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Ecological interpretations of early (Mammalia, Cervidae) from Ceyssaguet (Haute-Loire, France)

Thomas M. KAISER Zoological Institute and Museum, University Greifswald, J. S. Bach Str. 11-12, D-17489 Greifswald (Germany) [email protected]

Roman CROITOR Institute of Zoology, Academy of Sciences of Moldova, Academiei str. 1, Kishinau 2028 (Moldova) [email protected]

Kaiser T. M. & Croitor R. 2004. — Ecological interpretations of early Pleistocene deer (Mammalia, Cervidae) from Ceyssaguet (Haute-Loire, France). Geodiversitas 26 (4) : 661-674.

ABSTRACT The mesowear method is applied to reveal the type of forage eaten by the late Villafranchian deer of Ceyssaguet (Haute-Loire, France). Further, habitat properties of the paleoenvironment of Ceyssaguet are reconstructed. The most common deer , Eucladoceros ctenoides Nesti, 1841 and Metacervoceros rhenanus Dubois, 1904, are interpreted as abrasion-dominated mixed feeders with close to 45% grass in their forage which was similar to that KEY WORDS of the south east Asian hog deer and sambar. A third deer species, Mammalia, Cervidae, Praemegaceros obscurus Azzaroli, 1953, represented only by three speci- early Pleistocene, mens, is interpreted as a more attrition dominated mixed feeder. Combined France, with mesowear analysis, taphonomical and morpho-functional data allow the mesowear analysis, paleodiet reconstruction, reconstruction of the paleoenvironment of Ceyssaguet as an open grassland paleoenvironment. with wooded habitats near the lake, under cold climate.

GEODIVERSITAS • 2004 • 26 (4) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 661 Kaiser T. M. & Croitor R.

RÉSUMÉ Interprétations écologiques des cerfs (Mammalia, Cervidae) du Pléistocène inférieur de Ceyssaguet (Haute-Loire, France). L’étude de l’usure de niveau intermédiaire (« mesowear ») est appliquée ici pour rechercher le type d’alimentation consommée par les cerfs villafran- chiens de Ceyssaguet (Haute-Loire, France). Les espèces les plus communes, Eucladoceros ctenoides Nesti, 1841 et Metacervoceros rhenanus Dubois, 1904, sont interprétées comme brouteuses d’herbe, se nourrissant principalement d’herbes fraîches, comme les cerfs asiatiques tels que le cerf-cochon et le MOTS CLÉS sambar. Une troisième espèce de cerf, Praemegaceros obscurus Azzaroli, 1953, Mammalia, Cervidae, qui n’est représentée que par trois spécimens, est interprétée comme brouteur Pléistocène inférieur, mixte. Combinée à l’analyse de l’usure de niveau intermédiaire, les données France, taphonomiques et morphofonctionelles permettent de reconstruire le usure de niveau intermédiaire, reconstruction du paléorégime, paléoenvironnement de Ceyssaguet comme une prairie ouverte, associée à des paléoenvironnement. habitats forestiers au bord du lac, sous un climat froid.

INTRODUCTION Ferrand; M.-F. Bonifay pers. comm.). This is in agreement with the biochronological age THE CEYSSAGUET SITE (Bonifay 1991, 1995). The formation of the The paleoecological reconstruction of the termi- fossiliferous loess deposits underlied by basalt is nal Villafranchian period of Europe is of parti- estimated to cover 0.1 Ma (M.-F. Bonifay pers. cular interest as it allows to reveal the dramatic comm.). This results in an age estimate of climate shifts on the continent resulting in faunal 1.2 Ma for the Ceyssaguet fauna (Bonifay 1991). changes in the Quaternary period. Herbivorous The lower fossiliferous stratum yielded the are useful in paleoenvironmental majority of mammalian , many of which reconstruction and can provide unique informa- are represented by articulated skeletons. This tion on diet and thus allow reconstruction of suggests that the fossil remains belong to carcass- elements of the paleoflora and the character of a es that were initially buried in the lake deposits. fossil habitat (e.g., Vrba 1975; Cerling et al. The upper stratum has yielded isolated and often 1997; MacFadden & Cerling 1998; Kaiser fragmented bones with traces of predation 2003). (Bonifay 1986). In the present work, the mesowear method (Fortelius & Solounias 2000; Kaiser et al. 2000; MAMMALS ANALYSED Kaiser & Solounias 2003) is applied as a tool to The Cervidae are the predominant reconstruct the paleodiet of herbivorous mam- group in the Ceyssaguet mammalian fauna. mals in order to infer the type of forage available Among the four recognized deer species (Croitor at the late Villafranchian site of Ceyssaguet & Bonifay 2001), the largest number of speci- (Haute-Loire, France) and to infer the paleoenvi- mens belongs to a large-sized species, Eucladoceros ronment of this site. ctenoides Nesti, 1841 (Fig. 1A). A small-sized The fossil site of Ceyssaguet formed in a lake deer species, Metacervoceros rhenanus Dubois, within a volcano crater (Bonifay 1991, 1995), 1904 (Fig. 1B), is represented by 10 specimens dated by K/Ar at 1.3 Ma (unpublished data from (one shed antler, seven dentition remains and the Geochronology Laboratory of Clermont- two postcranial bones, a femur and a tibia) in the

