New Data On The Cainotheriidae (Mammalia, Artiodactyla) From The Early Oligocene of South-Western France
New Data On The Cainotheriidae (Mammalia, Artiodactyla) From The Early Oligocene of South-Western France
New Data On The Cainotheriidae (Mammalia, Artiodactyla) From The Early Oligocene of South-Western France
Zoological Journal of the Linnean Society, 2005, 144, 145–166. With 5 figures
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.
© 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 some episodes in the history of this family more pre-
the Quercy Phosphorites during the late nineteenth cisely, in particular around the time of the Eocene/
century. Plesiomeryx, represented by P. cadurcensis Oligocene transition, which corresponds to the
Gervais, 1873, and Caenomeryx, represented by Grande Coupure as originally defined by Stehlin
C. procommunis (Filhol, 1877), have been described (1910).
from undated specimens collected in karstic fissures in Based on material collected in the Escamps locality
Quercy. of Quercy (MP19), Legendre (1980) showed that the
Fieldwork in the Quercy Phosphorites, organized Oxacroninae were more diverse than had been previ-
by the Universities of Montpellier, Poitiers and Paris, ously recognized in the late Eocene. A study in prep-
has been carried out since 1965. It has led to the dis- aration by Legendre & Sudre on the Cainotheriidae
covery of many new cainotheriid remains. Analysis of from late Eocene localities in France supports the idea
the Quercy mammalian fauna, together with that of of a rapid diversification of this group. At least five lin-
southern Germany and some Palaeogene mammalian eages, including at least one member of the Cain-
lineages, shows that these lineages can be arranged otheriinae, were present in this period (Legendre &
chronologically according to a reference level scale Sudre, pers. comm., 1997).
(Mammalian Palaeogene, henceforth MP) defined at In order to assess the relationships of late Eocene
the Mainz Symposium and revised at BiochroM in and early Oligocene Cainotheriidae, it is necessary to
1997 (Schmidt-Kittler, 1987; BiochroM, 1997; Fig. 1). study the newly dated material from the Quercy Phos-
Using this newly dated material, we can now define phorites, comparing the taxonomic status of these
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 CAINOTHERIIDS 147
early Oligocene forms with those described from the ROQ2 Roqueprune 2
undated old collections. UMII Montpellier II University, France
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
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 (Tables 1–3). The Paroxacron material from Ravet,
is split, although this feature, which only can be Aubrelong (MP21) and from La Plante 2 (MP22) dif-
observed on unworn teeth, is variable. fers from that from Escamps (Legendre, 1980) in that
the P/2-P/4 row is a little shorter. Material is rare,
Lower dentition: The double-rooted P/1 is premolari-
however, so it is difficult to determine whether these
form, elongate and narrow on the mandible from
differences are simply due to variation or whether
Ravet (Fig. 2C). This premolar possesses a sharp
they represent a trend toward premolar reduction in
median cusp and a faint paraconid. The double-rooted
the evolution of this lineage. In any case, it is the first
and elongate P/2 is separated from P/1 by a diastema
record of a small species of Paroxacron that has been
which varies from 0.8 to 1.4 mm in length. The para-
found in early Oligocene levels.
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/ PAROXACRON BERGERI HEISSIG, 1978
2, its paraconid is more pronounced than that of P/2
1959 cf. Cainotherium? elongatum Filhol (? Berger);
and a crest extends from a median cusp to the disto-
Berger p. 45, pl. 4, fig. 5, pl. 5 fig. 3.
lingual part of the tooth. The stocky P/4 is less elon-
1959 Cainotherium? n. sp.; Berger, p. 47, pl. 5, figs 6-
gate than the first premolars. The paraconid is
10.
strongly pronounced and the protoconid and the meta-
conid are also developed. The latter are separated by a Holotype. A right mandibular fragment I/3-M/3 (BSP
notch, although the metaconid is situated distally 1879XV201), locality of Mouillac in Quercy Phospho-
(Fig. 2D). rites. Age and precise locality are unknown (Old
Quercy collections).
