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Early Pleistocene Feliformia from Palan-Tyukan (Azerbaijan)
Historical Biology
Abstract
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 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.
Recent field surveys performed in the newly-discovered Taurida Cave (Crimea, Russia, Late Villafranchian, 1.8–1.5 Ma) enabled us to unearth dentognathic remains of the dirk-toothed cat Megantereon. Here we describe in detail the recovered remains further comparing it anatomically and biometrically with coeval Eurasian and African sites. The performed analysis suggests the inclusion of the Taurida remains in the hypodigm of the Villafranchain European species Megantereon adroveri, a medium-sized saber-toothed cat poorly known in Europe as a consequence of the scarcity and fragmentary nature of the known remains. Our results and the putative transitional characters displayed by the Taurida specimen support the idea put forward by previous authors of a continuous European lineage of dirk-toothed cats.
This paper describes the maxillaries of adult specimens of saber-toothed cats Homotherium crenatidens (Fabrini, 1890) and Megantereon sp. from the Taurida Cave (Crimea, Late Villafranchian; age, 1.8–1.5 Ma). We discuss the morphometric variability in the morphology of the upper carnassials and the length of the С1–Р3 diastema.
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.
At present, numerous sites from Europe and Asia have yielded up remains of Panthera gombaszogensis. It is without doubt the one Felid most similar to Panthera onca but it is an ancestor, not the present day jaguar. Palaeogenetics estimates the divergence between the lion and the jaguar to 2 Myrs. Obviously, jaguars originate from Africa and they spread in Europe between 1.95 et 1.77 Myrs during the time of Olduvai polarity subchron. Recently Hemmer et al. (2010) demonstrated that the semi-mandible of Dmanissi (Georgia) corresponds with a new taxon Panthera onca georgica ssp. nov., the earliest known Asian member of the species. This taxon spread out through Asia and got to North America during a glacial period between 0.99 and 0.78 Myrs (between the Jaramillo polarity subchron and the end of the Matuyama magnetochron), and finally reached South America not before the Rancholabrean. The Château Breccia (Saône-et-Loire, France) has given an abundant paleontological material of Panthera gombaszogensis (about 400 remains, with a minimum of 6 individuals). It is the mid Middle Pleistocene European form. The pollen analysis indicates temperate climatic conditions but cooler than today and sometimes even quite cold. The landscape was open with grasses spread over the uplands, while woodlands of pine and fir, but also some broadleaved trees were found in the valley. The Château Breccia largely contributes to the story of Panthera gombaszogensis. It demonstrates that the species was at least adapted to a cold temperate climate and to a biotope highly different from that of the today's jaguar. It shows also the replacement of Panthera gombaszogensis by Panthera spelaea fossilis about 0.6 Myrs ago, after a period of coexistence of these two Felids.
A left lower molar of the European jaguar, Panthera onca gombaszoegensis (Kretzoi, 1938), collected ex situ at the locality 'De Kuilen' near Langenboom (province of Noord-Brabant, The Netherlands), is described and illustrated. This specimen is compared to Early Pleis-tocene material from Untermassfeld (Thüringen, eastern Germany) and from Tegelen (province of Limburg, The Netherlands). The three other records of P. onca gombaszoegensis from The Netherlands are briefly reviewed. P. onca gombaszoegensis is a fauna element of the Late Pliocene and Early Pleistocene in The Netherlands.
This article reviews the faunal composition, biochronology and succession of Late Pliocene through early Pleistocene mammalian localities and faunas of Greece. Some important faunal events are identified that permit stratigraphic subdivision within this great time span. The main faunal changes occur around the Pliocene-Pleistocene temporal boundary and at the end of Villafranchian s.l.