662 GEODIVERSITAS • 2004 • 26 (4) Ecology of Pleistocene deer

FIG. 1. — Habitus of Eucladoceros ctenoides Nesti, 1841 (A), Metacervoceros rhenanus Dubois, 1904 (B) and Praemegaceros obscurus Azzaroli, 1953 (C) from Ceyssaguet (Haute-Loire, France). The reconstruction of P. obscurus is based on material from the site of Pietrafitta (Italy). Scale bar: 50 cm.

GEODIVERSITAS • 2004 • 26 (4) 663 Kaiser T. M. & Croitor R.

Ceyssaguet. All deer species included in this study went extinct on the European mainland before the first waves of glaciation. Due to the limited material of fallow deer from Ceyssaguet, this species is not considered here. A brief description of the species is in progress. – Eucladoceros ctenoides (Ec) (Fig. 1A) is similar in size to a modern . Based on the male skull from Ceyssaguet, the body mass is esti- mated 250 kg applying methodology by Janis (1990). It is characterised by comparatively simply build, light antlers and a narrow, long braincase. Compared to E. dicranios Nesti, 1841, the preorbital fossae are reduced and very small. The antlers are comb-like with up to six tines of variable shape inserted on the antler beam in the same plane. Upper molars have no additional enamel folds beside the hypocone spur. The internal enamel of the FIG. 2. — Proximal epiphysis of the femur (both found in arti- culation with tibiae and metatarsals of deer): Eucladoceros hypocone of upper premolars is well folded. The ctenoides Nesti, 1841 (A), Praemegaceros obscurus Azzaroli, 1953 (B); posterior aspect of the femur head (1), proximal aspect lingual wall of the P2 is cleft. As a primitive of the femur head (2). Note the long and medially prominent character the p4 shows a low degree of molariza- caput femoris (cap. fem.) in E. ctenoides and the more sharp- ened and prominent lesser trochanter (tr. min.) in P. obscurus tion. In some cases, however, an advanced mor- (indicated by arrow). Scale bar: 10 cm. phology of the p4 has been reported (Azzaroli & Mazza 1992; De Vos et al. 1995; Croitor & Bonifay 2001). fossil assemblage. Praemegaceros obscurus Azzaroli, Croitor & Bonifay (2001) pointed out some 1953 (Fig. 1C) is another large-sized deer in the characters of E. ctenoides and P. obscurus that Ceyssaguet fauna, but is quite rare. Only two allow distinguishing the postcranial morphology specimens, a radiocubitus (No. 2-11010) and a of these two species, including the peculiar char- fragment of an upper mandible (No. 2-10332) acters of the femur. In E. ctenoides (Fig. 2A1, were ascribed to an ancient fallow deer sp. A2), compared to P. obscurus (Fig. 2B1, B2), the The paucity of the material does not allow a reli- caput femoris is comparatively long and more able species identification, however it has been prominent medially, while the lesser trochanter is supposed that the fallow deer from Ceyssaguet more prominent and extended posteriorly. may be conspecific with Dama vallonnetensis De Unlike P. obscurus and the closely related Lumley, Kahlke, Moigne & Moulle, 1988 from E. dicranios the distribution of E. ctenoides during the latest Villafranchian-Postvillafranchian of the latest Villafranchian seems to be limited to Italy and southern France (Croitor & Bonifay northwest Europe (France, Holland) (Croitor & 2001). Both late Villafranchian Praemegaceros Bonifay 2001). obscurus and Dama vallonnetensis are represented The material used in this analysis comes mainly by abundant fossil remains in the meridional from the second level of the Ceyssaguet strati- areas of Europe as southern France and Italy graphic column, which yielded the largest num- (Abbazzi 1995; Croitor 2001a; Croitor & ber of cervid remains in the fossil sample Bonifay 2001). The scanty remains of the above (Bonifay 1986). All individuals of E. ctenoides mentioned deer in the studied sample therefore from Ceyssaguet including the two preserved indicate that it was a less common species at skulls of male individuals, have their antlers shed.

