Early Pleistocene Locality Research Articles

Target Deformation of the Equus stenonis Holotype Skull: A Virtual Reconstruction

Equus stenonis is one of the most prevalent European Pleistocene fossil horses. It is believed to be the possible ancestor of all Old World Early Pleistocene Equus, extant zebras and asses, and as such provides insights into Equus evolution and its biogeography and paleoecology. The Equus stenonis holotype skull (IGF560) was first described by Igino Cocchi in 1867, from the Early Pleistocene locality of Terranuova (Upper Valdarno basin, Italy). IGF560 is a nearly complete, although medio-laterally crushed and badly compressed skull. Here we provide the first application of a new virtual reconstruction protocol, termed Target Deformation, to the Equus stenonis holotype. The protocol extends beyond classic retrodeformation by using target specimens as a guide for the virtual reconstruction. The targets used as a reference are two fragmentary, yet well-preserved E. stenonis skulls, coming from Olivola (Italy; IGF11023) and Dmanisi (Georgia; Dm 5/154.3/4.A4.5), both Early Pleistocene in age. These two specimens do not display any major deformation, but preserve different, only slightly overlapping portions of the skull. The virtual reconstruction protocol we carried out has shown its feasibility, by producing two 3D models whose final morphology is perfectly congruent with the natural variability of a comparative sample of E. stenonis specimens. This study shows the potential of using even broken or otherwise fragmentary specimens to guide retrodeformation in badly distorted and damaged specimens. The application of Target Deformation will allow us to increase the availability of comparative specimens in studies of fossil species morphology and evolution, as well as to the study of taphonomic processes.

First report of Sus strozzii (Suidae, Mammalia) from the Early Pleistocene of Hungary (Dunaalmás) and species distinction based on deciduous teeth.

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.

Northern Asian Pliocene–Pleistocene beremendiin shrews (Mammalia, Lipotyphla, Soricidae): a description of material from Russia (Siberia), Kazakhstan, and Mongolia and the paleobiology of Beremendia

AbstractBeremendiini is an extinct group of soricine shrews that were widely distributed during the Pliocene and Pleistocene. Their occurrence in China has been investigated, but their presence in northern Asian regions has remained poorly studied. This paper analyzes 56 fossil remains of Beremendiini collected from 16 early Pliocene to early Pleistocene localities in Russia (Siberia), Kazakhstan, and Mongolia and shows the presence of two beremendiin species:Beremendia fissidens(Petényi, 1864) andBeremendia minorRzebik-Kowalska, 1976. Northern AsianBeremendiaconsiderably vary in size and qualitative characteristics, although most of the different states have been identified in European or Chinese specimens. Through the application of geometric morphometric techniques, mandibular shape analyses reveal similarities between the members of the beremendiin generaPeisorexKowalski and Li, 1963,BeremendiaKormos, 1934, andLunanosorexJin and Kawamura, 1996. Shape analyses and comparisons of mandibular characteristics reveal ‘trophic’ analogies betweenBeremendiaandBlarinaspp. and a new model of ‘Mandible Swinging and Sliding’ (MSS-model) accounting for the similarities in mandibular morphology with implications for the understanding of the diet ofBeremendia.UUID:http://zoobank.org/ef92ee1a-3977-44b2-86d7-381665bcf78b

Establishing a neotype for Crocidura obtusa Kretzoi, 1938 (Mammalia, Soricidae): an emended description of this Pleistocene white-toothed shrew species

We establish a neotype in the collection of the Hungarian Natural History Museum, Budapest (Hungary) for Crocidura obtusa (Mammalia, Soricidae). The species was originally described by M. Kretzoi as a new species from the Early Pleistocene locality of Gombaszög (now Gombasek, Slovakia) in 1938, but the holotype was lost in 1956. The neotype is a complete left mandible from the Osztramos 8 site (Hungary, Early Pleistocene). Due to the incomplete original description given by Kretzoi, a new definition of this species also had to be composed. Kretzoi’s distinctive characteristics between C. obtusa and the recent Crocidura species are accepted here, but further differences were discovered in comparison with the contemporary C. kornfeldi. According to our taxonomic results, C. obtusa was present in Central Europe, mainly in the Carpathian Basin, from the Early Pleistocene (ca. 1.2 Ma) to the earliest Late Pleistocene (ca. 130–115 ka).