Ten new track-bearing boulders discovered in the Parco Naturale delle Dolomiti Friulane area (Carnic Prealps, NE Italy) during the last 15 years demonstrate that bipedal dinosaurs with a functionally tridactyl, digitigrade pes, within the size-class of the ichnogenus Anchisauripus Lull, 1904, were the most common trackmakers in the carbonate tidal flats of the Dolomia Principale during the latest Triassic. If the footprint morphology actually reflects the functional tridactyl condition of the trackmakers’ feet, theropods are the most suitable trackmakers. The ichnoassociation of the Parco delle Dolomiti Friulane area differs from coeval ichnoassociations in the dominance of mid-sized tridactyl footprints and the rarity of small tridactyl (Grallator Hitchcock, 1858; length <15 cm) and chirotheriid tracks. Tracks potentially produced by basal sauropodomorphs, which were the most common European dinosaurs in the late Norian-Rhaetian interval, are underrepresented.
The Villafranchian faunas of Greece were limited known for a long time, but during the last three decades several new fossiliferous sites have been discovered and new material has been unearthed. The carnivores constitute a remarkable part of this material and their study gave useful information. The Villafranchian carnivoran guild includes numerous taxa belonging to various families. Some of them are interesting as they are differentiated from the other Eurasian ones. They cover the entire Villafranchian but the mass is known from the middle- to epi- Villafranchian. Despite the discontinuous data they provided useful biochronological results. The coexistence of Pliocrocuta perrieri and Pachycrocuta brevirostris in the locality of Gerakarou 1 is a strong evidence for the middle/late Villafranchian boundary. The composition of the Villafranchian carnivores suggests that the canids predominate, while the felids, hyaenids and mustelids are well represented; the ursids are represented by a single taxon the well known Ursus etruscus, which seems to be common during the entire Villafranchian. The carnivoran guild structure and the multivariate analysis of the Greek Villafranchian carnivoran assemblages (middle, late and epi Villafranchian) in comparison with modern assembalges from open and closed environments indicate an open landscape, being in agreement with previously results for the Villafranchian of Greece. During Epi-Villafranchian there is a faunal renewal which is related to more open and dry environment like a savannah bushland/grassland. This article is protected by copyright. All rights reserved
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.
Carnivores of the Khapry faunal assemblage that is commonly considered as an analogue of the middle Villafranchian West European fauna (Late Pliocene, Zone MN17) were revised. Members of nine Carnivora
genera are described in detail, and five of them are first included in the list of assemblage taxa. Carnivores
of the Khapry assemblage are represented by Nyctereutes megamastoides (Pomel), Canis cf. senezensis
Martin, Lutra sp., Pannonictis nestii (Martelli), Pliocrocuta perrieri (Croizet et Jobert), Pachycrocuta brevirostris
(Aumard), Homotherium crenatidens (Fabrini), Acinonyx pardinensis (Croizet et Jobert), and Lynx issiodorensis (Croizet et Jobert). The Khapry carnivore assemblage includes taxa characteristic of the Late Pliocene Eurasian faunas. Its lower stratigraphic limit corresponds to the beginning of the middle Villafranchian, as it follows from presence of Homotherium crenatidens with more advanced dental characteristics than those of homotheres from the lower Villafranchian of Eurasia. The occurrence of
Nyctereutes megamastoides that was not encountered above the middle Villafranchian bounds the upper stratigraphic limit of the Khapry Carnivora. The “Canis -event” and “ Pachycrocuta -event” are considered to be important records around the middle-upper Villafranchian boundary in Europe. Presence of Canis senezensis is characteristic of the final stage of the middle Villafranchian in Western Europe, and dispersal of Pachycrocuta brevirostris is typical of the lowermost late Villafranchian, whereas the wolflike forms of
Canis and Pachycrocuta appeared somewhat earlier in Asia. The Khapry faunal assemblage as a whole is characterized by a certain similarity with Asian analogues of the middle Villafranchian faunas.
