Bollettino della Società Paleontologica Italiana, 52 (3), 2013, 177-186. Modena

The morphology of femur as palaeohabitat predictor in insular bovids

Roberto Rozzi & Maria Rita Palombo

R. Rozzi, Dipartimento di Scienze della Terra, Sapienza Università di Roma, Piazzale A. Moro 5, I-00185 Roma, Italy; [email protected] M.R. Palombo, Dipartimento di Scienze della Terra, Sapienza Università di Roma, Piazzale A. Moro 5, I-00185 Roma, Italy; CNR, Istituto di Geologia ambientale e Geoingegneria, Montelibretti (Roma); [email protected]

KEY WORDS - Bovidae, ecological morphology, femur, Pleistocene, Sardinia, Java.

ABSTRACT - A variety of methods have been developed with regard to the use of bovid postcranial elements in the functional morphology approach to palaeohabitat prediction. One postcranial element that has proven useful in past habitat reconstructions is the bovid femur. In this study we applied a biometrical method, combined with an eco-morphological analysis, on Nesogoral (Dehaut, 1911) from Sardinia and Duboisia santeng (Dubois, 1891) from Java. Both are insular fossil bovids of similar size but have originated from different ancestors and are believed to inhabit different environments. Measurements of Sardinian and Javanese specimens were processed in comparison with those of the main extant groups of Bovidae in order to predict the preferred habitat category (Forest, Heavy Cover, Light Cover, Open). A principal component analysis (PCA) was carried out to further investigate the structure of the data. This study highlighted the difficulty of inferring palaeohabitats of fossil bovids from the functional morphology of their long bones. In fossil bovids, the assignment of a taxon to a particular category is a “best fit” designation as a number of taxa could have ranged over several habitat types, as has been documented for most of the extant bovids. Bearing in mind this drawback, results confirm that D. santeng was a forest dweller, while Sardinian bovids belonged to different habitat categories. These findings support in part the hypothesis of an adaptive radiation of the Sardinian bovids of the endemic genus Nesogoral.

RIASSUNTO - [La morfologia del femore come strumento per dedurre il palaeohabitat di bovidi insulari] - Vari metodi sono stati proposti per dedurre l’habitat preferenziale di specie fossili di mammiferi sulla base delle caratteristiche morfofunzionali delle ossa, in particolare di quelle dello scheletro appendicolare. Nel caso dei bovidi, risulta complesso definire gruppi ecologici ben caratterizzati, individuati sulla base delle caratteristiche biometriche degli elementi scheletrici di specie attuali, dal momento che molte specie hanno ampia valenza ecologica. Ne consegue che la predizione dell’habitat preferenziale di una specie fossile e la sua assegnazione ad un determinato gruppo ecologico sono in alcuni casi problematiche e le indicazioni ricavate dall’analisi dei diversi elementi scheletrici di un singolo individuo possono essere discordanti tra loro. Pur tenendo conto di questi limiti, l’analisi morfometrica del femore è risultata essere fra quelle che forniscono le indicazioni più affidabili. In questo lavoro, lo studio biometrico di femori di Nesogoral (Dehaut, 1911) e Duboisia santeng (Dubois, 1891), due bovidi pleistocenici endemici, rispettivamente, della Sardegna e di Giava, di dimensioni simili ma discendenti da antenati diversi e adattati a differenti condizioni ambientali, è stato affiancato da un’analisi ecomorfologica qualitativa dei caratteri utili a designare ecomorfotipi. Una predizione dell’habitat preferenziale dei bovidi endemici della Sardegna e di Giava e una loro assegnazione ad uno dei quattro principali gruppi ecologici (Forest, Heavy Cover, Light Cover, Open) è stata effettuata confrontando i dati biometrici relativi ai taxa in studio con un database relativo a 38 specie attuali. Per una migliore valutazione critica delle indicazioni di predizione ottenute, è stata effettuata un’analisi delle componenti principali (PCA). Sulla base dei risultati delle analisi qualitative e quantitative, si può ragionevolmente affermare che i risultati confermano che Duboisia santeng prediligeva ambienti tipicamente forestali, mentre i bovidi sardi devono essere assegnati a differenti gruppi ecologici (Forest, Heavy Cover, Light Cover). Questo dato è in accordo con la presenza tra i femori nel campione sardo di tre diversi morfotipi e può essere considerato un supporto all’ipotesi di una radiazione adattativa dei bovidi sardi del genere endemico Nesogoral.

