Late Pliocene carnivores from western Macedonia (Greece)

Some carnivores from the late Pliocene (Villafranchian) locality of “Dafnero 1” (DFN) are studied. The site is in the Grevena basin, western Macedonia, Greece. The material is described and compared with other Greek and European samples. Three speciesNyctereutes megamastoides, Vulpes alopecoides andChasmaporthetes lunensis have been determined. The DFN carnivores are similar to those from St.-Vallier (France) and Puebla (Spain). The fauna is dated as Middle Villafranchian (latest Pliocene); more precisely it belongs to the St.-Vallier Faunal Unit, MN 17, with an absolute age of about 2. om. y.     

Earliest occurrence of Puma pardoides (Owen, 1846) (Carnivora,Felidae) at the Plio/Pleistocene transition in western Europe: New evidence from the Middle Villafranchian assemblage of Montopoli,Italy

The puma-like cat Puma pardoides (Owen, 1846) is reported from several Eurasian localities dated to the Late Pliocene–Early Pleistocene, but its Italian fossil record is really poor; in fact, the only possible occurrence comprises some fragmentary postcranial material from the Late Villafranchian locality of Pirro Nord (1.6–1.3 Ma). In the present paper, we describe an isolated left upper carnassial of a medium-sized felid belonging to the collection of fossil mammals from Montopoli (Tuscany, Italy), the type locality of the Middle Villafranchian Faunal Unit (∼2.6 Ma). All the morphological and morphometric characters of the tooth are consistent with an attribution to Puma pardoides (Owen, 1846), which comes out as the second felid species from Montopoli together with Acinonyx pardinensis (Croizet et Jobert, 1828) and is, possibly with the specimens from Perrier-Étouaries, the earliest occurrence in western Europe.     

The African species Megantereon whitei from the Early Pleistocene of Monte Argentario (South Tuscany,Central Italy)

A partial skull and articulated postcranial elements of Megantereon whitei have been recorded during the 1950s from a karst deposit in the Monte Argentario area (Grosseto). These fossils recently became available for study. The bones are quite well preserved, included in a hard reddish matrix with calcareous clasts. The fossil is part of a faunal assemblage referred to the Late Villafranchian (Early Pleistocene). M. whitei from Monte Argentario is characterised by elongated upper canines, stronger than those of the Upper Valdarno (Tuscany) specimens and similar to the Pirro Nord (Apulia) fossil. The manus is robust, the first phalanges are quite long in comparison to the metacarpals. The Early Pleistocene European M. whitei represents an African element that took part in the faunal dispersal from Africa to Europe that occurred at the Plio-Pleistocene transition.     

The presence ofsus strozzii in the villafranchian (villanyian) of macedonia (greece)

The locality of Gerakarou is a new Pleistocene one in Macedonia (Greece) found in 1978. Between the material recovered there is a suid skull, which is studied in this article. Its morphological characters and dimensions allow us to determine this asSus strozzii Forsyth Major. It seems to be similar to the Upper Valdarno and Olivola suids and different from the other Villafranchian suid,Sus minor. Thus it is dated to Late Villafranchian (Villanyian). The certain presence of the species in Greece, where it was known only from some isolated specimens referred asSus cf.strozzii, completes the knowledge of the geographic distribution of the species in Southern Europe.     

Plio-Pleistocene evolution of the genus Castor (Rodentia, Mammalia) in Europe: C. fiber plicidens of Pietrafitta (Perugia, Central Italy)