Tool-using hominids, as carnivorous animals, would have been part of the various carnivore guilds present in Plio-Pleistocene Africa. Hominid dietary strategies must be understood within the larger context of carnivore behavior and ecology, as carnivorans could have affected the abilities of hominids to procure meat and/or marrow. The functional anatomy of extant and fossil carnivorans was examined to infer behaviors in fossil carnivorans that would have impacted on hominid dietary strategies in terms of carcass availability. Comparisons of guild structure were carried out to examine changes in carnivoran interactions and their implications for hominid behavior. Plio-Pleistocene carnivorans engaged in a wider range of behaviors than modern carnivorans. The sabertoothed felids Dinofelis and Megantereon probably did not provide much larger carcasses than modern species. Another sabertooth, Homotherium generated larger carcasses, but may have disarticulated and transported these carcasses. Fossil representatives of modern taxa may not have been equivalent ecologically within the carnivoran guild. Overall, hominids in eastern Africa probably had a greater range of scavenging opportunities than did those of southern Africa during the Plio-Pleistocene. Local and continent-wide extinction events in large-bodied carnivoran guilds from 1 to 2 Ma had a substantial effect on carcass availability and the risk to hominid scavengers. These structural changes in the carnivore guild may have provided an opportunity for hominids to widen their niche with respect to dietary behavior.
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.
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.
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.
Felid material from the middle Early Pleistocene Haro River quarry, northern Pakistan, is described. Most of the material is referred to the Eurasian jaguar, Panthera gombaszogensis georgica , representing the first record of the jaguar in southern Asia. A systematic review of dental morphology of all living species of Pantherinae is made to provide a better background for studying the evolution of pantherine cats. A renewed dispersal scenario of the jaguar in Eurasia is provided. Panthera gombaszogensis georgica is suggested to be a distinct subspecies distributed in central, western and southern Asia during the Early Pleistocene, characterized by relatively small body size and slender dentition. The Eurasian jaguar does not possess the derived dental characters of the living jaguar Panthera onca (e.g. weak or no canine vertical groove, robust premolars), and presumably did not specialize on hard‐shelled prey, and therefore should be viewed as a distinct species from the living P. onca . Two additional felids from the Haro River Quarry, the large sabre‐toothed felid? Megantereon sp. and a small undetermined feline, are also reported. The Felidae faunal composition of southern Asia during the Early Pleistocene is generally similar to that of northern Eurasia, and the paleoenvironment is probably slightly more closed.
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 homotheres, although known from the Greek fossil record since the beginning of the 1930’s, have only been recorded from fragmentary material, sometimes undescribed and with questionable determinations. In 2017, the discovery of an almost complete cranium and a fairly complete humerus of a machairodontine in Dafnero-3, a lower Pleistocene site from Western Macedonia (Greece), confirm the presence of homotheres in Greece. The Dafnero-3 material is described and compared with Eurasian specimens. We conclude that in morphology and size the Dafnero-3 specimen coincides with Homotherium latidens (Owen, 1846). The taxonomy, geographic and stratigraphic distribution of Homotherium is also discussed. A biostratigraphic revision gives the distribution of the genus in Greece. Homotherium is referred from six Greek localities and its distribution covers the time span from the early Villafranchian (early Villafranchian, locality Milia, Western Macedonia; MN16, 3.0-2.5 Ma) to the middle Pleistocene (locality Tourkovounia near Athens, ~0.4 Ma).
