Palaeoloxodon and Human Interaction: Depositional Setting,Chronology and Archaeology at the Middle Pleistocene Ficoncella Site (Tarquinia,Italy)

The Ficoncella site in northern Latium (Italy) represents a unique opportunity to investigate the modalities of a short occupation in an alluvial setting during the Lower Palaeolithic. The small excavation area yielded a lithic assemblage, a carcass of Palaeoloxodon antiquus, and some other faunal remains. The main objectives of the study are to better characterize the depositional context where the Palaeoloxodon and the lithic assemblage occur, and to evaluate with greater precision the occupation dynamics. A 25 m-long well was drilled just above the top of the terrace of the Ficoncella site and faunal and lithic remains were analyzed with current and innovative techniques. The archaeological site contains floodplain deposits as it is located next to a small incised valley that feeds into a larger valley of the Mignone River. A tephra layer capping the site is ⁴⁰Ar/³⁹Ar dated to 441± 8 ka. Collectively, the geochronologic, tephrochronologic and geologic data, suggest the site was occupied during MIS 13. The new results should prompt further research at Ficoncella in order to improve our understanding of the dynamics of human settlement in Europe during the Early to Middle Pleistocene.     

The Crocuta crocuta spelaea (Goldfuss 1823) population and its prey from the Late Pleistocene Teufelskammer Cave hyena den besides the famous Paleolithic Neandertal Cave (NRW,NW Germany)

Remains from at least seven individuals of the Late Pleistocene Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) from the Teufelskammer Cave in the Neandertal valley (North Rhine-Westphalia, northwest Germany) are described. The small cave was a well-frequented hyena den of the Early to Middle Late Pleistocene which was only 100 m from the famous small Feldhofer Cave, where the first Neandertal human skeleton was found. The high amount of hyena bone material (37%) and its strongly chewed and incomplete prey remains of the mixed mammoth steppe and boreal forest megafauna prove one more of 11 recently known hyena den caves in the Rhenish Massif. Hyenas and cave bears have used the cave, but Neandertal humans lived possibly not at the same time in the same valley. Although hyenas occupied mainly the smaller caves such as the Teufelskammer Cave, humans preferred large portal cave entrances such as in the Neandertal valley with the Small Feldhofer Cave.     

The Xujiayao 14 Mandibular Ramus and Pleistocene Homo Mandibular Variation

The earlier Late Pleistocene mandibular ramus from Xujiayao (northern China) preserves traits that vary distributionally among western Old World Pleistocene Homo samples and between Early/Middle Pleistocene archaic humans and Late Pleistocene modern humans in eastern Eurasia. Xujiayao 14 presents a lateral mandibular notch crest, an open mandibular foramen, a wide ramus, an asymmetrical mandibular notch, an enlarged superior medial pterygoid tubercle, (probably) a retromolar space, and gonial eversion, as well as an unusual depression in the planum triangulare. The first two traits appear ancestral for Later Pleistocene and recent Homo and are dominant among modern humans. The second two traits largely separate Xujiayao 14 and archaic Homo from modern humans. The next two traits are found in the highest frequency among the Neandertals, although gonial eversion contrasts with Late Pleistocene Neandertals. Xujiayao 14, in the context of Pleistocene and recent Homo samples and the other Xujiayao human remains, therefore provides a morphological mosaic, highlighting regional variation through the Pleistocene.     

Oaks in the Kashmir Valley with Remarks on their History

The paper records the occurrence of indigenous oaks (Quercus semecarpifolia and Q. dilatata) in the Kashmir valley, which have hitherto been reported as absent there. It also gives an outline history of the oaks in the Kashmir valley since the beginning of the Pleistocene and discusses the historical status of the modern oaks in the valley.     

Notes on the systematics and distribution of Palaearctic blues (Lepidoptera,Lycaenidae)

Two new species and one new subspecies of blues are described from Central Asia, namely Polyommatus kuronjerus Korb, sp. n. (type locality: Kyrgyzstan, the Suusamyr valley, 7 km E of Suusamyr, 2800 m), Neolycaena turkomana Korb, sp. n. (type locality: Turkmenia, Badkhyz Nature Reserve, the Kyzyldzhar valley) and Satyrium acaudatum balasagyna Korb, ssp. n. (type locality: Kyrgyzstan, the Kyrghyz Mts., Tatyr, Golubinyi Waterfall, 1200 m). A new location for Plebejidea elvira (Eversmann, 1854) is determined for the Boomskoye valley in the Kyrghyz Mts. This new location is related to the late Pleistocene refuge in the Issyk-Kul valley.     

