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Early Pleistocene Equidae and Suidae from Palan-Tyukan (Azerbaijan)
Historical Biology
Abstract
The Early Pleistocene site of Palan-Tyukan is located in Transcaucasia, northwestern Azerbaijan. More than 300 mammalian bones were laid close to each other in a 25 m2 lens-like accumulation, in a stratum of normally magnetised (the upper part of the Olduvai subchron) yellowish-grey Lower Apsheron loams. The present study is based on the analysis of the Equidae and Suidae remains. The taxonomic scenario of the Early Pleistocene Equidae is intricate and has been a matter of long debate with a multitude of hypotheses. The small-sized horse remains from the locality are ascribed here to species Equus (Allohippus) senezensis. The species likely used here are mostly open landscapes for their main habitat. The main conclusion about the Suidae remains from Palan-Tyukan was reached after a detailed morphological and metrical analysis. The fossil material is here referred to Sus strozzii. Seemingly, during the initial phase of the Early Pleistocene the species lived in humid subtropical Transcaucasian riverine forests and swamps. An Equidae/Suidae community of the Palan-Tyukan type presents evidence of the wider variety of environments ranging from wooded areas including bodies of water to scrub and even savannas as a landscape in a relatively humid subtropical climate.
... Before the wild boar appeared in the European fossil record close to the Early-Middle Pleistocene boundary ( Van der Made et al., 2017;Cherin et al., 2020;Iannucci, 2022), S. strozzii Forsyth Major, 1881 occupied the territory for most of the Early Pleistocene, being recorded from several localities from Spain to western Asia (Mein et al., 1978;Van der Made and Moyà-Solà, 1989;Kostopoulos and Athanassiou, 2005;Cherin et al., 2018;Iannucci et al., 2020a;Iltsevich and Sablin, 2022). Sus strozzii was a large-sized suid closely related to the so-called warty pigs of Island Southeast Asia (Azzaroli, 1952(Azzaroli, , 1975Cherin et al., 2018;Iannucci et al., 2020a), which constitute a diverse and yet relatively poorly known contingent of species (Hardjasasmita, 1987). ...
... Unfortunately, there are not but a few mammal localities of this age in Europe, and as previously remarked even less document the occurrence of suid for the whole middle Villafranchian. Sus strozzii and S. arvernensis are usually regarded as species adapted to tropical to subtropical or even swampy environments (e.g., Pickford and Obada, 2016;Iltsevich and Sablin, 2022), a consideration that rests mainly on their phylogenetic proximity to warty pigs from Island Southeast Asia and on the presence, at least in S. strozzii, of some anatomical similarities with Potamochoerus (Azzaroli, 1952(Azzaroli, , 1975Faure and Guérin, 1984;Iannucci et al., 2020a). If this palaeoecological interpretation is correct, the onset of the Quaternary glaciations and the consequent spread of open environments at 2.6 Ma may have forced suid populations to cope with stressful conditions, eventually promoting the evolution of S. strozzii, a new species able to survive during the Early Pleistocene. ...
Some remains of suids were recovered during the second half of the 1800s from Quercia (Tuscany, Italy), at a close distance to, but from a lower stratigraphic position than the better-known mammal fauna of Olivola. Although they were collected a long time ago, Quercia suids are described for the first time in this work. This sample represents one of the earliest occurrences (middle Villafranchian) of Sus strozzii, a large-sized suid present in Europe during the Early Pleistocene, but only abundant ∼2.0–1.8 Ma. A biometric comparison with selected samples of Pliocene to early Middle Pleistocene European suids is carried out, showing some dimensional changes in S. strozzii as well as differences between S. strozzii and other species. Quercia has been somehow eclipsed by other historical samples from Tuscany, namely the extensive collection of the Upper Valdarno and the diverse fauna of Olivola, but it is a different and important palaeontological locality. Apart from S. strozzii, the local fauna of Quercia-Vaccareccia includes Anancus arvernensis, Canis sp., Stephanorhinus etruscus, Leptobos etruscus, ‘Pseudodama’ sp., and Castor sp., representing one of the few mammal assemblages referable to the Faunal Unit of Coste San Giacomo (late middle Villafranchian, MNQ 17b), corresponding to ∼2.2–2.1 Ma.
The species composition of the mid-Early Pleistocene fauna in northwestern Azerbaijan has been elucidated through the analysis of fossil material recovered from the Palan-Tyukan site. This research also included a review of the relevant literature. The paleontological material has been fossilized to a similar extent, indicating similar burial conditions. Animal bones at the site were accumulated during a single sedimentation cycle in sediments of what was once a shallow, heavily silted body of water with slow-flowing or standing water. We have identified 16 species of large mammals, whose presence has allowed us to determine the time period in which this faunal complex existed, as well as the characteristics of the natural environment at the time of its formation. The order Carnivora is represented by 10 individuals, the order Proboscidea by 2 individuals, the order Perissodactyla by 3 individuals, and the order Artiodactyla by 18 individuals. The Palan-Tyukan paleo-landscape can be described as a mosaic of different environments. Flat riverine areas with dense vegetation gradually transitioned into the hilly steppe with mixed vegetation of varying densities. Our reconstruction of the paleogeographic environment indicates that large mammals of Palan-Tyukan lived there under conditions of a relatively humid subtropical climate. The Palan-Tyukan faunal complex belongs to the Mediterranean palaeobiogeographical subregion, biozone MNQ18, at the beginning of the Late Villafranchian period, and falls within the time interval from 1.93 to 1.77 million years ago.
The Early Pleistocene site of Palan-Tyukan (MNQ18, ca 1.85 Mya) is located in Transcaucasia, northwestern Azerbaijan. In total, more than 300 mammalian bones were collected there by one of the authors (M.V. Sablin) in 1986 and 1990. The remains were laid close to each other in a 25 m2 lens-like accumulation, in a stratum of normally magnetized (the upper part of the Olduvai subchron) yellowish-grey Lower Apsheron loams. No any excavations have been carried here out since. The present study is based on the analysis of the Bovidae remains. The material is represented by fragments of skulls and horn cores, isolated teeth and bones of the postcranial skeleton. As a result of the revision of the collection, Leptobos (Smertiobos) cf. etruscus (Falconer, 1859), Gazellospira torticornis (Aymard, 1854) and Gazella cf. bouvrainae (Kostopoulos, 1996) were identified. The degree of the unworn M3 tooth hypsodonty of Leptobos, as well as the size and morphology of limb bones indicate a greater similarity with those of L. etruscus from Olivola and Tasso. The characteristics of the horn cores, as well as the upper teeth and limb bones of the medium-sized spiral-horned antelope from Palan-Tyukan indicate a greater similarity with those of the nominative subspecies G. torticornis torticornis. The dimensions and degree of flattening of the horn cores, as well as the parameters of the p4 tooth, are similar to those of the medium-sized gazelle G. bouvrainae, described from several Early Pleistocene localities of Greece. The taxonomic composition of the Bovidae association from Palan-Tyukan is consistent with the previously determined stratigraphic position of the locality, the fauna from which belongs to the Psekups faunal complex of Eastern Europe, correlated with the Late Villafranchian of Western Europe. The composition of these herbivores community testifies to the spread of savanna-like forest-steppe landscapes in the west of Transcaucasia during this period of the Early Pleistocene.