Comparison: The presence of a premolariform P/1, the
very elongate premolars and the occurrence of a Diagnosis. Species of the genus Paroxacron without
diastema between P/1-P/2 and P/2-P/3 enables the diastema between P/2-P/3. The edge of the mandible is
form described above to be attributed to Paroxacron, more strongly bent than in P. valdense. The angulus
although the dimensions of this specimen are more mandibulae is short, high and angular. It is limited by
comparable to those of Plesiomeryx cadurcensis 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
EARLY OLIGOCENE CAINOTHERIIDS 151
is falciform. The choanae are large. The horizontal (N = 12, K–S, d = 0.135, P > 0.20, W = 0.949,
portion of the lower jaw is slender (Hürzeler, 1936). 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,
Referred material: Aubrelong 1: left maxillary frag-
d = 0.104, P > 0.20, W = 0.968, P = 0.887).
ment with P2/-M3/ (ABL1 1452), right maxillary frag-
The weakness of variation coefficients (4 < V < 10)
ment with P3/-M3/ (ABL1 1453); isolated left P3/
confirms the homogeneity of samples of P. cadurcensis
(ABL1 1559); 12 mandibular fragments, one with left
from localities of the same reference level. For the
P/1-P/2 (ABL1 1457), one with left P/2-M/2 (ABL1
length of P/3 from Aubrelong 1 and Ravet (MP21):
1458), one with right P/2-M/1 (ABL1 1459), one with
N = 24, M = 3.40, s = 0.23, K–S, d = 0.897, P > 0.20,
left P/2-P/4 (ABL1 1461), one with right P/2-P/3 (ABL1
W = 0.96, P = 0.438, V = 6.87. P/3 from Mas de Got and
1463), one with right P/3-M/2 (ABL1 1465), one with
La Plante 2 (MP22): N = 21, M = 3.02, s = 0.21, K–S,
left P/2-P/3 (ABL1 1487), one with broken right P/2-M/
d = 0.084, P > 0.20, W = 0.97, P = 0.672, V = 6.93. P/3
2 (ABL1 1492), one with broken right P/2-M/1 (ABL1
from Pech Crabit, Mounayne, Itardies (MP23): N = 6,
1497), one with left P/1-P/3 (ABL1 1780), one with
M = 3.21, s = 0.16, K–S, d = 0.19, P > 0.20, W = 0.97,
right P/3-P/4 (ABL1 1786), one with right P/2-P/4
P = 0.897, V = 5.05.
(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 Description
right P2/ (RAV 597, RAV 973), an isolated left P2/
Upper dentition: A maxillary fragment from Aubre-
(RAV 598), an isolated left P3/ (RAV 840); five man-
long 1 (Fig. 3A) bears a premolariform P1/ that is
dibular fragments: with left P/2-M/2 (RAV 1017), one
short and biradicular. The elongate P2/ has a weakly
with left P/1-M/1 (RAV 1024), one with right P/3-M/1
developed distolingual tubercle. The triangular P3/
(RAV 1025), two with left P/2-M/2 (RAV 1026, RAV
has a well developed mediolingual cusp from which
1027); an isolated left P/2 (RAV 853), three isolated
two crests extend to the prominently strong parastyle
right P/2 (RAV 739, RAV 740, RAV 892), four isolated
and metastyle. The labial and lingual cusps of P4/ are
right P/3 (RAV 732, RAV 889, RAV 895, RAV 1037),
median, the parastyle and the metastyle are strong.
four isolated left P/3 (RAV 849, RAV 887, RAV 898,
The same characters are evident in specimens (PL2
RAV 901). Mas de Got: 13 mandibular fragments, one
1180) from La Plante 2 (MP22).
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 Lower dentition: On the mandibular fragment (MGT
right P/2-P/4 (MGT 151), one with left P/3-M/3 (MGT 104), the single-rooted P/1 is caniniform and the
154), one with left P/2-P/4 (MGT 190), one with left P/ anterior stylid is not clearly marked (Fig. 3B). The
2-P/3 (MGT 195), one with right P/3-M/1 (MGT 3185), diastema that separates P/1 from P/2 varies between
one with right P/3-P/4 (MGT 3196), one with right P/1- 0.5 and 1.9 mm in length. The double-rooted P/2 bears
P/2 (MGT 3199), one with left P/2-P/3 (MGT 3200), one a well developed paraconid, while the main cusp is
with right P/2-P/3 (MGT 3202), one with right P/2-P/4 directed backward and reaches the distolingual angle.