Fossil mega- and micro-flora from Bernasso (Early Pleistocene, southern France): A multimethod comparative approach for paleoclimatic reconstruction

Bernasso is an Early Pleistocene locality that has a well-dated floral fossil content including both pollen grains and megaremains. Pollen record shows vegetation changes corresponding to two glacial and one interglacial periods. The megafloral fossil assemblage was interpreted as an association of present-day Mediterranean and Caucasian taxa that developed during a late interglacial stage of the beginning of the Pleistocene. In the present study, climatic reconstruction using the Climate Leaf Analysis Multivariate Program (CLAMP) and the Leaf Margin Analysis (LMA) methods based on the megaflora on one hand and using the Climatic Amplitude Method (CAM) based on the pollen flora on the other hand have been used to check the qualitative climatic estimates based on the comparison between paleovegetation data and similar modern ecosystems. CLAMP and LMA analyses show that the Bernasso climate was colder than the present-day climate of Southern France and of the Hyrcanian and Euxinian regions. On contrary, CAM shows mean annual temperatures higher than the modern Bernasso temperature and equivalent to the temperature of the Hyrcanian and Euxinian regions. Mean annual precipitation and relative humidity reconstructed by CLAMP and CAM are in agreement and indicate that the climate was wetter than today at Bernasso. The differences between the three methods highlight important methodological biases that could be solved by including additional calibration sites in Europe and in the Mediterranean region.

AHystrixLinnaeus, 1758 incisor (Mammalia, Rodentia) from the Early Pleistocene of Senèze, France

ABSTRACT Excavations carried out between 2001 and 2006 in the Early Pleistocene locality Seneze (Haute-Loire, France) yielded the fragment of a large upper incisor with anteriorly grooved enamel band, documenting a large-bodied rodent in Seneze for the first time. A thorough study of this incisor and particularly of its enamel microstructure led to its assignment to Hystrix refossa Gervais, 1852, despite the presence of longitudinal grooves which are usually considered as characteristic of the castorid genus Trogontherium Fischer von Waldheim, 1809 and which have not been noticed in Hystrix Linnaeus, 1758 so far. The presence of H. refossa is in accordance with the partly steppic environment previously documented in the Seneze locality.

A fourth mandible and associated dental remains of Gigantopithecus blacki from the Early Pleistocene Yanliang Cave, Fusui, Guangxi, South China

Dentognathic remains of Gigantopithecus blacki from the newly discovered Early Pleistocene locality of Yanliang Cave, Guangxi, South China are described. These include an incomplete mandible, only the fourth discovered and the first known from a site other than Liucheng, as well as 25 isolated teeth. Comparisons of the Yanliang mandible show that the best preserved part of the right corpus is morphologically similar to the left side of the Liucheng Mandible III. In addition, the Yanliang mandible and the Liucheng Mandible III share a similar degree and pattern of wear on the premolars and molars. The partially resorbed alveolus for the right M2 in the Yanliang mandible indicates antemortem tooth loss, which is the first record of its kind for Gigantopithecus blacki. Comparisons of the enamel–dentine junction morphology show that the isolated upper premolars from Yanliang are similar to those of Gigantopithecus blacki from Early Pleistocene sites, and differ from the more specialised form from the Middle Pleistocene Hejiang Cave. This supports the biochronological evidence that Yanliang Cave is Early Pleistocene in age.