The Early Pleistocene site of Palan-Tyukan (MNQ18, ca 1.85 Mya) is located in Transcaucasia, northwestern Azerbaijan. More than 300 mammalian bones were laid close to each other in a 25 m² lens-like accumulation, in a stratum of normally magnetised (the upper part of the Olduvai subchron) yellowish-grey Lower Apsheron loams. The Palan-Tyukan fauna includes two species of sabertoothed cats. The medium-sized sabertoothed cat remains are ascribed here to Megantereon cf. cultridens. The large size of the massive fossil ulna from Palan-Tyukan is comparable to that of a broadly lion-sized felid. We attribute the bone to the species Homotherium cf. crenatidens. The morphological characters and the size of the Panthera remains from Palan-Tyukan suggest its similarities with the P. gombaszogensis remains from Olivola, Upper Valdarno and Dmanisi localities (the initial phase of Early Pleistocene). The occurrence of P. cf. gombaszogensis in Palan-Tyukan provides information on stratigraphic range of these taxa: it represents one of the earliest records of the genus Panthera in all of Eurasia. The hyaenid remains are ascribed here to Pliocrocuta perrieri. A Feliformia community of the Palan-Tyukan type presents evidence of the wider variety of environments ranging from wooded areas and savanna landscape.
The Early Pleistocene site of Palan-Tyukan (MNQ18, ca 1.85 Mya) is located in Transcaucasia, northwestern Azerbaijan. In total, more than 300 mammalian bones were collected there by M.V. Sablin in 1986 and 1990. The remains were laid close to each other in a 25 m2 lens-like accumulation, in a stratum of normally magnetized (the upper part of the Olduvai subchron) yellowish-grey Lower Apsheron loams. No any excavations have been carried here out since. The present study is based on the analysis of the Caniformia remains. The occurrence of raccoon dog in Palan-Tyukan provides knowledge about the Eurasian stratigraphic range of this taxon: it represents one of the latiest records of the species Nyctereutes megamastoides (Pomel, 1842) in all of Europe. The bear tooth (was found earlear by N.A. Lebedeva nearby) is ascribed here to the species Ursus etruscus Cuvier, 1823. The taxonomic scenario of the Plio-Pleistocene tribe Galictini of Eurasia is intricate and has been debated since the beginning of the last century. The dispute arose mainly due to the scantiness and the sparse nature of their fossil record. The main conclusion about the Pannonictis nestii (Martelli, 1906) remains from Palan-Tyukan was reached after a detailed morphological and metrical analysis. Palearctic badger remains were assigned to the species Meles thorali Viret, 1950 – the ancestor of the European and Asian badgers. The occurrence of M. thorali in Palan-Tyukan represents one of the latiest records of the species. The otter material from Palan-Tyukan is here referred to Lutraeximia cf. umbra Cherin et al., 2016. A Caniformia community of the Palan-Tyukan presents evidence of the wider variety of environments ranging from wooded areas with bodies of water to scrub and even savanna landscape in a relatively humid subtropical climate. The fauna of Palan-Tyukan is an important biostratigrafic reference fauna for mammal zonation in Europe. New discoveries can also help to clarify the taxonomic status of many of these extinct forms.
Evolution of the genus Equus has been a matter of long debate with a multitude of hypotheses. Currently, there is no consensus on either the taxonomic content nor phylogeny of Equus. Some hypotheses segregate Equus species into three genera, Plesippus, Allohippus and Equus. Also, the evolutionary role of European Pleistocene Equus stenonis in the origin of the zebra-ass clade has been debated. Studies based on skull, mandible and dental morphology suggest an evolutionary relationship between North American Pliocene E. simplicidens and European and African Pleistocene
Equus. In this contribution, we assess the validity of the genera Plesippus, Allohippus and Equus by cladistic analysis combined with morphological and morphometrical comparison of cranial anatomy. Our cladistic analysis, based on cranial and postcranial elements (30 taxa, 129 characters), supports the monophyly of Equus, denies the recognition of Plesippus and Allohippus and supports the derivation of Equus grevyi and members of the zebra-ass clade from European stenonine horses. We define the following evolutionary steps directly relevant to the phylogeny of extant zebras and asses:
E. simplicidens–E. stenonis–E. koobiforensis–E. grevyi -zebra-ass clade. The North American Pliocene species Equus simplicidens represents the ancestral stock of Old World Pleistocene Equus and the zebra-ass clade. Our phylogenetic results uphold the most recent genomic outputs which indicate an age of 4.0–4.5 Ma for the origin and monophyly of Equus.
The monodactyl horses of the genus Equus originated in North America during the Pliocene, and from the beginning of the Pleistocene, they have been an essential part of the large ungulate communities of Europe, North America and Africa. Understanding how body size of Equus species evolved and varied in relation to changes in environments and diet thus forms an important part of understanding the dynamics of ungulate body size variation in relation to Pleistocene paleoenvironmental changes. Here we test previously published body mass estimation equations for the family Equidae by investigating how accurately different skeletal and dental measurements estimate the mean body mass (and body mass range) reported for extant Grevy's zebra (Equus grevyi) and Burchell's zebra (Equus quagga). Based on these tests and information on how frequently skeletal elements occur in the fossil record, we construct a hierarchy of best practices for the selection of body mass estimation equations in Equus. As a case study, we explore body size variation in Pleistocene European Equus paleopopulations in relation to diet and vegetation structure in their paleoenvironments. We show a relationship between diet and body size in Equus: very large-sized species tend to have more browse-dominated diets than small and medium-sized species, and paleovegetation proxies indicate on average more open and grass-rich paleoenvironments for small-sized, grazing species of Equus. When more than one species of Equus co-occur sympatrically, the larger species tend to be less abundant and have more browse-dominated diets than the smaller species. We suggest that body size variation in Pleistocene Equus was driven by a combined effect of resource quality and availability, partitioning of habitats and resources between species, and the effect of environmental openness and group size on the body size of individuals.
