Mammal fauna during the Late Pleistocene and Holocene in the far northeast of Europe

The paper summarises materials on the mammal remains in northeastern Europe, dated by radiocarbon. Altogether, 23 local faunas of small mammals and 47 local faunas of large mammals were analysed. Multidimensional statistical analysis shows a strong correlation between changes in small mammal fauna composition and climate changes throughout time. The correlations with the spatial gradients, however, are less pronounced. The faunas are classified into three groups: (1) faunas of Holocene age; (2) Late Pleistocene ‘stadial’ assemblages; and (3) Late Pleistocene ‘interstadial’ assemblages. In some cases, changes in species abundance are better understood in terms of biotic interrelations rather than of climatic effects. The most pronounced change in small mammal fauna composition and structure occurred at the Preboreal/Boreal boundary, and a less conspicuous alteration took place at the LGM/Lateglacial transition. The most noticeable transformation in the large mammal fauna composition is dated to the early Holocene. Less significant changes are observed at the Middle Weichselian/LGM transition and at the LGM/Lateglacial transition. It is safely concluded that variations in the faunas of small and large mammals recorded in NE Europe during the last 35 000 years occurred synchronously and unidirectionally.     

泥河湾地区晚上新世一个新的地层单位——稻地组

本文记述了河北泥河湾盆地的一个新的地层单位——稻地组。在四个实测剖面上对稻地组进行了分层描述,根据其岩性、化石特征、及其与上下地层接触关系,确定了该地层组的含义,并对其时代(晚上新世)作了讨论。     

中国北方晚更新世哺乳类动物群的划分及其地理分布

<正> 新第三纪以来,由于青藏高原、喜马拉雅山脉与西部山脉的隆起,我国形成了西高东低的地形特点,导致了气候和环境的变迁,明显地影响了中国哺乳动物的分布。在晚更新世,中国南北动物群分化逐渐扩大,动物群更替尤为频繁。中国晚更新世地层及动物群的研究已有一段相当长的历史,发现了众多重要的遗址和化石地点,积累了丰富的资料。笔者综合有关资料,对我国北方晚更新世动物群,特别是猛犸象-披毛犀动物群的划分、地     

On the present habitats and ecology of Vertigo pseudosubstriata Ložek, 1954 (Mollusca,Gastropoda, Vertiginidea) in Central Asia and its distribution history in Central and Eastern Europe

The small terrestrial gastropod Vertigo pseudosubstriata Ložek, 1954 is one of the rarest glacial indicator species in the Pleistocene of Central and Eastern Europe. In all, this species has been found at only about 15 sites in Europe. V. pseudosubstriata was initially described as a fossil in Central Europe and was discovered only later alive in Central Asia. With regard to its modern distribution, 25 habitats with V. pseudosubstriata have been examined in Tien Shan and in the central and southern Altai. These findings seem to capture the contemporary distribution of the species and provide information on the boundaries of its ecological requirements. These data are of great significance for the interpretation of the fossil assemblages. Since the few fossil specimens in Europe date from very different glacial periods in the Elsterian, Saalian Complex and Weichselian, it can be concluded that V. pseudosubstriata apparently immigrated in at least three distinct waves. Most of the Pleistocene specimens in eastern Central Europe and Eastern Europe are reported from archaeological sites of the Upper Middle Weichselian (Gravettian), roughly between 33 and 29 ka cal bp. In this paper, we review all reported modern and fossil occurrences and discuss the species' ecological range.     

Late Pleistocene and Holocene small- and large-mammal faunas from the Northern Urals

Data from the mammal fauna of the North Urals during the Late Pleistocene and Holocene are synthesised. Analysis of differentiation the degree of small- and large-mammal faunas during this time has been undertaken. Only differences of mammal species composition were significant between the Late Pleistocene and Holocene complexes, and within these complexes, the distinction between faunas was insignificant. The transition from the Late Pleistocene to the Holocene complex small-mammal faunas occurred in the Middle Late Valdai due to expansion of the forest species. In large-mammal faunas, the process was recorded later (in Dryas 3-Early Holocene) because of the extinction of some species and others that changed their areas of occupation.     

