Large mammals of Fouvent-Saint-Andoche (Haute-Saône, France): a glimpse into a Late Pleistocene hyena den Author Fourvel, Jean-Baptiste Author Fosse, Philippe Author Fernandez, Philippe Author Antoine, Pierre-Olivier text Geodiversitas 2015 2015-06-26 37 2 237 266 http://dx.doi.org/10.5252/g2015n2a5 journal article 10.5252/g2015n2a5 1638-9395 4535125 urn:lsid:zoobank.org:pub:0117CBA4-4CE0-4431-B5F6-721F998C72C7 Vulpes vulpes Linnaeus, 1758 MATERIAL EXAMINED. — NISP =13; MNI =5. 1842 sample: 2 left humeri; 1 right tibia; 1 left tibia. 1989-1992 sample: 2 right c; 1 right mandible; 2 left mandibles; 1 left ulna; 1 left metacarpal II; 1 right calcaneus; 1 left metatarsal V. DESCRIPTION Among the small canids remains (n=76), some of them have been associated to the red fox Vulpes vulpes (n=13) corresponding to five adults. The low number of elements determined at species level is related both to the state of conservation of the material and to the difficulties to distinguish Vulpes and Alopex Kaup, 1829 . The red fox is a small common carnivore in Pleistocene bone assemblages (e.g., Gönnersdorf in Poplin 1976 ; Gerde in Clot 1980 ). This predator is ubiquitous and well-represented during different Pleistocene glacial and interglacial climatic phases. In France , the red fox is found at Nauterie II (layer 11: Mindel-Riss), at Montmaurin-la-Niche, Nestier, and Gerde (Riss); it then cooccurs with the arctic fox Alopex lagopus (Linnaeus, 1758) throughout Würmian times, before it becomes a common carnivore during Holocene times ( Clot & Duranthon 1990 ). Morphometrical similarities and potential co-occurrence of red and arctic foxes during the Oxygen Isotopic Stage 3 (OIS3; Cohen & Gibbard 2011 ) necessitate performing a careful analysis of the bone material. At Fouvent, cranial and dental elements are represented by a left mandibular branch with p2-m2, a fragment of left mandible with m1 and m2, and a broken right mandible with m1 and m2 with two canines. As regards the red fox, according to Poplin (1976) , the lower canines are longer and thinner in comparison with isatis. At Gerde, Clot (1980) also pointed out the great difficulty to make the distinction between Vulpes and Alopex but determined the canines of fox based on their general proportions. We attributed the material of Fouvent to the red fox on the base of the concerned morphometrical data. Our three m1s (G8.B.509b, D11.E.3 and G8.B.507) leave no doubt for their specific assignment. Their large dimensions (respectively B = 6.4-6-6 with L = 16- 16-16) match perfectly those of red foxes as provided at Gerde ( Clot 1980 : table 49) and at Gönnersdorf ( Poplin 1976: 48 fig. 28). Postcranial elements of red fox include two humeri, one ulna, one metacarpal II, two tibiae, one calcaneus and one metatarsal V. The measurements are summarized in Table 2 . According to Altuna (2004) , the breadth of the distal humerus of the red fox ranges between 17.2 and 23.8 mm while Clot (1980) gives an interval comprised between 19.2 and 22 mm . According to these studies, the distal breadth of isatis is not greater than 19.5 mm , consequently the humerus of Fouvent (1842.118) is quite similar in size to the red fox. Published metric data available for the ulna are rare but the dimensions of the specimen of Fouvent suggest without no doubt an attribution to Vulpes . The total length ( 47 mm ) of the left metacarpal II(G8.A/B.165) of Fouvent is larger in comparison to the measurements of red fox ( Altuna 2004 ). However, the left metacarpal falls into A FIG. 3. — Cave hyena Crocuta crocuta spelaea (Goldfüss, 1832) remains from Fouvent-Saint-Andoche: A -D , left mandibles 1842.16 ( A , B ) and G6.82 ( C , D ); E -J , right G7.168 ( E , F ), H9.B.72 ( G , H ) and left G6.9 ( I , J ) lower carnassials; K -P , left upper carnassial F11.C.281 ( K , L ), no number ( M , N ) and G9.B.444 ( O , P ); Q , R , left humerus 1842.10; S -T , left radius 1842.90; U -V , right tibia 1842.141. Scale bar: 20 mm. Pictures by JBF. TABLE 2. — Postcranial measurements of Vulpes vulpes Linnaeus,1758 from Fouvent-Saint-Andoche (in mm). Abbreviations: *, dimensions of the tuber calcanei; sin , left; dext , right. Other abbreviations: see text.