The Miocene-Pliocene (Turolian-Ruscinian) transition represents a fundamental interval in the evolution of Euro-Mediterranean paleocommunities. In fact, the paleoenvironmental changes connected with the end of the Messinian salinity crisis are reflected by a major renewal in mammal faunal assemblages. An important bioevent among terrestrial large mammals is the dispersal of the genus Sus, which replaced all other suid species during the Pliocene. Despite its possible paleoecological and biochronological relevance, correlations based on this bioevent are undermined by the supposed persistence of the late surviving late Miocene Propotamochoerus provincialis. However, a recent revision of the type material of this species revealed an admixture with remains of Sus strozzii, an early Pleistocene (Middle Villafranchian to Epivillafranchian) suid, questioning both the diagnosis and chronological range of P. provincialis. Here we review the late Miocene Suidae sample recovered from the Casino Basin (Tuscany, central Italy), whose taxonomic attribution has been controversial over the nearly 150 years since its discovery. Following a comparison with other Miocene, Pliocene, and Pleistocene Eurasian species, the Casino Suidae are assigned to P. provincialis and the species diagnosis is emended. Moreover, it is recognized that all the late Miocene (Turolian) European Propotamochoerus material belongs to P. provincialis and that there is no compelling evidence of the occurrence of this species beyond the Turolian-Ruscinian transition (MN13-MN14).
Herein we describe a fragment of a mandible with a deciduous fourth premolar (dp4) from the Early Pleistocene locality of Dunaalmás, representing the first confirmed report of Sus strozzii from Hungary. The comparison of dp4 measurements supports a statistically significant distinction between S. strozzii and Sus scrofa. The two species overlap in time during the late Early Pleistocene of Europe (Epivillafranchian), but suid remains of this time-span are seldom classified at a species level. The correct taxonomic identification of the Epivillafranchian suids, which are often associated with evidences of hominin presence, is of great palaeoenvironmental value because S. scrofa and S. strozzii possess different ecological requirements.
We present an inventory of the progress of recent research on the biostratigraphy and palaeoecology of the genus Equus sensu lato in Europe. Our discussion starts with the new hypotheses concerning the dispersal and evolution of non-caballine equids of the Early and the beginning of the Middle Pleistocene, focusing on recent discoveries, description of taxa, and revised diagnosis. In particular, we deal with the major debate surrounding the number of lineages, “stenonid horses,” related species and the new subgenus “Sussemionus.” Possible phylogenetic scenarios and relationships with extant species are also taken into consideration. There is consensus that the lineage of true horses emerged in Europe at the beginning of the Middle Pleistocene. Their extensive skeletal plasticity enabled them to survive under a wide range of climatic conditions and environments, from interglacial forests to grass-steppes during glacial episodes. They exhibit homogeneous overall morphology but variability in terms of dental and skeletal proportions. Patterns of general body size, muzzle shape, macro- and microdonty, proportions of the limbs, robustness of the metapodials, and breadth of the third phalanx are influenced by global climate, the characteristics of the vegetation (in particular the degree of openness) and substrate, and are governed by Bergmann's and Allen's rules or other environmental pressures. In addition to palaeoecological information, these ecomorphological adaptations paradoxically provide real biochronological details in a given geographical province. Throughout the Upper Pleistocene, horses underwent a size diminution initiated at the end of the Middle Pleistocene. After the Last Glacial Maximum (LGM), investigation of body size in different regions of Europe demonstrates the existence of latitudinal and longitudinal clines, and the fragmentation of the horse population. The so-called European wild ass, Equus hydruntinus, has a wide geographical distribution but seems to be less ecologically flexible, or, alternatively, more specialized than true horses, with which it is often found in sympatry. The latest palaeogenetic studies place the species into phylogenetic context within the Asiatic wild asses, however, palaeontological evidence points to E. hydruntinus as a separate species. Its tooth morphology varies between biogeographical areas.
The intermontane Anagni Basin (Frosinone, central Italy) is an important region for Italian biochronology and paleoecology due to the presence of two rich fossil assemblages dated to the Early (Coste San Giacomo) and Middle Pleistocene (Fontana Ranuccio). These sites have yielded a vast collection of large fossil mammals with a well-documented presence of fossil equids in both localities (represented mostly by isolated teeth). Coste San Giacomo is dated to around 2.1 Ma, thereby having recorded the effects of the onset of the Quaternary glacial cycles, which led to a gradual trend toward colder and more arid conditions in the Northern Hemisphere. The fossil equids of this site belong to the first group of grazing stenonid equids of the genus Equus that spread to the Italian Peninsula during the so called “Elephant- Equus” event, which marked the appearance of new large mammals living in herds in open and arid environments. The site of Fontana Ranuccio is dated to around 400 ka, close to the MIS 12–11 succession (the “Mid-Brunhes event”), which marked the end of the Middle Pleistocene Transition. The fossil horses from Fontana Ranuccio represent one of the oldest caballoid (or “true horses”) populations of the Italian Peninsula. The Anagni Basin, thus, provides important data to investigate paleoecological adaptations of these groups of equids in response to two critical environmental and climatic shifts of the Pleistocene. We explore their niche occupation by examining long-term dental wear patterns and tooth enamel carbon and oxygen stable isotopic composition. Both taxa appear to have exhibited a narrow dietary niche, displaying a clear abrasive (highly specialized) grass-rich diet. In particular, caballoid equids from Fontana Ranuccio show a more abrasion-dominated mesowear signature. Stenonid equids from Coste San Giacomo exploited broader and more diverse landscapes during the Early Pleistocene, whereas caballoid horses from Fontana Ranuccio appeared to have limited their dietary adaptations to a stricter grazing behavior in more closed environments.
The appearance of monodactyl equids in Eurasia and their dispersal towards South Europe is a significant event that marks the beginning of the Quaternary period. During the Pleistocene, horses were a common element in most European large mammal faunal assemblages, providing important palaeoecological clues, but their taxonomy, nomenclature and phylogeny, as well as their actual biochronological significance have been widely debated by scholars. The evolutionary history of horses here regarded as “stenonoid” ( Equus livenzovensis , Equus stenonis , Equus ex gr. E. senezensis , Equus altidens , Equus hydruntinus , the large-sized horses Equus major and Equus suessenbornensis , sometimes referred to as the so-called “ Equus bressanus-Equus suessenbornensis group”), is one of the most intriguing. This study, mainly focusing on the South European representatives of this group, aims to highlight and discuss the major debated issues, re-examining the taxonomy of some Early and early Middle Pleistocene horses. Alternative phylogenetic hypotheses will also be discussed with the aim of providing new clues on the chronology and palaeoecology of some key species/samples.