INTRODUCTION locomotor patterns and habitat preferences. Since then, various authors (e.g., Kappelman, 1988; Köhler, 1993; Several methods have been developed to infer Kappelman et al., 1997) explored and described these palaeohabitats of fossil bovids from the functional patterns, developing both qualitative and quantitative morphology of their bones (Kappelman, 1988; Köhler, methods to use femur in the functional morphology 1993; Plummer & Bishop, 1994; Kappelman et al., 1997; approach to palaeohabitat prediction. DeGusta & Vrba, 2003, 2005a, b; Kovarovic & Andrews, In this study we combined an eco-morphological 2007; Weinand, 2007; Plummer et al., 2008; Klein et al., analysis with the morphometric method developed by 2010). The femur is among the postcranial elements that Kappelman (1988) and improved by Kappelman et al. have been proved as useful in habitat reconstructions (1997) on the extinct insular bovids from Sardinia and of extant and extinct bovids, due the variation of some Duboisia from Java. This aiming to test the usefulness of functional features according to locomotor patterns across femur morphology as palaeohabitat predictor on insular the habitat spectrum (Kappelman, 1988; Kappelman fossil bovids of similar size but originated from different et al., 1997). Gentry (1970) already noted that some ancestors and believed to inhabit different environments peculiar morphologies of the femur represent functional (Van den Bergh, 1988; Palombo et al., 2006a, 2013; Rozzi complexes that demonstrate clear linkages between & Palombo, 2013a, b; Rozzi et al., 2013).

ISSN 0375-7633 doi:10.4435/BSPI.2013.19 178 Bollettino della Società Paleontologica Italiana, 52 (3), 2013

A SHORT ACCOUNT ON SARDINIAN genus and species Asoletragus gentryi Palombo, Valli, AND JAVANESE BOVIDS Arca & Tuveri, 2006. This bovid differs from other genera thus far known in the Mediterranean region in having The Sardinian Plio-Pleistocene bovids straight, almost conical horn cores, closely inserted, In Sardinia (Fig. 1), endemic bovids ascribed to the slightly divergent and strongly inclined backwards (about so called “Nesogoral group”, Nesogoral melonii (Dehaut, parallel to the glenoid surface of the temporal bone). 1911) have been first reported (Olbia, northeastern The peculiar shape and position of the horn cores make Sardinia) from breccias deposits of Capo Figari karstic phylogenetic relationships with Neogene European bovids network (Gliozzi & Malatesta, 1980; Palombo et al., 2013 difficult to ascertain. and references therein). Subsequently, bovid teeth found Different morphotypes have also been recognised in the same locality have been dated by ESR method at in teeth, mandibles and postcranial bones, but relating about 1.807.500 ± 20% years BP (Mtoji Ikeya fide Van der them to the respective skull morphotypes is extremely Made, 1999). Bovid remains have also been reported from difficult (Palombo, 2002a, b; Rozzi, 2010). The peculiar other Late Pliocene - Early Pleistocene sites on the island, features of long bones of Sardinian bovids, showing a such as Mandriola (Oristano, western central Sardinia: different combination of “Caprini” and “non-Caprini” Late Pliocene, Nesogoral sp.) and Capo Mannu (Oristano, characteristics, make the attempt to discriminate between western central Sardinia: Early Pleistocene, Nesogoral ecomorphological and taxonomical traits undoubtedly sp. and bovid remains of small size which do not fully challenging. Nonetheless, biometry and morphology of conform to the morphology of Nesogoral spp.), Monte astragalus and phalanges of Sardinian bovids indicate that Tuttavista (Orosei, eastern Sardinia: Early Pleistocene, at they inhabited a variety of environments, ranging from least two genus and three species) and from an unknown mountain areas to forests (Rozzi & Palombo, 2013a). locality in the Campidano area (south western Sardinia: unknown age, Nesogoral cenisae Van der Made, 2005) The Javanese Pleistocene bovids (Van der Made, 2005; Palombo et al., 2006a, b, 2013; Fossil bovids are very common in the Early-Middle Abbazzi et al., 2008; Palombo, 2009). Pleistocene of Java. The fossil record includes at least four The richest sample comes from Monte Tuttavista, species: Bubalus palaeokerabau (Dubois, 1908), Bibos where more than 1500 bovid specimens have been palaesondaicus (Dubois, 1908), Epileptobos groeneveldtii collected from nine karstic fissures. The preliminary (Dubois, 1908) and Duboisia santeng (Dubois, 1891) analysis of cranial specimens from three fissures (VI-3; (Fig. 3). B. palaeokerabau was a very large buffalo, X-ghiro; XI-antilope) shows the presence of different characterised by horn cores with a triangular cross section, morphotypes (Fig. 2) and demonstrates that more than long more than two metres. B. palaesondaicus, a banteng- one taxon occur in the same fossiliferous deposits. Two like bovine, shows horn cores with oval cross-section, morphotypes can be ascribed to as many species within curving backwards. E. groeneveldtii was a peculiar the genus Nesogoral (morphotype A = Nesogoral aff. N. leptobovine characterised by large horn cores, shifted melonii and morphotype B = Nesogoral sp. 1), while a farther behind the orbits, and a triangular occiput, with partial skull from fissure “VI-3” was assigned to the new an upper angle formed by the parieto-occipital eminence.

