Early Pleistocene Locality Research Articles

Etruscan Wolf Canis etruscus (Canidae, Carnivora) from the Early Pleistocene of Crimea (Taurida Cave).

The cranial and mandibular fragments of the medium-sized canid from the Early Pleistocene locality of Taurida cave (about 1.8-1.5 Ma) in Crimea are described and included in the hypodigm of the species Canis etruscus Forsyth Major, 1877, a common member of the Late Villafranchian fauna of the Circum-Mediterranean region. This species is reported for the first time from the Northern Black Sea region. The discovered evidence reinforce previous published idea regarding the periodically use of the Taurida cave by canids and other large predators as a den and shelter.

Cheetah Acinonyx pardinensis (Felidae, Carnivora) from the Early Pleistocene of Crimea (Taurida Cave).

A mandible fragment of Acinonyx pardinensis (Croizet et Jobert, 1828) is described from the Early Pleistocene locality in the Taurida cave (Crimea, Late Villafranchian, about 1.8-1.5 Ma). This is the first discovery of the genus Acinonyx in the Pleistocene of Crimea. Along with other felids, such as Homotherium, Megantereon, Panthera, Lynx, and Puma, cheetahs were typical representatives of the Late Villafranchian faunas of Eurasia. In the Taurida locality, Acinonyx pardinensis is co-occurred with Homotherium crenatidens, Megantereon adroveri, and Lynx issiodorensis.

Re-investigation of fossil Lemmini specimens from the early and Middle Pleistocene of Western and Central Europe: Evolutionary and paleoenvironmental implications

The current study focuses on the emblematic Myopus/Lemmus species complex (tribe Lemmini) in the European Pleistocene fossil record. The members of the two genera occupy distinct ecological niches and have different external appearances, but they are remarkably similar in their dental morphology, so that they were commonly thought of as undistinguishable in the fossil record. Thus, more or less all European Lemmini fossils have been assigned to the genus Lemmus. In the Early Pleistocene site of Schernfeld (Germany), the species Lemmus kowalskii had been described. It was thought by some authors that all Lemmini from Early to late Middle Pleistocene belong to this species.In the current study, we investigated Lemmini molar morphology from Western and Central European sites including Schernfeld (Early Pleistocene), Sackdillinger Höhle (Sackdilling Cave), and Koněprusy C718 (both early Middle Pleistocene), as well as other fossil localities with fewer specimens, formerly assigned to Lemmus kowalskii. Using an extensive modern referential material of Lemmus and Myopus, this study proposes to re-evaluate taxonomic status of the Middle and Early Pleistocene Lemmini. This modern referential also allows a better understanding of the morphology of Lemmus kowalskii specimens and its variability.Our results highlight the very high variation within fossil populations, as well as significant statistical differences between populations of the Early and Middle Pleistocene localities. A large part of these fossil specimens is firmly identified as Myopus sp., including the L. kowalskii holotype. Our identifications demonstrate that in most Early and Middle Pleistocene sites considered in this study, both genera (Lemmus and Myopus) are present. Possible interpretations and consequences for current view of lemming history are discussed, as well as some of the paleoecological and paleoenvironmental implications.

The Occurrence of Suids in the Post-Olduvai to Pre-Jaramillo Pleistocene of Europe and Implications for Late Villafranchian Biochronology and Faunal Dynamics