The paper presents zooarchaeological analysis of the remains of Stenon horse Equus (Allohippus) stenonis from the site Muhkai 2a (layer 2), Central Dagestan, Russia. They are of special interest because of their large number and can testify to one of the first visits of an ancient man in the North Caucasus – about 1.95 million years ago. As a result of the comparison of the composition and ratio of the Stenon horse bones, their taphonomic features, showed that the site is the result of natural death of the animals. It is assumed that the bulk of the bones were laid in the deposits of a once shallow, periodically drying out and strongly silted reservoir with weakly flowing or standing water. After complete drying of the reservoir, the paleontological material was sealed in a clay mass, which in turn was buried during further geological processes. Based on this, we suggest a possible scenario for the formation of the bone accumulation. Most likely, it was a watering place for various animals, where some of them perished for natural reasons, for example, they got bogged down, drowned, and someone of them became prey for predators. Later, there was a burial of whole corpses, or parts thereof, transported to the pond. Obviously, the bodies of recently dead animals could not help attracting an ancient man, being an easily accessible source of protein food. The presence of artifacts and cut marks on the femur of Stenon horse indicate the presence of an ancient man in Muhkai 2a (layer 2), but the human activity seems to have played a minimal role in the taphocenosis forming.
This paper summarizes knowledge on the Neogene–Quaternary terrestrial fossil record from the Middle Kura Basin accumulated over a century and aims to its integration into the current research. This fossil evidence is essential in understanding the evolution of the Eurasian biome, since this territory is located at the border of Eastern Mediterranean and Central Asian regions. The general biostratigraphic framework suggests existence of two major intervals of the terrestrial fossil record in the area, spanning ca. 10–7 Ma and ca. 3–1 Ma, and points to an important hiatus between the late Miocene and late Pliocene. General aspects of the paleogeographic history and fossil record suggest that the biogeographic role of the Middle Kura Basin has been changing over geological time from a refugium (Khersonian) to a full-fledged part of the Greco-Iranian province (Meotian–Pontian). The dynamic environmental changes during the Quaternary do not depict this territory as a refugium in its general sense. The greatest value of this fossil record is the potential to understand a detailed history of terrestrial life during demise of late Miocene Hominoidea in Eurasia and early Homo dispersal out of Africa. Late Miocene record of the Middle Kura Basin captures the latest stage of the Eastern Paratethys regression, and among other fossils counts the latest and the easternmost occurence of dryopithecine, Udabnopithecus garedziensis, while the almost uninterrupted fossil record of the late Pliocene–Early Pleistocene covers the time interval of the early human occupation of Caucasus and Eurasia.
Sus cf. strozzii from the Khapry strata of the northeastern part of the Azov Region is described. The parameters and measurements of the tooth system correspond to those of S. strozzii in Western Europe. The specimens from the Khapry and Liventsovka localities suggest that this species is a typical member of the Late Pleistocene faunas of Eastern Europe.
Palaeontological data are compatible with the current biomolecular views about the emergence of Equus 2.3 Ma and of Hemiones around 1.1 Ma. There is, however, no evidence for an early separation of the Caballine branch. Recognizable Caballines appear at the beginning of the Middle Pleistocene, not between 2.2 and 1.6 Ma. At that period, two groups are documented: one with a Zebra-like skull and Zebra-Ass-like upper and lower dentitions, another with a very peculiar lower dentition. Probable descendants of the latter (unformally named "Sussemiones" because of some resemblances between Süssenborn and Hemione cheek teeth) are extremely widespread, from North America to Ethiopia, and include dry-adapted (E. granatensis-like) and humid-adapted (E. coliemensis-like) species. Their extinction seems contemporary with the appearance of Caballines and other species closely related to extant: E. hydruntinus related to Hemiones, E. melkiensis related to Asses, and E. mauritanicus and E. capensis related to Plains Zebras. © E. Schweizerbart'sche Verlagsbuchhandlung (Nägele u. Obermiller), 2006.
The recording of tooth wear is essential for the investigation of age in zooarchaeological assemblages, but most tooth wear methodologies apply only to mandibular teeth, thereby neglecting potentially valuable maxillary data. The large sample of pig maxillary jaws and teeth recovered at Durrington Walls has provided the opportunity to design a new recording method for maxillary as well as mandibular jaws. Work on previously excavated animal bone material from Durrington Walls (Albarella and Payne, 2005) suggested the possibility of seasonal pig killing at the site, but the issue has not, until now, been explored in detail. This paper therefore has a dual purpose: to describe the new method for recording tooth wear on pig teeth; and to use the new information from both the mandibular and maxillary teeth to explore pig age at death and seasonality at Durrington Walls. The results provide evidence of differential deposition of pigs of different ages at Durrington Walls, with one midden context containing younger pigs brought to the site to provide meat for predominately winter-based feasting events, and other contexts containing remains of older pigs (mainly in their second year) deposited in both domestic and more public locales also predominantly in winter. The study highlights the usefulness of maxillary teeth for our understanding of past systems of pig exploitation as well as the desirability of recording their wear in animal bone assemblages.
Statistical description and comparisons of the equids recently collected at Saint-Vallier (LD2 and LD3) and of the old collections. Estimations of the weight and height at the withers. Size indices. The fossils of the lower level (LD2) are intermediate in size between those of El Rincon 1 and of the younger level of Saint-Vallier. Moreover, they are as slender as those of El Rincon 1. Most fossils from the younger level (LD3) may be referred to A. stenonis vireti. A few fossils belong to a smaller and slenderer form. Weight estimations of equids aged between 2.5 and 1.5 Ma. Tentative classification according to weight and morphology. Taxonomical and chronological remarks, in particular on Livenzovka.