In the lignite sediments of Pietrafitta (Tiberino Basin, Umbria, Central Italy), a rich fossil assemblage of vertebrate, invertebrate and plant remains belonging to the Farneta Faunal Unit (Late Villafranchian, Early Pleistocene) was collected. Among them fossil beaver cranial remains are relatively abundant. Western-Central Europe Villafranchian beaver remains were assigned to C. plicidens by some authors because molar occlusal surface patterns show complex enamel crenulations. Several recent authors have classified them as C. fiber while analysing other morphological patterns. Our samples have been compared to Plio-Pleistocene fossil remains and to living European populations of the genus Castor. New morphometric parameters on molar occlusal surfaces have been defined and statistical analyses (One-Way MANOVA, Principal Component Analysis and Canonical Discriminant Analysis) have been performed on them. The results point out a subspecific separation between the Late Villafranchian beaver of Pietrafitta, Quarata and San Giovanni in Valdarno (Umbria and Tuscany) and C. fiber populations. St. Vallier (France) Late Pliocene and Mosbach 2 (Germany) Middle Pleistocene beavers classified respectively by Viret and Friant as C. plicidens, show a C. f. fiber molar teeth pattern. Therefore, C. fiber plicidens did not occur in Central-Western Europe and this subspecific name may be used only for the local populations of Valdarno and Tiberino Basin (San Giovanni in Valdarno, Quarata, Pietrafitta and a few localities of the same area), at that time peripheral populations, probably semi-isolated during the Late Villafranchian, and at the southern limit of the geographic range of C. fiber.     

Lower Pleistocene stratigraphy of the Burdur Basin of SW Anatolia

The Burdur Basin is one of the NE-trending extensional orogen-top basins of SW Anatolia hosting alluvial-fan, fluvial and lacustrine deposits from the Late Miocene onward. The remains of Equus (Allohippus) sp., Paracamelus cf. gigas, and medium-sized deer (Cervidae gen. indet.) from the uppermost reach of the basin-fill succession show an interregional palaeobiogeographical linkage. The composition of fossil associations is typical for the Villafranchian of eastern Europe and central Asia. The architecture of the basin-fill stratigraphy and associated fossil taxa refines the Early Pleistocene regional palaeogeography and biodiversity of Anatolia drawn over the Eurasian migration pattern.     

The fossil American lion (Panthera atrox) in South America: Palaeobiogeographical implications

By the late 19th and early 20th centuries, authors described several specimens belonging to a very large felid, the size of a lion, from some Late Pleistocene localities at southern Chile and Argentina. These remains were considered as belonging to large and now extinct subspecies of jaguar. In the present contribution based on qualitative and quantitative postcranial and cranial characters, we conclude that that the “Patagonian Panthera” should be considered as representing remains of the American Cave Lion Panthera atrox. Evidence at hand indicates that Panthera atrox was a very large lion-like felid, with reddish skin, that inhabited dry open habitats. Furthermore, in contrast to living jaguars, transported and accumulated its preys in rocky dens. The presence of at least two pantherine lineages (i.e. P. onca and P. atrox) in the Pleistocene of South America indicates that the lineage has a long and complex history in the New World, and that the history of the clade in this landmass is still far from being well understood.     

Revision of the rhinoceros remains (Rhinocerotidae,Mammalia) from the late Pliocene of Etouaires (Auvergne,France) and the morphological distinction between the postcranial bones of Stephanorhinus elatus and S. etruscus

This work revises the rhinoceros remains from the well-known early Villafranchian locality of Étouaires (Auvergne, France), which have been collected on multiple occasions since the late 18th century. The species Stephanorhinus elatus and S. etruscus are present, both represented mainly by postcranial elements. To identify them, a detailed preliminary analysis of the morphological differences between the postcranial skeletons of the two species has been undertaken, using the material from Vialette (Haute-Loire, early Villafranchian) and Senèze (Haute-Loire, early late Villafranchian) as comparison for S. elatus and S. etruscus respectively, plus some specimens of S. etruscus from Upper Valdarno (Tuscany, middle Villafranchian). These localities have been chosen because they all yielded only one of the two species. The morphological distinction between the two species is not easy, since the interspecific variability often overlaps and includes polymorphic characters. However, a few diagnostic characters are described, and intra-specific variability has been investigated as much as possible with the fossil material under investigation.     

Premiere decouverte en Europe Sud-occidentale de Praeovibos priscus (Mammalia,Artiodactyla, Ovibovinae) dans le gisement pleistocene moyen ante-risseien de la Caune de l'Arago (Tautavel,Pyrenees-Orientales,France)

Teeth, cranial and postcranial skeletton of Praeovibos were sampled from the sediments of the Caune de l'AragoCave. This material is enough to recognize the anatomical relationships between Praeovibos and Ovibos. Praeovibos is discovered for the first time in France and in South Western Europe also. The problem of the correlation of the whole European Middle Pleistocene is approached from a different point of view.     