664 GEODIVERSITAS • 2004 • 26 (4) Ecology of Pleistocene deer

This suggests that E. ctenoides inhabited the local- the body mass is estimated 400 kg. Its size thus ity of Ceyssaguet mainly during the springtime more or less equals that of modern . Only nine and summer season when shed their remains were identified as belonging to P. obscu- antlers. Only three of 16 antlers show rodent rus. The species is characterized by large and gnawing marks. This suggests that most of the robust antlers with a strong basal tine and a very antlered individuals died during the winter sea- long and flat second tine inserted on the medial son, when their carcasses were not subject to side of the beam (Abbazzi 1995; Abbazzi et al. rodent activity. 1999). Compared to E. ctenoides the dentition is – Metacervoceros rhenanus (Mr) (Fig. 1B) is a more advanced. The p4 is well molarized, and rather small deer, comparable in size to the mod- there are no additional enamel folds in the upper ern fallow deer. Based on the male skull from cheek teeth. The caput femoris is short and Ceyssaguet, the body mass is estimated 70 kg rounded compared to E. ctenoides (Croitor & applying methodology by Janis (1990). It has a Bonifay 2001). Abundant finds of P. obscurus long neurocranium and a short orbitofrontal were discovered in England and areas of conti- region combined with long posteriorly sloping nental Europe, like Italy, Romania and Moldavia pedicles (Croitor & Bonifay 2001). The preor- (Abbazzi 1995; Abbazzi et al. 1999; Croitor & bital fossae are deep and large. The antlers are Bonifay 2001). slender, and three-pointed. The basal tine springs off high above the burr. Upper molars have a hypocone spur, a protocone fold and a weak cin- MATERIAL AND METHODS gulum. The lower premolars, including p4, are comparatively simply built. The distribution of MESOWEAR ANALYSIS M. rhenanus during the latest Villafranchian peri- The mesowear method was developed by od mainly coincides with the distribution area of Fortelius & Solounias (2000). Mesowear is based E. ctenoides and these two species are often found on facet development on the occlusal surfaces of together (Croitor & Bonifay 2001). the ungulate upper molar teeth. The degree of M. rhenanus is represented by individuals of vari- facet development reflects the relative propor- ous ontogenetic stages at Ceyssaguet. However, tions of tooth to tooth contact (attrition) and the fossil remains of this species are much less food to tooth contact (abrasion), attrition creat- numerous compared to E. ctenoides. Among ing facets and abrasion obliterating them. This 26 antlers, nine are connected with pedicles methodology defines ungulate tooth mesowear (35%). Ten of 17 shed antlers are gnawed by by two variables: 1) occlusal relief and 2) cusp rodents (about 60%), while only two of nine shape. Occlusal relief (OR) is classified as high antlers which are inserted in pedicles are gnawed. (h) or low (l), depending on how high the cusps It seems therefore that the majority of antlers still rise above the valley between them. Occlusal connected to the skull were unavailable to relief is used in the analyses as percentages % rodents after the death of their owners. This is high and % low (Table 1). The second mesowear possibly because these individuals drowned in the variable, cusp shape, includes three scored attrib- water during the winter season. It should be utes: sharp (s), round (r) and blunt (b) according noted, that the taxon Metacervoceros is often cited to the degree of facet development (see Fortelius as s.l. or Pseudodama Azzaroli, 1992 & Solounias 2000: fig. 1). Cusp shape is also (Croitor & Bonifay 2001). Croitor (2001a) used as a percentage and is given in Table 1 as the restricted the genus Pseudodama only to its type three variables % sharp, % round and % blunt. species Dama nestii nestii Azzaroli, 1947. When both (paracone and metacone) cusp apices – Praemegaceros obscurus (Po) is the largest deer were preserved, the sharpest of the two was species discovered at the Ceyssaguet site (Fig. 1C). selected for analysis as proposed by Fortelius & Based on the male skull from Pietrafitta (Italy), Solounias (2000).

GEODIVERSITAS • 2004 • 26 (4) 665 Kaiser T. M. & Croitor R.

TABLE 1. — Distribution of mesowear variables in the populations of Eucladoceros ctenoides Nesti, 1841 (Ec), Metacervoceros rhenanus Dubois, 1904 (Mr) and Praemegaceros obscurus Azzaroli, 1953 (Po) from Ceyssaguet (Haute-Loire, France). Abbreviations: n, number of available specimens; Mesowear variables: l, absolute scorings low; h, absolute scorings high; s, absolute scorings sharp; r, absolute scorings round; b, absolute scorings blunt; % l, percent low occlusal relief; % h, percent high occlusal relief; % s, percent sharp cusps; % r, percent rounded cusps; % b, percent blunt cusps.