The fossil site of Senèze, located in the Auvergne volcanic province of the French Massif Central, is well-known for its mammalian fauna and its role as the reference locality for the Late Villafranchian time interval, an important phase in the evolution of the fauna of Europe. This site is a filled-in volcanic explosion-crater lake (maar) where fossils are found in volcano-clastic sediments interfingering with lacustrine deposits at the lake margin. The Senèze mammals are generally considered to date between 2.2-1.5 Ma, but some authors have argued that this assemblage is not unitary, rather composed of two associations of different age. Moreover, little has been published about the geological setting or the processes of site formation. Our Franco-American research project, co-directed by the three first authors since 2000, is re-examining Senèze for the first time in 60 years. The team effort has three main goals: 1) to clarify the local geology (stratigraphy, mineralogy) of this complex site and determine the taphonomic processes involved; 2) to utilize a combination of methods (ESR and argon dating; tephrochronological and paleomagnetic correlation) to definitively establish the age of the site and its fauna; and 3) to collect additional mammalian fossils (especially of rare animals such as carnivores, primates and rodents) and samples of the non-mammalian biota, including vertebrates (e.g., birds and fishes), terrestrial and lacustrine invertebrates, pollen and diatoms from known points within the revised and dated stratigraphy. Several partial skeletons which appear devoid of carnivore disturbance have been recovered, and a new taphonomic hypothesis is proposed to explain this pattern. Some preliminary argon dates are presented for levels below the fossil mammals, while preliminary ESR and paleomagnetic results agree with an age broadly near 2 Ma. Pollen collected in situ and from coprolites allows a first step toward paleoenvironmental reconstruction. © E. Schweizerbart'sche Verlagsbuchhandlung (Nägele u. Obermiller), 2006.
Kurzfassung Die Fundstätte Gerakarou wurde 1978 in Mazedonien entdeckt. Das dort gesammelte Material enthielt den dieser Abhandlung zugrunde
liegenden Suiden-Schädel, der sich nach seiner Form und seinen Dimensionen alsSus strozzii Forsyth Major bestimmen läßt. Er scheint mit den Suiden der Fundorte Olivola und oberes Valdarno übereinzustimmen, sich dagegen
vonSus minor, dem anderen Suiden des Villafranchium, zu unterscheiden. Die Fundschicht unseres Schädels gehört dem oberen Villafranchium
an. Da aus Griechenland bisher nur isolierte Einzelfunde vonSus cf.strozzii bekannt waren, erweitert der nun gesicherte Nachweis der Art die Kenntnis ihrer geographischen Verbreitung in Südeuropa.
Several localities of Late Pliocene mammal remains from the territory of north-east Azov
region (East Europe, Russia) are considered. Their geological and taphonomical characteristics
are given. 30 mammal taxa of the Khapry faunal unit were described. Th ere are 33 taxa of large
mammals in the revised list of this fauna. It is proved that the Khapry theriocomplex is the
analog of West European Middle Villafranchian faunas of the Saint-Vallier level. The correlation
with some Late Pliocene faunas from Europe and Asia is given.
The taxonomy of the European stenonoid horses is revised using multivariate statistical techniques in this paper. The objectives are: to identify the taxonomic status of this group; to revise the specific and subspecific diagnoses, and the valid taxa; and to analyze their paleoecology and evolution. We conclude that: 1) there are six species among the Plio-Pleistocene European stenonoids: Equus major Deperet, in Delafond and Deperet 1893, ex Boule; Equus livenzovensis Bajgusheva, 1978; Equus stenonis Cocchi, 1867; Equus senezensis Prat, 1964; Equus altidens von Reichenau, 1915; and Equus sussenbornensis Wats, 1901; 2) There are seven subspecies, three of E. stenonis, two of E. senezensis, and two of E. altidens; 3) There are two monophyletic groups according to the 'neighbor-joining method,' one composed of the middle-to-small-size species, and the other of the biggest ones; 4) There is a trend toward decreasing body size through time in stenonoid horses. This trend, that follows the inverse direction of Cope's role, can he related to the climatic-environmental changes that occurred from the middle Villafranchian to the early Galerian; 5) Large stenonoids were recorded in the middle Villafranchian and were associated with open and dry habitats (steppes), whereas the small ones lived in more closed and wet habitats (savanna-mosaic, woodlands) from late Villafranchian to earliest Galerian.
The enzymatic activity; the microbial population; and the intensities of the nitrification, ammonification, CO2emission, and cellulose decomposition were studied in gray-brown, meadow-sierozemic, meadow-forest alluvial, and yellow (zheltozem)
gley soils in the subtropical zone of Azerbaijan under natural vegetation, crop rotation systems with vegetables, and permanent
vegetable crops. On this basis, the biological diagnostics of these soils were suggested and the soil ecological health was
evaluated. It was shown that properly chosen crop rotation systems on irrigated lands make it possible to preserve the fertility
of the meadow-forest alluvial and zheltozem-gley soils and to improve the fertility of the gray-brown and meadow-sierozemic
soils.
European continental mammal biochronology has been developed in Italy since the pioneering work of Augusto Azzaroli. The Villafranchian is a Mammal Age, a biochronological unit. Biochronology is a sub-division of geological time on the basis of the succession of evolutionary stage of faunal assemblages and dispersal events. The Villafranchian Mammal Age is based on European large mammals and corresponds, in the International Stratigraphic Scale, to a time span from Late Pliocene to most of the Early Pleistocene. Within these limits the Villafranchian spans from around 3.5 Ma to about 1.0–1.1 Ma. “Villafranchian” is a concept relatively common within the scientific literature of continental stratigraphers in Europe. Unfortunately, it is sometimes used in the wrong way or in wrong contexts, especially because the term Villafranchian has been used with the meaning of a continental stage, totally misunderstanding its significance as a biochronologic unit. The authors provide an update of the Villaranchian Mammal Age and its present significance.
A sample of 225 skulls belonging to all extant species of Equus and of 33 fossil skulls of ages comprised between 3.4 Ma and 0.7 Ma was studied by multivariate analyses of size, shape, and form on four axial lengths. A simple bivariate discrimination using the overall palatal length and the cranial length allows an almost complete discrimination between modern (extant and young) equids (subgenus Equus) and old plesippine and stenonine forms. The modern pattern, possibly related to a bigger brain case, seems to appear less than 1.5 Ma ago in fossil species that can already be recognized as closely related to the extant Grevy's zebras, plains zebras, and horses. This does not support separate branchings of Equus species from a common plesippine stem.
The equid material from the late Pliocene locality of Sesklo (Thessaly, Greece) is described and compared in this article. It belongs to a large and fairly stout Equus stenonis form, which shares many morphological characters with the species samples from Saint-Vallier, La Puebla de Valverde and Olivola, as well as with already known Equus stenonis samples from other Greek localities (Dafnero, Volax). Its main features are the big skull and limbs, the short protocones and the very simple enamel plication in the teeth. Large sized and relatively robust stenonid horses are common elements of the late Pliocene faunas of Greece.