Comparing two different Early Pleistocene microfaunal sequences from the caves of Atapuerca, Sima del Elefante and Gran Dolina (Spain): Biochronological implications and significance of the Jaramillo subchron

No ideal method can be used alone to provide an accurate and irrefutable chronology for dating European Early Pleistocene localities. The solution lies in the combination of different, but complementary, approaches for complementing their chronological framework. In that regard, the results of the present study show that biochronology may be a useful additional tool to further refine the chronology of these localities, by providing not only relative chronological information between the sites, but also giving some important indications about a younger or older Jaramillo age.Our study presents the revised faunal list of each Early Pleistocene levels from the localities of Gran Dolina (TD) and Sima del Elefante (TE), in Atapuerca. Both localities have very likely the richest associations of small mammal species in the Quaternary of Europe. Perhaps the most striking observation is the lack of similarities between the assemblages from each site, which can only be explained by a chronological gap between them. Although the Jaramillo subchron has not been directly identified at Atapuerca, the existing chronostratigraphical framework combined with similarities of the faunal assemblages between Gran Dolina and Vallparadís indicate with some confidence that Sima del Elefante levels are older than the Jaramillo subchron. Consequently, the Early Pleistocene levels of both Atapuerca sites TE and TD are the only sequence from Spain that allows the study of a continuous stratigraphic succession of the interval around the Jaramillo subchron in terrestrial sequences. The biostratigraphy of the Early Pleistocene from Europe, based mainly in the few isolated localities of the Olduvai–Jaramillo time interval, is reinforced by the faunal succession of the Atapuerca sites.

The insectivores (Soricidae, Erinaceidae; Eulipotyphla; Mammalia) from Cueva Victoria (Early Pleistocene, Murcia, Spain)

In the Early Pleistocene locality of Cueva Victoria, the insectivores are represented by several fossil teeth, mandibles and skull fragments. In the present work, we describe and provide measurements of these fossils for the first time. Insectivores show a rather low taxonomic diversity in Cueva Victoria, with only two different species. Considering the environmental requirements for their extant relatives, the presence of only one hedgehog (Erinaceus cf. praeglacialis Brunner, 1933), and one shrew (Crocidura kornfeldi Kormos, 1934) points to the existence of dry and warm conditions in the surrounding area of Cueva Victoria, unfavourable to other contemporary species of insectivores.

Resource partitioning and niche separation between mammoths (Mammuthus rumanus and Mammuthus meridionalis) and gomphotheres (Anancus arvernensis) in the Early Pleistocene of Europe

This paper presents the results from a tooth microwear analysis of three proboscideans (Mammuthus rumanus, Mammuthus meridionalis, and Anancus arvernensis) from four Early Pleistocene localities in Europe (Red Crag, Norwich Crag, Chilhac, and Eastern Scheldt). The particularity of these four localities is that mammoths and gomphotheres co-occur. The quantitative microwear data (numbers of pits and scratches) were informative about the broad feeding traits in each species. For both genera studied, diets range from strict browsing (including leaf and fruit browsing) to grass-dominated mixed feeding. These data reveal highly variable dietary traits in the mammoths and gomphotheres studied, but the qualitative results provide evidence of differences between the two sympatric genera. We were able to identify the consumption of fruits, seeds, bark and twigs in A. arvernensis, and the ingestion of high quantities of grit in M. rumanus and M. meridionalis which is most likely related to open or semi-open habitats. The data also support the hypothesis that resource partitioning existed between mammoths and gomphotheres when they co-occurred at a locality.

New 400–320 ka Gigantopithecus blacki remains from Hejiang Cave, Chongzuo City, Guangxi, South China

Gigantopithecus blacki is a typical member of the Stegodon–Ailuropoda faunal complex (sensu lato) that inhabited southern China or, more broadly, mainland Southeast Asia during the Early and Middle Pleistocene. Current evidence indicates that the giant ape became extinct during the Middle Pleistocene. Recently, new remains of G blacki and associated mammalian fossils have been unearthed from a karst cave site, Hejiang Cave, in Chongzuo City, Guangxi, South China. The age of the Gigantopithecus-bearing depositional unit is estimated to be 400–320 ka using 230Th–234U disequilibrium U-series dating of flowstone samples bracketing the deposits. These finds document the latest occurrence of Gigantopithecus and provide potential insights regarding its extinction. Comparisons of dental dimensions between the Hejiang G. blacki remains, more than four hundred isolated teeth from Early Pleistocene localities, and over ninety isolated teeth from local drugstores show that the Hejiang teeth are slightly larger in their buccolingual dimensions. In addition, the crowns of the three unerupted upper premolars differ from those of all of the other Gigantopithecus material in having more complex crenulations. The differences in dental dimensions and morphology are possibly reflective of dietary responses to environmental changes that eventually led to the extinction of Gigantopithecus.