Species n l h s r b % l % h % s % r % b

Eucladoceros ctenoides 17 1 16 3 14 0 6 94 18 82 0 Metacervoceros rhenanus 12 012290010018820 Praemegaceros obscurus 303110010050500

Ec, Mr, Po; h,s,r Chi-square = 1.2562, df = 4, p-value = 0.8688 Ec, Mr; h,s,r Chi-square = 0.0751, df = 2, p-value = 0.9632 Ec, Po; h,s,r Chi-square = 1.1592, df = 2, p-value = 0.5601 Mr, Po; h,s,r Chi-square = 0.9577, df = 2, p-value = 0.6195 Ec, Mr, Po; l,h Chi-square = 0.9108, df = 2, p-value = 0.6342 Ec, Mr; l,h Chi-square = 0.0317, df = 1, p-value = 0.8586 Ec, Po; l,h Chi-square = 1.0114, df = 1, p-value = 0.3146

Fortelius & Solounias (2000) restricted their (2000) were used. The set of fossil taxa was plot- study of ungulate mesowear to the labial wall of ted within a nested set of extant ungulate species. the M2. This methodology was modified in Extant species were classified into the three broad selecting all M1-M3s of the species investigated dietary categories: browser, mixed feeder and following the “extended” mesowear method grazer, following the “conservative” (CONS) introduced by Kaiser & Solounias (2003). This classification of Fortelius & Solounias (2000). All methodology was chosen because it allows to statistics were computed using Systat 9.0 and include tooth samples with small numbers of Axum 6 software. Hierarchical cluster analysis individuals. It is recognized that applying the with complete linkage (furthest neighbour) was modified method by Kaiser & Solounias (2003), applied following the standard hierarchical amal- that was originally developed for use in hyp- gamation method of Hartigan (1975). According sodont equids may not be immediately trans- to the default settings of Systat 9.0 the algorithm ferred to dentitions. Recent studies by of Gruvaeus & Weiner (1972) was used to order Kaiser & Franz-Odendaal (2003) however indi- the trees. The three mesowear variables % high, cate, that combined analysis of upper molars % sharp and % blunt were analysed in cluster sta- into mesowear investigation results in a valid tistics. We perform three cluster analysis using mesowear signal in . different sets of extant reference species. In Dental specimens included in this study are Figure 4A we used all 54 extant species classified upper cheek teeth showing wear stages 2 and 3 of as “normal” and “typical” by Fortelius & Solou- Kaiser et al. (2003). Unworn teeth, specimens in nias (2000), in Figure 4B we use the reduced very early wear, and very worn teeth were exclud- set of 27 “typical” species and in Figure 4C we ed. The tips of labial cusp apices of upper molars only employ nine extant Cervidae as reference are broken in some specimens. Those specimens species. were also excluded from the analysis. In the end, Chi-square corresponding probabilities were 17 specimens of Eucladoceros ctenoides, 12 speci- computed for each dataset giving the probability mens of Metacervoceros rhenanus and three speci- that the null hypotheses of independence should mens of Praemegaceros obscurus were available for be rejected (at an error probability of 0.05). The this investigation (Table 2). absolute frequencies of mesowear variables As comparative data for dietary classification, 54 (“low”, “high”, “sharp”, and “round”) were tested extant species reported by Fortelius & Solounias for significance.

666 GEODIVERSITAS • 2004 • 26 (4) Ecology of Pleistocene deer

TABLE 2. — Maxillary cheek teeth attributed to Eucladoceros ctenoides Nesti, 1841 (Ec), to Metacervoceros rhenanus Dubois, 1904 (Mr) and to Praemegaceros obscurus Azzaroli, 1953 (Po) from Ceyssaguet (Haute-Loire, France). Abbreviations: Spec-Id, specimen number (Cey, Maison méditerrannéenne des Sciences de l’Homme, Aix-en-Provence, France); Tooth, tooth position; Side, side (r, right; l, left); OR, occlusal relief mesowear variables scoring (h, high; l, low); CS, cusp shape mesowear scores (s, sharp; r, round; b, blunt); (P), paracone cusp; (M), metacone cusp.

Spec-Id Species Tooth Side OR CS (P) CS (M)

Cey-8194 Ec M1 r h r r Cey-4534 Ec M1 r h r r Cey-7217 Ec M1 l h – r Cey-10590 Ec M1 r h r r Cey-2047 Ec M1/2 l h – r Cey-12072 Ec M1/2 r l r r Cey-6303 (12398) Ec M1/2 l h r r Cey-2292 Ec M2 r h s s Cey-8194 Ec M2 r h r r Cey-4534 Ec M2 r h r r Cey-7217 Ec M2 l h r r Cey-10590 Ec M2 r h s r Cey-8194 Ec M3 r h r r Cey-4534 Ec M3 r h r r Cey-7217 Ec M3 l h s r Cey-10590 Ec M3 r h – r Cey-nn/a Ec M3 l h r r Cey-14393 Mr M1 l h – r Cey-9412 Mr M1 r h – – Cey-84-2176 Mr M1 r h r r Cey-93-10755-a Mr M1/2 l h r – Cey-93-10755-b Mr M1/2 l h – r Cey-14393 Mr M2 l h r r Cey-12303 Mr M2 r h s s Cey-9412 Mr M2 r h r r Cey-84-2176 Mr M2 r h – s Cey-n/n-b Mr M2 r h r r Cey-14393 Mr M3 l h r r Cey-n/n-b Mr M3 r h r r Cey-8949 Po M1 l h – – Cey-8949 Po M2 l h r r Cey-8949 Po M3 l h – s