Absolute and relative endocranial size in Neandertals and later Pleistocene Homo

Eurasian Neandertals encompass the entire observed range of recent and fossil Homo sapiens in absolute, but not relative endocranial volume, and Neandertals attest an average EQ significantly lower than their Upper Pleistocene successors. While the cognitive, social, and evolutionary implications of this phenomenon have been emphasised, the statistical basis of a mean inference of EQ in the Neandertal hypodigm has not been appropriately demonstrated. A demonstrable male bias in the available postcranial, not cranial, series has skewed perceptions of Neandertal brain-to-body size scaling towards a rejection of the null hypothesis. A simple resolution to this problem is a concise assessment of paired associated covariates against a suitable recent human comparator series. Permutations of Fisher's z and Student's t statistics are valid metrics in tests of significance in single datum hypotheses.     

The Pachycrocuta and Hyaena lineages (plio-pleistocene and extant species of the Hyaenidae). Their relationships with miocene ictitheres: Palhyaena and Hyaenictitherium

The large «crocutoïd hyaenas from the Plio-Pleistocenedeposits of Eurasia do not belong to the same lineage as the extant species Hyaena hyaena and must be referred to the genus PachycrocutaKretzoi, of which an emended diagnosis is given. This revision takes into account some undescribed or ill-known specimens from Russia, China and Africa. They definitely establish that Pachycrocuta ranged over Eurasia, from West Europe to East China, as early as the early Villafranchian at least, and that it was present in North Africa too; a large sample from the Odessa Catacombs affords an estimation of the intraspecific variation in the Ruscinian species, H. pyrenaica, known until now by only a few specimens from the western part of the Mediterranean basin, and it shows H. pyrenaica to be the ancestral form of the Villafranchian Eurasian species P. perrieri, from which derived P. brevirostris, the last species of the lineage, as previously shown by other authors.It appears that the “Hyaena lineage evolved simultaneously in Africa; we knew already that the root of this lineage is H. abronia, a species from the late Miocene of South Africa whose generic attribution is discussed relative to some Ictitheres from Shan-Si, Samos, Sahabi and Klein Zee. The hypothesis of a common African origin of the two lineages is not excluded, if not demonstrated. The relationships of the Pleistocene European species H. prisca and that of the extant African species H. brunnea are discussed.     

New fossils of Eucladoceros boulei (Artiodactyla,Mammalia) from Early Pleistocene Nihewan Beds,China

Recent excavations at Shanshenmiaozui site in Nihewan Basin of North China uncovered a rich collection of comb-antlered deer, which includes the first discovery of the yearling antler, complete upper and lower dentitions (both deciduous and permanent), associated limb bones including the longest metapodials ever recovered. Based on toothrow length and the dental characters as well as the postcranial bones, the new fossil materials can be referred to Eucladoceros boulei that is estimated to be 350 kg and represents the largest Pleistocene cervid ever recovered in China. Besides the large size, E. boulei is also characterized by the pronounced anterior cingulids and entostylid ribs on lower molars as well as the moderately pachyostosed mandibles. The present study shows that the body weight of large cervids can be estimated by the length of toothrow and metacarpal, with exception for Cervus elaphus, which has larger toothrow length, but shorter metacarpal and smaller body size. E. boulei is a typical element of the Early Pleistocene fauna in northern China. The early Villafranchian is a bottleneck period for cervid evolution in northern China, which is characterized by the following features: decrease of cervid diversity, disappearance of archaic groups, and the rise of the medium to large-sized three-tined cervini taxa. The sudden appearance of the very large and/or multi-tined cervids at the Pliocene–Pleistocene transition may represent a great migration event of mammals. The Early Pleistocene cervids from Nihewan Basin are very diverse, and are in need of more taxonomic work.     