In the fossil record hyaenids are both diverse and abundant, and nearly 100 species have been named. Hyaenids have an expanded ectotympanic and semi-recumbent to recumbent septum bullae, and all fossil taxa likely to display this morphology are reviewed herein. The systematics and cladistics of the group are described. The age of the Hyaenidae is suggested to be some 25 million years. This is compatible with biochemical dates. Macroevolutionary patterns in the Hyaenidae are examined on the basis of the established cladogram and are found to be overwhelmingly gradual. Taxic patterns show that the Hyaenidae were most diverse in the late Miocene. -from Authors
The Zanda Basin in the western Tibet Autonomous Region, China, produces fossils of Miocene–Pleistocene age. Proximity of the basin to the Himalaya Range makes Zanda an important region for understanding vertebrate evolution and dispersal in and around the Tibetan Plateau. Five field seasons of prospecting in the basin have resulted in a rich collection of fossil mammals, with highest abundance and taxonomic richness in the Pliocene section of the Zanda Formation. In this report, we describe the first Tibetan Plateau record of the bone-cracking hyaenine carnivoran Pliocrocuta perrieri from early Pliocene (4.1 Ma) deposits in Zanda. The Zanda Pliocrocuta specimen is the smallest known Pliocrocuta in China. In both morphology and size, it is similar to other early occurrences of Pliocrocuta in Spain and northern China. Materials from Zanda and Yushe represent the earliest and least robust Asian specimens of a highly variable Pliocene hyaenid species, and indicate that Pliocrocuta were already widespread across diverse environments at their first appearance. The presence of two fossil hyaenid species along with at least three canids and one felid in the Plio-Pleistocene Zanda fauna indicates a less deteriorated large predator guild in the Himalayan foothills compared to extant ecological communities in western Tibet.
African machairodont specimens previously referred to three species of Megantereon are considered to represent a single species in turn argued to be conspecific with the Eurasian species Megantereon cultridens (Cuvier). The area of origin of Megantereon remains unclear, but doubt is expressed about claims for an earliest appearance of the genus in North America. It is probable that the North American species M. hesperus is a junior synonym of M. cultridens. - Author
In this beautifully illustrated natural history that links extinct larger feline species with those still in existence, collaborators Alan Turner and Mauricio Anton weave together the evidence of modern feline behavior with that of the fossil record. Turner's clear, insightful prose and Anton's masterly illustrations combine to offer specialists and newcomers alike an accurate and accessible guide to the evolution of cats.
In addition to a rich, contemporaneous vertebrate faunal record, the fossil locality of Biśnik Cave in southern Poland, has yielded some Eurasian jaguar elements from layers 19b–d. Through placing the faunal assemblage from the examined layers at around MIS 9 or MIS 10, the corresponding jaguar remains appear to be the among the stratigraphically youngest indication of Panthera gombaszoegensis in Europe. Detail analysis of size variability among Early and Middle Pleistocene individuals from varies sites showed no increase in size during Eurasian jaguar evolution. Jaguar size cannot be use as a biochronological tool, and should be linked with geographical and individual variability as well as with sexual dimorphism.
The assemblage from Kanapoi represents the most diverse early Pliocene carnivore assemblage from sub-Saharan Africa. Carnivora from Kanapoi were originally described by Werdelin (2003a), but continuing field work has brought to light significant new material from the site, shedding new light on the earliest post-Miocene radiation of Carnivora in Africa. Most importantly, a second species of Enhydriodon has been recovered from the site, including the first specimen to include a large part of the neurocranium. This makes Kanapoi the first site to include two species of this genus. This addition to the fauna will be of prime significance to understanding the ecology and evolutionary radiation of these giant, extinct otters. Other significant new finds include additional material of a wildcat-sized felid. Finds of this group are rare, and the new Kanapoi material adds significantly to our knowledge of the stem lineage of the genus Felis, which is widespread in Africa today.
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 Pachycrocuta Kretzoi, 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.