RESULTS ables do not permit separation of the populations (p-values range between 0.31 and 0.86). In all three fossil cervid species compared in this The cluster diagrams (Fig. 4) depict the relations analysis (Ec, Mr and Po), occlusal relief is at of datasets by joining them in the same clusters. least 94% high. Cusp shape scorings ranged The closer the data are, the smaller is the between 18% (Ec and Mr) and 50% (Po) sharp, Euclidean distance (ED) at the branching point. 82% (Ec and Mr) and 50% (Po) round. There Figure 4A shows three main clusters, cluster one are no blunt cusps in either sample (Table 1; containing only grazers, cluster two containing Fig. 3). one part of the mixed feeders some of the grazers Chi-square analysis of all variables, however, and very few browsers, and cluster three corre- indicates low significance of differences (p = sponding to the attrition-dominated end of the 0.56-0.96). Eucladoceros ctenoides is most similar spectrum containing most of the browsers and to Metacervoceros rhenanus as indicated by p = the attrition-dominated mixed feeders, but no 0.96. Taken alone, also the occlusal relief vari- grazers.

GEODIVERSITAS • 2004 • 26 (4) 667 Kaiser T. M. & Croitor R.

Eucladoceros ctenoides (Ec) Metacervoceros rhenanus (Mr) Praemegaceros obscurus (Po) 100 A B C 80

60

40

20

0 lhsrblhsrblhsrb

Redunca redunca (rr) ellipsiprymnus (ke) (To) 100 D E F 80

60 G(G)G(G) M(M)

40

20

0 lhsrblhsrblhsrb

Cervus unicolor (Cu) Axis porcinus (Ap) scriptus (Ts) 100 GJH 80

60 M(M) M(G) M(M)

40

20

0 lhsrblhsrblhsrb

Axis axis (Ax) Ourebia ourebi (Oo) Cervus canadensis (Cc) 100 KML 80

60 M(G) M(G) M(M)

40

20

0 lhsrblhsrblhsrb

Cervus duvauceli (Cd) hemionus (OH) Alces alces (AA) 100 NPO 80

60 M(G) B(B) B(B)

40

20

0 lhsrblhsrblhsrb

FIG. 3. — Histograms of mesowear variables % low (l), % high (h), % sharp (s), % round (r) and % blunt (b). Histograms of Eucladoceros ctenoides Nesti, 1841 (A), Metacervoceros rhenanus Dubois, 1904 (B) and Praemegaceros obscurus Azzaroli, 1953 (C) are based on the data given in Table 1. Comparative histograms are based on published data by Fortelius & Solounias (2000). Redunca redunca (bohor ) (D), Kobus ellipsiprymnus (common ) (E), Taurotragus oryx (eland) (F), Cervus unicolor (sambar) (G), Axis porcinus (hog deer) (H), Tragelaphus scriptus (bushbuck) (J), Axis axis () (K), Ourebia ourebi () (L), Cervus canadensis (wapiti) (M), Cervus duvauceli (barashingha) (N), Odocoileus hemionus () (O), Alces alces (elk, ) (P). Letters in centre of graphs indicate dietary trait after Fortelius & Solounias (2000). Abbreviations: G, grazer; M, mixed feeder; B, browser. First character: CONS (conservative) classification; second character (in brackets): RADI (radical) classification after Fortelius & Solounias (2000).