The fossil Bovidae (Artiodactyla, Mammalia) from Gesher Benot Ya'aqov, Israel: Out of Africa during the Early-Middle Pleistocene transition

We report the study of the collection of fossil bovid specimens from the Early-Middle Pleistocene Acheulian site of Gesher Benot Ya‘aqov. This locality, situated in the Levantine Corridor (the bottleneck that connects Africa and Eurasia) is a key site to explain the faunal and human dispersals out of Africa during the Matuyama/Brunhes boundary around 0.8 Ma. Two species of bovine (Bos sp., and Bovini gen. et sp. indet. cf. Bison sp.), one antelope (Gazella sp. cf. G. Gazella), and another indeterminate Bovidae gen. et sp. indet., have been recorded. The largest species, Bos sp., is an African immigrant related to the species from the Eritrean site of Buia, Bos buiaensis, which evolved from the buffalo of Olduvai Pelorovis oldowayensis, and colonized the Eurasian continent in parallel with the dispersal of the Acheulian culture into the northern continent. Numerous important species first recorded in several localities of Early-Middle Pleistocene transition from Eurasia are included in this dispersal out of Africa, including the megaherbivore, Palaeoloxodon antiquus, and the carnivores Crocuta crocuta, and later, Panthera leo and Panthera pardus. This faunal turnover is coincident with the change to colder climates that dominated the Middle Pleistocene.     

Systematical position and paleoecology of the endemic deer Megaceroides algericus Lydekker, 1890 (Cervidae,Mammalia) from the late Pleistocene-early Holocene of North Africa

The unusual cranial morphology of the endemic extinct African deer Megaceroides algericus (Lydekker, 1890) is described. Some details of cranial and dental morphology suggest that M. algericus is closely related to the Eurasian giant deer Megaloceros giganteus (Blumenbach, 1799). The paper presents also a discussion on paleoecology and functional morphology of M. algericus, as well as its origin, phylogenetic and taxonomic position. Megaloceros mugarensis (Di Stefano, 1996) from the middle Pleistocene of Levant is regarded as a probable forerunner of M. algericus.     

Partial skeleton of Theropithecus brumpti (Primates, Cercopithecidae) from the Chemeron Formation of the Tugen Hills, Kenya

Here we describe a complete skull and partial skeleton of a large cercopithecoid monkey (KNM-TH 46700) discovered in the Chemeron Formation of the Tugen Hills at BPRP Site #152 (2.63 Ma). Associated with the skeleton was a mandible of an infant cercopithecoid (KNM-TH 48364), also described here. KNM-TH 46700 represents an aged adult female of Theropithecus brumpti, a successful Pliocene papionin taxon better known from the Omo Shungura Formation in Ethiopia and sites east and west of Lake Turkana, Kenya. While the morphology of male T. brumpti is well-documented, including a partial skeleton with both cranial and postcranial material, the female T. brumpti morphotype is not well-known. This skeleton represents some of the first associated evidence of cranial and postcranial female T. brumpti remains. In addition to the complete skull, postcranial material includes elements of the axial skeleton and lower limb. While aspects of the skeleton conform to those of specimens previously assigned to T. brumpti, other features on the femur and tibia appear to differ from those previously described for this species. It is unclear whether these differences represent general variation within the T. brumpti population, variation between the sexes in T. brumpti, or the incorrect assignment of previous isolated hindlimb specimens. In total, the observable morphological features of the hindlimb suggest that KNM-TH 46700 was a terrestrial quadruped similar to modern savannah baboons (Papio). From the available evidence, it is difficult to assess whether or not KNM-TH 46700 frequently engaged in the specialized squatting and shuffling behavior observed in extant geladas (Theropithecus gelada).     

Deer of the genus <Emphasis Type="Italic">Megaloceros</Emphasis> (Mammalia,Cervidae) from the Early Pleistocene of Ciscaucasia

A new deer species, Megaloceros stavropolensis sp. nov., from the pre-Apsheronian sandy–clayey deltaic deposits of the Georgievsk sand pit (village of Podgornoe, Stavropol Region) is described. The fauna of large and small mammals from the bone beds of this locality is dated terminal Villanian, end of the MNQ 17 Zone, beginning of the Late Villafranchian, and assigned to the beginning of the Psekups Mammalian Assemblage. This is the earliest known member of the genus, which possesses antlers with well-developed palmation. The middle tine is absent and the posterior tine is a part of the antler palmation. Other specimens of Early Pleistocene Megaloceros are represented by fragments.     

