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.     

Context and environment in taphonomic analysis: Examples from Alaska's Porcupine River Caves

Field investigations of caves along Alaska's Porcupine River document three major mechanisms which modify bone in patterns similar to alterations produced by man: (1) carnivore fracture; (2) rodent gnawing; and (3) rock fall and rubble scarring. A late Wisconsin faunal assemblage composed of Equus sp., Rangifer tarandus, Ovis dalli, Bison sp., proboscidean, numerous small mammal species, birds, and fish is well documented. This faunal assemblage suggests a mosaic environment of grassland-tundra-forest in the immediate vicinity of these caves and implies that the late Wisconsin environment in north-central Alaska may have been characterized by a number of microenvironments and colder, dryer, steppe conditions. Taphonomic data which have historically been interpreted to support human occupation of eastern Beringia during the Pleistocene are critically examined and the context of these discoveries (not the specimens themselves) provides the test essential to document the antiquity of man in North America prior to 12,000 yr ago.     

Wolverine in northern England at about 83,000 yr B.P.: Faunal evidence for climatic change during isotope stage 5

Cave sediments from Stump Cross Cave in northern England contain Pleistocene mammal remains. Uranium-series dating of calcium carbonate deposits closely associated with the fossiliferous horizons has established an absolute age of 83,000 ± 6000 yr B.P. for a faunal assemblage largely comprised of wolverines (Gulo gulo). This date lies firmly within the younger portion of oxygen-isotope stage 5. The occurrence of wolverines in the vicinity of Stump Cross Cave at ca. 83,000 yr B.P. indicates a significant climatic deterioration from ca. 120,000 yr B.P., when an Ipswichian interglacial fauna with hippopotamus was present in this part of northern England.     

<Emphasis Type="Italic">Mesopithecus</Emphasis> (Primates: Cercopithecoidea) from Villafranca d’Asti (Early Villafranchian; NW Italy) and palaeoecological context of its extinction

Remains of the cercopithecid Mesopithecus monspessulanus are relatively rare. Two previously unpublished mandibles of M. monspessulanus (housed in the Basel Naturhistorisches Museum), from the Italian locality of Villafranca d’Asti are described. These remains belong to the assemblage of the Triversa Faunal Unit, dated to the Early Villafranchian, that is to the unit MN16a (Middle Pliocene) of the European mammal biochronology. According to this recently revised biochronological attribution, Villafranca d’Asti records the last Mesopithecus occurrence in Europe. The NOW (Neogene Old World) database has been used as a basis to evaluate both the mammalian faunal and the palaeoenvironmental context at the time surrounding this last occurrence of Mesopithecus. The comparison (taxonomical composition and ungulate hypsodonty), between the Villafranca d’Asti assemblage and other Plio-Pleistocene mammal communities of Europe, shows that the extinction of Mesopithecus is related to a faunal turnover and a change toward more open landscapes during the Early–Middle Villafranchian transition. This is consistent with the “Elephant-Equus event”, that occurred in Europe at 2.5 Ma. Furthermore, the co-occurring but more terrestrial cercopithecid Macaca crossed this faunal turnover. This strengthens the assumption that the latest Mesopithecus species, M. monspessulanus, had woodland-oriented adaptations. Editorial handling: J.-P. Billon-Bruyat & M. Chiari (Guest)     

Small mammals from the early Pleistocene of the Granada Basin, southern Spain

The Pliocene and Pleistocene continental sedimentary records of the western sector of the Granada Basin, southern Spain, consist of alternating fluvial and lacustrine/palustrine sediments. Two Quaternary sections from this sector have been sampled: Huétor Tájar and Tojaire. They have yielded remains of rodents, insectivores and lagomorphs. The presence in the Huétor Tájar and Tojaire sections of Mimomys, Apodemus atavus, Castillomys rivas and two different species of Allophaiomys, indicates an Early Pleistocene age. These deposits, which are related to a fluvio-lacustrine system, can be differentiated from an older (Pliocene) braided fluvial system. Their dating has important repercussions on the paleogeographic reconstruction of the basin. The conditions inferred from the ecological preferences of the small mammal associations are wet and cold. These associations suggest a predominance of open herbaceous habitats, followed by forested habitats; semiaquatic habitats are the least represented.     