Dmanisi (Georgia) is one of the oldest Early Paleolithic sites discovered out of Africa. In addition, it is the best site to understand the first Homo deme out of Africa and the first hominin occupation of Central to Western Eurasia. It has produced more than 40 hominin remains, including several very informative skulls, found in direct association with faunal remains and numerous lithic artifacts. To date, fossil amphibians and reptiles are one of the few proxies that have been used to propose quantitative climate reconstructions for the time where the hominins were living at Dmanisi. The aim of the present study is to review and amplify the previous paleoclimatic interpretation given by Blain et al. (2014), with an enhanced methodology using geographic information system and based on uncertain distribution area-occupied distribution area discrimination technique. This procedure permits to approach to a more precise common species distribution area and then using the WorldClim v. 2.1 climate database to propose for the first time monthly reconstructions for temperature and rainfall, in addition to annual parameters. The same technique is used to infer dominant ecoregions through the study area and potential tree coverage. Dmanisi climate is newly reconstructed as warm and semi-arid, similar to the present-day Mediterranean climate. New estimates however suggest warmer temperatures than previously reconstructed, together with a slightly higher but much more irregular amount of rainfall. The aridity indexes suggest a six months dry period, from May to October. Associated regional paleoenvironment is mainly characterized by Mediterranean forests, woodlands, and scrub, and the potential tree coverage is around 25%, that is, much less forested than today. New estimates agree with the ‘Iberian’ hominin ecological model, and with other proxies (large mammals, small mammals and archaeobotanical remains) that indicate a period of increased aridity contemporaneous with human occupations of the site.
The Early Pleistocene locality Muhkai 2 was discovered in Central Dagestan, northeastern Caucasus, Russia, in 2006. Archaeological investigations of the site have been directed by Corresponding member of RAS H.A. Amirkhanov (Institute of Archaeology RAS, Moscow). The article presents the results of the study of osteological material from the excavations of 2006–2017. The cultural layers with stone tools and mammal bones are found within stratum of brownish-grey petrified loams with reversed magnetization. We identified 14 large species of mammals, which are indicators for the dating of the faunal assemblage and crucial in identifying the specific characteristics of the natural environment at the time of site formation. Stenon horse and Etruscan wolf dominate the spectrum of species at Muhkai 2. All large mammal species from the site are inhabitants of open and semi-open landscapes and animals indicative of closed biotopes are absent. The biotope appears to have been a dry savannah-steppe with small areas of forest vegetation. The climate at this time was warm and quite arid. Most likely, the animals died here due to natural causes and their carcasses were buried at the bottom of an ancient, slightly saline and shallow, temporary body of water. Data from the theriofauna place the age of the site Muhkai 2 within the chronological range of 2.1–1.77 Mya.
Résumé
Le présent article a pour objectif de présenter les résultats d’études de la collection paléontologique issue du site de Muhkai 2 (Daghestan central, Russie). Actuellement, elle est constituée de 2508 ossements déterminés, attribués à 19 espèces de mammifères. Le gisement résulte probablement d’une mort naturelle des animaux. Il est déterminé que toutes les espèces livrées par le site de Muhkai 2 caractérisent des paysages ouverts ou semi-ouverts. L’âge géologique estimé du site est compris entre 2,1 et 1, 77 Ma.
The Early and Middle Pleistocene were characterised by two major climatic events: the onset of the Quaternary glaciations and the Early-Middle Pleistocene Transition (EMPT) with glacial cycles changing their periodicity. The present study reviews recent research on the palaeoecological adaptations of Pleistocene ungulates from Mediterranean Europe following these global climatic changes, through an examination of dental wear patterns and hypsodonty indices of 27 fossil taxa from 4 key localities. Ungulates from Coste San Giacomo (2.1 Ma) adopted a wide range of feeding behaviours suggesting the presence of heterogeneous environments in the region. Following the gradual deterioration of climatic conditions and drier habitats ungulates display narrower diet ranges with no incidence of obligated browsers as attested at Olivola (~1.8 Ma). The diffusion of open habitats from the Early Pleistocene onwards is also reflected by the reduction of brachydont taxa. Discrepancy between dental mesowear and microwear patterns of Vallparadís Estació (~1.0–0.6 Ma) and Fontana Ranuccio (0.4 Ma) ungulates points to an increase in seasonality during the EMPT and after its consolidation. All examined taxa display dietary plasticity through the Early and Middle Pleistocene with even specialised groups, such as equids, showing a certain flexibility in their diet.
The sedimentary record of the Caspian Basin is an exceptional archive for the palaeoenvironmental, palaeoclimatic and biodiversity changes of continental Eurasia. During the Pliocene–Pleistocene, the Caspian Basin was mostly isolated but experienced large lake level fluctuations and short episodes of connection with the open ocean as well as the Black Sea Basin. A series of turnover events shaped a faunal record that forms the backbone of the Caspian geological time scale. The precise ages of these events are still highly debated, mostly due to the lack of well-dated sections. Here, we provide an integrated magneto-biostratigraphic age model for two sections from the Kura Basin – Goychay and Hajigabul. Our results reveal several major intervals with elevated salinity, associated with mesohaline faunas, and propose the following age constraints: 1) The Productive Series–Akchagylian boundary is marked by a marine transgression from the open ocean that occurred around the Gauss-Matuyama reversal (~2.7–2.6 Ma); 2) The Akchagylian–Apsheronian transition is characterized by a regression event and introduction of a new, "Pontocaspian" mollusc assemblage, dated within the Reunion sub-chron (~2.13 Ma). The ostracod assemblages of the two sections do not show a major faunal turnover here; 3) The early Bakunian transgression occurs after the upper Apsheronian lowstand (0.85–0.83 Ma). We conclude that major transitions in the age-indicative mollusc groups sometimes occur at different time intervals (i.e. diachronic) and are highly depended on the local depositional environments. A high-resolution interdisciplinary approach on sections outside the Kura Basin is required to better understand the potential diachroneity of these turnover events in the entire Caspian Basin.
