusp to the disto-
lingual part of the tooth. The stocky P/4 is less elon-
gate than the first premolars. The paraconid is
strongly pronounced and the protoconid and the meta-
conid are also developed. The latter are separated by a
notch, although the metaconid is situated distally
(Fig. 2D).
Comparison: The presence of a premolariform P/1, the
very elongate premolars and the occurrence of a
diastema between P/1-P/2 and P/2-P/3 enables the
form described above to be attributed to Paroxacron,
although the dimensions of this specimen are more
comparable to those of Plesiomeryx cadurcensis
(Tables 1–3). The Paroxacron material from Ravet,
Aubrelong (MP21) and from La Plante 2 (MP22) dif-
fers from that from Escamps (Legendre, 1980) in that
the P/2-P/4 row is a little shorter. Material is rare,
however, so it is difficult to determine whether these
differences are simply due to variation or whether
they represent a trend toward premolar reduction in
the evolution of this lineage. In any case, it is the first
record of a small species of Paroxacron that has been
found in early Oligocene levels.
PAROXACRON BERGERI HEISSIG, 1978
1959 cf. Cainotherium? elongatum Filhol (? Berger);
Berger p. 45, pl. 4, fig. 5, pl. 5 fig. 3.
1959 Cainotherium? n. sp.; Berger, p. 47, pl. 5, figs 6-
10.
Holotype. A right mandibular fragment I/3-M/3 (BSP
1879XV201), locality of Mouillac in Quercy Phospho-
rites. Age and precise locality are unknown (Old
Quercy collections).
Diagnosis. Species of the genus Paroxacron without
diastema between P/2-P/3. The edge of the mandible is
more strongly bent than in P. valdense. The angulus
mandibulae is short, high and angular. It is limited by
a short but strong incisura vasorum anteriorly. P/3

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
150 C. BLONDEL

and P/4 have a more complete talonid and the internal Table 2. Measurements and statistical data for teeth of
wall of the lower molars is higher than in P. valdense Paroxacron cf. bergeri. N, number of specimens; L, length; l,
(Heissig, 1978). width; LV, limit values; M ± SE, Mean ± standard error

N L/l LV M ± SE
PAROXACRON CF. BERGERI HEISSIG, 1978
Referred material: Mas de Got: a left maxillary frag- MGT
ment with P1/-M3/ (MGT 3210), a right maxillary P1/ 1 L 3.37
fragment with P3/-M3/ (MGT 3211). Pech Crabit: a 1 l 1.14
right maxillary fragment with P3/-M2/ (PCT 412). P2/ 1 L 4.17
1 l 1.54
Roqueprune 2: a left maxillary fragment with P2/-M1/
P3/ 2 L 3.89–3.94 3.92
(ROQ2 348). Mounayne: a left maxillary fragment
2 l 3.10–3.30 3.2
with P3/-P4/ (MOU 186).
P4/ 2 L 2.92–3.15 3.04
Measurements: See Table 2. 2 l 3.7–3.76 3.73
M1/ 2 L 4.22–4.4 4.31
Description: The upper premolars are elongate. The 2 l 4.78–5.11 4.95
P1/, borne by the maxillary (Fig. 2E), is premolari- M2/ 2 L 4.29–4.59 4.44
form. The P2/ bears a small median cusp but no lin- 2 l 5.52 5.52
gual cusp; the latter is replaced by a cingulum. The M3/ 2 L 4.2–4.45 4.33
well developed anterior lobe of P3/ possesses salient 2 l 5.07–5.5 5.29
styles on the ectoloph and a weakly developed distol- P3/-P4/ 2 L 6.5–7 6.75
ingual cusp. M1/-M3/ 2 L 12.4–12.5 12.45
Comparison: Some specimens from Mas de Got MOU
(MP22), Pech Crabit, Roqueprune 2 and Mounayne P3/ 1 L 3.48
(MP23) are of a size comparable to those referred to 1 l 3.18
P. huerzeleri (Berger, 1959). They differ from this spe- P4/ 1 L 3.1
cies, however, in P2/, which has a small median cusp 1 l 3.75
and a lingual cingulum and in P3/, which has a more P3/-P4/ 1 L 7.2
developed anterior lobe and a weaker distolingual ROQ2
cusp than those of P. huerzeleri. All these features are P2/ 1 L 4.59
similar to those of species belonging to Paroxacron. 1 l 2.04
Heissig (1978) attributed to Paroxacron (under the P3/ 1 L 4.13
name P. bergeri) a fragment of a mandible from 1 l 4.2
Quercy and a specimen from Bernloch, Switzerland P4/ 1 L 3.5
(MP23). This poorly known specimen, which was orig- 1 l 4.75
inally attributed to Cainotherium by Berger (1959), is M1/ 1 L 4.15
1 l 5.56
represented by a few isolated teeth. Among these,
P2/-P4/ 1 L 12.6
some examples of P3/ are similar to those described in
P3/-P4/ 1 L 8.7
this study. According to Heissig (1978), the P/2-P/3
diastema, typical of Paroxacron, is absent in P. bergeri. PCT
It was not possible, however, to distinguish elements P3/ 1 L 4.21
of the lower dentition in the material from the Quercy 1 l 3.46
localities. Thus it is with some reservations that this P4/ 1 L 3.29
material is attributed to P. bergeri. 1 l 4.8
M1/ 1 L 4.16
1 l 6.29
SUBFAMILY CAINOTHERIINAE CAMP & M2/ 1 L 4.8
VANDERHOOF, 1940 1 l 6.8
GENUS PLESIOMERYX GERVAIS, 1873 P3/-P4/ 1 L 7.7
PLESIOMERYX CADURCENSIS GERVAIS, 1873
Holotype: Right mandibular P/2-M/3 (MNHN-QU
1772). Mouillac locality, Quercy Phosphorites. Age and
C and between P/1 and P/2. P1/, P2/, P/2 and P/3 are
precise locality are unknown (Old Quercy collections).
short, P3/ with a strong lingual cone, P2/ with a weak
Diagnosis: Protocone and protoconule present (stage lingual cone. The premaxilla reaches the frontal. The
B, Hürzeler, 1936: 8). Long diastema between P1/ and nasal is long, narrow and straight. The ethmoidal slit