Plio-Pleistocene geological record and small mammal faunas, eastern shore of the Azov Sea, Southern European Russia

The shoreline of the Taganrog Gulf of the Azov Sea at the mouth of Don River provides a series of extended Upper Pliocene and Quaternary sections that have been actively studied in the last century. This extraglacial region had a complex sedimentary history combining subaerial aggradation with marine, fluvial, and deltaic sedimentation. The well-exposed stratigraphical sequence and abundant palaeontological record continuously attract geologists and highlight the region as one of international importance for the addressing of numerous problems of Late Pliocene and Quaternary stratigraphy and palaeogeography. Fossil mammalian faunas of the region include important Eurasian biostratigraphical markers such as Stenocranius ex gr. hintoni-gregaloides, Lagurini spp., and Mimomys savini. For many years, fossil remains of mammals provide decisive clues to the geological history of the region. Recent geological studies of reference sections have provided data on small mammals, palaeomagnetism and palaeogeographical reconstructions in the northeastern part of the Azovian Region. Mammalian assemblages indicate the presence of the Late Pliocene, late Early Pleistocene, Middle Pleistocene and Late Pleistocene levels and, in addition, provide a clear biostratigraphical context for the Early Middle Pleistocene transition.     

临夏盆地晚新生代哺乳动物群演替与青藏高原隆升背景

临夏盆地的晚新生代沉积中富含哺乳动物化石,以晚渐新世巨犀动物群、中中新世铲齿象动物群、晚中新世三趾马动物群和早更新世真马动物群的化石最为丰富。晚新生代是青藏高原快速隆升的时期,临夏盆地的4个主要哺乳动物群在构造剧烈变化的背景下发生了显著的更替。通过对不同动物群所代表的生态特征的分析,恢复了临夏盆地晚新生代以来的气候环境演变过程:晚渐新世以温暖湿润的森林环境为主,间杂有一些开阔地带;中中新世的森林更加茂密,水体更加丰富;晚中新世演变为炎热半干旱的稀树草原环境,季节性变化加强;早更新世气候寒冷而干燥,并伴有显著的海拔升高。青藏高原在晚渐新世的隆升幅度还不足以阻挡大型哺乳动物在高原南北的交流,但到中中新世已成为明显的障碍,至晚中新世对动物迁徙的阻碍作用更加突出,而临夏盆地在早更新世已经达到相当大的高度,产生了一个高原或高山的动物群     

The stratigraphical significance of deer species in the cromer forest-bed formation

Deer species provide a valuable biostratigraphical tool through Cromer Forest-bed times, due particularly to species turnover between the Early and early Middle Pleistocene. This study is based on the reidentification of 348 fossil antlers. The provenance of most large mammal fossils from the CF-bF, collected over 150 years, was recorded only by the nearest coastal village. None the less, analysis of the cervid taxa by these ‘localities’ reveals interesting patterns. The fauna of the West Runton Freshwater Bed is of early Middle Pleistocene complexion, and that from the foreshore at East Runton is of Early Pleistocene. Pure or nearly pure early Middle Pleistocene assemblages also occur at Kessingland-Pakefield and at Trimingham. At Overstrand, Sidestrand, Mundesley, Bacton and Happisburgh, there is a mixture of Early and early Middle Pleistocene elements. Analysis of Savin's data shows that fossils of earlier species were generally found further down the beach than those of later ones. Late nineteenth century geological surveys, made when bone collecting was at its peak, give additional information about fossil horizons, which in several cases can be related to modern stratigraphical units. Most Early Pleistocene large-mammal bones came from Pastonian conglomerates, in contrast to small-mammal and molluscan assemblages mostly extracted from Pre-Pastonian Crag. However, the diversity of Early Pleistocene CF-bF cervid species in comparison with continental faunas, and their pattern of distribution between sites, suggests they may span more than one chronostratigraphic stage. Early Middle Pleistocene assemblages came from strata now referred to the Cromerian, and the differing proportions of taxa between sites provide limited evidence of time-transgression.     