Re-Defining <Emphasis Type="Italic">Canis etruscus</Emphasis> (Canidae,Mammalia): A New Look into the Evolutionary History of Early Pleistocene Dogs Resulting from the Outstanding Fossil Record from Pantalla (Italy)

An outstanding sample of Canis etruscus has been found within the faunal assemblage from the early Pleistocene site of Pantalla (Italy), which is referred to the early late Villafranchian. Canis etruscus appeared in Europe about 2 Ma ago. It is regarded as an important taxon for biochronology, as its first occurrence (the “wolf event”) has been used to define one of the Villafranchian faunal turnovers. The discovery of four crania from Pantalla prompted a revision of C. etruscus, in order to better describe its cranial morphology. Since early studies, the distinction between C. etruscus and the coeval C. arnensis has been based mainly on mandibular traits. For this reason, our study is aimed at highlighting differences in craniodental characters between the two species. Canis arnensis has been conventionally considered a jackal-like dog, while C. etruscus is regarded as a wolf-like dog. Consequently, we decided to use jackals for comparison, in addition to C. lupus. Although the jackal group has been traditionally considered as quite homogenous (different species are partially sympatric and similar in both size and ecology), recent genetic studies demonstrate that jackals are not monophyletic. Considering the model offered by extant species, our goal is to delineate the degree of intra- and interspecific variability among the basal forms of the genus Canis.     

Fossil penguin (Aves: Sphenisciformes) cranial material from the Eocene of Seymour Island (Antarctica)

Our knowledge of the cranial morphology of early penguins remains poor, particularly for Paleogene taxa. This paper describes a partial penguin skull and additional isolated cranial elements from the Eocene La Meseta Formation of Seymour Island, Antarctica. These specimens cannot be assigned to named taxa at present, but there is a strong possibility they belong to La Meseta penguins known only from postcranial elements. The skull shares extensive dorsal development of the temporal fossae with extant and fossil Spheniscus and the fossil penguins Paraptenodytes and Marplesornis, indicating the adductor complex was powerful in early penguins. Partial mandibles belonging to a much larger penguin are similar to Paraptenodytes and differ from all living penguins in the lack of a hooked medial process of the articular and the presence of a foramen anterior to the mandibular cotyles. Given the rarity of penguin cranial remains, these specimens provide important new insight into early penguin evolution.     

Allodaposuchus palustris sp. nov. from the Upper Cretaceous of Fumanya (South-Eastern Pyrenees,Iberian Peninsula): Systematics,Palaeoecology and Palaeobiogeography of the Enigmatic Allodaposuchian Crocodylians

The controversial European genus Allodaposuchus is currently composed of two species (A. precedens, A. subjuniperus) and it has been traditionally considered a basal eusuchian clade of crocodylomorphs. In the present work, the new species A. palustris is erected on the base of cranial and postcranial remains from the lower Maastrichtian of the southern Pyrenees. Phylogenetic analyses here including both cranial and postcranial data support the hypothesis that Allodaposuchus is included within Crocodylia. The studied specimen suggests little change in postcranial skeleton along the evolutionary history of crocodylians, except for some bone elements such as the axis, the first caudal vertebra and the ilium. The specimen was found in an organic mudstone corresponding to a coastal wetland environment. Thus, A. palustris from Fumanya is the first Allodaposuchus reported in lacustrine-palustrine settings that expand the ecological range for this genus. The S-DIVA palaeobiogeographic reconstruction of ancestral area suggests that early members of Crocodylia rapidly widespread for the Northern Hemisphere landmasses no later than the Campanian, leading the apparition of endemic groups. In that way “Allodaposuchia” represents an endemic European clade probably originated in the Ibero-Armorican domain in the late Campanian and dispersed by the Southern European archipelago prior to the early Maastrichtian.