Phylogeographic Patterns of mtDNA Variation Revealed Multiple Glacial Refugia for the Frog Species Feirana taihangnica Endemic to the Qinling Mountains

Diversification patterns and demography of montane species are affected by Pleistocene climate fluctuations. Empirical cases from the Qinling Mountains (QM) region, which is a major biogeographic divider of East Asia, are few. We used DNA sequence data of the complete mitochondrial ND2 gene to detect effects of the Pleistocene glaciations on phylogeographic profiles of a frog species, Feirana taihangnica, which is endemic to the QM. Four distinct lineages consisting of seven sublineages were revealed. The strongest signal of biogeographical structure (F _ct = 0.971, P < 0.01) was found when populations were grouped according to these seven sublineages. One narrow secondary contact zone was detected in the middle QM between the lineage from middle QM and the lineage from eastern QM. Coalescent simulations indicated that this species colonized the QM region by a stepping-stone model. Divergences among lineages had likely been influenced by the uplift of the Tibetan Plateau during the late Miocene-to-late Pleistocene, as well as by the Pleistocene climatic cycles. Coalescent simulations also suggested that F. taihangnica populations have persisted through the Pleistocene glacial periods in multiple refugia across the QM region. Demographic analyses indicated that all lineages, except the lineage in the Funiu Mountains, have been experienced postglacial expansion of population size and distribution range. In conclusion, Pleistocene climate fluctuations and tectonic changes during the late Miocene-late Pleistocene have profoundly influenced the phylogeography and historical demography of F. taihangnica.     

New Oldowan localities at high level within Kilombe Caldera,Kenya

New occurrences of early artefacts ascribed to the Oldowan tradition come from localities at high level within the caldera of the extinct Kilombe volcano, located in the central rift valley of Kenya. The trachyte cone and caldera of Kilombe volcano formed at ca. 2.5 Ma, and the record of >130 m of sediment-fill indicates that the caldera subsequently held a lake for long periods during the Early Pleistocene. The Oldowan artefact localities, dated by ⁴⁰Ar/³⁹Ar and palaeomagnetism to ～1.78 Ma, lie east of the centre of the caldera, on the west side of an ancient small lake, which later drained away as a gorge formed on the east side of the mountain. The artefacts are dominantly made of Kilombe trachyte, and are associated with a fauna of large animals including Hippopotamus gorgops. These are the first Oldowan localities to be discovered in a new area of the Kenyan rift valley in the last thirty years, and their presence at high level in rugged landscape indicates that the associated hominins were exploiting a full range of environments.     

The geographical and chronological pattern of herpetofaunal Pleistocene extinctions on the Iberian Peninsula

Despite the postulation of the “herpetofaunal stability hypothesis”, which suggests that the herpetofauna underwent relatively few changes during the Pleistocene, the amphibian and squamate faunas of western Europe are known to have progressively diminished during the Pliocene and the beginning of the Pleistocene. Iberian Early Pleistocene sites continue to document the presence of “exotic” herpetofaunal elements that are supposed to have disappeared from the mainland, such as Oriental vipers, agamid lizards, the anguids Pseudopus and Dopasia, and a representative of the green toad group (Bufo viridis sensu lato), as well as possibly gekkonids, a scincid lizard (Lygosominae), the anguid Ragesaurus and blind snake (Scolecophidia). The geographical and temporal pattern of these progressive southward withdrawals and extirpations on the Iberian Peninsula shows that extirpation events occurred in northern Spain at roughly the Olduvai paleomagnetic event and in southern Spain just before the Jaramillo paleomagnetic reversal, thus permitting us to establish a framework that can be used to complement the biochronological zonation of the Iberian Early Pleistocene.     

Villafranchian-Galerian mammal faunas transition in South-Western Europe. The case of the late Early Pleistocene mammal fauna of the Frantoio locality,Arda River (Castell’Arquato,Piacenza, Northern Italy)

We investigated the fossil mammal assemblage from the late Early Pleistocene (early Galerian) site of the Arda River. Seven taxa were identified: one carnivore (Ursus dolinensis), five artiodactyls (Sus strozzii, Pseudodama farnetesis, Praemegaceros sp., Bison sp., and Hippopotamus sp.) and one perissodactyl (Stephanorhinus hundsheimensis). The Arda River faunal assemblage includes taxa already present in the Villafranchian, such as Pseudodama and Sus strozzii. The bear U. dolinensis and the bison, a more evolved form than Bison (Eobison), are taxa typical of the beginning of the Galerian mammal age. Paleontological, sedimentological and paleobotanical data point towards the presence of woodlands settled on an alluvial plain with patches of grassland dated around 1.0 Ma. The continental sediments of the Arda River represent one of the first steps of the forming of the alluvial Po valley.     