中国晚侏罗世多瘤齿兽类哺乳动物的发现及其意义

多瘤齿兽是一类已绝灭的哺乳动物,其以后部牙齿上有很多小瘤(小的隆起,或者小尖)来适于咀嚼植物为特征。总的来说,它们的门齿和臼齿与那些啮齿类相似,但在演化时间上多瘤齿兽比新生代的啮齿类早的多。多瘤齿兽是杂食者,或者说是成功开拓了其它脊椎动物无法开拓的植食生境的植食者(可以树叶、种子、蕨类、裸子植物,再加上蠕虫和昆虫为食)。由于它们多种食性和运动方式,多瘤齿兽成为中生代数量最多的哺乳动物,几乎占侏罗纪和白垩纪时期所有哺乳动物种类的一半。它们的支系是哺乳动物历史上生存时间最长的支系,出现于170 Ma前,大约绝灭于35 Ma。但是,因为多瘤齿兽具有很多独特的、高度特化的牙齿和头骨特征,古生物学家们长期困惑于它们的进化起源。最近,中国辽西发现了一件产自晚侏罗世髫髻山组的多瘤齿兽类哺乳动物。由于其牙齿上发育明显的纹饰,如很多小的脊、沟、凹槽等,新发现的哺乳动物被命名为皱纹齿兽(Rugosodon)。另一方面,由于与皱纹齿兽亲缘关系最近的哺乳动物产自西欧的侏罗纪地层,因此其种名被命名为欧亚皱纹齿兽(Rugosodon eurasiaticus)。新发现为晚侏罗世时期欧洲和亚洲的哺乳动物具有高度的相似性提供了最新的证据。皱纹齿兽是一种夜行的哺乳动物,生活于气候温和的湖滨,与长羽毛的恐龙近鸟龙、翼龙类达尔文翼龙、大量节肢动物、其它几种哺乳动物共同生活于一起。根据与现生哺乳动物的比较,并依据欧亚皱纹齿兽手部比例、末端指节的形态等,推论欧亚皱纹齿兽很可能是地栖哺乳动物,或者说地栖比攀爬的可能性更大。这一发现对于研究多瘤齿兽早期的演化、食性分异、运动适应起源等具有重要的科学意义。     

辽宁西部第四纪哺乳动物及其地层时代

以大、小凌河流域为中心的辽西地区第四纪地层分布广泛, 奥陶纪石灰岩为主的基岩使该地区喀斯特地貌发育, 并在水溶蚀作用下形成洞穴, 为哺乳动物栖息提供了理想的场所。20世纪70年代以前曾发现过建平、凌源西八间房、喀左鸽子洞和锦县沈家台等化石产地, 之后陆续发现了喀左帽儿山洞穴、二布尺洞穴、朝阳马山洞和凌源沟门洞化石产地等。根据动物化石组合及其对比, 辽西发现的这些化石群分别代表中更新世晚期、晚更新世早期、晚更新世中期和晚更新世晚期4个地质时代。这些材料的发现, 对辽西乃至整个东北地区第四纪哺乳动物研究及区域第四纪地层的划分对比等具有重要意义。     

Effects of Pleistocene environmental changes on the distribution and community structure of the mammalian fauna of Mexico

Biological communities in Mexico experienced profound changes in species composition and structure as a consequence of the environmental fluctuations during the Pleistocene. Based on the recent and fossil Mexican mammal checklists, we determine the distribution, composition, diversity, and community structure of late Pleistocene mammalian faunas, and analyze extinction patterns and response of individual species to environmental changes. We conclude that (1) differential extinctions occurred at family, genus, and species level, with a major impact on species heavier than 100 kg, including the extinction all proboscideans and several ruminants; (2) Pleistocene mammal communities in Mexico were more diverse than recent ones; and (3) the current assemblages of species are relatively young. Furthermore, Pleistocene relicts support the presence of biogeographic corridors; important refugia existed as well as centers of speciation in isolated regions. We identified seven corridors: eastern USA–Sierra Madre Oriental corridor, Rocky Mountains–Sierra Madre Occidental corridor, Central United States–Northern Mexico corridor, Transvolcanic Belt–Sierra Madre del Sur corridor, western USA–Baja California corridor, Tamaulipas–Central America gulf lowlands corridor, and Sonora–Central America Pacific lowlands corridor. Our study suggests that present mammalian assemblages are very different than the ones in the late Pleistocene.     