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166

is falciform. The choanae are large. The horizontal
portion of the lower jaw is slender (Hürzeler, 1936).
Referred material: Aubrelong 1: left maxillary frag-
ment with P2/-M3/ (ABL1 1452), right maxillary frag-
ment with P3/-M3/ (ABL1 1453); isolated left P3/
(ABL1 1559); 12 mandibular fragments, one with left
P/1-P/2 (ABL1 1457), one with left P/2-M/2 (ABL1
1458), one with right P/2-M/1 (ABL1 1459), one with
left P/2-P/4 (ABL1 1461), one with right P/2-P/3 (ABL1
1463), one with right P/3-M/2 (ABL1 1465), one with
left P/2-P/3 (ABL1 1487), one with broken right P/2-M/
2 (ABL1 1492), one with broken right P/2-M/1 (ABL1
1497), one with left P/1-P/3 (ABL1 1780), one with
right P/3-P/4 (ABL1 1786), one with right P/2-P/4
(ABL1 1787); three isolated left P/2 (ABL1 1488, ABL1
1492¢, ABL1 1668), three isolated right P/3 (ABL1
1462, ABL1 1777, ABL1 1794). Ravet: two isolated
right P2/ (RAV 597, RAV 973), an isolated left P2/
(RAV 598), an isolated left P3/ (RAV 840); five man-
dibular fragments: with left P/2-M/2 (RAV 1017), one
with left P/1-M/1 (RAV 1024), one with right P/3-M/1
(RAV 1025), two with left P/2-M/2 (RAV 1026, RAV
1027); an isolated left P/2 (RAV 853), three isolated
right P/2 (RAV 739, RAV 740, RAV 892), four isolated
right P/3 (RAV 732, RAV 889, RAV 895, RAV 1037),
four isolated left P/3 (RAV 849, RAV 887, RAV 898,
RAV 901). Mas de Got: 13 mandibular fragments, one
with right P/1-M/3 (MGT 104), one with left P/2-P/4
(MGT 129), one with left P/2-M/2 (MGT 135), one with
right P/2-P/4 (MGT 151), one with left P/3-M/3 (MGT
154), one with left P/2-P/4 (MGT 190), one with left P/
2-P/3 (MGT 195), one with right P/3-M/1 (MGT 3185),
one with right P/3-P/4 (MGT 3196), one with right P/1-
P/2 (MGT 3199), one with left P/2-P/3 (MGT 3200), one
with right P/2-P/3 (MGT 3202), one with right P/2-P/4
(MGT 3203). La Plante 2: five maxillary fragments,
two with left P3/-M2/ (PL2 1180, PL2 1191), one with
left P1/-P2/ (PL2 1201), two with left P2/-P3/ (PL2
1211, PL2 1213), 10 mandibular fragments, one with
right P/2-M/3 (PL2 1168), three with left P/3-M/3 (PL2
1169, PL2 1242, PL2 1275), one with right P/4-M/3
(PL2 1173), one with left P/2-M/3 (PL2 1174), two with
left P/3-M/3 (PL2 1175, PL2 1176), one with left P/2-P/
4 (PL2 1177), one with right P/2-P/3 (PL2 1250). Pech
Crabit: three right mandibular fragments with P/3-M/
1 (PCT 366) and one with P/2-P/3 (PCT 367, PCT 385).
Mounayne: two left mandibular fragments, one with
P/2-M/2 (MOU 209) and one with P/2-P/4 (MOU 148),
a right maxillary fragment with P2/-P3/ (MOU 187).
Itardies: a right mandibular fragment with P/2-M/3
(ITD 941).
Measurements: See Table 3. For P. cadurcensis, the
Kolmogorov-Smirnov and Shapiro-Wilk tests confirm
normality for L P/3 from Aubrelong 1 (N = 12, K–S,
d = 0.163, P > 0.20, W = 0.955, P = 0.721), Ravet
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
EARLY OLIGOCENE CAINOTHERIIDS 151
(N = 12, K–S, d = 0.135, P > 0.20, W = 0.949,
P = 0.629), La Plante 2 (N = 9, K–S, d = 0.244, P > 0.20,
W = 0.903, P = 0.269) and Mas de Got (N = 12, K–S,
d = 0.104, P > 0.20, W = 0.968, P = 0.887).
The weakness of variation coefficients (4 < V < 10)
confirms the homogeneity of samples of P. cadurcensis
from localities of the same reference level. For the
length of P/3 from Aubrelong 1 and Ravet (MP21):
N = 24, M = 3.40, s = 0.23, K–S, d = 0.897, P > 0.20,
W = 0.96, P = 0.438, V = 6.87. P/3 from Mas de Got and
La Plante 2 (MP22): N = 21, M = 3.02, s = 0.21, K–S,
d = 0.084, P > 0.20, W = 0.97, P = 0.672, V = 6.93. P/3
from Pech Crabit, Mounayne, Itardies (MP23): N = 6,
M = 3.21, s = 0.16, K–S, d = 0.19, P > 0.20, W = 0.97,
P = 0.897, V = 5.05.
Description
Upper dentition: A maxillary fragment from Aubre-
long 1 (Fig. 3A) bears a premolariform P1/ that is
short and biradicular. The elongate P2/ has a weakly
developed distolingual tubercle. The triangular P3/
has a well developed mediolingual cusp from which
two crests extend to the prominently strong parastyle
and metastyle. The labial and lingual cusps of P4/ are
median, the parastyle and the metastyle are strong.
The same characters are evident in specimens (PL2
1180) from La Plante 2 (MP22).
Lower dentition: On the mandibular fragment (MGT
104), the single-rooted P/1 is caniniform and the
anterior stylid is not clearly marked (Fig. 3B). The
diastema that separates P/1 from P/2 varies between
0.5 and 1.9 mm in length. The double-rooted P/2 bears
a well developed paraconid, while the main cusp is
directed backward and reaches the distolingual angle.
On P/3, the paraconid is situated lingually and P/4 has
a well developed style. The protoconid and the meta-
conid form large tubercles separated by a deep notch
(e.g. ABL1 1458). There are no differences between the
material from Aubrelong 1 and that from La Plante 2
or Pech Crabit.
Comparison: The presence of a single-rooted canini-
form P/1, a diastema between this premolar and P/2,
the presence of a well developed mediolingual cusp
and a short anterior lobe on P/3 indicate that this
small cainotherine from Aubrelong 1 (MP21) belongs
to Plesiomeryx. In specimens from the localities of
Aubrelong 1, La Plante 2, and Pech Crabit, the dimen-
sions of teeth in the lower dental rows (Table 3), and
the features of the lower premolars agree with the
description of P. cadurcensis. The same is also true for
the upper tooth rows, which correspond in their mor-
phology and dimensions to material of P. cadurcensis
described by Hürzeler (1936) and Berger (1959). The
general similarity of the morphological patterns and
152 C. BLONDEL

Figure 3. Plesiomeryx cadurcensis. A & B, occlusal view. A, left P2/-M3/ (ABL1 1452). B, right C-M/3 (MGT 104).
P. huerzeleri. C & D, occlusal view. C, right P2/-M3/ (MGT 3209). D, left P/2-M/3 (MGT 3187). Scale bar = 1 mm.

dimensions of the material reflect a degree of homoge-
neity despite the different ages of the studied
material.
PLESIOMERYX HUERZELERI BERGER, 1959
1937 Caenotherium gracile Pomel; Dehm, 1937: 352.
Holotype: Skull BSP1952 II 1149. Gaimersheim fis-
sure filling (Germany) (MP28).
Diagnosis: Species of Plesiomeryx that appears similar
to P. cadurcensis, but is larger. The fronto-nasal
suture is short. The lingual cingulum of P/3 is missing.
The lingual and labial cones of P/4 reach the same
level (Berger, 1959).
Referred material: Mas de Got: four maxillary frag-
ments, one with right P2/-M3/ (MGT 3209), one with
right P3/-M2/ (MGT 3208), one with left P1/-M1/ (MGT
3212), and one with left P4/-M3/ (MGT 3213); 14 man-
dibular fragments, two with left P/2-M/3 (MGT 131,
MGT 3187), three with left P/3-M/3 (MGT 103, MGT
115, MGT 134), two with left P/2-M/1 (MGT 105, MGT
138), two with right P/3-M/2 (MGT 107, MGT 3183),
one with left P/1-P/3 (MGT 124), two with right P/3-P/
4 (MGT 191, MGT 3177), one with right P/3-M/1 (MGT
3193), one with left P/2-P/3 (MGT 195). La Plante 2:
three mandibular fragments, two with right P/3-M/1
(PL2 1171, PL2 1221), one with left P/2-P/3 (PL2
1216). Pech Crabit: four left maxillary fragments P1/-
P2/ (PCT 382, PCT 391, PCT 431, PCT 1000); 11 man-
dibular fragments, two with left P/2-P/3 (PCT 348),