(MGT 3203). La Plante 2: five maxillary fragments, On P/3, the paraconid is situated lingually and P/4 has
two with left P3/-M2/ (PL2 1180, PL2 1191), one with a well developed style. The protoconid and the meta-
left P1/-P2/ (PL2 1201), two with left P2/-P3/ (PL2 conid form large tubercles separated by a deep notch
1211, PL2 1213), 10 mandibular fragments, one with (e.g. ABL1 1458). There are no differences between the
right P/2-M/3 (PL2 1168), three with left P/3-M/3 (PL2 material from Aubrelong 1 and that from La Plante 2
1169, PL2 1242, PL2 1275), one with right P/4-M/3 or Pech Crabit.
(PL2 1173), one with left P/2-M/3 (PL2 1174), two with
Comparison: The presence of a single-rooted canini-
left P/3-M/3 (PL2 1175, PL2 1176), one with left P/2-P/
form P/1, a diastema between this premolar and P/2,
4 (PL2 1177), one with right P/2-P/3 (PL2 1250). Pech
the presence of a well developed mediolingual cusp
Crabit: three right mandibular fragments with P/3-M/
and a short anterior lobe on P/3 indicate that this
1 (PCT 366) and one with P/2-P/3 (PCT 367, PCT 385).
small cainotherine from Aubrelong 1 (MP21) belongs
Mounayne: two left mandibular fragments, one with
to Plesiomeryx. In specimens from the localities of
P/2-M/2 (MOU 209) and one with P/2-P/4 (MOU 148),
Aubrelong 1, La Plante 2, and Pech Crabit, the dimen-
a right maxillary fragment with P2/-P3/ (MOU 187).
sions of teeth in the lower dental rows (Table 3), and
Itardies: a right mandibular fragment with P/2-M/3
the features of the lower premolars agree with the
(ITD 941).
description of P. cadurcensis. The same is also true for
Measurements: See Table 3. For P. cadurcensis, the the upper tooth rows, which correspond in their mor-
Kolmogorov-Smirnov and Shapiro-Wilk tests confirm phology and dimensions to material of P. cadurcensis
normality for L P/3 from Aubrelong 1 (N = 12, K–S, described by Hürzeler (1936) and Berger (1959). The
d = 0.163, P > 0.20, W = 0.955, P = 0.721), Ravet general similarity of the morphological patterns and
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
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- Referred material: Mas de Got: four maxillary frag-
neity despite the different ages of the studied ments, one with right P2/-M3/ (MGT 3209), one with
material. 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,
PLESIOMERYX HUERZELERI BERGER, 1959 MGT 3187), three with left P/3-M/3 (MGT 103, MGT
1937 Caenotherium gracile Pomel; Dehm, 1937: 352. 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),
Holotype: Skull BSP1952 II 1149. Gaimersheim fis- one with left P/1-P/3 (MGT 124), two with right P/3-P/
sure filling (Germany) (MP28). 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:
Diagnosis: Species of Plesiomeryx that appears similar three mandibular fragments, two with right P/3-M/1
to P. cadurcensis, but is larger. The fronto-nasal (PL2 1171, PL2 1221), one with left P/2-P/3 (PL2
suture is short. The lingual cingulum of P/3 is missing. 1216). Pech Crabit: four left maxillary fragments P1/-
The lingual and labial cones of P/4 reach the same P2/ (PCT 382, PCT 391, PCT 431, PCT 1000); 11 man-
level (Berger, 1959). dibular fragments, two with left P/2-P/3 (PCT 348),
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
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
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
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
one with broken left P/2-M/1 (PCT 911), one with left one with right P/2-M/1 (MOU 146), two with left P/2-P/
P/2-M/3 (PCT 916), one with left P/2-M/1 (PCT 362), 4 (MOU 141, MOU 152), one with right P/2-P/4 (MOU
two with left P/3-P/4 (PCT 364, PCT 939), two with left 168), and one with right P/2-P/3 (MOU 145).
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 Measurements: See Table 4. For P. huerzeleri, the Kol-
right P/3-M/2 (PCT 904). Roqueprune 2: a left maxil- mogorov-Smirnov and Shapiro-Wilk tests confirm
lary fragment with P2/-M1/ (ROQ 353); three mandib- normality for the length of P/3 from Mas de Got
ular fragments, one with left P/2-M/2 (ROQ 342), one (N = 14, K–S, d = 0.166, P > 0.20, W = 0.912, P = 0.169)
with left P/2-M/1 (ROQ 373), one with right P/2-P/4 and Pech Crabit (N = 11, K–S, d = 0.154, P > 0.20,
(ROQ 345). Mounayne: five mandibular fragments, 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
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
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
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 CAENOMERYX FILHOLI (LYDEKKER, 1885A)
developed distolingual talon. The anterior lobe of the 1885a Caenotherium Filholi Lydekker, p. 63.