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
We review the larger pattern of appearance of the Hyaenidae in Europe and outline their part in the turnover of the guild of larger Carnivora that occurs across the Miocene–Pliocene boundary. The earliest record of the family is in MN4, although the patchy nature of the earliest records makes it difficult to be certain about the continent of origin. There is a clear pattern of morphological evolution over that long timespan, from the earliest viverrid-and herpestid-like forms through dog-like and more cursorial taxa to the larger, bone-crunching animals of the later Miocene and the Pliocene–Pleistocene epochs. Miocene dog-like hyaenas may indicate that social hunting had emerged by that time, while the appearance of larger species means that hyaena-accumulated bone assemblages may potentially occur in any late Miocene to Pleistocene locality. # 2008 Elsevier Masson SAS. All rights reserved. Résumé Nous révisons le modèle d'apparition des Hyaenidae en Europe et nous soulignons leur rôle dans le renouvellement des communautés de grands carnivores au cours du Miocène et du Pliocène. Le premier enregistrement de cette famille est situé dans la MN4, même si la rareté des premiers enregistrements ne permet pas d'établir avec certitude leur continent d'origine. Il existe un modèle clair d'évolution morphologique pendant cette période, des premières formes apparentées aux viverridés et aux herpestidés en passant par des formes de type dog-like et des taxons plus cursoriaux, jusqu'aux grands animaux broyeurs d'os du Miocène terminal et du Plio-Pléistocène. Les hyènes miocènes qualifiées de dog-like peuvent indiquer que la chasse sociale, en groupe, ait émergé à cette époque, alors que l'apparition des espèces de grande taille signifie que les hyènes accumulatrices de vestiges osseux, ont pu potentiellement exister dans les localités du Miocène terminal au Pliocène.
Comparative evaluation of fossil remains of the lower dentition of the jaguar, Panthera onca (Linnaeus, 1758) in the light of recent DNA assessment allows a comprehensive phylogeographic interpretation. The speciation process led a jaguar stem population, clearly of African origin, to disperse over Europe during the time of the Olduvai polarity subchron (1.95-1.77 Myr) (Panthera onca toscana). Based on a hemimandible from Dmanisi, Republic of Georgia, dated to about 1.77 Myr, a new taxon Panthera onca georgica ssp. nov. is proposed for the earliest known Asian member of the species. Its generalized, more cutting dentition mediates between the contemporaneous P. onca toscana and the two later sister subspecies, the Eurasian P. onca gombaszoegensis, which is characterized by a specialized cutting and crushing dentition, and the North American P. onca augusta. Placing the tooth differences within geographic coordinates indicates a central Asian evolutionary node between the latter two forms. Transcontinental dispersal probably brought the jaguar to North America during a glacial period between the Jaramillo polarity subchron and the end of the Matuyama magnetochron (0.99-0.78 Myr), to finally reach South America not before the Rancholabrean. Divergent taxonomic concepts for Pleistocene jaguars are discussed: single species P. onca with several subspecies, two species P. gombaszoegensis and P. onca, or three species P. toscana, P. gombaszoegensis and P. onca.
Megantereon cultridens was a derived, Late Pliocene to Early Pleistocene sabrecat, and although fossils of animals referred to the genus and species have been found at several locations throughout Eurasia, most are fragmentary. However, the specimen SE311 from Senéze in France represents an almost complete and well-preserved skeleton, and this is only known from very few other sabrecats, thus providing a rare glimpse into the full anatomy of an unusual and derived sabrecat. In this monograph, we provide a complete overview of the anatomy of Megantereon cultridens SE311, and compare it with extant large felids, and the few other derived sabrecats from which ample fossil material is known, although most frequently representing several specimens. SE311 was a large specimen of M. cultridens and would have had a body mass of 100–110 kg and a head–body length or around 160 cm, which is similar to a small lioness or large male jaguar. Megantereon sp. were sexually dimorphic, and the size of SE311 suggests that it was a male. As with several other derived sabrecats, it was powerfully built, and had particularly robust forequarters with very well-developed muscle attachment sites, indicating a powerful forelimb and shoulder musculature. The neck was proportionally much longer than in extant felids, and the thoracic and particularly lumbar region was proportionally shorter, mimicking the condition in other derived sabrecats from which large parts of the vertebral column is known. Megantereon probably lived in open-forest environments and preyed on cervids, which were dispatched with a shearing bite from the hypertrophied and blade-like upper canines to the throat of the prey, while the prey was held immobile with the massive forelimbs, thus minimizing the risk of damage to the fangs. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151, 833–884.