Sorex bifidus n. sp. and the rich insectivore mammal fauna (Erinaceomorpha, Soricomorpha, Mammalia) from the Early Pleistocene of Żabia Cave in Poland

Thirteen species of insectivore mammals, among them the new taxon Sorex bifidus n. sp.and twelve others are described from one of the few middle Early Pleistocene localities of Poland, Żabia Cave. They are: Erinaceus sp. (Erinaceidae), Talpa minor and T. europaea (Talpidae), Paenelimnoecus pannonicus, Sorex minutus, S. runtonensis, S. (Drepanosorex) praearaneus, S. (D.) margaritodon, Asoriculus gibberodon, Neomys newtoni, Petenyia hungarica and Beremendoia fissidens (Soricidae). Measurements, systematic positions, palaeoecological requirements and illustrations are given. The four species of Sorex indicate a humide sylvan paleoenvironment. The abundance of moles is also connected with relatively humid conditions and more or less open areas with soft soils. On the other hand, S. runtonensis is probably an indicator of arid and relatively open biotope and the hedgehog, Erinaceus sp., of forest, shrubs and open areas. Neomys newtoni is connected with open water bodies. This shows a mosaic landscape – forest, open habitat patches and water bodies in the vicinity of the studied cave. The impoverishment of the shrew fauna of Europe during the Late Pliocene and Early Pleistocene caused by the extinction of the Pliocene taxa and the appearance of more numerous species of the genus Sorex are presented.

Reconsideración de la sistemática de <i>Cervavitus</i> (Cervidae, Artiodactyla, Mammalia) del Pleistoceno Inferior

La revisión sistemática de Cervivatus sugiere que deriva del principal clado de los cérvidos posteriores a los muntiacinos, e implica que Procervulinae, Dicrocerinae y la primeras formas de Munticiacinae serían holometacarpales, como también lo es Cervivatus, originario en los bosques de Moldavia (Europa del Este) durante el Vallesiense final (MN 10), a partir de un antecesor braquiodonto y holometacarpal, con astas con dos o tres candiles y pliegue paleomerícido, y que da lugar a C. novorossiae. Este se dispersó a Europa occidental durante el comienzo del Turoliense, y más al oeste a Francia durante el Rusciniense. Su dispersión en los bosques del norte de China se produjo también a comienzos del Turoliense, estando representado por C. shanxius. Existe una gran cantidad de ejemplares de C. shanxius con metápodos laterales completos, que debían ser útiles para equilibrar el cuerpo en las ramas de los árboles. Las áridas mesetas loésicas actuales fueron bosques húmedos durante el Mioceno final. C. shanxius emigró hacia el sur durante el Plio-Pleistoceno probablemente debido al ambiente más seco del norte de China, como consecuencia de la elevación de los Himalayas, dando lugar a C. ultimus y C. fenqii, que sobrevivieron en el sur de China hasta el Pleistoceno inicial (MNQ18).

Hylopetes magistri, new name for Hylopetes debruijni Reumer & Van den Hoek Ostende, 2003, preoccupied

In 2003, we published a new species of Hylopetes, H. debruijni, from the early Pleistocene locality of Tegelen, The Netherlands (Reumer and Van den Hoek Ostende, 2003). This publication appeared i...