Fig. 1 - Map of localities of bovid findings in Sardinia. R. Rozzi & M.R. Palombo - Femur as palaeohabitat predictor in insular bovids 179

D. santeng, regarded as a typical endemic species due to its small size, was a small boselaphine with short and keeled horn cores (Hooijer, 1958), whose remains characterise the Trinil H.K. and Kedung Brubus Faunal units (Van den Bergh et al., 2001). This species had a body mass of about 54 kg, significantly smaller than the one of its putative ancestor Boselaphus of the Siwaliks (Lomolino et al., 2013; Rozzi et al., 2013). The rich sample of D. santeng comes from several Javanese localities, such as Trinil, Kedung Brubus, Mojokerto, Sangiran and Bumiayu (Fig. 4). A few remains of this species were described from the late Middle Pleistocene site of Tambun (peninsular Malaysia) (Tougard, 2001).

MATERIALS AND METHODS

The sample used for this study includes 51 femora, 12 belonging to Duboisia santeng and 39 to Sardinian bovids, collected from the karstic fissures of Monte Tuttavista, being no femora known from other localities. The Monte Tuttavista specimens are stored in the Operating Office of the “Soprintendenza per i Beni Fig. 2 - Skulls of Early Pleistocene endemic bovids from Sardinia: Archeologici per le province di Sassari e Nuoro” and in a) Incomplete skull still embedded in the bone breccias block, the Museo Archeologico of Nuoro. The specimens of D. holotype of the species Nesogoral melonii (Dehaut, 1911) (Capo santeng belong to the Dubois Collection of the Naturalis Figari, northeastern Sardinia) (specimen PU29864 stored at the Museo Regionale di Scienze Naturali di Torino); b) Nesogoral Biodiversity Center (Leiden, The Netherlands) and were aff. N. melonii (quarry XI-Antilope, Monte Tuttavista, Orosei, collected during the last century from various sites on eastern Sardinia) (specimen 1 stored at the Operating Office of the Java (Trinil H.K. and Kedung Brubus Faunal units). “Soprintendenza per i Beni Archeologici per le province di Sassari Despite their disparate provenance the sample of femura e Nuoro”); c) Nesogoral sp. 1 (quarry X-Ghiro, Monte Tuttavista, of D. santeng is rather homogeneous regarding both Orosei, eastern Sardinia) (specimen 1000 stored at the Operating morphology and biometry. Office of the “Soprintendenza per i Beni Archeologici per le province di Sassari e Nuoro” ); d) Asoletragus gentryi (Palombo, Measurements of the femur useful for distinguishing Valli, Arca & Tuveri, 2006) (quarry VI-3, Monte Tuttavista, Orosei, among bovids from different habitats were taken on the eastern Sardinia) (specimen 1179 stored at the Operating Office specimens of Sardinian bovids and D. santeng following of the “Soprintendenza per i Beni Archeologici per le province di Kappelman (1988) and Kappelman et al. (1997). In Sassari e Nuoro”). addition to FHA (= femoral head area), which shows a

