A new early Late Oligocene (MP 26) continental vertebrate fauna from Saint-Privat-des-Vieux (Alès Basin, Gard, Southern France) Author Vianey-Liaud, Monique Author Comte, Bernard Author Marandat, Bernard Author Peigné, Stéphane Author Rage, Jean-Claude Author Sudre, Jean text Geodiversitas 2014 2014-12-26 36 4 565 622 http://dx.doi.org/10.5252/g2014n4a4 journal article 6390 10.5252/g2014n4a4 8bc26961-9853-4113-a63d-4cf691aa7100 1638-9395 4538625 Issiodoromys pauffiensis Vianey-Liaud, 1976 HOLOTYPE . — Left dentary with p4-m3, MPF-1 (Vianey- Liaud 1976)); coll. UM2 . TYPE LOCALITY . — Mas-de-Pauffié, Lot, Quercy, MP 26 (Chattian). EMENDED DIAGNOSIS. — (Completing Vianey-Liaud 1976: 55 and Schmidt-Kittler & Vianey-Liaud 1987): Evolutionary stage intermediate between I. minor (Schlosser, 1884) and I. quercyi (Schlosser, 1884) . Cement rarely present at the bottom of sinus and sinusid. Horizontal uppermasseteric crest strongerthan in I. minor , especially in its posterior part. Differs from I. minor by the more elongated anterior lobe of dp4, including two well-developed oblique ridges, mesially to the mesolophid (Ldp4 of I. pauffiensis from Mas-de-Pauffié = 2.71 mm to 3.01 mm ); antesinusid generally absent on molars, anteroconid reduced to a weak vertical ridge when present, and in some specimens preceded by a small additional inflection of the crown on dp4; enamel cover thinner; simplified occlusal patterns in some individuals. Differs from I. quercyi by itssmaller size and relatively frequent remnants of the antesinusid-anteroconid on molars. Differs from I. limognensis Schmidt-Kittler & Vianey- Liaud, 1987, in having the anterior lobe of dp4 less stretched, including less pronounced antesinusids (Ldp4 of I. limognensis from Pech-Desse = 2.96 mm to 3.67 mm ), cement limited to the bottom of sinussinusid when present, whereas it reaches the occlusal surface – including teeth slightly to moderately worn – in I. limognensis . MATERIAL EXAMINED. — 29 DP4 ( SPV 200-228), 40 P4 ( SPV 229-266), 113 M1-2 ( SPV 267-389), 26 M3 ( SPV 390-414); 29 dp4 ( SPV 415-442), 21 p4 ( SPV 443- 463), 101 m 1-2 ( SPV 464-564), 23 m 3 SPV 565-589). MEASUREMENTS. — To the variations in size between the first and second molars (the first being generally smaller than the second) must be added those due to effects of wear on the occlusal surface orientation of hypsodont teeth (Vianey-Liaud et al. 1995: 274, fig. 12), and finally those due to individual variability. Ŋe size variations are expressed by the Pearson coefficient: v = 100Sd / Lmoy ( Table 2 ). Ŋe low size variability observed at Saint-Privat-des-Vieux reinforces the hypothesis on the taphonomy of the fossil assemblage from this locality ( Bonnet et al. 2011: 12 ). DESCRIPTION Lower teeth dp4 ( Fig. 4 ). Ŋe morphology of dp4 posterior lobe is constant, while some variations are observed in the anterior lobe. Ŋis is actually the part of dp4 that most changes during the evolution of the lineage, extending mesially the abrading surface of the tooth row. All dp4 show 2-3 oblique ridges between the mesolophid and the prelobe tip. Ŋe protoconid is split by a small labial notch, already sketched in populations of I. minor and present in the type population of I. pauffiensis (e.g., Schmidt-Kittler et al. 1997: 385, fig. 7; Vianey-Liaud et al. 1995: 267, fig. 5). Ŋe elongated central cuspid, which occupies the mesolophid position, runs obliquely towards the anterior arm of the protoconid to which it may be connected or not. Once upon 20 well preserved teeth, this central cusp reaches the tip of the prelobe. In all other cases, it is preceded by two oblique crestids parallel to it. Ŋe first is usually reduced to a small cusp. A continuous cingulid, as high as other ridges, follows the outline of the anterior lobe of teeth from the front flank of the TABLE 2. — Measurements of Issiodoromys pauffiensis Vianey-Liaud, 1976 from Saint-Privat-des-Vieux, compared to that (for M1-2 and m1-2) of I. pauffiensis from Mas-de-Pauffié, and m1-2 of I. quercyi from Ehrenstein7. Abbreviations: Sd , standard deviation; antcd , number of m2 with anteroconid present.