The Cenozoic on-shore basins of Northern Vietnam: Biostratigraphy,vertebrate and invertebrate faunas

A first account of paleontological data from three Cenozoic on-shore basins in Northern Vietnam, i.e. the Na Duong, Cao Bang, and Hang Mon basins, reveals a rich fossil fauna and flora of supposed Oligocene age, offering a great potential for taxonomic, paleoenvironmental, and paleobiogeographic studies. Two excavation campaigns unearthed well-preserved fossil remains of mammals, crocodiles, at least six turtle species, some 20 fish taxa, some other 20 mollusc species, and different plant remains. The majority of these taxa are regarded as new to science. However, close affinities to modern faunas of northern Southeast Asia demonstrate the importance of these fossils for an evaluation of the biological history of this modern biodiversity hot spot. Moreover, the fossil assemblages may help to disentangle the intricate Cenozoic tectonic evolution of Southeast Asia by application of paleobiogeographic modelling. Finally, the discovery of complex paleo-food-webs and the presence of several taxa indicative of certain ecological conditions provide a solid base for autecologic, synecologic and paleoclimatic studies. The potential biostratigraphic value of the macrofauna has to be demonstrated yet, as evolutionary concepts for most of the respective groups have not been proposed to date.     

Late Sarmatian mammal localities of the Eastern Paratethys: Stratigraphic position,magnetochronology, and correlation with the European continental scale

Analysis of geological sections, paleogeography, and paleomagnetic data is used to reveal succession of the middle to late Sarmatian mammal localities of the Eastern Paratethys and their correlation with the continental stratigraphic scale of Western Europe. Until recently, the late Sarmatian localities were correlated with MN10 and even MN11 zones. As is proved, all the known late Sarmatian localities should be correlated with the upper half of Zone MN9. The terminal late Sarmatian faunas only, which are correlative with the lowermost Chron C4Ar3r and older than 9.6 Ma, can be referred to Zone MN10. According to essential changes in taxonomic composition of faunas from continental localities around the Eastern Paratethys, which are recorded in the mid-late Sarmatian, Zone MN9 can be divided in two subzones MN9a and MN9b separated by boundary at ～10.5 Ma. The refined correlation can be used to establish difference between faunas of the Southeastern, Central, and West European paleozoogeographic provinces and to assess diachronism in dispersal of mammals. In the Southeastern province, many forms characteristic of the Turolian in Central and Western Europe first appeared as early as in the mid-Vallesian. The results obtained indicate that faunal criteria used to define boundaries of MN zones in Western Europe are of a regional importance being inapplicable to the entire North Eurasia and even to Europe as a whole that is unfortunately ignored by many paleontologists. Criteria of distinction should be worked out for each paleozoogeographic province. As geochronological levels of faunal changes are identical throughout the northern Palearctic, they suggest impact of global factors despite variable manifestation in different regions.     

周口店地区第四纪哺乳动物群的演化与环境变迁探讨

讨论了Ａｌｉｌｅｐｕｓａｎｎｅｃｔｅｎｓ、ＨｙｐｏｌａｇｕｓｂｒａｃｈｙｐｕｓＬｅｐｕｓｓｐ．、Ｓｃｉｕｒｏｔａｍｉａｓｐｒａｅｃｏｘ、ＡｌｌｏｃｒｉｃｅｔｕｓｔｅｉｌｈａｒｄｉＣｒｉｃｅｔｉｎｕｓｖａｒｉａｎｓ、Ｈｙｐｅｒａｃｒｉｕｓｙｅｎｓｈａｎｅｎｓｉｓ、Ａｌｌｏｐｈａｉｏｍｙｓｃｆ．ｐｌｉｏｃａｅｎｉｃｕｓ、Ｐｒｏｅｄｒｏｍｙｓｃｆ．ｂｅｄｆｏｒｄｉ、Ｃｈａｒｄｉｎｏｍｙｓｎｉｈｅｗａｎｉｃｕｓ、Ｍｕｓｃｆ．ｍｕｓｃｕｌｕｓ和Ａｐｏｄｅｍｕｓｄｏｍｉｎａｎｓ的地史分布。根据哺乳动物群的分析与对比，得出第１８地点为早更新世早期，第１２地点、东洞地点为早更新世中期，第９地点、西洞地点为早更新世晚期，第１３地点为早更新世末期至中更新世初期，第１、２、５、６、１１、１０、２１、２３及上店洞地点为中更新世，新洞为中更新世晚期至晚更新世早期，东岭子洞为晚更新世，山顶洞为晚更新世晚期。还讨论了周口店地区一些主要第四纪哺乳动物群的生态类型特点，第四纪的气候波动及生态环境变迁。     