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 EARLY OLIGOCENE CAINOTHERIIDS 153

Table 3. Measurements and statistical data for teeth of Plesiomeryx cadurcensis. N, number of specimens; L, length; l,
width; LV, limit values; M ± SE, Mean ± standard error; s, standard deviation, V, variation coefficient

N L/l LV M ± SE s V

ABL1
P2/ 1 L 3.47
1 l 1.62
P3/ 3 L 3.00–3.27 3.11 ± 0.1 0.14 4.56
3 l 2.47–2.78 2.64 ± 0.1 0.16 5.95
P4/ 2 L 2.71–2.87 2.79
2 l 3.48–3.49 3.47
M1/ 2 L 3.18–3.24 3.21
2 l 4.15–4.22 4.19
M2/ 2 L 3.47–3.48 3.46
2 l 4.56–4.57 4.57
M3/ 2 L 2.95–3.15 3.05
2 l 4.40–4.48 4.44
P2/-P4/ 1 L 9.4
P3/-P4/ 2 L 5.6–5.8 5.7
M1/-M3/ 2 L 10.2–10.3 10.3
P2/-M3/ 1 L 18.9
P/1 2 L 2–2.3 2.15
2 l 1.08–1.3 1.19
P/2 12 L 2.25–3.45 2.95 ± 0.09 0.31 10.4
12 l 1.02–1.36 1.13 ± 0.03 0.1 9.11
P/3 12 L 3.07–3.63 3.37 ± 0.05 0.16 4.63
12 l 1.29–1.78 1.53 ± 0.04 0.14 9.17
P/4 7 L 2.91–3.48 3.26 ± 0.08 0.2 6.04
7 l 1.86–2.29 2.07 ± 0.07 0.16 7.65
M/1 5 L 2.85–3.38 3.15 ± 0.1 0.2 6.34
5 l 1.86–2.69 2.52 ± 0.09 0.18 7.27
M/2 3 L 3.25–3.64 3.44 ± 0.14 0.2 5.66
3 l 2.74–2.9 2.82 ± 0.06 0.08 2.84
P/2-P/4 3 L 9–9.7 9.43 ± 0.27 0.38 4.61
P/3-P/4 4 L 6.4–6.9 6.73 ± 0.14 0.24 3.51
M/1-M/3 1 L 11.2
RAV
P2/ 3 L 3.19–3.3 3.25 ± 0.04 0.06 1.7
3 l 1.5–1.65 1.59 ± 0.06 0.08 4.9
P3/ 1 L 3.5
1 l 2.92
P/1 1 L 1.98
1 l 0.98
P/2 8 L 2.74–3.15 2.88 ± 0.06 0.17 5.85
8 l 1–1.33 1.13 ± 0.04 0.11 10
P/3 12 L 3.03–3.97 3.44 ± 0.09 0.29 8.57
12 l 1.30–1.77 1.55 ± 0.05 0.15 9.79
P/4 5 L 2.62–3.22 3 ± 0.12 0.23 7.61
5 l 1.77–2.10 1.93 ± 0.09 0.17 8.93
M/1 5 L 2.65–3.11 2.91 ± 0.09 0.17 5.81
5 l 2.14–2.61 2.26 ± 0.1 0.2 9
M/2 3 L 3.29–3.51 3.44 ± 0.09 0.13 3.7
3 l 2.58–2.85 2.67 ± 0.14 0.2 5.84
P/2-P/4 3 L 8.2–9.2 8.7 ± 0.35 0.5 5.75
P/3-P/4 4 L 5.6–6.4 6.1 ± 0.21 0.36 5.92

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154 C. BLONDEL

Table 3. Continued

N L/l LV M ± SE s V

PL2
P1/ 1 L 2.48
1 l 0.93
P2/ 3 L 2.62–2.98 2.85 ± 0.13 0.19 6.72
3 l 1.46–1.81 1.62 ± 0.13 0.18 11
P3/ 4 L 2.62–3.02 2.82 ± 0.1 0.17 5.88
4 l 2.66–2.85 2.74 ± 0.05 0.08 2.93
P4/ 2 L 2.27–2.45 2.36
2 l 2.88–3.3 3.31
M1/ 2 L 2.89–2.99 2.94
2 l 3.87–4.14 4.01
M2/ 2 L 3.24–3.45 3.36
2 l 4.17–4.48 4.33
P3/-P4/ 3 L 5.1–5.7 5.4 ± 0.22 0.31 5.69
M1/-M3/ 1 L 9.7
P/2 5 L 2.47–3.04 2.68 ± 0.11 0.21 8
5 l 1.15–1.26 1.19 ± 0.02 0.04 3.48
P/3 9 L 2.49–3.18 2.85 ± 0.06 0.18 6.3
9 l 1.37–2.02 1.62 ± 0.07 0.19 11.4
P/4 9 L 2.72–3.1 2.89 ± 0.04 0.12 4.32
9 l 1.86–2.16 2.01 ± 0.04 0.12 6.19
M/1 8 L 2.53–2.98 2.81 ± 0.06 0.15 5.2
8 l 2.3–2.76 2.48 ± 0.06 0.17 6.97
M/2 8 L 2.78–3.18 3.04 ± 0.05 0.13 4.43
8 l 2.25–3.26 2.75 ± 0.11 0.3 11
M/3 8 L 3.8–4.62 4.2 ± 0.12 0.33 7.92
8 l 2.06–2.81 2.36 ± 0.1 0.26 10.9
P/2-P/4 4 L 8.3–9.2 8.7 ± 0.21 0.37 4.3
P/2-M/3 3 L 19–20.5 19.7 ± 0.54 0.76 3.88
P/3-P/4 8 L 5.2–6.4 5.83 ± 0.15 0.41 7.03
M/1-M/3 8 L 10–11.1 10.7 ± 0.13 0.35 3.26
MGT
P/1 2 L 2.07–2.23 2.15
2 l 0.97–1.13 1.05
P/2 10 L 2.34–2.99 2.76 ± 0.06 0.18 6.41
10 l 1.05–1.42 1.23 ± 0.05 0.14 11
P/3 12 L 2.96–3.36 3.15 ± 0.04 0.13 4.02
12 l 1.3–1.92 1.6 ± 0.06 0.19 11.8
P/4 9 L 2.82–3.57 3.13 ± 0.08 0.23 7.4
9 l 1.92–2.19 2.02 ± 0.05 0.14 6.87
M/1 4 L 2.93–3.36 3.2 ± 0.11 0.19 5.87
4 l 2.27–2.77 2.61 ± 0.13 0.23 8.85
M/2 3 L 3.25–3.85 3.56 ± 0.35 0.5 8.45
3 l 2.88–2.96 2.9 ± 0.04 0.05 1.69
M/3 2 L 4.08–4.10 4.09
2 l 2.32–2.34 2.33
P/2-P/4 3 L 8.3–8.4 8.37 ± 0.04 0.06 0.69
P/3-P/4 8 L 5.4–6.1 5.7 ± 0.09 0.25 4.45
M/1-M/3 2 L 10.8–11.4 11.1
MOU N VL M ± sm s V
P2/ 1 L 3.34
l 1.64
P3/ 1 L 3.72
l 2.74