It has been proposed that suids were absent from Europe during the post-Olduvai to pre-Jaramillo Early Pleistocene (from less than 1.8 to more than 1.2 Ma) and that their “re-appearance” in the late Early Pleistocene would mark the end of the late Villafranchian and the beginning of the Epivillafranchian. Arguments enumerated in favor of this “suid gap” are the lack of suid remains from extensively sampled fossil localities of this age and the high reproductive potential (r-strategy) of suids, which would translate in a high commonness of their remains in the fossil record. However, here it shown that while suids’ reproductive potential is certainly exceptional within artiodactyls, there is no direct relationship between the reproductive strategy and preservation rate of a taxon in the fossil record. In Early Pleistocene localities of Europe and adjoining areas, where suids are present in a fossil assemblage, they are always rare. In terms of number of occurrences (frequency), suids range from being moderately common (~2.0–1.8 Ma) to moderately rare (~1.1–1.0 Ma). Suid material is also described herein from Peyrolles (Issoire, France; reference locality for MNQ 19), a site dated at 1.47 Ma, providing direct evidence for the presence of suids within the purported “suid gap”. The case of suids underlines an important source of caveat in inferring faunal dynamics of the late Early Pleistocene of western Europe—including the dispersal of hominins—i.e., the unequal geographical distribution of the paleontological sites of post-Olduvai to pre-Jaramillo age. Indeed, Peyrolles is the only large mammal site in western Europe located outside the Iberian and Italian Peninsulas reliably dated around 1.5 Ma. In the post-Olduvai to pre-Jaramillo period, there is a paucity of radiometric estimates (or they have too coarse a resolution) and of paleomagnetic excursions detectable in continental deposits. Basically, for this time span, there is a high dependence on biochronological correlations, although, at the same time, these correlations are less reliable—because these are based on a few sites not covering the entire spectrum of environments present in Europe and the sites are not independently dated with methods that outperform biochronology—than those for other periods.

Seasonality and Lithic Investment in the Oldowan

Seasonality is a critical driver of resource availability within individual generations and is therefore likely to have exerted selective pressures on hominin evolution. Nonetheless, it has frequently been overlooked in discussions of early technological variability. This paper examines the relationship between seasonality and lithic investment in the Oldowan of eastern Africa, to broaden our understanding of hominin behavioural adaptability. We develop a framework for the identification of seasonal signatures in the archaeological record and examine the nature of lithic variability across 12 Early Pleistocene localities. In combining these insights, we suggest that seasonality played a pivotal role in shaping the pattern of lithic investment in the Oldowan, with peak investment related to greater wet season processing of plant resources and mobility across the landscape. The productivity of cores, retouching of flakes, and use of percussive implements may each play a role in this seasonal patterning of tool use. Our analysis provides further evidence that Oldowan hominins were able to adapt their technological behaviours to changing environmental conditions, even at the scale of intra-seasonal fluctuations in resource availability, and how this interacts with landscape familiarity over longer timescales.

Water, gravity, and trophic dynamics in the Misiam palimpsest, Olduvai Gorge

Misiam is a modern leopard-generated wildebeest bone accumulation located on the slope of a steep ravine within Olduvai Gorge (Tanzania). We present a taphonomic assessment of how leopard feeding behaviour shaped the bone assemblage, accumulated during the wet season, as well as contextualising the extent of secondary hyaenid scavenging. The spatial formation of this taphonomic palimpsest has resulted from the temporal sequence of interactions with the bone remains. The impact of additional agents and processes—such as fungi, insects, weathering, and, most importantly, gravitational displacement driven by water flows generated during the rainy season—adds to the complexity of this locality. This last factor may have played a decisive role in the density-mediated osteological attrition documented in the bone assemblage, perhaps to an even greater extent than hyaenid activity. Given the critical importance for analogical studies of documenting all relevant taphonomic factors at work in modern assemblages, our perspective allows for a comprehensive interpretive assessment of the site. The Misiam surface assemblage, a leopard-generated bone accumulation palimpsest with multiple taphonomic agents and processes at work within a short time span, has therefore significant implications for nuancing or complementing the interpretation of Early Pleistocene localities, such as FLK-North, FLK-Zinj or TK in Olduvai (Tanzania), Ubeidiya (Israel) or Barranco León in Orce (Spain), among others.

Carnivora from the Early Pleistocene locality of Karnezeika (Southern Greece)

Karnezeika is an Early Pleistocene locality in the Peloponnese, South Greece. More than 3000 specimens have been unearthed and studied, revealing the presence of a large mammal assemblage, which corresponds to the middle Villafranchian biochronological unit. The bovid fauna of the site has already been presented, as well as the scarce remains of a large cercopithecid, which characterizes Karnezeika as a primate-bearing fossil site. Herein, we describe the Carnivora of this locality. The carnivore guild of Karnezeika was found to consist of seven different taxa including Vulpes alopecoides, Felis sp., Megantereon cultridens, Ursus etruscus, Pachycrocuta brevirostris, Baranogale helbingi, and a Mustelidae indet. Interestingly, the presence of the giant, short-faced hyena P. brevirostris among the assemblage shows that this species was present in SE Europe before the Olduvai subchron, similarly to Western Europe. Furthermore, based on the new and relatively well-preserved specimens belonging to the rare mustelid B. helbingi, we discuss some possible taxonomic interpretations of this species. Finally, the majority of the carnivora taxa present in Karnezeika indicate a mixed habitat which agrees with previous palaeoenvironmental results concerning the site.