Detection and characterization of Hyaloklossia kasumiensis (Apicomplexa: Sarcocystidae) from Nagoya Daruma pond frog,Pelophylax porosus brevipodus

The Nagoya Daruma pond frog (Pelophylax porosus brevipodus) is an endangered anuran species endemic to the semi-aquatic environment of Japan. On June 11, 2021, four of the six Nagoya Daruma pond frogs captured from the wild and kept in an aquarium died in rapid succession. Pathological and/or parasitological examinations of two of the four carcasses revealed protozoan infection in their kidneys. The pathogen identified as Hyaloklossia kasumiensis (Apicomplexa: Sarcocystidae: Hyaloklossiinae) based on their morphological and molecular characterization. This is a new host record of H. kasumiensis. Mitochondrial cox1 sequence analysis suggest that the present species detected from Nagoya Daruma pond frog belongs to the same lineage as those found in the Tokyo Daruma pond frog (P. p. porosus) from Ibaraki Prefecture, Japan. This particular lineage may be prevalent among Pelophylax porosus ssp. distributed in Japan.     

Gene flow across secondary contact zones of the Emys orbicularis complex in the Western Mediterranean and evidence for extinction and re‐introduction of pond turtles on Corsica and Sardinia (Testudines: Emydidae)

European pond turtles represent a phylogeographically deeply structured complex of distinct taxa. Here, we use mitochondrial DNA sequences (cytochrome b gene) and eight polymorphic microsatellite loci to investigate genetic differentiation and gene flow of Sicilian, Corsican and Sardinian pond turtles and of subspecies involved in two secondary contact zones in the Pyrenean region and Southern Italy. Mitochondrial and microsatellite differentiation is largely concordant in populations from the core regions of the distribution ranges of the studied taxa. Both marker systems provide no evidence for gene flow between Sicilian pond turtles (Emys trinacris) and Southern Italian subspecies of E. orbicularis. By contrast, in the contact zones limited gene flow occurs between distinct subspecies of E. orbicularis. Although the Southern Italian contact zone is significantly older than the Pyrenean contact zone of Holocene age, patterns of asymmetric introgression are similar. Introgressive hybridization leads to the exchange of mitochondria, but microsatellite data indicate only a few individuals with mixed ancestry. This suggests that incipient isolating mechanisms maintain largely discrete nuclear genomic gene pools. Furthermore, this implies that Southern Italy acted as a hotspot rather than as a melting pot of genetic diversity during the last glacial. Pond turtles from Corsica and Sardinia are not differentiated from continental populations of the subspecies E. o. galloitalica, neither in the mitochondrial nor in the quickly evolving microsatellite markers. As the fossil record argues for a continuous presence of pond turtles on both islands since the Middle Pleistocene, this suggests that the native island populations became extinct and the extant turtles were later introduced by prehistoric settlers. The lack of genetic differentiation of pond turtles from Corsica and Sardinia supports the view that the subspecies described from these islands are not valid.     

Dispersal of Pleistocene Equus (Family Equidae) into South America and Calibration of GABI 3 Based on Evidence from Tarija,Bolivia

The dispersal of Equus into South America during the Great American Biotic Interchange (GABI) represented a major event for Pleistocene land-mammal age chronology on that continent. It has been argued that this dispersal occurred during the late Pleistocene, ∼0.125 Ma, and it defines the base of the Lujanian South American Land Mammal Age (SALMA). In this scenario, Equus dispersed during the fourth and latest recognized phase of the interchange, i.e., GABI 4. Although Equus was widely distributed in South America during the Pleistocene, only a few localities are calibrated by independent chronostratigraphic data. In this paper, new biostratigraphic evidence documents that Equus occurs from 15 superposed faunal horizons or zones throughout the Tolomosa Formation at Tarija, Bolivia. This biostratigraphic sequence is independently calibrated to occur between ∼0.99 to <0.76 Ma during the middle Pleistocene Ensenadan SALMA and coincident with GABI 3, not GABI 4. Tarija remains the only well calibrated Ensenadan locality at which Equus is found. The new biostratigraphic data presented here are unambiguous and document the earlier (pre-Lujanian) occurrence of this genus in South America. The hypothesized dispersal of the genus Equus into South America at ∼0.125 Ma is no longer supportable in light of the new biostratigraphic evidence presented here. The new data from Tarija thus have continent-wide implications for the origins and biogeography of Equus in South America as well as the calibration of GABI 3.     