Fig. 3 - Skull of the endemic bovid Duboisia santeng (Dubois, 1891) from Java, lectotype of the species in frontal (a) and lateral view (b) (Trinil, Java). Specimen 2037, Naturalis Biodiversity Center (Leiden, The Netherlands). 180 Bollettino della Società Paleontologica Italiana, 52 (3), 2013

clear increase (FHSS = femoral head shape score; LTR = lesser trochanter anteroposterior/mediolateral ratio; PSR = proximal anteroposterior: mediolateral shaft ratio) or decrease (PHR = patellar lip height ratio; MPLR = medial patellar lip ratio) in magnitude with increasingly closed conditions, were used in this study (Tab. 1). These measurements highlight some functional features mirroring different locomotor patterns across the habitat spectrum (see below). To predict habitat preference of Sardinian bovids and D. santeng, the biometrical data obtained for these species were compared by means of univariate and multivariate analyses with those of 38 species of extant bovids (Kappelman et al., 1997). Four general categories of habitat (Forest, Heavy Cover, Light Cover, and Open) were used, as in Kappelman (1988) and Kappelman et al. (1997). The Forest category includes taxa inhabiting scrub-forest, forest, and rainforest; Heavy Cover category includes taxa which frequent bush, woodland, swamp, and near-water habitats; Light Cover category gathers taxa which frequent light bush, tall grass, and hilly areas; Open category is that of taxa which frequent edge or ecotone, open country, and arid country (Kappelman et al., 1997; DeGusta & Vrba, 2003, 2005a). To further investigate the structure of the data, we carried out a multivariate analysis (principal component analysis, PCA) by means of JMP (version 9; SAS Institute Inc., Cary, NC, 2009) software, using as variables the focal measurements of D. santeng and Sardinian bovids and those in the database of extant bovids completed by Kappelman et al. (1997). Fig. 4 - Map of the main localities of Duboisia santeng (Dubois, 1891) findings across the Sundaic province. Modified from Rozzi et al. (2013). RESULTS Palaeoecological information obtained by means of quite general pattern whose variations seem to be more morphology and morphometric methods are in agreement related to body size (Kappelman et al., 1997), only the with each other and enabled us to infer habitat preferences most significant femoral variables, showing either a of the focal species.

Variable Description

Femoral head shape was scored on the distribution of articular area in the cranial projection and Femoral head shape score (FHSS) measured the taper of the femoral head along the mediolateral axis.

Area (cm2) was calculated from a simple geometric construction of the superior view of the Femoral head area (FHA) femoral head articular surface (see Kappelman, 1988).

The anteroposterior and mediolateral dimensions at the lesser trochanter were measured with Lesser trochanter anteroposterior: mediolateral ratio (LTR) dial calipers. This ratio describes the morphology of the proximal femur at the distal margin of the intertrochanteric fossa.

The anteroposterior and mediolateral dimensions of the proximal shaft at the base of the lesser Proximal anteroposterior: mediolateral shaft ratio (PSR) trochanter were measured with dial calipers. The ratio provides an estimate of the loading forces that operate through the proximal shaft.

The anteroposterior heights of the medial and lateral patellar lips to the respective femoral Patellar lip height ratio (PHR) condyles were measured perpendicular to the shaft with an osteometric board. The ratio describes the relative symmetry of the distal femur.

The distance from the medial patellar lip to the femoral condyle in the anteroposterior plane, and the distance from the center of the patellar groove to the intercondylar notch were measured Medial patellar lip ratio (MPLR) perpendicular to the shaft with an osteometric board and a dial calipers, respectively. The ratio of the medial patellar lip height to the distal AP shaft diameter estimates the relative moment arm of the extensor muscles that cross the knee.

