Crocuta crocuta spelaea (Goldfüss, 1832)

Fourvel, Jean-Baptiste, Fosse, Philippe, Fernandez, Philippe & Antoine, Pierre-Olivier, 2015, Large mammals of Fouvent-Saint-Andoche (Haute-Saône, France): a glimpse into a Late Pleistocene hyena den, Geodiversitas 37 (2), pp. 237-266 : 248-250

Crocuta crocuta spelaea (Goldfüss, 1832)

MATERIAL EXAMINED. — NISP =850; MNI =121.

1842 sample: 2 left mandibles; 1 atlas; 3 cervical vertebrae; 3 thoracic vertebrae; 1 left humerus; 1 left radius; 1 left ulna; 1 left metacarpal II; 1 right metacarpal V; 1 right innominate; 1 left innominate; 1 left femur; 2 right tibiae; 1 right calcaneus; 1 left calcaneus; 1 left talus; 1 right metatarsal IV; 1 right metatarsal V.

1989-1992 sample: 6 skull fragments (occipital); 2 right maxillae; 1 left maxilla; 4 right DI3; 6 left DI3; 8 right DC; 6 left DC; 5 right D2; 3 left D2; 4 right D3; 14 left D3; 3 right D4; 6 left D4; 1 right I1; 2 left I1; 2 right I1-2; 2 left I1-2; 15 right I2; 17 left I2; 16 right I3; 9 left I3; 8 right C; 8 left C; 7 right P1; 13 left P1; 8 right P2; 6 left P2; 9 right P3; 14 left P3; 14 right P4; 20 left P4; 11 right mandibles; 8 left mandibles; 5 mandibles?; 2 right dc; 7 left dc; 5 right d2; 9 left d2; 10 right d3; 11 left d3; 11 right d4; 20 left d4; 2 right d1; 5 left i1; 1 left i1-2; 5 right i2; 5 left i2; 1 right i2-3; 1 left i2-3; 20 right i3; 21 left i3; 22 right c; 21 left c; 7 left p2; 21 left p3; 20 right p3; 34 right p4; 35 left p4; 33 right m1; 23 left m1; 140 tooth fragments; 1 atlas; 1 cervical vertebra; 2 lumbar vertebrae; 4 caudal vertebrae; 3 right humeri; 4 left humeri; 2 left radius; 1 radius; 1 left ulna; 1 right scapholunar; 1 left scapholunar; 1 left pisiform; 1 pisiform; 2 right metacarpals II; 1 femur?; 1 patella?; 1 left tibia; 1 tibia?; 1 right fibula; 1 left fibula; 2 right tali; 1 left cuboid; 1 right metatarsal II; 1 left metatarsal II; 1 left metatarsal III; 1 left metatarsal V; 3 sesamoids?; 2 metapodials?; 9 first phalanges; 6 second phalanges; 6 third phalanges.

DESCRIPTION Cave hyena is represented by 850 bone remains originating from both 1842 (NISP=24) and 1989-1992 (NISP=826) samples. This material represents about 6% of the complete collection,

about 25% of the NISP and 77% of the carnivores.Cave hyena remains are related to 121 individuals, ranging from juvenile to old adults (Fourvel et al. submitted); this series is among the largest ones for Late Pleistocene cave hyenas in Europe. Morphological characters ( Fig. 3A, V ) and measurements of both teeth and postcranial elements ( Table 8) recovered in Fouvent closely match the descriptions of late Pleistocene Crocuta crocuta spelaea published in the literature ( Reynolds 1902; Clot 1980; Testu 2006; Barycka 2008). The extreme morphological variability of referred dental remains most probably coincides with ecomorphotypic variations, thus precluding any use of cave hyenas in a biochronological purpose (e.g., Kurtén 1963; Kurtén & Poulianos 1977; Klein & Scott 1989; Baryshnikov 1999). However, Würmian cave hyenas seem to differ from other representatives of Crocuta in having a large size, especially as regards dentition. In Fouvent, upper and lower carnassials are at the same time large, robust, and highly metrically variable. Figure 7 shows breadth/length ratios for upper and lower carnassials (P4: Fig. 7A; m 1 : Fig. 7B ) from Fouvent, compared to various Late Pleistocene and modern spotted hyena populations. Hyena teeth from Fouvent reveal strong size variability. However, this material has more large dimensions (B/L) than Middle Pleistocene hyenas ( Crocuta crocuta intermedia from Lunel-Viel 1 in Bonifay 1971; Crocuta crocuta praespelaea from Petralona in Kurtén & Poulianos 1977, 1981) and similar proportions than many OIS3 populations (Jaurens in Ballesio 1979; Cueva de las Hienas in Fosse 1997; Labeko Koba in Altuna & Mariezkurrena 2000; Conives in Fourvel 2008). Other teeth and postcranial elements are in perfect agreement with the observation as detailed here above. Morphological and metric features of the Fouvent cave hyena coincide with the large and robust hyenas as recorded in the OIS3 (between more or less 57 and 29 ky; Fourvel 2012).

Super-Order UNGULATA Linnaeus, 1766 [description by JBF, PFos, PFer, POA]

Eight ungulate species have been recognized (2219 NISP, 192 MNI). The collection is truly remarkable with a high frequency of mega-herbivores (8 mammoths and 39 woolly rhinoceroses) and large species (16 giant deers, 31 large bovids and 62 horses) ( Figs 8 , 9 ).

Order PROBOSCIDEA Illiger, 1811

Family ELEPHANTIDAE Gray, 1821

Subfamily ELEPHANTINAE Gray, 1821

Genus Mammuthus Brookes, 1828

Figures

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.

FIG. 7. — Cave hyena C. c. spelaea (Goldfüss, 1832): Upper (P4) and lower (M1) carnassial Breadth and Length ratios compared to extant and extinct populations. Measurements in mm. Confidence ellipses with confidence interval (0.95) are figured.

FIG. 1. — Geographical location of Fouvent-Saint-Andoche, NE France. Map realized using Geoatlas software and Geoportail open-access site.

FIG. 8. — Woolly rhinoceros Coelodonta antiquitatis (Blumenbach,1799) remains from Fouvent-Saint-Andoche (Coelodonta antiquitatis): A, D1; B-E, D2; F-H, D3; I-N, D4; O. d2; P, Q, d3; R-T, d4; U, V, M1; W, X, M2; Y, M3; Z, AA, humerus shaft cylinders; AB, AC, tibia shaft cylinders. Scale bar: 20 mm. Pictures by JBF.

FIG. 9.— Ungulates (other than rhinoceros) remains from Fouvent: A, B, giant deer Megaloceros giganteus Blumenbach,1799:right 1842.14 (A) and left 1842.13 (B) mandibles; C-E, red deer Cervus elaphus Linnaeus, 1758: basal part of right antler 1842.109 (C) and ingested left talus 1842.97 (D, palmar view; E, dorsal view); F-H, horse Equus germanicus Nehring,1884:right metacarpal 1842.2 (F), first 1842.6 (G) and third H9.A.1 (H) phalanges;I-L, large bovid Bovidae:right calcaneum 1842.78 (I), first phalange G10.E/A.9 (J, dorsal view; K, lateral view) and metapodial distal portion F11.A.50 (L); M-O reindeer Rangifer tarandus (Linnaeus, 1758): basal part of left antler 1842.106 (M), portion of right antler F10.B.329 (N) and right metatarsal G9.24 (O). Scale bar: 20 mm. Pictures by JBF.