Early Pleistocene Bovidae from Palan-Tyukan (Azerbaijan)

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.

To den or not to den. Contributions to the taphonomic history of the Early Pleistocene site of Venta Micena 4 (Orce, Guadix-Baza Basin)

Venta Micena (Orce, Guadix-Baza Basin, Spain) is an Early Pleistocene locality renowned for the richness and quality of its palaeontological record. VM is spread over an area of 2.5 km2, where several exposed fossil outcrops are visible amidst its gorges and ravines. The best known of these sites, VM3, has been interpreted as a hyaena den. In addition, a new site, named VM4, has recently been the focus of fieldwork and taphonomic studies. The publication by Luzón et al. (2021) pointed out that VM4 presents a more complex history than VM3. First, two different sub-levels were identified: VM4-I and VM4-II. Secondly, the preliminary taphonomic analysis showed conspicuous differences with regard to VM3. Nevertheless, such interpretation has been challenged by Palmqvist et al. (2022) who proposed that VM3 and VM4 are both the result of a single depositional process, entailing the selective transport of skeletal parts by the giant extinct hyaena Pachycrocuta brevirostris back to its den. Using well-preserved faunal elements whose depositional context and provenance are reliable, in this paper we show that: 1) there are two clearly defined sub-levels in VM4 with some shared taphonomic characteristics as well as some notable differences; 2) VM3 and VM4 exhibit enough divergence to support differences in site formation processes; 3) The interpretation of both VM4-I and VM4-II is more consistent with their characterisation as open-air sites in which multiple agents and depositional processes contributed to its formation, rather than with hyaena dens. Nevertheless, excavations are still in progress at VM4 and therefore any results and interpretations ought to be considered as provisional.

Chasmaporthetes lunensis (Hyaenidae, Carnivora) from the Early Pleistocene of Crimea

A maxillary fragment of the extinct hyena Chasmaporthetes lunensis (Del Campana, 1914) is described from the Early Pleistocene locality of the Taurida cave (Crimea, Late Villafranchian, 1.8-1.5 Ma). The species was a typical representative of the Villafranchian faunas of Eurasia. This is the first record of the genus Chasmaporthetes in the Pleistocene of Crimea.

"Oldowan" in the northern Caucasus – archaeology from the seabed or just geology

The authors discuss the geoarchaeological data indicating that the Oldowan-age assemblages, which are reported during the last years on the Taman peninsula, in the north-western Caucasus, and in Dagestan, in the north-eastern Caucasus, are found in the marine coast environment near the contact with the sea basin, with evidence for the presence of shallow water (in Muhkai II and other localities in Dagestan) or wave activity (in Kermek and other localities on Taman). The marine environment is not reported in any Early Pleistocene archaeological site in either Europe or south-western Asia, or Africa. Moreover, the geological contexts identified in the Early Pleistocene localities in the northern Caucasus correspond to typical conditions for the formation of eoliths or geofacts. The combined archaeological and geoarchaeological indicators strongly suggest that these North Caucasian assemblages most likely represent collections of naturally broken stones that were selected by researchers among fragments of the same rock that is present naturally in these areas.

Early Pleistocene Lynx issiodorensis (Felidae, Carnivora) from the Taurida Cave, Crimea.

The cranial and mandibular remains of two adult individuals of Lynx issiodorensis (Croizet et Jobert, 1828) are described from the Early Pleistocene locality of the Taurida cave (Crimea, Late Villafranchian, 1.8-1.5 Ma). This lynx species was a typical representative of the Villafranchian fauna of the Eastern Mediterranean. A high craniological variability of L. issiodorensis is noted.

Pleistocene foxes (Vulpes, Canidae, Carnivora) from the Taurida Cave, Crimea.