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 EARLY OLIGOCENE CAINOTHERIIDS 155

Table 3. Continued

N L/l LV M ± SE s V

P/2 2 L 2.89–2.99 2.94

2 l 1.35–1.37 1.36
P/3 2 L 2.96–3.41 3.19
2 l 1.67–2.64 2.16
P/4 2 L 2.95–2.99 2.97
2 l 2.07–2.14 2.11
M/1 1 L 3.06
1 l 3
M/2 1 L 3.22
1 l 2.8
P/2-P/4 2 L 8.8–9.6 9.2
P/3-P/4 2 L 6.1–6.7 6.4
PCT
P/2 2 L 2.93–2.97 2.95
2 l 1.31–1.45 1.38
P/3 3 L 3.22–3.32 3.27 ± 0.04 0.05 1.54
3 l 1.63–1.92 1.74 ± 0.11 0.16 9.18
P/4 1 L 2.89
1 l 2.18
M/1 1 L 3.33
1 l 2.53
P/3-P/4 1 L 6.2
ITD
P/2 1 L 2.94
1 l 1.22
P/3 1 L 3.09
1 l 1.32
P/4 1 L 3.4
1 l 1.92
M/1 1 L 3.65
1 l 2.85
M/2 1 L 3.6
1 l 2.85
M/3 1 L 5.06
1 l 2.6
P/2-M/3 1 L 21.3
P/2-P/4 1 L 9.6
P/3-P/4 1 L 6.7
M/1-M/3 1 L 11.7

one with broken left P/2-M/1 (PCT 911), one with left
P/2-M/3 (PCT 916), one with left P/2-M/1 (PCT 362),
two with left P/3-P/4 (PCT 364, PCT 939), two with left
P/3-M/3 (PCT 349, PCT 915), one with right P/3-M/3
(PCT 491), one with right P/3-M1 (PCT 496), one with
right P/3-M/2 (PCT 904). Roqueprune 2: a left maxil-
lary fragment with P2/-M1/ (ROQ 353); three mandib-
ular fragments, one with left P/2-M/2 (ROQ 342), one
with left P/2-M/1 (ROQ 373), one with right P/2-P/4
(ROQ 345). Mounayne: five mandibular fragments,
one with right P/2-M/1 (MOU 146), two with left P/2-P/
4 (MOU 141, MOU 152), one with right P/2-P/4 (MOU
168), and one with right P/2-P/3 (MOU 145).
Measurements: See Table 4. For P. huerzeleri, the Kol-
mogorov-Smirnov and Shapiro-Wilk tests confirm
normality for the length of P/3 from Mas de Got
(N = 14, K–S, d = 0.166, P > 0.20, W = 0.912, P = 0.169)
and Pech Crabit (N = 11, K–S, d = 0.154, P > 0.20,
W = 0.971, P = 0.896).

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
156 C. BLONDEL

Table 4. Measurements and statistical data for teeth of Plesiomeryx huerzeleri. N, number of specimens; L, length; l,
width; LV, limit values; M ± SE, Mean ± standard error; s, standard deviation, V, variation coefficient

N L/l LV M ± SE s V

MGT
P1/ 1 L 3.46
1 l 1.2
P2/ 1 L 4.29
1 l 2.62
P3/ 3 L 2.88–3.6 3.99 ± 0.15 0.21 5.17
3 l 3.83–4.22 3.33 ± 0.28 0.39 11.8
P4/ 4 L 2.82–3.09 3.04 ± 0.12 0.21 6.89
4 l 4.18–4.52 4.35 ± 0.10 0.18 4.07
M1/ 4 L 4.10–4.30 4.2 ± 0.05 0.08 1.96
4 l 4.18–5.42 4.8 ± 0.40 0.69 14.5
M2/ 3 L 4.02–4.64 4.36 ± 0.23 0.32 7.23
3 l 4.99–6.06 5.47 ± 0.38 0.54 9.96
M3/ 2 L 4.09–4.86 4.48
2 l 5.21–5.25 5.23
P3/-P4/ 2 L 7.1–7.3 7.2
M1/-M3/ 2 L 12.3–13.1 12.7
P/1 1 L 2.3
1 l 1.19
P/2 6 L 3.04–3.87 3.56 ± 0.13 0.28 7.82
6 l 1.41–1.6 1.49 ± 0.03 0.07 4.78
P/3 14 L 3.33–4.30 3.97 ± 0.08 0.28 7.09
14 l 1.62–2.07 1.86 ± 0.04 0.15 8.01
P/4 12 L 3.53–4.12 3.87 ± 0.06 0.2 5.21
12 l 2.13–2.79 2.46 ± 0.07 0.22 9.1
M/1 10 L 3.65–4.27 3.99 ± 0.06 0.19 4.72
10 l 2.63–3.68 3.26 ± 0.11 0.33 10.1
M/2 7 L 3.98–4.44 4.21 ± 0.06 0.15 3.5
7 l 3.18–4.08 3.5 ± 0.14 0.35 10.1
M/3 5 L 5.19–6.01 5.58 ± 0.15 0.3 5.46
5 l 2.83–3.85 3.13 ± 0.22 0.44 14
P/2-P/4 4 L 10.2–11.1 10.6 ± 0.23 0.39 3.67
P/3-P/4 10 L 7.1–7.8 7.4 ± 0.08 0.23 3.05
M/1-M/3 5 L 12.6–13.8 13.2 ± 0.24 0.47 3.59
PL2
P/2 1 L 3.15
1 l 1.23
P/3 3 L 3.35–3.85 3.58 ± 0.18 0.25 7.08
3 l 1.74–2.02 1.9 ± 0.11 0.15 7.7
P/4 2 L 3.54–3.83 3.69
2 l 2.13–2.48 2.31
M/1 2 L 3.46–3.74 3.6
2 l 3.10–3.37 3.24
P/3-P/4 2 L 7.6–7.7 7.65
MOU
P/2 5 L 3.54–3.80 3.62 ± 0.05 0.1 2.9
5 l 1.36–1.69 1.54 ± 0.07 0.13 8.38
P/3 4 L 3.85–4.75 4.11 ± 0.25 0.43 10.4
4 l 1.74–2.12 1.96 ± 0.09 0.16 8.21
P/4 4 L 3.46–3.99 3.70 ± 0.13 0.23 6.29
4 l 2.41–2.67 2.51 ± 0.06 0.11 4.46
M/1 1 L 3.99
1 l 3.08