Tab. 1 - Description and abbreviations of measurements of femora, following Kappelman (1988) and Kappelman et al. (1997). R. Rozzi & M.R. Palombo - Femur as palaeohabitat predictor in insular bovids 181

Fig. 5 - Morphotypes recognized in the femur of bovids from Monte Tuttavista (Orosei, eastern Sardinia): a) morphotype 1, specimen ORVIB0030; b) morphotype 2, specimen ORXB1005; c) morphotype 3, specimen ORX4blocco. Modified from Rozzi (2010).

Ecomorphology al., 2013). The Javanese bovid shows a marked vertical Three morphotypes have been recognized in the inclination of the cranial articular surface of the femoral femur of Sardinian bovids, independently from their size head with respect to the vertical axis of the shaft, a quite (Rozzi, 2010), which mirror different locomotor patterns rounded, pear-shaped caput femoris with a proximally across the habitat spectrum. The morphotype 1 (F1+F3 in and centrally situated fossa of the ligamentum, while the Rozzi, 2010) is characterised by a week indent between distal articular surface shows an overall symmetry of the caput femoris and great trochanter, and a pear-shaped patellar lips. These features characterise taxa requiring caput femoris in proximal view. The morphotype 2 (F2 maneuverability and a great degree of abduction and axial in Rozzi, 2010) is still characterised by a week indent rotation at the hip and they are shared, for instance, by between caput femoris and great trochanter, but by a B. tragocamelus and typical forest dwellers (Fig. 6). In rather spherical caput femoris in proximal view and distal particular, the morphology of caput femoris of D. santeng condyls of similar size. The morphotype 3 (F4 in Rozzi, can be related to morphological types A as defined by 2010) is characterised by a strong indent between caput Köhler (1993), which are typical of taxa living in from femoris and great trochanter, a top of great trochanter moderately to very humid wooded, and semiaquatic wide compared with its own narrower lower part (in habitats (Fig. 6). These features, despite being markedly lateral view) and a pear-shaped caput femoris in proximal ecomorphological, can be also used to distinguish femora view (Fig. 5). Bovids with a more rounded caput femoris and a more proximally and centrally situated fossa of the ligamentum, as shown by morphotype 2, have a pattern of locomotion requiring maneuverability and a great degree of abduction and axial rotation at the hip. This morphological trait can be found in bovids living on rocky terrains, and taxa inhabiting close canopy settings, often facing a complex nearly three-dimensional substrate of tree trunks, shrubs, roots, and bushes, all of which could affect high-speed cursorial locomotion at a great extent. It is worth noting that among Sardinian femora, no one shows a femoral head cylindrical in shape, as that characterising species inhabiting open environments where ground-level obstacles could be only occasionally present. Indeed a cylindrical caput femoris acts to limit movements to the parasagittal plane (Kappelman, 1988; Köhler, 1993; Kappelman et al., 1997). Accordingly, the different combinations of features exhibited by femura of Sardinian bovids can be related to various patterns of locomotion on uneven ground as exemplified, for instance by types A and C described in Köhler (1993), which are respectively characterised by a pear-shaped and a spherical caput femoris. Taxa referred to type A, such as the cervids Dama dama (Linnaeus, 1758) and Axis axis (Erxleben, 1777) and the bovid Boselaphus Fig. 6 - Morphological comparison among femora in caudal (a1, b1, tragocamelus (Pallas, 1766), include forest dwellers and c) and proximal (a2, b2, d, e) views of Duboisia santeng (a1, a2) species living on moist ground. Taxa referred to type C, (specimen 5668 stored at the Naturalis Biodiversity Center, Leiden), Boselaphus tragocamelus (b1, b2) (specimen 1949.12.30.1 stored at such as Capra, are mostly related to rocky grounds. NHM Natural History Museum, London), a typical forest dweller Morphological comparison shows that the femur of D. (c) as in Kappelman (1988) and the types A1 (d) and A2 (e) defined santeng shares several features with typical extant forest by Köhler (1993). Continuous and dashed lines show the orientation dwellers, including species of the genus Boselaphus, to of the cranial articular surface and the shape of the caput femoris, which its putative ancestor likely belongs (see Rozzi et respectively. Modified from Kappelman (1988) and Köhler (1993). 