P3/ is elongate and the metastyle is well developed 1885b Caenotherium filholi Lydekker, p. 176.
(ITD 961, PCT 420) (Fig. 4A). The styles of the P4/ 1936? Caenotherium Filholi Lydekker; Hürzeler,
are strong and clearly prominent on the ectoloph. p. 102.
1937 Caenotherium commune aff. elegans Pomel;
Lower dentition: P/1 is caniniform, single-rooted and Dehm, p. 353.
separated from P/2 by a diastema of 3.8 mm (ROQ 1959 Caenomeryx filholi (Lyddeker) Berger, 1959: 3.
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, Holotype: Skull (BMNH M1399). Age and precise
the features are more pronounced and the para- locality unknown (Old Quercy collections).
stylid is situated lingually (Fig. 4B). The corpus Diagnosis: Diastema between P/1 and P/2 and
mandibulae is bulky and very wide beneath the between C and P1/. Premolars are not shortened. Pre-
molar area. maxillo-frontal suture variably present. The ethmoi-
dal slit is irregular. The mandible is high, massive and
Comparison: The preserved upper and lower teeth
long (Berger, 1959).
are in both their features and their measurements
similar to undated specimens from Old Quercy col- Referred material: Pech Crabit: two maxillary frag-
lections and attributed to Caenomeryx procommunis ments, one with left P2/-M3/ (PCT 417), and one with
by Hürzeler (1936). The definition of the genus Cae- right P3/-P4/ (PCT 390), two isolated left P3/ (PCT
nomeryx is based on this species. The new material 392, PCT 418), 12 mandibular fragments, two with left
provides evidence of the presence of a similar spe- P/2-P/4 (PCT 437, PCT 453), two with left P/2-M/1
cies in early Oligocene levels. Specimens from the (PCT 928, PCT 933), one with broken left P/2-M/2
younger locality of Le Garouillas, MP25 reference (PCT 445), one with right P/2-P/3 (PCT 936), one with
level (Sudre, 1995), described under the name of right P/3-M/3 (PCT 449), three with left P/3-P/4 (PCT
C. cf. procommunis, belong to this same Caenomeryx 456, PCT 959, PCT 984), one with left P/3-M/2 (PCT
lineage. 971), and one with right M/1-M/3 (PCT 299).
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
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.
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
162 C. BLONDEL
Measurements: See Table 6. For C. filholi, the Kolmog- 6) and similar to the Gaimersheim species (MP28)
orov-Smirnov and Shapiro-Wilk tests confirm nor- C. filholi in its characters. Moreover, it is very similar
mality for P/3 from Pech Crabit (N = 10, K–S, to C. filholi forma b described by Berger (1959) in its
d = 0.201 P > 0.20, W = 0.928 P = 0.386,V = 6.18). upper teeth dimensions. However, the mandibular
teeth are somewhat smaller.
Description
Upper dentition: The material displays all the fea- Discussion
tures that characterize the genus: the double-rooted The older age of the Pech Crabit and Itardies locali-
P2/ is short. It has a wide lingual talon with a well ties (MP23) might explain the size difference
developed lingual cusp. The P3/ is triangular and between the specimens from Pech Crabit and from
short. The median protocone is well developed, while Gaimersheim, although this does not justify the dis-
the parastyle and the metastyle are prominent on the tinction of this form at the specific level. The
ectoloph (Fig. 4C). C. filholi lineage may have evolved between MP23
and MP28 concurrently with C. procommunis. Noth-
Lower dentition: The single-rooted P/1 is separated ing in the collection indicates the presence of Cae-
from P/2 by a diastema 3.3 mm long (ITD 944d). The nomeryx before MP22. I consider that the C. filholi
median tubercle of P/2 is poorly developed, and the lineage present near MP23 originated from that of
short P/3 displays a paraconid strongly carried P. cadurcensis because of numerous similar features.