Macrobenthic communities from wetland impoundments and adjacent open marsh habitats in South Carolina

Forty-eight core and grab samples were taken from two impoundments and an adjacent tidal creek and salt marsh during each of six sampling periods (January, June and November 1983; and January, April and July 1984). Habitats sampled within the impoundments included the perimeter ditch and shallow vegeted areas dominated byRuppia maritima, Spartina alterniflora, andScirpus robustus. The adjacent tidal creek bottom and low marsh ofS. alterniflora were sampled for comparison with the impoundment sites. Major differences in faunal composition and density of macrobenthic invertebrates were observed between habitats in this study. Macrobenthic density was highest (475 individuals 0.05 m^?2) at the impoundment site dominated byScripus robustus, where oligochaetes were abundant. The open marsh site had a density of 254 individuals 0.05 m^?2. Among unvegetated sites, density for all sampling periods was higher in Chainey Creek than in the perimeter ditches of the impoundments. The total number of taxa was highest for the open marsh and tidal creek sites. The impoundments contained vegetated sites which were inhabited by fewer species than nonimpounded sites, while the perimeter ditch sites were comparatively depauperate. Cluster and nodal analyses identified four broad assemblages based on habitat: 1) an open marsh assemblage, 2) a creek assemblage, 3) a eurytopic assemblage, and 4) an impoundment assemblage. The separation of faunal assemblages by sampling site rather than sampling period suggests that physical differences between habitats were important factors determining distribution patterns.     

Early Neogene Radiolarian faunal turnover in the northern Indian Ocean: Evidence from Andaman-Nicobar

Two intervals of faunal turnover are revealed by the study of radiolarians from the Early to Middle Miocene sequence of Andaman-Nicobar belonging to Stichocorys wolffii-Calocycletta costata-Dorcadospyris alata zones. These faunal changes are reflected in the values of species diversity, change in abundance of taxa, origination and extinction events and change in radiolarian assemblages. One such faunal change is identified in the latest Early Miocene. The time of this faunal change is marked by the extinction of species like Carpocanopsis cingulata and appearance of Calocycletta costata, Giraffospyris toxaria, Acrocubus octopylus and Liriospyris parkerae, an increasing trend in percentage of cold water species and a decreasing trend in species diversity upwards. The interval coincides with the time of initiation of cooling of sea surface water. Another, and the most prominent faunal turnover of radiolarians is recognized in the Middle Miocene Dorcadospyris alata Zone at about 14.8–12.7 Ma and is characterized by almost complete disappearance of an earlier dominant assemblage and an increase in abundance of an assemblage that was practically absent in the older sequence. The time of this turnover can be correlated with the time of Middle Miocene cooling identified in the examined sequence.     

Pleistocene vertebrates of the Yukon Territory

Unglaciated parts of the Yukon constitute one of the most important areas in North America for yielding Pleistocene vertebrate fossils. Nearly 30 vertebrate faunal localities are reviewed spanning a period of about 1.6 Ma (million years ago) to the close of the Pleistocene some 10 000 BP (radiocarbon years before present, taken as 1950). The vertebrate fossils represent at least 8 species of fishes, 1 amphibian, 41 species of birds and 83 species of mammals. Dominant among the large mammals are: steppe bison (Bison priscus), horse (Equus sp.), woolly mammoth (Mammuthus primigenius), and caribou (Rangifer tarandus) – signature species of the Mammoth Steppe fauna (Fig. 1), which was widespread from the British Isles, through northern Europe, and Siberia to Alaska, Yukon and adjacent Northwest Territories. The Yukon faunas extend from Herschel Island in the north to Revenue Creek in the south and from the Alaskan border in the west to Ketza River in the east. The Yukon holds evidence of the earliest-known people in North America. Artifacts made from bison, mammoth and caribou bones from Bluefish Caves, Old Crow Basin and Dawson City areas show that people had a substantial knowledge of making and using bone tools at least by 25 000 BP, and possibly as early as 40 000 BP. A suggested chronological sequence of Yukon Pleistocene vertebrates (Table 1) facilitates comparison of selected faunas and indicates the known duration of various taxa.     