182 Bollettino della Società Paleontologica Italiana, 52 (3), 2013 of D. santeng from those of Axis lydekkeri (Martin, 1886), D. santeng to wooded environments (Fig. 7b). The value a cervid similar in size that coexisted on Java with the of PSR, unfortunately available only for one specimen, endemic bovid (Van den Bergh, 1988). falls within the overlapping range of extant Forest, Heavy Cover and Light Cover dwellers (Fig. 7b). Biometry As regards the PCA analysis, in both the score plots Results obtained by biometrical analysis are reasonably where measurements of femur of Sardinian bovids and consistent with evidence resulting from morphology. It is those of D. santeng were respectively processed with worth noting that, as regards femur of extant bovids, measurements of the extant bovid sample (Fig. 8a), the ranges of variation of each score for every habitat ecomorphospace of each habitat category of extant taxa group highlight a morphological trend across the habitat is quite well defined, despite a few overlapping areas. As spectrum from closed to open landscape (Kappelman, regards Sardinian bovids, the Monte Tuttavista specimens 1991). In particular, the need of a great manoeuvrability form a quite dispersed group, falling into the second and of typical Forest dwellers, in functional terms, is marked the third quadrants, which contain small to medium sized by a spherical shape of caput femoris (high FHSS), a extant Forest, Light Cover and Heavy Cover dwellers greater moment arm for the external rotators of the hip (Fig. 8a). Conversely, results obtained for D. santeng (Fig. (high LTR), a cross-section of the diaphysis that in part 8b) would suggest a marked adaptation of this taxon to reflects great anterior-posterior directed loading forces Forest environments. The first two principal components transmitted through the femoral head to the proximal shaft explained the 99.957% of the variation in the PCA (high PSR), and a lower moment for extension at the knee including the Sardinian bovids and the 99.879% in that (low MPLR, PHR). including the Javanese bovid. It is worth noting that the Comparing scores obtained for Sardinian bovids variables affecting the first component the most are the and D. santeng with the ranges of variation of each femoral head shape score, which counts more than 80% habitat group of extant bovids, it appears that most of in the femora of D. santeng, and the caput femoris area, the values (e.g., MPLR, PHR, PSR, LTR) overlap with which counts about 78% in the femora of Sardinian bovids all the habitat categories (Fig. 7). The values obtained (see Tab. 2). Both dimensions are strictly related with the for FHA fall within the ranges of extant Forest, Light body size of the species. Therefore, the largest species tend Cover and Open dwellers, while FHSS values fall within to set quite apart, independently from the habitat category the ecomorphospaces of Forest, Heavy Cover and Light to which they are predicted to belong. Cover dwellers. As regards D. santeng, most of the values obtained for FHA, LTR and PHR falls within the range of extant Forest dwellers, although a few values scatter DISCUSSION AND CONCLUSION within the ecomorphospace of all habitat categories (Fig. 7b). Most values obtained for FHSS and MPLR fall just The Bovidae family includes more species than any outside the ecomorphospace of extant Forest dwellers, other extant family of large mammals, many of them hypothetically suggesting a more extreme adaptation of characterised by a high ecological amplitude. Indeed, the