towards the lingual side (Fig. 4D). The presence of P. cadurcensis and C. filholi near
MP23 contradicts the idea that these two species are
Comparison: The material reported here represents a present at chronologically distinct levels (Berger,
species that is larger than C. procommunis (Tables 5, 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
EARLY OLIGOCENE CAINOTHERIIDS 163
Material: Pech Crabit: a left maxillary fragment bear- The first representatives of the Cainotheriidae, such
ing P2/-M2/ (PCT 416) (Blondel, 1996). as Oxacron courtoisi, appear in western Europe
c. MP18. Oxacronines appeared around MP19, with
Measurements: See Table 7. the species Paroxacron valdense from Mormont-Entr-
Description: This maxillary fragment (Fig. 4E) is dis- eroches, Switzerland (Hooker & Weidmann, 2000) and
tinguished from other material found at Pech Crabit Paroxacron sp. from Escamps (Legendre, 1980). An
by its larger size, in particular the length of the tooth apparently new form from the Palembert locality (MP
rows and the presence of an elongate P2/ with a low 19, Quercy Phosphorites) may be added to these spe-
main cusp. This premolar has a lingual talon that is cies (Legendre pers. comm., 1997).
reduced to a lingual cingulum from the parastyle to Representatives of the Cainotheriinae appear
the metastyle. On P3/ the lingual talon is weakly around MP 19; their presence in Escamps and Ros-
developed and bears a tiny tubercle. The metastyle is ières 2 is inferred from large isolated molars that can-
more prominent than the parastyle. not be identified at the generic level. These molars
may represent Cainotherium commune, described by
Comparison: This maxillary fragment resembles Berger (1959) and reported from Escamps. In any
those referred to Cainotherium by its characters. Its case, this record, consisting of at least three distinct
dimensions agree with the corresponding part of the genera, indicates a remarkable degree of diversity for
skull of C. commune figured by Berger (1959: pl. 4, this family in the late Eocene.
fig. 2) and supposedly from Escamps (Old Quercy col- Among the Oxacroninae, Paroxacron seems to be the
lections). In the new Quercy collections, no specimen only genus that persists beyond the Grande Coupure
referred to Cainotherium has yet been found on the (Fig. 5). It is present at Coyrou (MP20/21; Legendre
Escamps site. The Berger specimen has a P3/ with a et al., 1995), Aubrelong 1, Ravet (MP21) and La Plante
lingual talon that is more developed than in the spec- 2 (MP22). Few Cainotheriinae are present in MP21,
imen from Pech Crabit (MP23). The Pech Crabit max- which is the first post-Eocene reference level. Plesi-
illary has a P3/ that is comparable to that of the much omeryx cadurcensis forms a lineage that persisted
younger species C. laticurvatum from the Aquitanian through to MP25; it may continue after this level,
of Chavroches, in Allier, France (Hürzeler, 1936). The where it is represented by a large indeterminate cain-
scarcity of material does not permit checking the diag- otherid from Quercy Phosphorites. The Cainotheriidae
nostic characters of the cranium given by Berger exhibit great diversity near MP22, with the appear-
(1959) and the genus diagnosis cannot be emended. ance of new lineages of Paroxacron (P. cf. bergeri),
Plesiomeryx (P. huerzeleri) and Caenomeryx
(C. cf. procommunis). All of these lineages, with the
Table 7. Measurements of teeth of Cainotherium sp. from
exception of the small Paroxacron, survived beyond
Pech Crabit. L, length; l, width
this reference level, while a new lineage of Caenom-
P2/ P3/ P4/ M1/ M2/ P3/-P4/ P2/-P4/
eryx (C. filholi) appeared near MP23.
The late Oligocene material from Quercy has not yet
L 4.82 4.24 3.4 4.94 5.07 8.5 13.3 been studied, so it is not known when these lineages
l 2.26 3.37 5.24 6.37 6.79 became extinct. However, some lineages are near
more recent reference levels, such as C. filholi and
© 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 145–166
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
EARLY OLIGOCENE CAINOTHERIIDS 165
P. huerzeleri, which are represented in Gaimerscheim Montpellier II) and the anonymous reviewers for their
(MP28) according to Berger (1959: 52, fig. 6). An comments.
unpublished study (Douzery, 1991) suggests the pres-
ence of C. cf. filholi and C. filholi b in Pech Desse and
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