Hemipelagic cephalopods from the Maastrichtian (late Cretaceous) Parras Basin at La Parra,Coahuila, Mexico,and their implications for the correlation of the lower Difunta Group

Few biostratigraphic data exist from the Parras and La Popa basins, mainly due to the absence of index fossils. This paper describes 19 ammonoid species from 15 genera and 1 nautilid from La Parra, southeastern Coahuila, Mexico. The assemblage consists of Tethyan [(Baculites ovatus, Brahmaites (Anabrahmaites) vishnu, Fresvillia constricta, Hauericeras rembda, Pachydiscus (P.) ex gr. neubergicus, Solenoceras reesidei, Tetragonites cf. superstes], cosmopolitan (Anagaudryceras politissimum, Desmophyllites diphylloides, Diplomoceras cylindraceum, Gaudryceras kayei, Phyllopachyceras forbesianum, Pseudophyllites indra), and cold water taxa [Fresvillia teres, Hypophylloceras (Neophylloceras) surya, H. (N.) hetonaiense, Pachydiscus (P.) cf. egertoni]. Eutrephoceras sp. and Menuites juv. sp. were not determined to species level. A similar assemblage was recently described from the coeval Méndez Formation at Cerralvo, Nuevo León. Species endemic to North America, particularly the Western Interior Seaway, are absent at La Parra. The ammonoid assemblage and associated planktonic foraminifers allow for precise biostratigraphic assignation to the early Maastrichtian planktonic foraminiferal zone CF 5, and thus provide an important marker level for correlation of the lower Difunta Group. The new biostratigraphic data presented herein allow for the first time precise dating of the Cañon del Tule Formation of the Difunta Group. Their combination with existing sequence- and magnetostratigraphic data improve the correlation of the lower Difunta Group with time-equivalent lithostratigraphic units such as the Cárdenas Formation in Mexico. They also provide new insight into ammonoid migration patterns induced by sea-level changes. Baculites ovatus migrated into the La Popa Basin as a result of the sea-level highstand documented at La Parra.     

Bearing up well? Understanding the past,present and future of Australia's koalas

The modern Koala Phascolarctos cinereus is the last surviving member of a once diverse family Phascolarctidae (Marsupialia, Phascolarctomorphia). Nine genera and at least 16 species of koala are known. Late Oligocene sediments of central Australia record the oldest fossils and highest species diversity. Five species are known from the early to middle Miocene rainforest assemblages of the Riversleigh World Heritage Area, Queensland. With the onset of dryer conditions after the middle Miocene climatic optimum (~ 16 Ma), rainforest habitats contracted resulting in the apparent extinction of three koala lineages (Litokoala, Nimiokoala, Priscakoala). Phascolarctos first appears in the fossil record during the Pliocene and the modern species around 350 ka. Despite a dramatic decline in taxonomic diversity to a single extant species, the fossil record indicates that at most only three koala species coexisted in any given faunal assemblage throughout their 24 million year history. Within these assemblages, the vast majority of extinct koalas are extremely rare (some known from only a single specimen) which may reflect a general rarity within their palaeohabitats compared with the modern species which is represented by an estimated 400,000 individuals spread over most of eastern mainland Australia. Be that as it may, P. cinereus, although once geographically more widespread, occurring for example in Western Australia in the Pleistocene, underwent significant range contractions and localized population extinctions during the stressful climatic conditions of the late Pleistocene and more recently through human-induced habitat destruction. Combined with threats of disease, reduced genetic diversity and climate change, the survival of this iconic Australian marsupial is arguably a cause for concern.     

Effect of the Red Sea brine-filled deeps (Shaban and Kebrit) on the composition and abundance of benthic and planktonic foraminifera

Composition and abundance of benthic and planktonic foraminifera in surface sediments of the brine-filled Shaban and Kebrit Deeps and some bathyal-slope environments in the northern Red Sea were examined for correlation with environmental conditions (e.g., bathymetry, sediment grain-size, organic matter, and carbonates) of the brine-filled deeps and normal Red Sea water. About 67 benthic foraminiferal species were recorded in these sediments. The lowest faunal density and diversity were recorded in the Shaban and Kebrit Deeps, whereas the highest density and diversity were recorded in the bathyal-slope sediments. Cluster analysis divided the benthic foraminiferal species into three major faunal assemblages. Buccella granulata–Gyroidinoides soldanii–Bolivina persiensis assemblage dominated the 650–1,300 m depth due to predominance of oligotrophic, highly oxygenated bottom waters. The Melonis novozealandicum–Spirophthalmidium acutimargo assemblage was recorded in the deep and bathyal-slope sediments indicating its tolerance for wider ranges of environmental conditions. The deeps were only dominated by the Brizalina spathulata assemblage indicating existence of un-totally anoxic conditions. The deeps yielded also very low planktonic foraminiferal density that may be attributed to occurrence of the seawater–brine interface which not only minimized the deposition of high buoyancy, large-test species (Globigerinoides sacculifer, Globigerinella siphonifera, and Orbulina universa), but also overestimated the small-test species (Globigerinoides ruber, Globoturborotalita rubescens, and Globigerinita glutinata) in the sediments. These findings should be taken into consideration when reconstructing paleoceanographic conditions of the Red Sea using core sediments from the brine-filled deeps.     