The Vallparadís composite section (VCS) includes the nearby paleontological sites of Cal Guardiola and Vallparadís Estació (Vallès-Penedès Basin, northeastern Iberian Peninsula). The section spans from before the Jaramillo subchron to the early Middle Pleistocene (ca. 1.2-0.6 Ma). In this study, we describe the suid record from VCS and we review those from several other European sites, in order to refine the taxonomic identity and chronostratigraphic range of Quaternary suids in Europe. The VCS sample in- cludes a nearly complete skull, several teeth, and postcranial elements, and stands out as the richest European suid collection from the latest Early Pleistocene. Suid remains have been unearthed from both Cal Guardiola and Vallparadís Estació layers, whose age spans from the Jaramillo subchron (ca. 1.07-0.99 Ma; MIS31) to post-Jaramillo time (ca. 0.86 Ma; MIS21). Several craniomandibular and dental morphological features support an attribution to Sus strozzii. These features include a low and very deep preorbital fossa, a narrow nuchal crest, a well-developed longitudinal swelling in the middle of the mandibular corpus, the presence of styles/stylids in the upper/lower premolars, and especially the “verrucosic” morphology of the lower canine. The attribution to S. strozzii is also sustained by a cladistic analysis. These results provide interesting clues on the chronological occurrence of Quaternary suids. Sus strozzii is relatively common in Europe during the middle and early late Villafranchian (ca. 2.5-1.8 Ma), but it almost completely disappears during the latest Villafranchian (ca. 1.8-1.2 Ma). During and slightly after the Epivillafranchian (ca. 1.2-0.8 Ma), S. strozzii reappears in Europe although with relatively small samples, at VCS and several other sites including Untermassfeld (Germany; ca. 1.0 Ma), Le Vallonnet (France; ca. 1.2-1.1 Ma), Taman Peninsula (Russia; ca. 1.1-0.8 Ma), Arda River (Italy; ca. 0.99 Ma), and Slivia (Italy; ca. 0.8 Ma), among others. Consequently, in contrast to previous knowledge, we conclude that (1) S. strozzii survived in Europe (or returned there with a second dispersal event from Asia during the Epivillafranchian) at least until the end of the Early Pleistocene and (2) the arrival of Sus scrofa into that continent is not older than the Early-Middle Pleistocene boundary.
Well-defined stages of paleogeographical evolution with principally different natural conditions are characteristic of the long history of the Greater Caspian Sea. Nevertheless it is an indivisible period of evolution which began in the Akchagylian stage and continued during the Apsheronian and Pleistocene. A possible cause is location of the Caspian water bodies within the vast area in the centre of Eurasia where the epochs of warming and cooling and humidification and aridization were constantly alternating.
The skull and limb bones collected at Elandsfontein, Cape indicate that E. capensis was different from a Grevy's zebra. The body proportions were similar to those of an extant draft horse (E. caballus) and the skull resembled those of true Cape quaggas and a fossil Algerian plains zebra, E. mauritanicus.
Suids are found in Europe before and during the Olduvai magnetostratigraphic subchron, including the
Fonelas P-1 site in the Guadix Basin (Andalusia, Southern Spain, ~2.0 Ma), in which the remains have
been ascribed to Potamochoerus magnus, and many other localities that record the presence of Sus strozzii
(e.g., Saint Vallier and Senèze in France). However, there is no pig record in the biochronological range
comprised between the post Tasso Faunal Unit, which marks the base of the Late Villafranchian (~1.8 Ma),
and their arrival in Western Europe at layer TE9 from Sima del Elefante, Atapuerca, Northern Spain
(~1.2 Ma), where pigs are recorded under the name of Sus sp., and at the sites of Untermassfeld (Germany),
Vallonnet (France) and layer EVT12 of Vallparadís Estació (Spain), dated 1.1-1.0 Ma (MIS31),
which suid remains have been ascribed to Sus scrofa priscus in the first site and to Sus sp. in the other two.
Later, the genus Sus is recorded everywhere in Europe as a ubiquitous member of the Epivillafranchian/
Galerian and posterior faunas. When pigs are in an ecosystem, they use to be abundant in the large
mammal community given their opportunistic feeding behavior and high reproductive success. Their
short gestation periods and elevated offspring numbers allow them to colonize new and varied environments
and territories faster than other ungulates, which use to display a K-reproductive strategy, with
a single pup per birth. For this reason, suids are usually preserved in the fossil assemblages after their
dispersal and colonization of a geographic region. The arrival of suids phylogenetically related to Sus gr.
scrofa into Europe marks the end of the Late Villafranchian and the beginning of the Epivillafranchian,
which is approximately dated at ~1.2 Ma. Given that suids are omnivorous, generalist mammals with
bunodont teeth that do not tolerate very low temperatures, this suggests that their colonization of
Europe can be related to a change in the ecosystems and climate. In addition, the arrival of suids
postdates the earliest arrival of hominins into Western Europe, which is documented at the Orce sites of
Barranco León-D and Fuente Nueva-3 (~1.5-1.2 Ma). In these sites, rich faunal assemblages, abundant
lithic artifacts and one human tooth have been unearthed after more than twenty years of excavations,
but no evidence of pigs has yet been detected.
An extraordinary sequence of fossiliferous levels at the locality of Gran Dolina in the Atapuerca Hills (Burgos, Spain) records the Early–Middle Pleistocene transition. These levels are well dated by a variety of methods, including palaeomagnetism, which locates Lower–Middle Pleistocene boundary at the top of level TD7. Level TD6 is the type site of the species Homo antecessor and yielded over 90% of the European Early Pleistocene human record, while other levels have an archaeological record. The present paper deals with the earliest Middle Pleistocene ungulates of TD8, but we plan to describe the faunas, or at least the ungulates, of levels TD4 to TD8 in the context of the faunal changes that took place around the Early–Middle Pleistocene transition.
The Suidae are a family of Cetartiodactyla composed of 17 species classified in a minimum of five extant genera that originated at least 20 million years ago. Their success is evident in the multitude of habitats in which they are found as both natural and feral populations in tropical Island Southeast Asia, the high plateau of the Himalayas, Siberia, North Africa, the Pacific Islands, Australia, and the Americas. Morphological and molecular analyses of these species have revealed numerous aspects of their biology, including the ease with which many lineages have and continue to hybridize. This trait has made them an ideal model for evolutionary biologists. Suid species have also shared a deep history with humans, from their association with early hominids in Africa to their domestication. Here we review the current knowledge of this fascinating group and provide a comprehensive evolutionary history from the Oligocene to the present day. Expected final online publication date for the Annual Review of Animal Biosciences Volume 4 is February 15, 2016. Please see http://www.annualreviews.org/catalog/pubdates.aspx for revised estimates.
European large mammal assemblages in the 1.2–0.9 Ma timespan included Villafranchian taxa together with newcomers, mostly from Asia, persisting in the Middle Pleistocene. A number of biochronological schemes have been suggested to define these “transitional” faunas. The term Epivillafranchian, originally proposed by Bourdier in 1961 and reconsidered as a biochron by Kahlke in the 1990s, is at present widely introduced in the literature. This contribution, after selecting the most representative European large mammal assemblages within this chronological interval, provides a new definition proposal for the Epivillafranchian as a biochron included within the Praemegaceros verticornis FO/Bison menneri FO, and Crocuta crocuta FO.