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 EARLY OLIGOCENE CAINOTHERIIDS 157

Table 4. Continued

N L/l LV M ± SE s V

P/2-P/4 3 L 11.2–11.6 11.5 ± 0.16 0.23 2.01

P/3-P/4 3 L 7.4–7.9 7.7 ± 0.18 0.25 3.28
ROQ2
P2/ 1 L 3.89
1 l 2.4
P3/ 1 L 3.73
1 l 3.76
P4/ 1 L 3.01
1 l 4.63
M1/ 1 L 4.08
1 l 5.95
P2/-P4/ 1 L 11.4
P3/-P4/ 1 L 7.32
P/2 3 L 3.32–3.57 3.87 ± 0.09 0.13 3.87
3 l 1.54–1.63 1.59 ± 0.04 0.05 2.97
P/3 3 L 3.81–4.12 3.96 ± 0.11 0.16 3.93
3 l 1.83–2.07 1.94 ± 0.08 0.12 6.31
P/4 3 L 3.76–4.28 3.99 ± 0.17 0.24 5.91
3 l 2.36–2.95 2.6 ± 0.21 0.3 11.7
M/1 2 L 3.83–3.99 3.91
2 l 3.11–3.4 3.3
M/2 1 L 4.3
1 l 3.48
P/2-P/4 3 L 11–12.1 11.6 ± 0.4 0.56 4.8
P/3-P/4 3 L 7.7–8.5 8.2 ± 0.31 0.44 5.32
PCT
P1/ 4 L 3.4–3.89 3.65 ± 0.12 0.21 5.8
4 l 1.44–1.62 1.5 ± 0.05 0.09 8.84
P2/ 4 L 4.35–4.69 4.55 ± 0.09 0.15 3.19
4 l 2.36–2.62 2.47 ± 0.06 0.11 4.4
P/1 1 L 3.13
1 l 1.34
P/2 4 L 3.89–4.21 4 ± 0.09 0.16 3.87
4 l 1.48–1.79 1.62 ± 0.08 0.13 7.91
P/3 11 L 3.44–4.53 4.13 ± 0.11 0.34 8.31
11 l 1.76–2.33 2.07 ± 0.05 0.17 8.08
P/4 9 L 3.73–4.10 3.86 ± 0.07 0.19 4.97
9 l 2.11–2.79 2.6 ± 0.10 0.29 11
M/1 7 L 3.50–4.19 3.83 ± 0.10 0.25 6.54
7 l 3.09–3.47 3.26 ± 0.06 0.15 4.66
M/2 5 L 3.84–4.51 4.18 ± 0.14 0.28 6.59
5 l 3.15–3.98 3.54 ± 0.16 0.32 9.08
M/3 4 L 5.01–5.86 5.37 ± 0.21 0.36 6.66
4 l 3.01–3.76 3.26 ± 0.20 0.34 10.4
P/2-P/4 1 L 12.1
P/3-P/4 6 L 7.5–8.3 7.8 ± 0.13 0.29 3.7
M/1-M/3 4 L 13.5–14.2 13.8 ± 0.17 0.29 2.13

The weakness of variation coefficients confirms the s = 0.31, K–S, d = 0.14, P > 0.20, W = 0.92, P = 0.149,
homogeneity of P. huerzeleri samples from localities of V = 7.97. P/3 from Pech Crabit, Roqueprune 2,
the same reference level. For the length of P/3 from Mounayne (MP23): N = 18, M = 4.10, s = 0.33, K–S,
Mas de Got and La Plante 2 (MP22): N = 17, M = 3.90, d = 0.14, P > 0.20, W = 0.94, P = 0.282, V = 8.05.

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
158 C. BLONDEL
Description
Upper dentition: A maxillary fragment from Mas de
Got (MGT 3212) has a diastema that separates the
canine from the short, premolariform and double-
rooted P1/, which has a high parastyle. P2/ is con-
structed on the same pattern as P1/ but is more elon-
gate (Fig. 3C), and the posterior lobe has a small
lingual cusp. The triangular P3/ is three-rooted and
the lingual root bears a strong cusp. The anterior lobe
is only weakly developed, and the parastyle and meta-
style are not very prominent on the labial wall. P4/
bears a median protocone that is remarkably straight
and its parastyle and metastyle are well developed.
The maxillary fragments from Pech Crabit and Roque-
prune 2 exhibit the same features as the specimens
from Mas de Got.
Lower dentition: The mandible from Mas de Got bears
a single-rooted caniniform P/1. A marked diastema
(3.5 mm) separates P/1 from P/2, and the latter exhib-
its a well developed paraconid and a main median
cusp. P/3 is more developed than P/2, and bears a lin-
gually situated paraconid as in P. cadurcensis. Distal
to the main cusp, there is a small tubercle which cor-
responds to the beginning of a crest that extends to the
posterolingual angle of the crown (Fig. 3D).
Comparison: The Quercy specimens whose features
are generally similar to those of other species of
Plesiomeryx are larger than remains assigned to
P. cadurcensis (Tables 3, 4). They are most similar to
material of P. huerzeleri from Gaimersheim (MP28) on
the basis of its measurements: the length of the
diastema between P/1 and P/2, premolars with higher
crowns than those of P. cadurcensis, a P3/ with a main
cusp, a talon, a labial wall that is more developed than
that of P. cadurcensis, and with proportions that lie
between P/2-P/4 and M/1-M/3. Although Gaimersheim
is more recent than the localities of La Plante 2, Mas
de Got (MP22), Mounayne, Pech Crabit and Roque-
prune 2 (MP23), the features of P. huerzeleri are not
significantly different from these forms from Quercy.
Discussion
This attribution confirms Sudre’s (1995) suggestion,
based on the remains from Le Garouillas, that Plesi-
omeryx contains various lineages that evolved concur-
rently. This idea challenges the hypothesis of Berger
(1959), who considered P. cadurcensis to be ancestral
to P. huerzeleri.
GENUS CAENOMERYX HÜRZELER, 1936
CAENOMERYX PROCOMMUNIS (FILHOL, 1877)
1877 Cainotherium procommune Filhol: 212, figs 269,
270.
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
1936 Caenomeryx procommunis (Filhol), Hürzeler,
1936: 100.
Holotype: Fragment of mandible bearing P/2-M/3 (Fil-
hol collections, MNHNP). Age and precise locality
unknown (Old Quercy collections).
Diagnosis: Upper molars are wider than long. Proto-
cone and protoconule present (stage B, Hürzeler, 1936;
p. 8). Diastema between P1/ and C is short. Diastema
very short or missing between P/1 and P/2. Premolars
reduced, P3/ with a strong lingual cone, P2/ with a
weaker lingual cone, P1/ very small. The premaxilla
does not reach the frontal. The wide nasal bones
broaden behind the posterior extremities of the pre-
maxilla. The ethmoidal gap is irregular, the choanae
are large and the mandible is high, stocky and mas-
sive (Hürzeler, 1936).
CAENOMERYX CF. PROCOMMUNIS (FILHOL, 1877)
Referred material: Mas de Got: a right maxillary frag-
ment P3/-P4/ (MGT 3178), five mandibular fragments,
one with left P/2-M/1 (MGT 118), one with left P/2-P/4
(MGT 155), one with right P/2-P/3 (MGT 3207), and
one with left P/3-P/4 (MGT 127, MGT 3206). Pech
Crabit: three maxillary fragments, one with right P2/
-P3/ (PCT 420), two with right P3/-P4/ (PCT 350, PCT
398); seven mandibular fragments, one with right P/2-
M/2 (PCT 938), one with left P/2-M/2 (PCT 951), one
with right P/2-M/1 (PCT 441), one with right P/2-P/4
(PCT 486), one with left P/2-P/4 (PCT 982), and two
with left P/2-P/3 (PCT 995, PCT 997). Itardies: three
maxillary fragments, one with left P1/-M1/ (ITD 952),
one with right P1/-P2/ (ITD 959), and one with left P1/
-P2/ (ITD 961). Mounayne: two left maxillary frag-
ments with P3/-P4/ (MOU 210, MOU 211), and a left
mandibular fragment P/3-P/4 (MOU 143). Roque-
prune 2: three mandibular fragments, one with right
P/1-P/4 (ROQ2344), one with left P/3-M/2 (ROQ2341),
and one with left P/3-P/4 (ROQ2383).
Measurements: See Table 5. For C. procommunis, the
Kolmogorov-Smirnov and Shapiro-Wilk tests confirm
normality for P/3 from Pech Crabit (N = 7, K–S,
d = 0.246, P > 0.20, W = 0.905, P = 0.368).
The weakness of variation coefficients confirms the
homogeneity of Caenomeryx procommunis samples
from localities of the MP23 reference level. For
the length of P/3 from Pech Crabit, Mounayne,
Roqueprune2: N = 11, M = 4.40, s = 0.29, K–S,
d = 0.19, P > 0.20, W = 0.95, P = 0.697, V = 6.54.
Description
Upper dentition: A number of maxillary fragments
bear a short premolariform P1/, a P2/ with a wide
 EARLY OLIGOCENE CAINOTHERIIDS 159