The second find of a primate from the early Middle Pleistocene locality of Mauer (SW Germany): a molar of Macaca (Mammalia, Cercopithecidae)

The second find of a primate from the early Middle Pleistocene locality of Mauer (SW Germany): a molar of Macaca (Mammalia, Cercopithecidae)

Presence of the Asian origin Bovini, Hemibos sp. aff. Hemibos gracilis and Bison sp., at the early Pleistocene site of Venta Micena (Orce, Spain)

The assemblages of large bovids from the European Late Villafranchian are dominated by the presence of the genus Bison, a descendant of the Early-Middle Villafranchian forms of Leptobos. Chronologically, the oldest record of this genus at the gates of Europe is at Dmanisi, Georgia (1.77 Ma), where it was firstly named Dmanisibos. It is recorded until the Holocene. It is the most abundant Quaternary buffalo of the continent and is nearly the only genus recorded in most post-Olduvai Early Pleistocene sites of Europe. The revision of all bone and tooth remains of large bovids from the site of Venta Micena (∼1.5 Ma) shows that two species of buffaloes are present at this Early Pleistocene locality. One is clearly a form of Bison, while the other corresponds to other different Bovini of small size that may be ascribed to the genus of Indian origin Hemibos, thus representing the first record of this genus in the Late Villafranchian of Europe. This paper presents the cranial material of both buffaloes.

A re-evaluation of the diversity of Megantereon (Mammalia, Carnivora, Machairodontinae) and the problem of species identification in extinct carnivores

ABSTRACT In this article, hypotheses about the origin, evolution and dispersal of Megantereon are reviewed using the fossil specimens included in previous comparative studies as well as the remains identified in the late Pliocene site of Fonelas (Spain) and the early Pleistocene localities of Lantian, Lingyi, Longdan, Renzidong (China), and Untermassfeld (Germany). The validity of the two species proposed by Martínez-Navarro and Palmqvist (1995), Megantereon cultridens and M. whitei, is evaluated using tooth measurements and multivariate statistical methods. The hypothesis of sexual dimorphism as an explanation for the morphological variability of Megantereon is tested with a large sample of sexed individuals of Panthera pardus and Panthera leo. Results obtained indicate similar or even smaller differences in tooth dimensions between M. cultridens and M. whitei than between sexes in both leopards and lions, except in the case of the lower fourth premolar. However, in spite of a substantial overlap between both Megantereon species in the size of the upper canine, this tooth reverses the differences found for other tooth measurements, because M. cultridens shows larger cheek teeth on average than M. whitei but smaller sabers. This is confirmed by principal components and discriminant analyses, which reveal that sexual dimorphism in leopards and lions is a matter of tooth size and not of relative proportions and argues against the interpretation of M. cultridens and M. whitei as the sexes (males and females, respectively) of a single species. These results indicate that M. cultridens and M. whitei are valid species, because the differences in tooth measurements exceed those expected from sexual dimorphism and do not reveal the effects of biased sampling. Finally, an analysis of jaw anatomy reveals biomechanical differences between both Megantereon species, related to the relative efficiency of the biting muscles at the level of the lower carnassial.

Réévaluation de la présence humaine au Pléistocène inférieur dans le Sud de l'Espagne

The Early Pleistocene locality at Venta Micena (Orce, Guadix-Baza basin, province of Granada, Spain) has provided four fossil remains – skull fragment VM-0, and long bone diaphyses VM-1960, VM-3691, and VM-12000 – which have been tentatively attributed to the hominids. Although several methodologies have been used to ascertain the human affinities of these specimens – including anatomical, morphometric and immunological analyses – the results obtained have not been conclusive, instigating a persistent debate. A taphonomic approach is used here for estimating the probability that a taxon the size of Homo sp. (~ 50 kg) could be represented in the fossil assemblage by four bone fragments and no tooth remain. A least-squares regression analysis between the percentage of teeth and the body mass estimated for each taxon of large mammals ( N = 20) predicts a raw abundance of six teeth for Homo sp. in the assemblage. Given that up to the present moment no tooth remains attributable to the hominids has been unearthed during systematic excavations in the Venta Micena quarry, which has provided more than 15,000 fossils of large mammals, this argues strongly against the possibility that the three bone specimens could belong to Homo sp. The phalanx CV-0 from the Early Pleistocene site of Cueva Victoria (Cartagena, Spain) has also been attributed to the genus Homo. The taxonomic assignment of this specimen is biased, however, because it was not compared with Theropithecus oswaldi, the only primate species actually recorded from this karstic locality. A comparative anatomical and morphometric analysis of fossil and modern specimens of Theropithecus suggests that CV-0 can be attributed to T. oswaldi. As a result, Cueva Victoria does not contribute additional information concerning the first human settlements in Europe. By these reasons, apart from the paleoanthropological and archaeological findings from Atapuerca (TD lower levels and Sima del Elefante), the rich archaeological assemblages from Barranco León and Fuente Nueva-3 in Orce, dated 1.3-1.2 Myrs, which include fourteen hundred stone tools of Oldowan technology, constitute at present the only unequivocal evidence of human presence in Southeast Spain during Early Pleistocene times.