A mandible fragment and four isolated teeth of the fossil foxes, Vulpes alopecoides (Del Campana, 1913), Vulpes cf. vulpes (Linnaeus, 1758) and Vulpes sp., are described from the Early Pleistocene locality of Taurida cave (Crimea, Late Villafranchian, 1.8-1.5 Ma). Based on the size and morphological features, a mandible fragment and two M1 are attributed to V. alopecoides. The structure of m1 of Vulpes cf. vulpes is similar to that of V. vulpes and V. alopecoides, but the size greatly exceeds the size limits for the latter species. Although m1 of Vulpes sp. fits the size of the lower carnassials of V. alopecoides, its morphology is definitely unique among the Early Pleistocene representatives of the genus Vulpes.

New taxonomical, biochronological and palaeoenvironmental data from the Middle Pleistocene site of Cúllar de Baza 1 (Granada, Spain)

The intra-montane Guadix-Baza Basin is one of the few continental basins in Europe that hosts a well-dated set of fossiliferous sites spanning from the latest Miocene to the late Middle Pleistocene. The Cúllar de Baza 1 (CB-1) represents a key site to investigate the effects of the Early-Middle Pleistocene Transition, considered a fundamental transformation in the Earth’s climate state. Our review and update of the large mammal assemblage, and particularly equids, is of paramount relevance to understand the systematic affinities and the evolution of the Early and Middle Pleistocene European horses. We confirm the occurrence of two different taxa, the medium sized Equus altidens and the larger E. suessenbornensis. Moreover, we illustrate that CB-1 is essential for the biochronological studies of the latest Early Pleistocene/Middle Pleistocene transition (Epivillafranchian/Galerian ELMA); in particular with regard to the Last Occurrences of the Etruscan rhino Stephanorhinus etruscus and the large deer Megaloceros savini and the First Occurrence of the water-rat Arvicola mosbachensis in the Iberian peninsula. Finally, a development of a mosaic environment characterised the CB-1 site contrasting with the conditions reported for other Iberian late Early and Middle Pleistocene localities.

Faunal complex of the Early Pleistocene Muhkai 2 locality

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.

Giant Hyena Pachycrocuta brevirostris (Hyaenidae, Carnivora) from the Lower Pleistocene of Taurida Cave, Crimea.

The dental remains of a giant hyena Pachycrocuta brevirostris (Gervais, 1850) from the Early Pleistocene locality of the Taurida cave (Crimea, Late Villafranchian, 1.8-1.5 Ma) are described. This species was a typical representative of the Villafranchian fauna of the Eastern Mediterranean. The Taurida cave was occasionally used by hyenas and other carnivorans as a den and retreat.

The history of bears (Ursidae, Carnivora, Mammalia) from Silesia (southern Poland) and the neighbouring areas

Revision of the Silesian bear fauna, based on material from 152 sites, mainly cave and karstic localities, and also archaeological and open-air sites, shows the presence of 13 forms and species. These records encompass the last 16.5 Ma and may be divided into five main morphophyletic groups. The oldest bears, represented by the genera Ballusia and Ursavus , are dated to 16.5–11 Ma, and belonged to the stem forms of the subfamily Ursinae. After a break of 6 My, the earliest members of the genus Ursus appeared, and all known Pliocene bears in Silesia belong to the genus Ursus were identified as Ursus minimus . They represent one or two migration events. Additionally, between 3.6 and 3.2 Ma, a single occurrence of Agriotherium insigne was recorded from the Weze 1 site. A few Early Pleistocene bear records are represented by U. etruscus , which was a probable ancestor of both arctoid and spelaeoid bear lineages. The oldest representative of Ursus ex gr. arctos known so far and assigned to U. a. suessenbornensis is known from one latest Early Pleistocene (1.2–0.9 Ma) locality, while other Polish records of this form require confirmation. The first occurrence of U. deningeri , the oldest taxon within the U . ex gr. deningeri–spelaeus lineage, was recorded from ~700 ka deposits in Silesia. During the pronounced cold period of MIS 12, the Scandinavian ice sheet covered almost the entire modern territory of Poland, with the exception of the Sudetes and the Carpathians. The accompanying drastic faunal turnover led to the formation of the pan-Eurasian Mammoth Fauna at ~460 ka. At that time a characteristic member of this fauna, the steppe brown bear Ursus arctos priscus , a specific ecomorph adapted to live in open grasslands, appeared in this region. It survived until the beginning of MIS 1, when modern Ursus arctos arctos appeared in Silesia and survived to the present day. U. deningeri was the most common bear during the Middle Pleistocene, while the first records of U. spelaeus spelaeus appeared since MIS 7. The latter form was replaced by U. ingressus during the Late Pleistocene (~110–100 ka). Spelaeoid bears totally dominated the cave assemblage, and finally vanished between 27 and 24 ka.