Fig. 7 - Plots of dimensions of femora for the specimens of (a) Sardinian bovids (each specimen is denoted by a black triangle in the graphs) and (b) D. santeng (each specimen is denoted by a black sphere). The bars represent range of each habitat group’s distribution in the modern bovid reference sample (from the top to the bottom: first line = Forest; second line = Heavy Cover; third line = Light Cover; fourth line = Open). See Tab. 1 for measurement abbreviations. R. Rozzi & M.R. Palombo - Femur as palaeohabitat predictor in insular bovids 183

Fig. 8 - Results of the PCA analysis for femora of the Sardinian bovids (a) and Duboisia santeng (b). Specimens of these taxa are denoted by spheres and referred to mean values of typical representatives of Forest: Cephalophus natalensis Smith, 1834, Heavy Cover: Tragelaphus strepsiceros (Pallas, 1766), Light Cover: Redunca redunca (Pallas, 1767), and Open: Gazella granti (Brooke, 1872).

inclusion of these taxa in few habitat categories, as those have determined a change in the original bauplan, and defined by Kappelman (1988), is not trivial, especially 3) ecomorphological traits depending on environmental considering that populations of the same species can be factors. found in different environments, sometimes following It is generally accepted that insular artiodactyls, an ecogeographical gradient. Therefore, inferring whatever their phylogenetic relationships, share some palaeohabitat of fossil bovids from the functional peculiar traits, such as “low-gear” locomotion, dental morphology of their bones is not a simple task because reduction and marked hypsodonty, as pronounced as of the methodological limitations that could affect their degree of endemism. Although Sardinian bovids the consistency of the results obtained. Limitations of are markedly hypsodont and have simple horn cores, morphometric methods are mainly related to a number of the morphological traits characterising their slender factors such as the ecological amplitude of the focal taxa, limb bones partially contradict the evolutionary pattern the body size bias affecting the results of the statistical generally assumed as typical of insular endemic bovids. analysis (see above), and the problems related to the Sardinian taxa possibly preserved (even not developed) definition of habitat categories that could deal more slender and long limb bones and a cursorial aptitude specifically with the range of environments inhabited by due to the presence in Sardinia of the top predator “eury” bovids (Odum, 1968; Rozzi & Palombo, 2013a). Chasmaporthetes melei (Rook, Ferretti, Arca & Tuveri, These factors, affecting our biometrical analysis of femora 2004), that must have conditioned their evolution as well, stress the importance of integrating biometry (Palombo et al., 2006a, 2013). These features make with a qualitative, ecomorphological approach. Dealing the application to Sardinian bovids of morphometric with insular taxa, the scenario is further complicated by methods for palaeohabitat prediction - developed on extant the difficulty in discriminating among 1) plesiomorphic continental taxa - more reliable than in the case of typical features - inherited by either known or even unknown insular artiodactyls (see above). ancestors, maybe living in habitats different from those of This is also the case of D. santeng. In fact, the their insular descendants, 2) apomorphic features acquired slenderness of metapodials of the Javanese boselaphine because of the insular evolutionary process that might suggests that the moderate cursorial aptitude shown by 184 Bollettino della Società Paleontologica Italiana, 52 (3), 2013

Sardinian bovids

Variables Axis 1 Axis 2 Axis 3 Axis 4 Axis 5 Axis 6 Axis 7 Axis 8 FHSS -0.6211 0.7837 -0.007775 0.0005835 0.0005805 -0.0007719 0.0004228 -0.001251 FHA 0.7837 0.6211 0.008763 -0.004287 -0.000342 -0.002232 -0.001221 -0.0017 LTR -0.005933 0.0004201 0.6231 0.444 0.4343 -0.4737 0.007724 0.0399 PSR -0.003357 0.003097 0.4767 0.3521 -0.3159 0.6904 0.0705 0.2596 PHR 0.0035 0.001455 -0.2472 0.02933 0.009161 -0.2082 0.2006 0.9243 PGR 0.005894 0.0004843 -0.3055 0.5387 -0.26 -0.1477 0.6717 -0.2753 PMR -0.001311 -0.00006374 0.1192 -0.3881 0.5652 0.3194 0.6426 -0.0289 MPLR 0.0077 0.001232 -0.4645 0.4871 0.5697 0.363 -0.301 0.001703

Duboisia santeng

Variables Axis 1 Axis 2 Axis 3 Axis 4 Axis 5 Axis 6 Axis 7 Axis 8 FHSS 0.8312 0.556 0.005998 -0.005022 -0.0007751 0.0001338 -0.002095 0.00008165 FHA -0.5558 0.8312 -0.008453 0.01092 0.0001843 -0.000002757 -0.0008878 -0.002814 LTR 0.005313 -0.004173 -0.06256 0.5106 0.5171 0.371 -0.5662 0.09896 PSR 0.00471 -0.0001292 -0.04679 0.3307 0.5276 -0.1843 0.711 0.2654 PHR -0.004162 0.003205 0.06574 -0.1643 -0.1976 0.1032 -0.1017 0.9532 PGR -0.005571 0.004215 -0.1323 -0.7204 0.4882 0.4602 0.1031 -0.05272 PMR 0.002054 -0.0009609 0.1108 0.2811 -0.3799 0.7782 0.3902 -0.08055 MPLR -0.01003 0.003958 0.9796 -0.06952 0.1804 -0.01791 -0.02557 -0.04301 Tab. 2 - PCA loadings for femora of Sardinian bovids and Duboisia santeng. See Tab. 1 for measurement description and abbreviations.