668 GEODIVERSITAS • 2004 • 26 (4) Ecology of Pleistocene deer

The two species (Ec and Me) are in a sub-cluster The cluster models show a quite distant position of cluster two (ED = 24), containing mostly abra- of the deer from Ceyssaguet and the modern hol- sion-dominated mixed feeders and grazers, but arctic species like capreolus (Linnaeus, only two browsers (Fig. 4A). Within this cluster, 1758) (OL), Alces alces (Linnaeus, 1758) (AA) the two deer species are closely linked to Ourebia and Odocoileus hemionus (Rafinesque, 1817) ourebi Zimmermann, 1783 (Fig. 3L), an African (OH), which are linked with browsers. However, bovid, whose diet consists of up to 80% grass E. ctenoides and M. rhenanus share remarkable (Van Wieren 1996). Praemegaceros obscurus is in similarities with extant deer species from South- the major cluster three (ED = 42). There it shares East Asia (Fig. 4C). a sub-cluster of fourth order (ED = 14.4) with 12 mixed feeders and one browser. It is closest OBSERVATIONS ON TOOTH TIP DAMAGE linked to the mixed feeding bovid Taurotragus The tips of labial cusp apices of upper molars are oryx Pallas, 1766 (To) (Fig. 3F). broken in some specimens. This type of damage If cluster analysis is restricted to those extant of cusp tips is often recorded in M. rhenanus, comparison species, recognised to represent “typ- both in upper and lower cheek teeth. Damaged ical” dietary categories by Fortelius & Solounias cusp tips are also found in some specimens of (2000), the pattern becomes even clearer E. ctenoides. The broken tooth tips were freshly (Fig. 4B). Clusters one and two contain only damaged or slightly polished by food, a fact grazers. Ec and Mr are in cluster two. All mixed that suggests that the tips were damaged during feeders share cluster three, which also contains the ’s life shortly before its death. Since Po. All browsers are within cluster four. Within a damaged cusp tips are not the result of stable sub-cluster of ED = 16, the common waterbuck mesowear equilibriums, those cusps were exclud- (Kobus ellipsiprymnus Ogilby, 1833; Fig. 3E) (ke), ed from mesowear analysis. (Redunca redunca Pallas, 1767; Fig. 3D) (rr), roan (Hippotragus equinus Desmarest, 1804) (he) and DISCUSSION (Hippotragus niger Harris, 1838) (hn), all grazers, are the extant species classified next to Ec The mesowear signal of E. ctenoides and (Fig. 3A) and Mr (Fig. 3B). P. obscurus (Fig. 3C) M. rhenanus approaches the signals of the mod- remains closely linked to mixed feeders as eland ern African bovids Redunca redunca and Kobus (Taurotragus oryx; Fig. 3F) (To), and ellipsiprymnus, which live near rivers and feed on (Aepyceros melampus Lichtenstein, 1812) (Me). grass (Kingdon 1979). In both species grass Reducing the set of extant species to the makes up 80% of the diet (Van Wieren 1996). Cervidae, results in only two major clusters Gagnon & Chew (2000) classify these species as (Fig. 4C). Cluster one comprises all the browsers, obligate or variable grazers respectively. It is one mixed feeder (wapiti) and Po from interesting to note that among the extant Ceyssaguet. Cluster two comprises four extant Cervidae the mixed feeding (wapiti; Fig. 3M) cervids and Ec and Mr from Ceyssaguet. Here Ec and not the grazing species as the hog deer (Axis and Mr are closest to the sambar (Cervus unicolor porcinus Zimmermann, 1780; Fig. 3H), the chi- Kerr, 1792; Fig. 3G) (Cu) and the hog deer (Axis tal (Axis axis Erxleben, 1777; Fig. 3K) (70% porcinus Zimmermann, 1780; Fig. 3H) (Ap). grass) or the barashingha (Cervus duvauceli Following the “conservative” (CONS) classifica- Cuvier, 1823) (80% grass) are the extant dietary tion by Fortelius & Solounias (2000) all extant analogues. However, it should be noted here, that species in cluster two are mixed feeders. If the all these species are classified as “normal” by “radical” (RADI) classification scheme is applied, Fortelius & Solounias (2000), which indicates the sambar is the only mixed feeder, while the that they found their mesowear signatures to not remaining three taxa are classified as grazers. always correctly correspond to the known dietary

GEODIVERSITAS • 2004 • 26 (4) 669 Kaiser T. M. & Croitor R.

bb A cs 1 eb eg dl ab ct Cd Ax hn al Cu rr Ap he Rf Tr 2 ke TT Sc Ec 24 Oo Mr Lg EI Cl Me Ta LW Tq St Bt 42 Lv BE Ca Cc Po To Ts 14 Ti Ci Oc 3 Gg Gt Om GC Ma OL OH Ru DS AM OV OJ DB AA RS

0102030405060708090100 ED

FIG. 4. — Hierarchical cluster diagram based on the reference tooth positions M1-M3 according to the “extended” mesowear method (Kaiser & Solounias 2003). ED, Euclidean distance (root-mean-squared difference). Clusters are based on a set of 54 “normal” and “typical” extant species (A), on 27 “typical” extant species (B) from Fortelius & Solounias (2000) and on nine “normal” and “typical” Cervidae (C) from Fortelius & Solounias (2000). CONS, conservative classification; RADI, radical classification (after Fortelius & Solounias 2000); % grass, percent grass in the diet of extant species (after a compilation by Van Wieren 1996). Browsers (CONS): AA, Alces alces; EI, Ammodorcas clarkei; AM, Antilocapra americana; BE, Boocercus eurycerus; DB, Diceros bicornis; DS, Dicerorhinus sumatrensis; GC, Giraffa camelopardalis; LW, Litocranius walleri; OH, Odocoileus hemionus; OJ, Okapia johnstoni; OL, Capreolus capreolus; OV, Odocoileus virginianus; RS, Rhinoceros sondaicus; TT, Tragelaphus strepsiceros. Grazers (CONS): ab, Alcelaphus buselaphus; al, Alcelaphus lichtensteinii; bb, bison; cs, Ceratotherium simum; ct, Connochaetes taurinus; dl, lunatus; eb, Equus burchelli; eg, Equus grevyi; he, Hippotragus equinus; hn, Hippotragus niger; ke, Kobus ellipsiprymnus; rr, Redunca redunca. Mixed feeders (CONS): Ap, Axis porcinus; Ax, Axis axis; Bt, Budorcas taxicolor; Ca, Capricornis sumatraensis; Cc, Cervus canadensis; Cd, Cervus duvauceli; Ci, ibex; Cl, Camelus dromedarius; Cu, Cervus unicolor; Gg, Gazella granti; Gt, Gazella thomsoni; Lg, guanicoe; Lv, Lama ; Ma, Antidorcas marsupialis; Me, Aepyceros melampus; Oc, canadensis; Om, Ovibos moschatus; Oo, Ourebia ourebi; Rf, Redunca fulvorufula; Ru, Rhinoceros unicornis; Sc, Syncerus caffer; St, Saiga tatarica; Ta, Tragelaphus angasi; Ti, Tragelaphus imberbis; To, Taurotragus oryx; Tq, Tetracerus quadricornis; Tr, Boselaphus tragocamelus; Ts, Tragelaphus scriptus. Fossil species: Ec, Eucladoceros ctenoides; Mr, Metacervoceros rhenanus; Po, Praemegaceros obscurus.