Quaternary glacial/interglacial alternations, influenced by orbital obliquity cycles with a 41-ka long periodicity, started in the northern hemisphere around 2.6 Ma ago. Such alternations affected the terrestrial ecosystems, especially those of the Mediterranean region, with changes in the floristic communities and the dispersal and radiation of a number of large mammal open dwellers. Analyses of tooth wear patterns of ungulates from the Early Pleistocene site of Coste San Giacomo allow for a more objective reconstruction on the paleoenvironments and the climate in the Italian Peninsula during this epoch. Our results show that this area was composed by a mosaic of biomes, in particular by steppe and woodlands/wetlands. Evidence of such heterogeneity is provided by the wide spectrum of feeding behaviours found among the numerous ungulate herbivores here recorded, with cervids (Axis cf. lyra, Croizetoceros cf. ramosus and Eucladoceros sp.) exhibiting browser diets, most of the bovids (Gazella borbonica and Leptobos sp. and Gallogoral meneghinii) being intermediate feeders and the equid Equus stenonis showing a strict grazer behaviour. These results provide new insights for a timing of changing ecosystems in Southern Europe and reveal the environmental legacy of this global climatic shift, which is essential for understanding the early occupation of Homo in Europe. Thus, our data provide new evidence that such an environmental heterogeneity and a wide spectrum of available food resources could have been the main factors favouring the settlement of early species of Homo in this area.
New excavations at Saint-Vallier yielded a single last lower milk molar of Sus strozzii, a species new for the site. Sus strozzii is known from the Lower Villafranchian onwards, but mainly from the Middle and Upper Villafranchian of Italy, France, Spain, Great Britain, Netherland, Romania, Greece, Turkey and Israel-Palestine. That large boar is related to the present long-snouted wild pigs of East Asia like Sus verrucosus and Sus celebensis, whose classical biotope is dense forest under a wet and rather warm climate.
Equus stenonis has become a catch-all for stenonid (=zebroid) horses from the Late Pliocene to the Early Pleistocene, even Middle Pleistocene, of Eurasia. Non-significant variation between local samples referred to E. stenonis has resulted in a host of named ‘subspecies’, lacking morphometric and biological meaning; neither do they have fine-stratigraphic utility as commonly believed. Some finds identified as E. stenonis do not belong to that species. Because of its frequent use (or disuse) as a stratigraphic marker, a review of E. stenonis is necessary.
Etude biométrique concernant les espèces sauvagesactuelles (E. grevyi, E. burchelli boehmi, E. zebra hartmannae, E. africanus, E. hemionus et E. przewalskii) et quelques formes fossiles (E. stenonis vireti et cf. vireti du Villafranchien de France et d'Espagne; E. tabeti et E. mauritanicus du Pléistocène inférieur et moyen d'Afrique du Nord; E. mosbachensis du Pléistocène moyen d'Allemagne). Remarques sur les interprétations fonctionnelles et évolutives de certains caractères: quille et largeurs distales articulaires et sus-articulaires, proportions proximales.
The isolation of the Caspian Sea took place in the latest Miocene coinciding with a significant lowering of sea level and the deposition of a massive lowstand deltaic system. This so-called Productive Series is the main reservoir unit of the SouthCaspian oil-province. The Productive Series is overlain by marine clays from the Akchagylian and Apsheronian regional stages. During the Plio-Pleistocene, the Caspianbasin experienced several short periods of intermittent connectivity with other marine basins. This work aims to create integrated, high-resolution, bio-magnetostratigraphic dating of these regional transgressions in key sections of the SouthCaspianBasin in Azerbaijan. We sample two long sections, the Lokbatan section in the Palaeo-Volga delta and the Xocashen section in the Kura Basin. Palaeoenvironmental reconstructions are derived from characteristic ostracod species. Rock magnetic analyses combined with thermal demagnetisation data indicate that the magnetic signal is carried dominantly by the iron oxide magnetite in the Productive Series of Lokbatan section and in the Xocashen section. The marine Akchagylian and Apsheronian of Lokbatan are characterised by the iron sulphide greigite, which appears to be of (near-) primary origin. The most logical correlation of the magnetic polarity patterns to the Geomagnetic Polarity Time Scale dates the Akchagylian transgression at ~ 3.2 Ma, a major transgression during the Apsheronian at ~ 2.0 Ma and the Bakunian transgression at 0.85–0.89 Ma. Ostracod assemblages indicate increasing salinities during these transgressions, from fresh water lacustrine to brackish-marine species. This implies that marine connections have been created with an adjacent basin that has a higher salinity, most likely the Black Sea
Sus strozzii is a large boar known from ten late early Villafranchian to early Middle Pleistocene European and Middle Eastern sites. During the Pleistocene, it was replaced by several subspecies of Sus scrofa. Associated faunas were studied using Fleming's ecological diagrams in the Andrews et al. version, slightly modified by us. Recent boar ecology and the osteological and odontological characters of recent and fossil forms and their functional implications allow a reconstruction of the biotopes of the fossil Sus strozzii and the fossil Sus scrofa: •more or less open forests are dominant, nearly always interrupted by grassy areas;•humidity is usual but much more so in the case of Sus strozzii, some modern Sus scrofa live in semi-arid conditions;•temperature may be quite different.RésuméSus strozzii est un grand sanglier connu en Europe et au Proche-Orient dans dix gisements de la fin du Villafranchien inférieur jusqu'au début du Pléistocène moyen. Il est remplacé par Sus scrofa représenté au Quaternaire par plusieurs sous-espèces.Nous avons étudié les associations de faune qui les accompagnent, utilisant en particulier les diagrammes écologiques de Fleming revus par Andrews et al., dont certains sous une forme légèrement modifiée. En tenant compte de ce qu'on connaît de l'écologie des sangliers actuels, des caractères ostéologiques et odontologiques des formes actuelles et fossiles et de ce qu'ils impliquent sur le plan de l'anatomie fonctionnelle, nous en déduisons que l'environnement de Sus strozzii et des formes fossiles de Sus scrofa se caractèrise par: •la dominance de la forêt, qui peut toutefois être plus ou moins dense, plus ou moins ouverte, et presque toujours entrecoupée d'étendues herbeuses;•une note humide, plus marquée pour Sus strozzii mieux adapté au terrain mou que Sus scrofa dont certaines formes actuelles s'accommodent bien de conditions de vie en milieu semi-aride;•une indifférence marquée à la température.
Palynology of the upper pliocene to middle pleistocene deposits in the southern part of the Caspian Region. Moscow: GEOS; p
- N Y Filippova