Issiodoromys pauffiensis , Saint-Privat-des-Vieux
N	L min	L max	L mean	v	Sd	w min	w max	w mean	Sd
dp4	20	2.81	3.1	2.94	2.99	0.088	1.03	1.46	1.31	0.115
p4	17	1.51	1.91	1.74	6.37	0.111	0.85	1.59	1.32	0.211
m1-2	96	1.72	2.48	2.07	7.53	0.156	1.03	2.11	1.49	0.227
m3	12	1.79	2.19		1.9	9	6.22 0.124 1.19
DP4	25	2.29	2.64 2.40 3.48 0.084 1 1.51 1.32
P4	28	1.59	1.9 9		1.72 6.06 0.104 0.74 2.15
M1-2	102	1.52	2.16	1.76	7.13	0.126	1.1	2.98	2.15	0.446
M3	17	1.08	1.95	1.41	13.43	0.189	1.17	1.44	1.55	0.321
Issiodoromys pauffiensis , Mas-de-Pauffié
N	L min	L max	L mean	v	Sd	w min	w max	w mean	Sd
dp4	28	2.68	3.26	2.97	9.76	0.290	1.05	1.47	1.26	0.21
p4	50	1.73	2.1	1.91	7.41	0.142	0.98	1.77	1.42	0.213
m1-2	177	1.51	2.48	2	24.25	0.485	0.93	2.3	1.62	0.685
m3	51	1.45	2.44	1.95	25.38	0.495	1	1.89	1.45	0.445
DP4	43	2.14	2.67	2.40	11.04	0.265	0.97	1.6	1.29	0.315
P4	57	1.56	2.04	1.8	13.33	0.24	0.98	2.5	1.48	0.31
M1-2	185	1.4	2.05	1.73	18.79	0.325	1	2.95	2.28	0.090
M3	28	1.14	1.78	1.46	21.92	0.32	1.09	2.07	1.58	0.49
Issiodoromys pauffiensis , St-Privat-des-Vieux					N	H min.	H max	H mean	Sd	antcd
m1-2	96	0.77	3.32	2.51	0.508	32
M1-2	102	0.95	3.76	3.05	0.636
Issiodoromys pauffiensis , Mas de Pauffié					N	H min.	H max	H mean	Sd	antcd
m1-2	124	0.97	3.72	2.35	1.375	79
M1-2	124	1.14	4.05	2.6	1.455
Issiodoromys quercyi , Ehrenstein 7
N	L min.	Lmax	Lmean	v	H min.	H max	Sd
m1-2	127	2	3.13	2.57	21.16	0.97	4.81	0.565

1.31 0.138 1.42 protoconid to the antero-lingual side of synclinid III (SIII). Labially, this cingulid shows two constant inflections: the antesinusid ahead of the protoconid, and a generally shallow pre-antesinusid.Ŋe lingual portion of the cingulid is vertically flat, and starts in continuity with the anterior cuspid of prelobe. Ŋis cuspid is prominent on unworn teeth. Ŋe size and morphology of these dp4 are very similar to that of dp 4 type population ( Table 2 ). p4 ( Fig. 5 A-C). Ŋere is a low morphological variability. A short mesio-distal expansion starting from the anterolophid inward SII is rarely observed (4/20). A small cingulid may be present at the base of the lingual opening of SIII (5/20). A p4 (1/20) bears only a remnant of the anteroconid. m1-2. Ŋe vestigial anteroconid / antesinusid – as a mesiolabial vertical bulge – seems more frequent on molars than on p4, but this may be due to the more important number of m1-2. On little worn m1-2, 18/77 teeth have this vestige, while it is absent (or lost due to abrasion) on 23 worn teeth. Fourteen teeth exhibit a short mesiodistal spur in SII. On a single specimen, a mesiodistal spur is added to that in the SIV. Another specimen shows a lingual interruption of the mesolophid, and displays two small lin- gual protrusions, the most distal one blocking the lingual opening of the SIII ( Fig. 5E ). Ŋe only very worn tooth (H = 0.77 mm ) exhibits a thin layer of cement at the bottom of the sinusid ( Fig. 5I ); this cement is not visible on the other teeth. FIG. 4. — Variations of the occlusal surface of lower dp4 of Issiodoromys Bravard in Gervais, 1848 : A -Q , I. pauffiensis Vianey-Liaud, 1976 , from Saint-Privat-des-Vieux (Gard; Lower Chattian, MP 26): A , SPV415; B , SPV423; C , SPV433; D , SPV424; E , SPV417; F , SPV430; G , SPV425; H , SPV418; I , SPV426; J , SPV419; K , SPV437; L , SPV420; M , SPV439; N , SPV438; O , SPV429; P , SPV427; Q , SPV422; R -AF , I. quercyi (Schlosser,1884) from Ehrenstein 7 (Germany; MP 27): R -X , Z -AD , occlusal views: R , no. 1983-XVIII 470; S , no. 1983-XVIII 457; T , no. 1983-XVIII 466; U , no. 1983-XVIII 460; V , no. 1983-XVIII 459; W , no. 1983-XVIII 458; X , no. 1983-XVIII 462; Y , no. 1983-XVIII 465, Y1 , occlusal view, Y2 , labial view; Z , no. 1983-XVIII 468; AA , no. 1983-XVIII 463; AB , no. 1983-XVIII 464; AC , no. 1983-XVIII 461; AD , no. 1983-XVIII 467. Scale bar: 1 mm. FIG. 5. — Occlusal views of lower teeth of Issiodoromys pauffiensis Vianey-Liaud, 1976 from Saint-Privat-des-Vieux (Gard; Lower Chattian, MP 26): A , left p4, SPV461; B , left p4, SPV460; C , left m1, SPV475; D , right m2, SPV481; E , right m1-2, SPV486; F , left m1- 2, SPV507; G , right m1-2, SPV519; H , left m1-2, SPV539; I , much worn right m1-2, SPV563; J , unworn right m3, SPV574; K , unworn right m3, SPV565; L , moderately worn right m3, SPV577. Scale bar: 1 mm. m3 ( Fig. 5 J-L). Ŋe mesiodistal spur ( Fig. 5L ) is present in the SII of 5/14 teeth and the anteroconid is detectable on one tooth only. Upper teeth DP4( Fig. 6 ). Again morphological variations are minimal ( Fig. 6 A-O). Ŋe anticline 4 may be continuous (21/24) or split into two parts (3/24). We observe a light variability in the labial cingulum blocking the opening of the median syncline (SII) (e.g., Fig. 6C ). It is sometimes absent and sometimes high, resulting of proximity of two spurs coming from the labial ends of the anticlines 2 and 3, or only one labial spur from the anticline 2, mesiodistal or oblique. Ŋe buccal cingular edge of the syncline III is usually swollen as a cusp that can spawn a spur extending into the syncline. P4 ( Fig. 7 A-C). Ŋe lingual end of the anticline 4 is free (12/29) or connected to the posterior cingulum (17/29). Ŋe labial portion of the anterior cingulum is rarely complete (1/29), it is either vestigial (11/29) or absent. Ŋe paracone is sometimes isolated from the protocone (7/29) due to the interruption of the protoloph (anticline 2). M1-2 ( Fig. 7 D-K). the morphological variations are minor and mainly concern SIV, which is mostly closed (98/101), or rarely mesially opened (2/101). A small mesiodistal spur runs backwards from the anticline 3 in the SIII (1/101). M3 ( Fig. 7L, M ). Ŋe size and shape of the M3 are more variable than those of M1-2, especially FIG. 6. — Variations of the occlusal surface of Upper DP4 of Issiodoromys Bravard in Gervais, 1848 : A -O , I. pauffiensis Vianey-Liaud, 1976 from Saint-Privat-des-Vieux (Gard;Lower Chattian,MP 26): A , SPV200; B , SPV209; C , SPV219; D , SPV206; E , SPV201; F , SPV204; G , SPV205; H , SPV216; I , SPV208; J , SPV210; K , SPV213; L , SPV211; M , SPV202; N , SPV212; O , SPV222; P -X2 , I. quercyi from Ehrenstein7 (Germany; MP 27): P , no.1983-XVIII 471, P1 , occlusal view, P2 , lingual view; Q -W , occlusal views: Q , no. 1983-XVIII 479; R , no. 1983-XVIII 472; S , no. 1983-XVIII 473; T , no. 1983-XVIII 474; U , no. 1983-XVIII 475; V , no. 1983-XVIII 476; W , no. 1983-XVIII 477; X , no. 1983-XVIII 478: X1 , occlusal view, X2 , lingual view. Scale bar: 1 mm. FIG. 7. — Occlusal views of upper teeth of Issiodoromys pauffiensis Vianey-Liaud, 1976 from Saint-Privat-des-Vieux (Gard; Lower Chattian,MP 26): A , left P4, SPV236; B , left P4, SPV235; C , left P4, SPV234; D , left M1-2, SPV302; E , left M1-2,SPV270; F , right M1- 2, SPV276; G , right M1-2, SPV269; H , left M1-2, SPV278; I , worn right M1-2, SPV362; J , worn left M1-2, SPV366; K , much worn left M1-2, SPV387; L , unworn left M3, SPV390; M, moderately worn right M3, SPV396. Scale bar: 1 mm. because of the greater or lesser reduction in the posterior lobe. On one of the smaller specimens, it only remains the anticline 3 at the back of the tooth. DISCUSSION Since the description of I. pauffiensis (Vianey-Liaud 1976) , we collected additional material for its type locality; for the present work, it has been measured, observed and compared to that of Saint-Privat-des-Vieux. Molars from Saint- Privat-des-Vieux are similar in size to those of I. pauffiensis from Mas-de-Pauffié. Ŋeir size is significantly smaller than that of I. quercyi (Ehrenstein 7, Germany [ Fig. 6 P-Z; Fig. 8B ], and Sarèle, France ; MP 27) ( Table 2 ). Ŋe size variability (length and width of teeth) of I. pauffiensis from Saint-Privat-des-Vieux is not higher than that observed in extant populations of brachyodont rodents or other mammals (e.g., Vianey-Liaud & Legendre 1986: 921, table 1). Ŋis variability is also less than that observed in fossil rodent teeth populations displaying a strong asymmetric hypsodonty, from both fissure fillings and stratified sedimentary deposits (e.g Vianey-Liaud & Legendre 1986: 921, table 3). It is thus a little lower than that of the I. pauffiensis type population from Mas-de-Pauffié, or that of I. quercyi from Ehrenstein 7 ( Table 2 ; Fig. 8B ). Ŋe range of crown height is within that of species exhibiting similar to close hypsodonty (the maximum and minimum heights being slightly lower than those from Mas-de-Pauffié, and slightly higher than in I. minor from Le Garouillas). Ŋis range is clearly wider for I. quercyi from Ehrenstein 7, and also for I. limognensis from Pech Desse, both species being significantly more hypsodont ( Fig. 8A ). Morphological variability observed in Mas-de- Pauffié, especially for deciduous teeth, is the same as described above for Saint-Privat-des-Vieux. Minor differences can be noted with regard to the dp4. Morphological variation in 25 dp4 from Mas-de-Pauffié shows the same characteristics as in the population of Saint-Privat-des-Vieux, except the presence of a low cingulid which partially or completely closes the bottom of the opening of the SIII on five specimens. Ŋis cingulid is also occasionally present on p4 from both localities or on molars: its absence on these five dp4 may be a statistical bias. Postcranial elements of I. pauffiensis from Saint-Privat-des-Vieux – in particular astragali, calcanei and humeri fragments – are quite similar to those of Mas-de-Pauffié (Vianey-Liaud et al. in press).