670 GEODIVERSITAS • 2004 • 26 (4) Ecology of Pleistocene deer

B bb cs 1 eb eg dl ct ab hn he 2 rr ke Ec 16 Mr Me Ca Cc Po 3 To Ts Gg Gt Om GC OH DS 4 OJ OV DB AA RS

0102030405060708090100 ED C CONS RADI % grass B B 2 AA

B B 9 OV

B B 11 OH 1 B B 9 OL

Po M M (47) Cc

Mr

Ec 2 M M 45 Cu M G Ap

M G 70 Ax M G 80 Cd

0 10 20 30 40 50 60 ED

GEODIVERSITAS • 2004 • 26 (4) 671 Kaiser T. M. & Croitor R.

trait. Nevertheless, this observation may shed Agrostis and Phalaris species are frequent in extant some light on the problem of inferring diet based central European forests and near lake shores, on mesowear analogues belonging to different where they constitute large parts of the herb major taxa such as the and the Cervidae. layer. This applies also for relatively cold climate All extant grazers have at least a small percentage conditions that Bonifay (pers. comm.) deduces of low occlusal reliefs, and with only few excep- for the Ceyssaguet site. The loess deposits, where tions (common waterbuck; Fig. 3E and sable the fossils were buried, are good evidence for antelope) some blunt cusps. In having only 4% these dry habitat conditions. low occlusal reliefs, the waterbuck is an exception Based on the bovid and cervid species that we among the species that are consistently classified identify as extant analogues, we estimate the ratio as grazers. The three cervids classified as mixed of grass eaten by E. ctenoides and M. rhenanus to feeders (CONS) or grazers (RADI), the hog deer range between 45% and 80%. The observation of (Fig. 3H), the chital (Fig. 3K) and the barashing- difference in mesowear signatures between bovids ha (Fig. 3N) more often have low reliefs, and the and cervids, however, should have an impact on latter two species have more than 20% blunt our interpretation. We therefore infer that the cusps. Nevertheless, none of these species are con- grass ration was probably closer to 45% (the sam- sistently considered pure grazers. The oribi bar ratio) than to 80% (the waterbuck ratio). (Bovidae; Fig. 3L) and the barashingha (Cervidae; The fact that there are no significant differences Fig. 3N) have almost exactly the same observed in the mesowear signature between grass/browse ratio of 80/10% and 80/15% E. ctenoides and M. rhenanus supports the respectively (after compilation by Van Wieren hypothesis that these two species were dependent 1996). When we compare mesowear signatures upon the same source of forage. Therefore it is we find that both have more round cusps then concluded that the ecological resource partition- sharp. The oribi, however, has almost no low ing between these two species was achieved by reliefs and no blunt cusps while the barashingha differential habitat preferences rather than by dif- dentitions are frequently low and blunt. This ferent foraging strategies as found, for instance, observation leads us to conclude, that deer denti- in the modern African bovids (Kingdon 1979; tions are more sensitive to increased abrasion con- Spencer 1995). trol in the mesowear equilibrium compared to This hypothesis is supported by the postcranial bovids. This biomechanical sensitivity is probably anatomy and some other eco-morphological the reason why the abrasive feeding deer species observations. The postcranial morphology of are all found in the “normal” group of Fortelius E. ctenoides shows peculiar characteristics indica- & Solounias (2000). And additionally this illus- tive for a quite particular habitat preference. The trates why in their bovid dominated model they long and medially prominent caput femoris found the mesowear of these cervids to not cor- found in E. ctenoides from Ceyssaguet (Fig. 2A) rectly correspond to the known dietary trait. We seems to control the movements of the hind-limb have not yet investigated the biomechanical back- in the parasagittal plane, as it was shown for ground in detail, but adaptive strategies towards open-landscape bovids (Sokolov et al. 1964; grazing are certainly less advanced in extant deer Kappelman 1991; Kostopoulos 2000). Some dif- compared to the Bovidae. ferences in the femoral morphology are also evi- That grazing bovids on the one hand and mixed dent in the shape of the lesser trochanter, which feeding and grazing cervids on the other hand is less rounded and prominent in P. obscurus both classified as “typical” are closest to (Fig. 2B2) than in E. ctenoides (Fig. 2A2) E. ctenoides and M. rhenanus indicates, that quite (Croitor & Bonifay 2001). It is evident that the abrasive food components were a readily available difference in lesser trochanter shape transmits the forage for the Ceyssaguet deer. Grass is mostly functional load of the muscles (m. quadriceps rich in abrasive phytoliths. C3 grasses as, e.g., femoris and m. iliopsoas) inserted in this area