The East Side Story – The Transylvanian latest Cretaceous continental vertebrate record and its implications for understanding Cretaceous–Paleogene boundary events

The latest Cretaceous continental vertebrate faunas of the wider Transylvanian area figured prominently in discussions concerning the Cretaceous–Paleogene Boundary (K-Pg) events when they were first described by Nopcsa between 1897 and 1929, because they were assumed to be late Maastrichtian in age. Subsequently their age was reconsidered as early Maastrichtian, and were thus regarded of lesser importance in understanding the K-Pg boundary events in Europe and worldwide. Moreover, Transylvanian continental vertebrate assemblages (the so-called ‘Haţeg Island’ faunas) were often lumped together as a temporally restricted assemblage with a homogenous taxonomic composition. Recent fossil discoveries and more precise dating techniques have considerably expanded knowledge of the Transylvanian vertebrate assemblages, their ages, and their evolution. A synthesis of the available stratigraphic data allows development of the first comprehensive chronostratigraphic framework of the latest Cretaceous Transylvanian vertebrates. According to these new data, expansion of continental habitats and emergence of their vertebrate faunas started locally during the latter part of the late Campanian, and these faunas continued up to the second half of the Maastrichtian. During this time, long-term faunal stasis appears to have characterized the Transylvanian vertebrate assemblages, which is different from the striking turnovers recorded in western Europe during the same time interval. This suggests that there was no single ‘Europe-wide’ pattern of latest Cretaceous continental vertebrate evolution. Together, the available data shows that dinosaurs and other vertebrates were relatively abundant and diverse until at least ca. 1 million years before the K-Pg boundary, and is therefore consistent with the hypothesis of a sudden extinction, although this must be tested with future discoveries and better age constraints and correlations.     

Southern dispersal and Palaeoecological implications of woolly rhinoceros (Coelodonta antiquitatis): review of the Iberian occurrences

Cold-adapted large mammal populations spread southward during the coldest and driest phases of the Late Pleistocene reaching the Iberian Peninsula. Presence of woolly rhinoceros (Coelodonta antiquitatis) can be identified from 23 Iberian sites, which is compiled and analyzed herein, and the fossil specimens from seven of these sites are described here for first time.Morphological and biometrical analyses demonstrate that the Iberian woolly rhinoceros did not significantly differ from individuals of other European populations, but represent the westernmost part of a continuous Eurasian belt of distribution.The first presence of woolly rhino in the Iberian Peninsula has been identified during the late Middle Pleistocene and early Late Pleistocene. However, the highest abundance of this species is recorded during MIS 3 and 2. The latest Iberian occurrences can be dated around 20 ka BP. The presence of woolly rhinoceros in the Iberian Peninsula correlates with periods of extreme dry and cold climatic conditions documented in Iberian terrestrial and marine sediment sequences.From a palaeobiogeographic point of view, the maximum southern spread of C. antiquitatis on the Iberian Peninsula was registered during the late Middle Pleistocene or early Late Pleistocene, reaching the latitude of Madrid (about 40°N). Subsequently, during MIS 3 and 2, all Iberian finds were restricted to the Northern regions of Iberia (Cantabrian area and Catalonia). The southern expansion of C. antiquitatis during the Late Pleistocene in the Iberian Peninsula reached similar latitudes to other Eurasian regions.The ecological composition of fossil assemblages with presence of woolly rhinoceros was statistically analyzed. Results show that temperate ungulate species are predominant at Iberian assemblages, resulting in a particular mixture of temperate and cold elements different of the typical Eurasian cold-adapted faunal associations. This particular situation suggests two possible explanations: a) Eventual migrations during the coldest time spans, resulting in a mixing of cold and temperate faunas, instead a faunal replacing; b) Persistence of woolly rhinoceros populations in the Iberian Peninsula during interglacial episodes confined at cryptic southern refugia.     