Stephanorhinus hundsheimensis (Mammalia,Rhinocerotidae) from the late early Pleistocene deposits of the Denizli Basin (Anatolia,Turkey)

Pleistocene rhinoceroses are poorly documented in Turkey where they have been reported only from the late early Pleistocene (1.3–1.1 Ma) travertine deposits of the Denizli Basin. In this work, new rhinoceros remains collected from this basin are assigned to a relatively large-sized Stephanorhinus hundsheimensis on the basis of their morphology and morphometry. The first Turkish record of this species is approximately coeval with the first appearance of S. hundsheimensis in Europe, chronologically referred to the late early Pleistocene, ca. 1.2 Ma. During that time, S. etruscus still survived in Iberian Peninsula, central Italy and Dacian Basin. The presence of two successive evolutionary morphs for S. hundsheimensis during the Pleistocene is not confirmed.     

Mitochondrial Analysis of the Most Basal Canid Reveals Deep Divergence between Eastern and Western North American Gray Foxes (Urocyon spp.) and Ancient Roots in Pleistocene California

Pleistocene aridification in central North America caused many temperate forest-associated vertebrates to split into eastern and western lineages. Such divisions can be cryptic when Holocene expansions have closed the gaps between once-disjunct ranges or when local morphological variation obscures deeper regional divergences. We investigated such cryptic divergence in the gray fox (Urocyon cinereoargenteus), the most basal extant canid in the world. We also investigated the phylogeography of this species and its diminutive relative, the island fox (U. littoralis), in California. The California Floristic Province was a significant source of Pleistocene diversification for a wide range of taxa and, we hypothesized, for the gray fox as well. Alternatively, gray foxes in California potentially reflected a recent Holocene expansion from further south. We sequenced mitochondrial DNA from 169 gray foxes from the southeastern and southwestern United States and 11 island foxes from three of the Channel Islands. We estimated a 1.3% sequence divergence in the cytochrome b gene between eastern and western foxes and used coalescent simulations to date the divergence to approximately 500,000 years before present (YBP), which is comparable to that between recognized sister species within the Canidae. Gray fox samples collected from throughout California exhibited high haplotype diversity, phylogeographic structure, and genetic signatures of a late-Holocene population decline. Bayesian skyline analysis also indicated an earlier population increase dating to the early Wisconsin glaciation (~70,000 YBP) and a root height extending back to the previous interglacial (~100,000 YBP). Together these findings support California’s role as a long-term Pleistocene refugium for western Urocyon. Lastly, based both on our results and re-interpretation of those of another study, we conclude that island foxes of the Channel Islands trace their origins to at least 3 distinct female founders from the mainland rather than to a single matriline, as previously suggested.     

Pleistocene and Holocene rhinocerotids (Mammalia,Perissodactyla) from the Indochinese Peninsula

Rhinocerotids were abundant and diverse in southern Asia during the Pleistocene and the Holocene epochs, as shown by palaeontological and archaeological discoveries published throughout the last century, whereas the only living rhinoceros in the Indochinese Peninsula is Rhinoceros sondaicus (Cat Loc Reserve, Vietnam). The Pleistocene-Holocene Indochinese rhinocerotid record consists of the extinct species Dicerorhinus gwebinensis (Early Pleistocene, Myanmar) and representatives of the Recent Asian Species Rhinoceros unicornis (Middle-Late Pleistocene), R. sondaicus (Middle Pleistocene-Recent), and Dicerorhinus sumatrensis (Middle Pleistocene-Holocene). This fossil record is synthesized, mapped for Early/Middle/Late Pleistocene and Holocene/Recent times, and then compared with coeval rhinocerotid assemblages from the adjacent areas (South China), subregions (Indian, Sundaic, Philippine, and Wallacean), and region (Palearctic), from a biochronological and biogeographical perspective.     

First discovery of colobine fossils from the early to middle Pleistocene of southern Taiwan

Here we report on two kinds of cercopithecid fossil monkeys (Cercopithecinae and Colobinae) from the early to middle Pleistocene sediments of the Chochen (=Tsochen) area (Tsailiao-chi or Shinhua Hill), southern Taiwan. The fossil specimens include the first fossil record of colobine monkeys from Taiwan, where only macaque monkeys now occur. All cercopithecine fossils were identified as Macaca cf. Macaca cyclopis, the extant Taiwan macaque, except for one extremely large isolated upper molar, which may belong to another macaque species. On the other hand, all colobine specimens fall within the size variation of extant and extinct Rhinopithecus, but its specific status cannot be determined because of the scantiness of the fossil material. In Taiwan, Rhinopithecus presumably became extinct in the late Pleistocene, probably owing to global cooling and vegetation change, whereas macaques, which are of almost the same body size as Rhinopithecus, survived as M. cyclopis to the present. The contrasting history of survival between the two kinds of monkeys may be due to ecological/behavioral differences between them or as a result of accidental events that occurred in the Pleistocene of Taiwan.