When beremendiin shrews disappeared in East Asia, or how we can estimate fossil redeposition

ABSTRACT The current paper first time describes a small Beremendia from the late Pleistocene deposits in the Koridornaya Cave locality (Russian Far East), which associated with the extinct Beremendia minor. The paper is the first attempt to use a comparative analytical method to evaluate a possible case of redeposition of fossil remains of this shrew. In this case, the 'comparative' part of the method corresponds to the 'faunistic approach' of Jelle Reumer, and the 'analytical' part of the method corresponds to his extended 'morphological approach. ’ The analytic method uses morphospace size estimation (MSE) based on principal component analysis and estimation of the variance to evaluate redeposition of Beremendia minor from the late Pleistocene deposits in the Koridornaya Cave. This result allows us to suggest the significant environmental contribution to the East Asian beremendiin extinction, such as climatic fluctuation, landscape changes and shrew community changes between the late Pliocene and early Pleistocene. We discuss the last remains of East Asian Beremendia (Peisorex) described from the Chinese early Pleistocene localities (Chiachiashan, Haimao, Renzidong) and the last remains of Lunanosorex described from the Chinese late Pliocene localities (Qipanshan Hill and Houxushan Hill). MSE was performed for soricine tribes with the ‘elongated’ condylar process.

Fossil bear material from the oldest deposits in the Jasna Strzegowska cave (Silesia, southern Poland)

Fossil bear material from the lowermost deposits of the Jasna Strzegowska cave is described and its taxonomic status assessed. Comparison with bear remains from other Early and early Middle Pleistocene localities of Eurasia shows the presence of two bear species: Ursus etruscus and U. deningeri , based on morphological and size characters of the teeth and postcranial bones. The teeth of U. deningeri from the Jasna Strzegowska cave are larger, wider and have much more complicated occlusal surfaces when compared with the teeth of U. etruscus . Both bear species are characterized by relatively short and robust metapodials, although those of U. etruscus are on average less massive. Some differences in muscle attachments and articular surfaces have also been found. Metapodials of primitive arctoid bears, that co-occurred with U. etruscus and U. deningeri , are much longer and slimmer. U. etruscus and U. deningeri probably did not live in the Jasna Strzegowska cave at the same time. Based on its time range in Eurasia and morphological characteristics, the age of U. etruscus is estimated as Early Pleistocene, between 1.8–1.3 Ma. This is its first description from Poland, and the remains described are in size and morphology indistinguishable from material from other European localities. For U. deningeri , an early Middle Pleistocene age is suggested.

The latest Early Pleistocene amphibians and reptiles from Kaiafas (Greece) and the first record of fossil Ophiomorus (Squamata, Scincidae)

Together with the rest of the Balkan Peninsula, Greece acted as a refuge for thermophilic amphibians and reptiles during the Quaternary Glaciations. In this work we provide new herpetological data on the latest Early Pleistocene locality of Kaiafas, in western Peloponnese. The site yielded one indeterminate salamandrid urodelan (Salamandridae indet.), one tree frog (Hyla gr. H. arborea), one indeterminate testudinid (Testudinidae indet.), at least two lacertid lizards (Lacertidae indet. A and B), one scincid lizard (Ophiomorus sp.), one anguid lizard (Pseudopus sp.), and one indeterminate “colubrine” snake (“Colubrinae” indet.). This assemblage is indicative of a surrounding environment with water bodies, wooded areas, and possibly loose soils with rocks and stones. In spite of being based on few remains, the study of the amphibians and reptiles from Kaiafas resulted in the identification of the first fossil of Ophiomorus known worldwide. Moreover, the two lacertid taxa displayed very peculiar morphological features which are either previously unknown or rare in that clade.