Magnetostratigraphy and Pliocene-Pleistocene boundary in Italy

Analysis of some sedimentary series near Rome and comparison with other series in northern and southern Italy give evidence that Arctica islandica, considered a marker species for the beginning of the Pleistocene, appears during the Re´union paleomagnetic event (about 2 my ago). A period of erosion, called the Aullan erosional phase, is evident in the lower part of the late Villafranchian (= early Pleistocene, Olivola faunal unit).     

Biodiversity,taxonomic richness and specialists-generalists during Late Pleistocene/Early Holocene times in Pampa and Patagonia (Argentina,Southern South America)

Human subsistence systems in the Pampa and Patagonia regions evolved from generalists during the Late Pleistocene/Early Holocene to specialists during the Middle Holocene, according to diversity and relative taxonomic richness counts. The general hypothesis is summarized as follows: 1) at the end of the Pleistocene and beginning of the Holocene (interval between 13.0 and 8.5 ka), the Pampean and Patagonian landscape was different from the present one, with a mammal biodiversity larger than in the Middle Holocene; 2) during the Middle Holocene, the highest mammal taxonomic richness corresponds to one species, Lama guanicoe. Although available biomass was the same for the hunter-gatherers, it was due to the larger abundance of individuals of that single species; 3) the emerged continental surface during the Late Pleistocene/Early Holocene was more extensive than during the Middle Holocene. In ecological terms, this greater land mass is reflected in an equally larger biodiversity, even assuming that populations of Pleistocenic megafauna were diminishing in number since ca. 13,500 years BP.Based on the analysis of our own information and published data, we state that the human groups which colonized the Pampean and Patagonian regions towards the end of the Pleistocene and beginning of the Holocene used generalist strategies encompassing a wide range of faunal resources (birds and mammals, especially of terrestrial habitats). Plasticity of hunter-gatherer societies, plus a complex technology, a higher social mobility and unfilled territories, allowed them to rapidly replace one resource for another in conditions of environmental stress. This lifestyle resulted in the occupation of different ecological zones (niches) and the evolution to specialized systems, based on one or a few mammal species, once the colonization and support in the different environments were successful and the resources of the Late Pleistocene/Early Holocene disappeared.     

Preliminary investigation of Late Pleistocene fauna from Ryonggok Cave No. 1, Sangwon County,North Hwanghae Province,Democratic People's Republic of Korea

The investigation of a limestone cave on the Ryonggok-ri of Sangwon County, North Hwanghae Province, DPRK, has yielded many faunal remains including Late Pleistocene deer, brown bears and horses. Uranium series dating of fossil teeth yielded dates of ~72ka at the base of Layer 2 – the bone-bearing unit – and ~44ka at the top. The rich and diverse mammal remains of Ryonggok Cave No. 1 are indicative of warm and humid temperate conditions during the Late Pleistocene, and reflect the presence of both dense forests and open grassland. In addition, pollen and spores from the site include two families and 24 genera of trees, six families and a genus of grasses and herbs, and two families and six genera of ferns, corroborating the environments inferred from the faunal remains.     