D. santeng, would relay with the presence on Java of All in all, the morphological and biometrical analysis various predators (Rozzi et al., 2013). At the time of performed on femur of Sardinian and Javanese insular Trinil H.K. faunal unit, in fact, the Javanese carnivore bovids, on the one hand highlighted the difficulties of guild included a tiger - Panthera tigris (Linnaeus, 1758) discriminating palaeohabitat preferences on the basis of - and a canid - Mececyon trinilensis Stremme, 1911. In morphometric methods and the importance of integrating the subsequent Kedung Brubus faunal level, despite the them with ecomorphology in order to obtain more reliable disappearance of M. trinilensis, a marked increase of the results. On the other hand it confirmed the usefulness of predatory pressure was produced by the occurrence of the femur analysis in inferring the palaeohabitat of insular large hyena Pachycrocuta brevirostris (Aymard, 1846) fossil bovids, which is in turn a key issue for a better (Hertler & Volmer, 2008). Moreover, not only carnivores understanding of their evolutionary processes. fed on D. santeng, but also reptiles and Homo erectus (Dubois, 1894), as testified by cut marks observed on long bones (Bouteaux & Moigne, 2010). Results obtained ACKNOWLEDGEMENTS for the femur of D. santeng confirm what stated in Rozzi et al. (2013), that this species was a forest-dweller, in Thanks are due to P. Bover and T. Kotsakis for their useful agreement with a palaeohabitat that, despite being strongly comments to the previous version of the manuscript. We are grateful heterogeneous at the time of Trinil H.K. and Kedung to the following individuals for access to collections and advice: R. Portela Miguez and R.C. Sabin (Natural History Museum, London), Brubus Faunal units, likely included relicts of rainforest J. de Vos and R. van Zelst (Naturalis Biodiversity Center, Leiden), (Sémah et al., 2010; Sémah & Sémah, 2012). N. Tuveri and and M. Arca (Soprintendenza per i Beni Archeologici Accordingly, despite the limitations mentioned above, delle Province di Sassari e Nuoro, sede operativa di Nuoro). on the one hand results obtained support the hypothesis Work supported by Italian PRIN project 2008 that the evolutionary processes undergone by both (n.2008RTCZJH_002, unit 2, PI M.R. Palombo). Sardinian and Javanese endemic bovids in environments with predators likely did not change their limb bones REFERENCES proportion; on the other hand, based on habitat predictions, we can confidently hypothesize that the contemporaneous Abbazzi L., Carboni S., Delfino M., Gallai G., Lecca L. & Rook presence in Sardinia of bovids (Asoletragus and at least L. (2008). Fossil vertebrates (Mammalia and Reptilia) from two species belonging to the genus Nesogoral) having Capo Mannu formation (Late Pliocene, Sardinia, Italy), with about the same size, but inhabiting different environments, description of a new Testudo (Chelonii, Testudinidae) species. supports the previously advanced hypothesis that Sardinian Rivista Italiana di Paleontologia e Stratigrafia, 114: 119-132. representatives of the genus Nesogoral might originated Aymard A. (1846). Communication sur le gisement de Sainzelles. Annales de la Société d’Agriculture, Science, et Arts Commerce from a still unknown ancestor by an evolutionary process du Puy, 13: 153-155. of adaptive radiation (Palombo et al., 2006a, 2013; Rozzi Bouteaux A. & Moigne A.-M. (2010). New taphonomical & Palombo, 2013a), similar to the one underwent by the approaches: The Javanese Pleistocene open-air sites (Sangiran, Cretan deer (de Vos, 2000). central Java). Quaternary International, 223-224: 220-225. R. Rozzi & M.R. Palombo - Femur as palaeohabitat predictor in insular bovids 185

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