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Cynotherium Sardous CRANIUM 23,1 - 2006 Adaptations of the Pleistocene island canid Cynotherium sardous for small (Sardinia, Italy) hunting prey and Alexandra der George Lyras van Geer Summary small canidthatlived the island of Sardinia-Corsica the Pleistocene. Once the Cynotherium sardous is a on during on island, the species gradually adapted, and became specialized in hunting small prey like the lagomorph Prolagus. Moreover, in order to fulfil mass-related energetic requirements, the species had to reduce body size compared to its which than the wolf. carried its head much in the foxes and ancestor Xenocyon, was larger grey Cynotherium way do, hold the when could head better than was able to its body low to ground stalking. Inaddition, it move its laterally any living canid. Samenvatting Cynotherium sardous is een kleine hondachtige, die leefde op heteiland Sardinië-Corsica gedurende het Pleistoceen. Eenmaal het eiland de zich zich in het kleine zoals de op paste soort aan en specialiseerde jagen op prooi haasachtige de kleiner Prolagus. Om aan de energiebehoeften, gerelateerd aan lichaamsgewicht, te voldoen, moest soort worden, vergeleken met zijn voorouder, Xenocyon, die groter was dan de huidige grijze wolf. Cynotherium hield zijn hoofd zoals hieldhet lichaam de het de kon ongeveer vossen doen, en laag bij grond bij besluipen van prooi. Daarbij hij zijn kop verder zijwaarts bewegen dan alle nu levende hondachtigen. Introduction When we hear about insular island mammals, we immediately make associations with pig-sized hippo's, mini-mammoths, giant rodents, deer adapted for mountain climbing, from the scientific of several apart names Plio-Pleistocene insular ungulates and micro- mammals. We all think of the well-established model according to which mammals reached the islands by swimming or rafting. Once there, due to limited resources and the absence of terrestrial mammalian their descen- predators, Fig 1 The studied skeleton of Cynotherium sardous dants became to the island The gradually adapted (missing parts in white). background outline re- environment by a change in body plan in presents the size of its ancestor Xenocyon lycaonoides several on data from ways, e.g. by reducing or increasing (based Sotnikova, 2001). their size, fusion of limb bones, etcetera. skelet Het bestudeerde van Cynotherium sardous That is in general true, but not in all cases. (ontbrekende delen in wit). Het achtergrondprofiel There for of mammalian stelt de are, example, cases grootte van zijn voorouder, Xenocyon lycao- on cases noides, voor data uit Sotnikova, predators that lived islands. Such in (gebaseerd op 2001). the first instance disturb the nice picture. However, their study provides us with a better The most promising place to study an insular and more complete picture of island mammal mammalian predator is provided by the islands evolution, not only because more and different of Sardinia (Italy) and Corsica (France). From taxa are involved, but also because of their rela- the Late Miocene to the Late Pleistocene, these tion with the other endemicanimals on the two islands, which were for most of their island. It appears from such studies that also geological history connected with each other, mammalian predators follow the earlier obser- were repeatedly colonized by mammals vations, and thus that also they show adapta- der (Sondaar & Van Geer, 2005), and as a conse- tions to the peculiar island environment. the fossil record of quence, Sardinia-Corsica is 51 the Pleistocene island canid sardous small Adaptations of Cynotherium (Sardinia, Italy) for hunting prey faunal became small and characterized by more changes and a isolated, it a specialized than other island with hunter et al. in greater diversity any small-prey (Lyras r press). insular mammals (table 1). Of particular interest of all the Sardinian-Corsican extinct mammals, the carnivores. Fossils of several carnivore are Materials and Methods from groups have beenrecovered of the Herewe the results morphofunc- Sardinia-Corsica: the hunting hyaena present tional analysis of a Cynotherium skeleton that Chasmaporthetes melei, the canid Cynotherium was excavated in CobedduCave (Lanaittu sardous, and several otters (Algarolutra majori, Nuoro, Sardinia, see also Van der Sardolutra ichnusae, Enhydrictis galictoides, Valley, Italy; Geer & Van der Geer, 2001). The skeleton is Megalenhydris barbaricina). fairly complete (fig. 1) andin an excellent state sardous fossils are found in relati- Cynotherium of preservation. The fact that the material numbers, including not skulls vely large only belongs to one individual, provides an excellent and dentition, but also This postcranials. opportunity for morphofunctional analysis. For can therefore be studied in more detail. species the