Table 5. Measurements and statistical data for teeth of Caenomeryx cf. procommunis. N, number of specimens; L, length;
l, width; LV, limit values; M ± SE, mean ± standard error; s, standard deviation, V, variation coefficient

N L/l LV M ± SE s V

MGT
P3/ 1 L 4.25
1 l 3.63
P4/ 1 L 3.23
1 l 4.36
P3/-P4/ 1 L 7.5
P/2 3 L 3.84–4.41 4.05 ± 0.22 0.31 7.67
3 l 1.57–1.66 1.63 ± 0.04 0.05 3.03
P/3 5 L 4.07–4.73 4.43 ± 0.14 0.28 6.38
5 l 2.09–2.35 2.2 ± 0.06 0.11 5.14
P/4 4 L 4.1–4.23 4.18 ± 0.03 0.06 1.44
4 l 2.37–3.02 2.72 ± 0.16 0.28 9.84
M/1 1 L 4.09
1 l 3.91
P/2-P/4 2 L 11.5–12 11.8
P/3-P/4 4 L 7.5–8.3 7.9 ± 0.22 0.38 4.75
MOU
P3/ 2 L 3.43–3.47 3.45
2 l 2.87–3.64 3.25
P4/ 2 L 3.2–3.21 3.21
2 l 4.34–4.96 4.65
P3/-P4/ 2 L 8.2–8.3 8.25
P/3 1 L 4.3
1 l 1.88
P/4 1 L 4.51
1 l 2.34
P/3-P/4 1 L 9.4
PCT
P2/ 1 L 4.71
1 l 2.5
P3/ 3 L 4.36–4.97 4.69 ± 0.22 0.31 6.58
3 l 3.95–4.62 4.28 ± 0.23 0.33 7.81
P4/ 2 L 3.7–3.79 3.75
2 l 5.5–5.88 5.69
P3/-P4/ 2 L 8.3–8.5 8.4
P/2 7 L 3.69–4.73 4.17 ± 0.13 0.31 7.42
7 l 1.56–2.29 1.8 ± 0.1 0.24 13.4
P/3 7 L 4.02–4.8 4.51 ± 0.10 0.25 5.64
7 l 2.03–2.92 2.29 ± 0.12 0.29 12.7
P/4 5 L 4.05–4.54 4.3 ± 0.11 0.21 4.88
5 l 2.41–2.88 2.65 ± 0.11 0.21 7.97
M/1 3 L 4.09–4.5 4.27 ± 0.15 0.21 4.88
3 l 3.48–3.52 3.51 ± 0.01 0.02 0.66
M/2 2 L 4.17–4.51 4.34
2 l 3.57–3.98 3.78
P/2-P/4 3 L 12.4–12.6 12.5 ± 0.08 0.11 0.93
P/3-P/4 4 L 8.4–8.8 8.7 ± 0.16 0.27 3.17
ITD
P1/ 3 L 3.39–3.6 3.49 ± 0.08 0.11 3
3 l 1.5 1.5
P2/ 3 L 4.26–4.5 4.49 ± 0.16 0.22 4.91
3 l 2.4–2.9 2.63 ± 0.18 0.25 9.56
P3/ 1 L 4.5
1 l 4.09

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160 C. BLONDEL

Table 5. Continued

N L/l LV M ± SE s V

P4/ 1 L 3.9
1 l 4.78
M1/ 1 L 5.15
1 l 6.26
P1/-P4/ 1 L 17
P2/-P4/ 1 L 12.4
P3/-P4/ 1 L 8.4
ROQ2
P/1 1 L 3.15
1 l 1.6
P/2 1 L 3.82
1 l 1.67
P/3 3 L 3.88–4.5 4.18 ± 0.22 0.31 7.44
3 l 1.91–2.09 2.02 ± 0.07 0.1 4.88
P/4 3 L 4.25–4.7 4.52 ± 0.17 0.24 5.23
3 l 2.53–3.12 2.81 ± 0.21 0.3 10.6
M/1 1 L 4.13
1 l 3.53
M/2 1 L 4.83
1 l 3.67
P/1-P/4 1 L 19.6
P/2-P/4 1 L 12.4
P/3-P/4 3 L 8.6–9.1 8.8 ± 0.18 0.25 2.85