Akhalkalaki: The Taphonomy of An Early Pleistocene Locality in the Republic of Georgia

As part of our investigations into the potential of the Republic of Georgia for providing information on early hominin occupation of Eurasia, we report here on Akhalkalaki, a large late Early Pleistocene locality located along the lower slopes of a Miocene andesitic cone. Originally excavated in the 1950s as a palaeontological site, it was re-opened in the 1990s and stone tools were found associated with the fauna, suggesting that it is also an archaeological occurrence. Excavations in the 1950s and 1990s uncovered thousands of bones of an early Galerian fauna, including the remains of new species of Hippopotamus, Equus , and Canis (Vekua, 1962, 1987) and dominated by the remains of Equus süssenbornensis. We present the stratigraphy of the site, which together with faunal correlations and reversed paleomagnetics indicates an age most likely in the late Matuyama Chron, probably between 980,000 and 780,000 years ago. Taphonomic analysis suggests that the fauna was deposited and buried over a short time period, and was heavily modified by carnivores, but we cannot demonstrate involvement by hominins. Based on evidence of abundant krotovina (animal burrows filled with sediment) and the lack of definitive evidence for hominin modification to the bones, the stone tools at the site may have been mixed in with the older fauna. The taphonomic characteristics of the Akhalkalaki bone assemblage are not readily explained with reference to assemblage formation processes developed with actualistic studies that have been mostly conducted in Africa, including carnivore dens, predator arenas, human hunting and scavenging, mass deaths, or attritional bone deposition. Because of extreme anthropogenic modification of the present environments, the temperate setting, and the presence of mainly extinct taxa, local models based on actualistic studies cannot approximate the mammalian ecology reflected in the Akhalkalaki bone assemblage. A few comparisons are made with preliminary taphonomic observations from Dmanisi, an Early Pleistocene Homo ergaster site not far away.

Taxonomic revision and phylogeny of Paedotherium and Tremacyllus (Pachyrukhinae, Hegetotheriidae, Notoungulata) from the Late Miocene to the Pleistocene of Argentina

ABSTRACT Two genera of the subfamily Pachyrukhinae (Notoungulata, Hegetotheriidae), Paedotherium and Tremacyllus, are revised. The material comes from late Miocene (Chasicoan) to early Pleistocene (Marplatan) localities in Argentina and Bolivia. The genus Raulringueletia is here considered a synonym of Paedotherium. Three valid species of Paedotherium are recognized and re-characterized: P. bonaerense, P. typicum, and P. minor. Two species of Tremacyllus are recognized: T. impressus in the Pampean region, and T. incipiens in Catamarca Province. The presence of Tremacyllus sp. into the late Chasicoan extends its known temporal range. Pachyrukhos occurs slightly earlier within the Chasicoan, coexisting with Paedotherium. Paedotherium is dominant in frequency in the Pampean area (Buenos Aires and La Pampa) and extremely scarce in the Subandean region (Catamarca), while Tremacyllus has an opposite occurrence. Cladistic analysis of the subfamily shows Paedotherium as a monophyletic group, where P. bonaerense is the sister taxon of the clade P. minor plus P. typicum. Pachyrukhos is the sister group of Paedotherium, and Tremacyllus appears to be closer to Pachyrukhos or Propachyrucos than to Paedotherium.