The Plio-Pleistocene site of Tegelen in the Netherlands contains some of the oldest evidence for the presence of the medium-sized pantherine felid of the European Pleistocene, Panthera gombaszoegensis, based on published fragmentary dental specimens. Measurements of these specimens, together with those from other localities, suggest that earlier representatives of the species are small, but one of a small number of previously unstudied specimens from the site now indicates the presence of a larger pantherine individual. Such a size difference could point to the presence of a second species, or suggest that the Tegelen deposits encompass a longer time span than has been supposed. Either of these would have serious implications; the first would increase the complexity of the early Pleistocene carnivore guild, and the second would threaten the status of Tegelen as a type site in the European Pleistocene biostratigraphic scheme. However, consideration of the size range in sexed samples of extant pantherines in conjunction with a re-examination of the size distributions in the hypodigm of P. gombaszoegensis suggests that sexual dimorphism offers a more plausible interpretation.
Human occupants of Europe shared food resources with a number of larger Carnivora, and their coexistence with two lion-sized felids, the lion and the scimitar-toothed machairodont Homotherium latidens, poses intriguing paleoecological problems. We investigate the ecology of Homotherium latidens using an exceptional sample of postcrania from the Spanish Early Pleistocene site of Incarcal, making comparisons with modern cats and with other machairodont species. Evidence of cursorial adaptations in Homotherium suggests a hunting technique different from modern cats or smilodontine sabre-tooths. Some, like reduction of the claws, would have limited the ability of individual homotheres to bring down large prey, implying group action. Homotherium would also have been disadvantaged in direct confrontation with Pleistocene lions by smaller body mass, reduced forepaw muscle strength, smaller claws and more fragile dentition. Its hunting technique would have worked best in more open habitats, but competition from lions would have forced it to seek moderate cover. Among factors that could de-stabilise coexistence of the two big cat species in Pleistocene Europe we invoke a decrease in environmental mosaicism associated with stepped climatic change over the last million years, and the increased importance of humans within the larger predator guild.
Large carnivores structure the character of scavenging opportunities in any environment. In Pliocene and early Pleistocene Africa there were three large sabertooth cats sympatric with the ancestors of the modern felid community, and scavenging opportunities were presumably different from those in modern Africa. An understanding of a possible scavenging niche for early hominids must articulate knowledge gained from actualistic research with detailed reconstructions of extinct carnivore paleoecology.
Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historiche Geologie
- D S Kostopoulos
- S Sen
Geological position of the remains of terrestrial mammals of Khapry, Taman’ and Tiraspol faunistical complexes in the cut of marine layers of Akchagyl and Apsheron of eastern Transcaucasia
- N A Lebedeva
Transactions of the institute for the history of material culture RAS
- M V Sablin
Zur Charakterisierung und stratigraphischen Bedeutung von Panthera gombaszogensis (Kretzoi, 1938)
- H Hemmer
Correlation between anthropogenic strata of the Ponto-Caspian
- N A Lebedeva
Machairondontinae [Megantereon cultridens roderici n. ssp. and Homotherium latidens (Owen, 1846)] at the Fonelas p-1 site (Guadix basin, Granada)
- A Arribas
- G Garrido
Die Feliden aus dem Epivillafranchium von Untermaßfeld bei Meiningen (Thüringen), Teil 3. Monographien des Römisch-Germanischen Zentralmuseums Mainz
- H Hemmer
A re-evaluation of the diversity of Megantereon (Mammalia, Carnivora, Machairondontinae) and the problem of species identification in extinct carnivores
- P Palmqvist
- V Torregrosa
- J A Pérez-Claros
- B Martínez-Navarro
- A Turner
Remains of carnivores and ungulates from the Lower Absheron deposits of Azerbaijan
- M V Sablin
On the new discovery fossil remains of the Equus stenonis Cocchi in the Caucasus
- I E Kuzmina
- M V Sablin
New cranial remains of Pliocrocuta perrieri (Carnivora, Hyaenidae) from the Villafranchian of the Iberian Peninsula. Bollettino della Società Paleontologica Italiana
- V Vinuesa
- J Madurell-Malapeira
- M Ansón
- D M Alba