The bivalve Anopaea (Inoceramidae) from the Upper Jurassic–lowermost Cretaceous of Mexico

In Mexico, the Upper Jurassic to lowermost Cretaceous La Casita and coeval La Caja and La Pimienta formations are well-known for their abundant and well-preserved marine vertebrates and invertebrates. The latter include conspicuous inoceramid bivalves of the genus Anopaea not formally described previously from Mexico. Anopaea bassei (Lecolle de Cantú, 1967), Anopaea cf. stoliczkai (Holdhaus, 1913), Anopaea cf. callistoensis Crame and Kelly, 1995 and Anopaea sp. are rare constituents in distinctive Tithonian–lower Berriasian levels of the La Caja Formation and one Tithonian horizon of the La Pimienta Formation. Anopaea bassei was previously documented from the Tithonian of central Mexico and Cuba, while most other members of Anopaea described here are only known from southern high latitudes. The Mexican assemblage also includes taxa which closely resemble Anopaea stoliczkai from the Tithonian of India, Indonesia and the Antarctic Peninsula, and Anopaea callistoensis from the late Tithonian to ?early Berriasian of the Antarctic Peninsula. Our new data expand the palaeogeographical distribution of the high latitude Anopaea to the Gulf of Mexico region and substantiate faunal exchange, in the Late Jurassic–earliest Cretaceous, between Mexico and the Antarctic Realm.     

Variation and pattern in the responses of mammal faunas to Late Pleistocene climatic change in southeastern South Australia

We examined mammal occurrence and variability through the Late Pleistocene vertebrate fossil deposit of Grant Hall in Victoria Fossil Cave, Naracoorte, South Australia. To determine long‐term patterns of change, we compared the composition and relative abundance trends of the assemblage with a nearby Middle Pleistocene deposit in Cathedral Cave. Total species richness did not change through the Grant Hall sequence, dated from 93 ± 8 to 70 ± 5 ka. However, species relative abundances varied between ecologically divergent species, and in some cases between species that demonstrate similar environmental preferences. For some species this variation is comparable to that recorded in Cathedral Cave. Of those showing similar trends between the two deposits, the forest inhabitant, Pseudomys fumeus, recorded an 8.6% decline through Grant Hall, coincident with a 9.7% increase in the dry heath/mallee dweller Pseudomys apodemoides. These patterns indicate that climatic transition from relatively warm, moist to cooler, drier conditions impacted some species in similar ways through climatic cycles of the past. However, the majority of the fauna demonstrated complex responses that are individual and variable through time. Statistical tests of species trends from the Grant Hall assemblage caution that large fossil samples are required to validate patterns observed. Copyright © 2012 John Wiley & Sons, Ltd.     

甘肃省东乡族自治县龙担黄土底部新发现的哺乳动物群初步报道

龙担哺乳动物群发现于甘肃省东乡族自治县的早更新世黄土底部。此前我国早更新世的哺乳动物化石大多发现于河-湖相地层中,在黄土中极少发现,而且主要在六盘山以东。在龙担哺乳动物群中还发现了灵长类化石,这在我国西北地区尚属首次。龙担哺乳动物群产于黄土底部约30m的地层中,最低的层位距“底板”(积石组砾岩)仅约3m。化石相当丰富,其中有不少新的种类。目前已发现的化石共21种,大体可以分为3类:1)与泥河湾哺乳动物群相同或很接近的;2)与泥河湾哺乳动物群相比,在进化水平上稍低,因而可能代表较早地质时代的;3)在泥河湾哺乳动物群中没有发现过的。龙担哺乳动物群的地质时代可能比泥河湾者稍早,可能约2MaB.P.。其组成倾向于表明,当时该地区周围还有相当面积的森林覆盖。     

The Olduvai buffalo Pelorovis and the origin of Bos

The origin of the genus Bos is a debated issue. From ∼ 0.5 Ma until historic times, the genus is well known in the Eurasian large mammal assemblages, where it is represented by Bos primigenius. This species has a highly derived cranial anatomy that shows important morphological differences from other Plio-Pleistocene Eurasian genera of the tribe Bovini such as Leptobos, Bison, Proamphibos-Hemibos, and Bubalus. The oldest clear evidence of Bos is the skull fragment ASB-198-1 from the middle Pleistocene (∼ 0.6-0.8 Ma) site of Asbole (Lower Awash Valley, Ethiopia). The first appearance of Bos in Europe is at the site of Venosa-Notarchirico, Italy (∼ 0.5-0.6 Ma). Although the origin of Bos has traditionally been connected with Leptobos and Bison, after a detailed anatomical and morphometric study we propose here a different origin, connecting the middle Pleistocene Eurasian forms of B. primigenius with the African Late Pliocene and early Pleistocene large size member of the tribe Bovini Pelorovis sensu stricto. The dispersal of the Bos lineage in Western Europe during middle Pleistocene times seems to coincide with the arrival of the Acheulean tool technology in this continent.