N°	Element	Side	GL	H incis	B incis	Bp	Lp	Bd	Ld
1842.118	Humerus	Sin	–	–	–	–	–	20.0	15.0
E10.B.10	Ulna	Sin	–	14.0	10.0	–	–	–	–
G8.A/B.165	2nd Metacarpal	Sin	47.0	–	–	5.0	7.0	6.5	5.5
1842.119	Tibia	Sin	–	–	–	–	–	16.0	12.0
1842.92	Tibia	Dext	–	–	–	23.5	25.0	–	–
G8.E.26	Calcaneus	Dext	29.0	–	–	6.5*	9.8*	–	–
F9.B.135	5th Metatarsal	Sin	56.0	–	–	4.0	8.0	6.0	6.0

TABLE 3. — Measurements of the Vth metatarsal of red and polar foxes, from Fouvent-Saint-Andoche ( Fvt ) and various localities (fossil and modern), in mm. Abbreviations see in text.

Site	GL	Bp	Lp	Bd	Ld	Reference
Taxon	Fvt 90-92 G8.B.288	51.0	6.0	8.0	5.7	5.0	this study
Aurensan inférieur	50.6	6.6	7.1	–	4.9	Clot 1980
Predmost	46.0-48.0 (n=2)	5.0-5.7 (n=2)	6.5-6.4 (n=2)	5.2-5.7 (n=2)	5.0-5.0 (n=2)	Clot 1980
A. lagopus	Prolom 2 (OIS3)	42.7-47.5 (n=5)	4.5-6.6 (n=13)	–	4.7-5.8 (n=5)	4.1-4.7 (n=5)	Baryshnikov 2006
Siuren 1 (OIS3)	44.9-50.0 (n=3)	5.0-6.1 (n=3)	–	5.6-5.8 (n=3)	4.4-4.9 (n=3)	Baryshnikov 2006
Modern	42.0-51.0 (n=9)	–	–	–	–	Altuna 2004
Coulet des Roches	56.8-66.5 (n=7)	6.7-8.0 (n=8)	7.6-9.0 (n=8)	5.7-7.3 (n=8)	5.5-6.2 (n=8)	Crégut-Bonnoure
V. vulpes	(OIS2) Gerde (OIS3)	56.5-65.2 (n=7)	6.0-7.8 (n=14)	6.3-8.3 (n=14)	5.9-6.8 (=8)	5.7-6.2 (n=8)	et al. 2010 Clot 1980
Modern	49.0-68.0 (n=26)	–	–	–	–	Altuna 2004

the variation range as provided by Clot (1980) for Pleistocene populations of Vulpes . The same is true for the measurements of both a proximal and a distal tibiae (respectively n°1842.119 and 1842.92) that we also assigned to the red fox. In the Table 3 , the maximum length ( 56 mm ) of the left metatarsal V (F9.B.135) is roughly similar to the mean value observed in living foxes ( Altuna 2004 ) corresponding to their smaller pleistocene representatives ( Clot1980 ).We observed the same thing with the maximum length of the calcaneus G8.E.26 (only 29 mm ) that could be associated to a small red fox, while populations of isatis only reach a maximum of 28.4 mm .