description of the skeleton we follow the is a small canid with dental charac- Cynotherium nomenclatureof Evans (1993). Data on the that are for canids ters typical hypercarnivorous myology of the living canid species have been canids that include significant quantities of (i.e., taken from Langguth (1969) and Evans (1993). meat in their diet and on prey mostly large To check these data and to see the nature of the Its cranial characters, however, indi- animals). different muscles and tendons, we dissected a cate that it was not able to hunt A large prey. red fox (Vulpes vulpes) and a domestic dog recent et al. (in phylogenetic analysis by Lyras (Canis familiaris). press), led to the conclusion that Cynotherium In order to investigate the functional signifi- originates from Xenocyon lycaonoides, a cance of the observed features we followed the large-sized dog, related either to some extinct form-function correlation method of analysis. wolves (Wang et al., 2005) or to the Cape This method extrapolates the correlation that is hunting dog, Lycaon pictus (Martinez-Navarro & observed between anatomical forms and certain Rook, 2003). Xenocyon reached Sardinia-Corsica functions or behaviours of living animals to the at the end of the early Pleistocene and once (mm) end distal humerus of width Greatest Cynotherium sardous Cynotherium sp. Xenocyon lycaonoides CORBEDDU CAPO FIGARI UNDERMASSFELD CORBEDDU DRAGONARA CAPO FIGARI STRANSKA SKALA UNDERMASSFELD Fig 2 Greatest width of the distal end of the humerus of Xenocyon lycaonoides from Stránska Skála, Czech Repu- and from with from Sardinia: blik, Undermassfeld, Germany (data Sotnikova, 2001), compared Cynotherium spp. C. NMB and from der C. sp. Ty-5362 from Capo Figari (data photograph van Made, 1999), sardous from Dragona- ra Cave (data from Malatesta, 1970) and from Corbeddu Cave (CB 84-8022). Grootste breedte van het onderuiteinde van het opperarmbeen van Xenocyon lycaonoides van Stránska Skála, Duitsland uit met Sardinië: Tsjechië, en Undermassfeld, (data Sotnikova, 2001), vergeleken Cynotherium spp. van C. NMB uit Van sp. Ty-5362 van Capo Figari (data en foto der Made, 1999) en C. sardous van de grot Dragonara (data uit Malatesta, 1970) en de grot Corbeddu (CB 84-8022). 52 CRANIUM 23,1 - 2006 order do et al., A fossil species (Radinsky, 1987). In to 2005). stratigraphically younger locality the of the is from which material so, we compared morphology Dragonara Cave, Sassari, Corbeddu skeleton with specimens from most belonging to several Cynotherium, individuals living canid species (see table 2). has been excavated (Malatesta, 1970). The material from Corbeddu Cave youngest comes Material of the living and of the fossil species near Oliena, Nuoro (hall II and hall IV, see Van was studied at and belongs to the following der Geer & Van der Geer, 2001). The skeleton institutes: AMNH(M): American Museumof from hall II, under study here, is the youngest Natural History, Deparment of Mammamology Cynotherium specimen (Klein Hofmeijer, 1996) New York, USA; CB: Corbeddu Cave, Sardinia, known till now. Its postcranial is of a smaller Italy; FMNH: Field Museum of Natural History, size than any specimen from Dragonara (fig. 2). Chicago, USA; MNHN: MuseumNational The latter specimens are in turn smaller than d'Histoire Naturelle, Paris, France; NMB: the humerus from Capo Figari. Naturhistorisches Museum Basel, Switzerland; NNML: Nationaal Natuurhistorisch Museum Although the data are limited, it is tempting to that with indication (Naturalis), Leiden, the Netherlands. suggest they provide us an of size reduction of Cynotherium through time. The phenomenon of size reduction becomes evident when Size reduction even more we compare Cynothe- rium with its probable mainland ancestor. As Findings of Cynotherium in Corsica are limited shown in figure 2, the humeri of Xenocyon (data & but Sardinia has (Robert Vigne, 2001), many from Sotnikova, 2001) are considerably larger localities with fossil remains of that canid. The than those of Cynotherium. oldest remains found till now come from be estimated with the In theory, body mass can breccia deposits on Capo Figari, Sassari (Van of a number of dental and skeletal der Made, 1999) and from fissure fillings at use measure- ments (Damuth & MacFadden, 1990). In dwarf Monte Tuttavista near Orosei, Nuoro (Abbazzi forms, however, dental measurements are not reliable in such estimations, as their relation with body proportions differs essentially from those of their mainland relatives (e.g. Sondaar, 1977; Lister & Davies, 2003). Thus, the body mass of Cynotherium can only be estimated based on postcranial elements. The method based on the circumference of the femur at the midshaft, developed by Anyonge (1993), gives the best estimation of body mass in living
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