tuberculate talon and a triangular P3/ with a well
developed distolingual talon. The anterior lobe of the
P3/ is elongate and the metastyle is well developed
(ITD 961, PCT 420) (Fig. 4A). The styles of the P4/
are strong and clearly prominent on the ectoloph.
Lower dentition: P/1 is caniniform, single-rooted and
separated from P/2 by a diastema of 3.8 mm (ROQ
344). The main cusp of the P/2 is low. The P/3 is con-
structed on the same pattern as P/2, but is wider,
the features are more pronounced and the para-
stylid is situated lingually (Fig. 4B). The corpus
mandibulae is bulky and very wide beneath the
molar area.
Comparison: The preserved upper and lower teeth
are in both their features and their measurements
similar to undated specimens from Old Quercy col-
lections and attributed to Caenomeryx procommunis
by Hürzeler (1936). The definition of the genus Cae-
nomeryx is based on this species. The new material
provides evidence of the presence of a similar spe-
cies in early Oligocene levels. Specimens from the
younger locality of Le Garouillas, MP25 reference
level (Sudre, 1995), described under the name of
C. cf. procommunis, belong to this same Caenomeryx
lineage.
CAENOMERYX FILHOLI (LYDEKKER, 1885A)
1885a Caenotherium Filholi Lydekker, p. 63.
1885b Caenotherium filholi Lydekker, p. 176.
1936? Caenotherium Filholi Lydekker; Hürzeler,
p. 102.
1937 Caenotherium commune aff. elegans Pomel;
Dehm, p. 353.
1959 Caenomeryx filholi (Lyddeker) Berger, 1959: 3.
Holotype: Skull (BMNH M1399). Age and precise
locality unknown (Old Quercy collections).
Diagnosis: Diastema between P/1 and P/2 and
between C and P1/. Premolars are not shortened. Pre-
maxillo-frontal suture variably present. The ethmoi-
dal slit is irregular. The mandible is high, massive and
long (Berger, 1959).
Referred material: Pech Crabit: two maxillary frag-
ments, one with left P2/-M3/ (PCT 417), and one with
right P3/-P4/ (PCT 390), two isolated left P3/ (PCT
392, PCT 418), 12 mandibular fragments, two with left
P/2-P/4 (PCT 437, PCT 453), two with left P/2-M/1
(PCT 928, PCT 933), one with broken left P/2-M/2
(PCT 445), one with right P/2-P/3 (PCT 936), one with
right P/3-M/3 (PCT 449), three with left P/3-P/4 (PCT
456, PCT 959, PCT 984), one with left P/3-M/2 (PCT
971), and one with right M/1-M/3 (PCT 299).

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 EARLY OLIGOCENE CAINOTHERIIDS 161

Figure 4. Caenomeryx cf. procommunis. A & B, occlusal view. A, right P2/-P3/ (PCT 420). B, left P/2-M/2 (PCT 951).
C. filholi. C & D, occlusal view. C, left P2/-P4/ (PCT 417). D, left P/2-M/1 (PCT 933). Cainotherium sp. E, left P2/-M3/ (PCT
416). Scale bar = 1 mm.

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162 C. BLONDEL

Measurements: See Table 6. For C. filholi, the Kolmog-
orov-Smirnov and Shapiro-Wilk tests confirm nor-
mality for P/3 from Pech Crabit (N = 10, K–S,
d = 0.201 P > 0.20, W = 0.928 P = 0.386,V = 6.18).
6) and similar to the Gaimersheim species (MP28)
C. filholi in its characters. Moreover, it is very similar
to C. filholi forma b described by Berger (1959) in its
upper teeth dimensions. However, the mandibular
teeth are somewhat smaller.

Description
Upper dentition: The material displays all the fea-
tures that characterize the genus: the double-rooted
P2/ is short. It has a wide lingual talon with a well
developed lingual cusp. The P3/ is triangular and
short. The median protocone is well developed, while
the parastyle and the metastyle are prominent on the
ectoloph (Fig. 4C).
Lower dentition: The single-rooted P/1 is separated
from P/2 by a diastema 3.3 mm long (ITD 944d). The
median tubercle of P/2 is poorly developed, and the
short P/3 displays a paraconid strongly carried
towards the lingual side (Fig. 4D).
Comparison: The material reported here represents a
species that is larger than C. procommunis (Tables 5,
Discussion
The older age of the Pech Crabit and Itardies locali-
ties (MP23) might explain the size difference
between the specimens from Pech Crabit and from
Gaimersheim, although this does not justify the dis-
tinction of this form at the specific level. The
C. filholi lineage may have evolved between MP23
and MP28 concurrently with C. procommunis. Noth-
ing in the collection indicates the presence of Cae-
nomeryx before MP22. I consider that the C. filholi
lineage present near MP23 originated from that of
P. cadurcensis because of numerous similar features.
The presence of P. cadurcensis and C. filholi near
MP23 contradicts the idea that these two species are
present at chronologically distinct levels (Berger,
1959).

Table 6. Measurements and statistical data for teeth of Caenomeryx filholi. N, number of specimens; L, length; l, width;
LV, limit values; M ± SE, Mean ± standard error; s, standard deviation,: V, variation coefficient

N L/l LV M ± SE s V

PCT
P2/ 1 L 4.12
1 l 2.6
P3/ 4 L 4.22–4.53 4.4 ± 0.08 0.13 2.93
4 l 4.07–4.51 4.29 ± 0.11 0.2 4.64
P4/ 2 L 3.87–3.91 3.89
2 l 5.41–5.47 5.44
M2/ 1 L 5.17
1 l 2.72
M3/ 1 L 4.87
1 l 6.25
P/2 6 L 3.62–4.53 4.2 ± 0.15 0.33 7.9
6 l 1.77–1.94 1.88 ± 0.03 0.07 3.52
P/3 10 L 4.34–5.16 4.74 ± 0.1 0.29 6.18
10 l 1.92–2.5 2.33 ± 0.06 0.17 7.14
P/4 9 L 4.52–5.07 4.79 ± 0.07 0.2 4.22
9 l 2.85–3.42 3.16 ± 0.07 0.19 5.99
M/1 6 L 4.42–5 4.69 ± 0.1 0.22 4.73
6 l 3.65–4.45 4.05 ± 0.13 0.3 7.49
M/2 4 L 4.22–5.27 4.95 ± 0.28 0.49 9.86
4 l 3.69–4.47 4.05 ± 0.2 0.34 8.3
M/3 2 L 6.61–7.25 6.98
2 l 3.31–3.83 3.57
P/2-P/4 4 L 12.3–13.4 13 ± 0.28 0.48 3.7
P/3-P/4 9 L 8.5–9.5 9.1 ± 0.11 0.32 3.47
M/1-M/3 2 L 16.1–17.6 16.9