Tephrochronology and tephrostratigraphy of two Pleistocene continental fossiliferous successions from central Italy

The stratigraphical context of two Middle Pleistocene fossiliferous palaeosols from Central Italy (Abruzzo and Tuscany) have been studied. Small mammals and molluscs occur in both palaeosols, which are covered by tephra layers that were analysed using an interdisciplinary approach. Application of fission‐track dating to apatites separated from the Case Picconetto tephra (Pescara, Abruzzo), yielded an age of 0.48 ± 0.04 Ma, indistinguishable from those previously determined for the Campani Quarry (Lower Valdarno, Tuscany) (0.46 ± 0.05 Ma and 0.48 ± 0.05 Ma). Geochemical and petrographic investigations indicate that these tephra originated from different volcanoes, the Alban Hills Volcanic Complex and the Vico Volcano (Latium) respectively. Small mammal and mollusc assemblages indicate different palaeoclimatic and palaeoenvironmental conditions for the Case Picconetto and Campani Quarry palaeosols. Warm and humid conditions can be inferred for the Campani Quarry site, whereas open and cold conditions can be inferred for Case Picconetto. On the basis of faunal data, fission‐track dates and attribution of tephra to specific volcanic eruptions, we suggest a correlation of these faunas with marine oxygen isotope stage 14 (Case Picconetto) and with marine oxygen isotope stage 11 (Campani Quarry), respectively. Copyright © 2003 John Wiley & Sons, Ltd.     

An Early Pleistocene hippopotamus from Westbury Cave,Somerset, England: support for a previously unrecognized temperate interval in the British Quaternary record

Although fossil assemblages from the late Early Pleistocene are very rare in Britain, the site of Westbury Cave in Somerset, England, has the potential to address this gap. The mammal fossils recovered previously from the Siliceous Member in Westbury Cave, though few in number, have hinted at an age for the deposits that is as yet unparalleled in Britain. Here, we describe the first bona fide occurrence of Hippopotamus in the British Early Pleistocene, discovered during recent reinvestigation of the Siliceous Member. The hippo fossil indicates a refined biochronological age of ca. 1.5–1.07 Ma for the Siliceous Member and a palaeoclimate that was warm and humid, which accords well with previous palaeoenvironmental inferences. A synthesis of late Early Pleistocene hippo occurrences suggests that the Siliceous Member hippo may have been part of an early colonization of north-west Europe by these megaherbivores, possibly during MIS (Marine Oxygen Isotope Stage) 31. Alternatively, it evidences a currently cryptic northward migration during an even earlier temperate phase. In either case, the Siliceous Member is likely to represent a warm period that has not been recognized previously in the British Quaternary record.     

甘肃广河地区中中新世哺乳动物化石的发现及其地层学意义

本文记述了采自甘肃广河石那奴中中新世地层的哺乳动物化石6目10科12属16种类（以小化石为主）,其中新种两个:Alloptox guangheensis sp. nov., Pseudaelurus guangheensis sp. nov.。通过对动物群性质的研究及与其它地区的详细对比,认为下化石层的时代应为中中新世早期,上化石层应为中中新世晚期。二者分别与山旺期、通古尔期大体相当。 广河地区中中新世早期车头沟组和中中新世晚期咸水河组的建立,表示该地区存在着中中新世两个时期的不同环境沉积。     

Reconstructing the palaeoenvironments of the early Pleistocene mammal faunas from the pollen preserved on fossil bones

We carried out a systematic investigation on the pollen content of sediment adhering to skeletal elements of large mammals which originate from the long lacustrine record of Leffe (Early Pleistocene of the Italian Alps). Three local faunas were discovered during mining activities along the intermediate part (spanning from 1.5 to 0.95 Ma) of the basin succession. The excellent pollen preservation allowed testing the reproducibility of the pollen signal from single skeletons. A clear palaeoenvironmental patterning, consistent with the ecological preferences of the considered mammal species, emerged from the canonical correspondence analysis of pollen types diagnostic for vegetation communities. Edaphic factors related to seasonal river activity changes and to the development of swamp forests in the riverbanks are significantly associated to the occurrences of Hippopotamus cf. antiquus, whereas finds of Mammuthus meridionalis belong to fully forested landscapes dominated by conifer or mixed forests of oceanic, warm to cool-temperate climate. Rhinoceros habitats include variable forest cover under different climate states. Distinct cool-temperate, partially open vegetation could be recognized for large deer included Cervalces cf carnutorum.A palynostratigraphic correlation between individual spectra and a reference palynostratigraphic record allowed assignment of many fossil remains to a precise stratigraphic position. This procedure also shown that the Leffe local faunas include specimens accumulated under different environmental and climate states, as a consequence of high-frequency climate changes characterizing the Late Villafranchian Early Pleistocene.