672 GEODIVERSITAS • 2004 • 26 (4) Ecology of Pleistocene deer

(Kappelman 1991). According to Kappelman grass forage in a rather dry and cold open-coun- (1991) the more pronounced posterior projection try setting which had wooded areas nearby. The of the lesser trochanter observed in closed-habitat ecological resource partitioning among the two bovids provides a larger moment arm for the best represented deer species was based on differ- external rotator muscles. These peculiarities of ent habitat choice rather then on differential for- the femoral morphology indicate E. ctenoides to age selection. M. rhenanus probably preferred the be a cursorial runner inhabiting more open habi- dense and tall grass and woods near the water and tat settings. The reduced preorbital fossae and the seems to have lived in small groups. E. ctenoides extremely high number of fossil remains also give was a larger gregarious species that inhabited evidence that E. ctenoides from Ceyssaguet was more open habitats in the Ceyssaguet lake area. rather a gregarious open-landscape species. Among deer, the reduced preorbital fossae are observed only in the open-landscape species like Acknowledgements Praemegaceros cazioti (Deperet, 1897) from the The authors are deeply indebted to Marie- late Pleistocene of Sardinia and Françoise Bonifay for access to the fossil collections giganteus (Blumenbach, 1803) from the late of the Maison méditerranéenne des Sciences de Pleistocene of Eurasia (Croitor 2001b). l’Homme (Aix-en-Provence, France) and for the M. rhenanus presents a different ecological type, valuable suggestions during the work upon the which, according to Köhler’s classification article. RC thanks all the German colleagues who (Köhler 1993), may be assigned to an inhabitant helped him during his work and stay at the of wooded habitats. This type of adaptation is Zoological Institute and Museum of the University indicated by the small body size, the simple of Greifswald. We further wish to thank Denis three-pointed antlers, the simple morphology of Geraads and Elmar Heizmann for their reviews teeth and the well developed preorbital fossae. and improvements to the manuscript and Fay The antlers preserved still attached to the pedicles Geisler for improving the English. The research suggest that M. rhenanus inhabited the area near was supported by the State of Mecklenburg- the lake during the winter season. During this Vorpommern Kap. 0710/68109, MG 04 to RC season the diet of M. rhenanus would have and the German Research Foundation (KA- included higher proportions of rough and abra- 1525/4-1) for funding related work by TMK. sive food components like dry grass, tree shoots and bark. This seasonal change in diet could have caused the observed damage of tooth tips with REFERENCES subsequent polishing by abrasive material. ABBAZZI L. 1995. — Megaceroides obscurus from the Compared to E. ctenoides and M. rhenanus, Val di Chiana (Cava Liberatori, Tuscany, Central P. obscurus is consistently classified as a mixed Italy, f. u., early Pleistocene). Remarks on the early feeder, closer to the browse dominated part of the evolution and systematical of Megaceroides. spectrum. However, the difference in the Bollettino Società Paleontologica Italiana 34 (2): 223-234. mesowear signature is not statistically significant ABBAZZI L., CROITOR R. & DAVID A. 1999. — Large because of the small sample available. P. obscurus sized deer Megaceroides obscurus (Azzaroli, 1953) is represented in Ceyssaguet by only nine remains from Eastern Moldova. Acta Zoologica Cracoviensia that probably belong to very few individuals. 42 (3): 377-392. AZZAROLI A. & MAZZA P. 1992. — The cervid genus Eucladoceros in the early Pleistocene of Tuscany. Palaeontographia Italica 79: 43-100. CONCLUSIONS BONIFAY M.-F. 1986. — Intérêt des études taphonomiquess au Pleistocène ancien (Soleilhac et Ceyssaguet, Blanzac, Haute-Loire). Bulletin du The Villafranchian of the area near the ancient Muséum national d’Histoire naturelle, 4e sér., lake from Ceyssaguet was an important source of Section C, 8: 269-281.

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Submitted on 19 May 2003; accepted on 3 June 2004.

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