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166

GENUS CAINOTHERIUM BRAVARD, 1828
(= CAENOTHERIUM AGASSIZ, 1846)
Type species: Cainotherium commune Bravard, 1835.
Diagnosis: The upper molars are quadrangular. Small
diastemata between C and P1/ and between P/1 and
P/2 are variably present. P2/, P/2 and P/3 are elongate.
P2/ lacks a lingual cusp. The premaxilla reaches the
frontal. The nasal is long, narrow and rectangular.
Presence of a depression over the nose root. A long
narrow crescent-shaped ethmoidal slit variably
present. The opening of choanae is narrow or large.
Presence or absence of a canalis alaris. The mandible
is long and thin (Berger, 1959: 39).
CAINOTHERIUM SP.
Material: Pech Crabit: a left maxillary fragment bear-
ing P2/-M2/ (PCT 416) (Blondel, 1996).
Measurements: See Table 7.
Description: This maxillary fragment (Fig. 4E) is dis-
tinguished from other material found at Pech Crabit
by its larger size, in particular the length of the tooth
rows and the presence of an elongate P2/ with a low
main cusp. This premolar has a lingual talon that is
reduced to a lingual cingulum from the parastyle to
the metastyle. On P3/ the lingual talon is weakly
developed and bears a tiny tubercle. The metastyle is
more prominent than the parastyle.
Comparison: This maxillary fragment resembles
those referred to Cainotherium by its characters. Its
dimensions agree with the corresponding part of the
skull of C. commune figured by Berger (1959: pl. 4,
fig. 2) and supposedly from Escamps (Old Quercy col-
lections). In the new Quercy collections, no specimen
referred to Cainotherium has yet been found on the
Escamps site. The Berger specimen has a P3/ with a
lingual talon that is more developed than in the spec-
imen from Pech Crabit (MP23). The Pech Crabit max-
illary has a P3/ that is comparable to that of the much
younger species C. laticurvatum from the Aquitanian
of Chavroches, in Allier, France (Hürzeler, 1936). The
scarcity of material does not permit checking the diag-
nostic characters of the cranium given by Berger
(1959) and the genus diagnosis cannot be emended.
Table 7. Measurements of teeth of Cainotherium sp. from
Pech Crabit. L, length; l, width
P2/ P3/ P4/ M1/ M2/ P3/-P4/ P2/-P4/
L 4.82 4.24 3.4 4.94 5.07 8.5 13.3
l 2.26 3.37 5.24 6.37 6.79
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
EARLY OLIGOCENE CAINOTHERIIDS 163
Discussion
A number of isolated molars indicate the presence of a
large cainotheriid in Escamps, Rosières 2 and Ravet
that is older than that of Pech Crabit. However, the
lack of material makes it difficult to determine
whether they should be referred to Cainotherium or
Caenomeryx. Material found in Pech Desse and Pech
du Fraysse, late Oligocene localities that are younger
than Pech Crabit, has not yet been studied and it is
not clear whether Cainotherium was present. The
study of all such material is necessary before the spe-
cies of Cainotherium from Pech Crabit can be identi-
fied. Thus, we consider that derived cainotherines
probably existed during the lower Oligocene.
DISCUSSION
The first representatives of the Cainotheriidae, such
as Oxacron courtoisi, appear in western Europe
c. MP18. Oxacronines appeared around MP19, with
the species Paroxacron valdense from Mormont-Entr-
eroches, Switzerland (Hooker & Weidmann, 2000) and
Paroxacron sp. from Escamps (Legendre, 1980). An
apparently new form from the Palembert locality (MP
19, Quercy Phosphorites) may be added to these spe-
cies (Legendre pers. comm., 1997).
Representatives of the Cainotheriinae appear
around MP 19; their presence in Escamps and Ros-
ières 2 is inferred from large isolated molars that can-
not be identified at the generic level. These molars
may represent Cainotherium commune, described by
Berger (1959) and reported from Escamps. In any
case, this record, consisting of at least three distinct
genera, indicates a remarkable degree of diversity for
this family in the late Eocene.
Among the Oxacroninae, Paroxacron seems to be the
only genus that persists beyond the Grande Coupure
(Fig. 5). It is present at Coyrou (MP20/21; Legendre
et al., 1995), Aubrelong 1, Ravet (MP21) and La Plante
2 (MP22). Few Cainotheriinae are present in MP21,
which is the first post-Eocene reference level. Plesi-
omeryx cadurcensis forms a lineage that persisted
through to MP25; it may continue after this level,
where it is represented by a large indeterminate cain-
otherid from Quercy Phosphorites. The Cainotheriidae
exhibit great diversity near MP22, with the appear-
ance of new lineages of Paroxacron (P. cf. bergeri),
Plesiomeryx (P. huerzeleri) and Caenomeryx
(C. cf. procommunis). All of these lineages, with the
exception of the small Paroxacron, survived beyond
this reference level, while a new lineage of Caenom-
eryx (C. filholi) appeared near MP23.
The late Oligocene material from Quercy has not yet
been studied, so it is not known when these lineages
became extinct. However, some lineages are near
more recent reference levels, such as C. filholi and
 164

Age MP
Mya O
30
C. BLONDEL

23.5 L ?
29
I ?
28
G
27 ?
O ?
26 ?
28.5
C 25 ?
Plesiomeryx
E 24 huerzeleri Caenomeryx
filholi Cainotherium
Plesiomeryx Caenomeryx sp.
N 23 cadurcensis cf. procommunis
Paroxacron
22 cf. bergeri
E
21
33.7
E 20 Paroxacron
O sp.
19 Oxacroninae Oxacron Paroxacron Cainotherium
C indet. commune?
courtoisi valdense
E 18
36
N
E

Figure 5. Chronological extension of the early Oligocene Cainotheriidae lineages. MP: Mammalian Palaeogene reference levels from Schmidt-Kittler (1987). Numer-
ical ages follow Odin (1994).

© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166

P. huerzeleri, which are represented in Gaimerscheim
(MP28) according to Berger (1959: 52, fig. 6). An
unpublished study (Douzery, 1991) suggests the pres-
ence of C. cf. filholi and C. filholi b in Pech Desse and
Pech du Fraysse, respectively (MP28). However,
rather than C. filholi, a preliminary examination
of this material reveals the remains to be
C. procommunis in Pech Desse and of the genus Cain-
otherium in Pech du Fraysse.
Thus it appears that two phases of diversification
occurred at the beginning of the history of the Cain-
otheriidae. The first occurred during the late Eocene
(MP19) with the first appearance of Oxacroninae, and
the second during the early Oligocene (MP22). Two
lineages of Plesiomeryx (P. cadurcensis–P. huerzeleri)
and Caenomeryx (C. procommunis–C. filholi) evolved
concurrently during the Oligocene. These two phases
around the Eocene/Oligocene transition correspond to
phases of diversification of artiodactyls in western
Europe during the late Eocene (e.g. with the Ano-
plotheriidae) and early Oligocene (e.g. with the rumi-
nants) (Blondel, 2001).
A turnover of terrestrial and marine biotas linked to
significant changes in ocean circulation and global cli-
mate occurred around the time of the transition. The
early Oligocene is marked by a cooling and aridity
(Cavelier, 1979; Berggren & Prothero, 1992; Prothero,
1994). In western Europe, the palaeobotanical data
indicate a radical floral change from the middle
Eocene to the early Oligocene with the replacement of
tropical/subtropical vegetation by more a temperate
and seasonal deciduous vegetation (Ollivier-Pierre
et al., 1987, Collinson, 1992; Cavagnetto & Anadón,
1996).
During this period there were important palaeogeo-
graphical changes in Europe, including a land conne-
ction between Europe and Asia caused by the closure
of the Turgaï strait and the formation of a corridor in
south-eastern Europe (Heissig, 1979; Ducrocq, 1995).
Thus many Asiatic immigrants, particularly small
sized ruminants, appeared in western Europe and
some endemic artiodactyl lineages such as the
Dacrytheriidae, the Xiphodontidae and the
Amphimerycidae became extinct (Blondel, 1997,
2001). In contrast, the Cainotheriidae persisted
through the transition period and rapidly diversified
during the early Oligocene. These small artiodactyls,
with their brachydont selenodont molars, were
adapted to important environmental modifications of
the early Oligocene.
ACKNOWLEDGEMENTS
I thank J. Sudre (EPHE, Montpellier II), B. Marandat,
S. Ducrocq (CNRS, Poitiers) and L. Meslin (CNRS,
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
EARLY OLIGOCENE CAINOTHERIIDS 165
Montpellier II) and the anonymous reviewers for their
comments.
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