Siwalik synopsis: A long stratigraphic sequence for the Later Cenozoic of South Asia

The Tertiary sediments of northern Pakistan are an exceptional record of terrestrial sedimentation and represent most of Neogene time. Foremost is the Siwalik Group of the Potwar Plateau, for which multiple, superposed fossil levels span ∼18–6 Ma. Well-developed magnetostratigraphic control provides secure dating so that Siwalik fossil horizons may be interpolated into a time scale with resolution to 100,000 years. We describe the geographic setting of the Potwar, give an overview of the temporal distribution of faunas, and discuss changes in paleohabitat and paleoecology with coinciding faunal change, as seen from the Siwalik viewpoint. The long Siwalik biostratigraphy of many successive assemblages with its resolved time scale may be compared directly with other well-dated sequences. Immigrant arrival and timing of faunal change may be traced. The basins of the Iberian Peninsula show somewhat different timing of introduction of hipparionine horses, and faunal turnover in the Siwaliks clearly precedes the Vallesian crisis in Spain. In contrast to the increasingly seasonal precipitation of the late Miocene Potwar, the paleohabitat of coeval North China appears to have been moist and equable, with high diversity faunas. Continued development and comparison of resolved Neogene records allow increasing resolution of the patterns of faunal change on regional to global levels.     

Neogene–Quaternary Mammalian Paleobiogeography of the Indian Subcontinent: An appraisal

A biostratigraphic evaluation of the Indian subcontinent Neogene–Quaternary mammal genera at 1 Ma intervals reveals a strong correlation between faunal turnovers (first and last appearances and immigrations), diversity and major tectonic and climate-induced sea level and vegetation changes at 23–22, 20–18, 17–16, 15–14, 11–10, 9–8, 3–2 and 1–0 Ma. The Early Miocene collision of Afro-Arabian and Eurasian Plates, followed by the Middle Miocene Climate Optima and the presence of evergreen rainforests, most likely facilitated the dispersals into and out of the subcontinent from the west along the coast and included African proboscideans, primates, creodonts, thryonomyid rodents and Asian chalicotheres, rhinos, suids, anthracotheres, carnivores and muroid rodents. The Late Miocene and Early Pleistocene cooler and drier conditions, the spread of grasslands and sea level drops may have influenced the exchange of African monkeys, Eurasian carnivores, bovids, equids, camelid, and South Asian suids, leporids and murids. Immigrations from the east (East and SE Asia) occurred mostly during the Miocene and involved primarily small mammals.     

On the status of “Urotherium antiquum” (Ameghino) (Xenarthra,Glyptodontidae)

Within the Late Neogene Glyptodontidae of the Pampean region of Argentina, “Urotherium antiquum” was described on the basis of some associated osteoderms of the dorsal carapace, which in addition include a partial skull and left hemimandible. The diagnostic characters are located on the exposed surface of the osteoderms of the dorsal carapace which somewhat resembles those of the Pleistocene genus Neuryurus. Although the relationship of “U. antiquum” to the remaining Glyptodontidae has never been clarified, some cladistic analyses suggest a close phylogenetic affinity with the clade composed of Plohophorus + (Glyptodon + Doedicurus). A careful comparison of “U. antiquum” to well-characterized taxa with similar stratigraphic and geographic provenance reveals that no significant skull differences are observed from Plohophorus figuratus Ameghino. It seems likely that the alleged ornamentation pattern that characterizes this species was produced by a taphonomic alteration of the exposed surface of the osteoderms, a process broadly distributed in glyptodonts having a “rosette” ornamentation pattern. Finally, some characters present in the osteoderms of the type specimen of “U. antiquum” suggest that it may be a juvenile specimen. In summary, “U. antiquum” should be considered a junior synonym of P. figuratus.     

Step-wise change of Asian interior climate preceding the Eocene–Oligocene Transition (EOT)

Understanding the global climate change from greenhouse to icehouse conditions at the Eocene–Oligocene Transition (EOT) 34 million years ago requires climatic records from oceanic as well as continental realms of the key Late Eocene “doubthouse” period preceding this switch. Here, we report integrated stratigraphic results from well-dated Late Eocene continental mudflat to saline lake paleoenvironments of the Xining Basin (northeastern Tibetan Plateau, western China) recording regional and global change. Cyclostratigraphic analysis strongly suggests continuous dominance of the 41-kyr obliquity cycle in the whole late Eocene interval down to the base of polarity chron C18n.2n at 39 Ma with additional input of the ~ 100-kyr eccentricity cycle up to the base of chron C13r at ~ 34.7 Ma. This might imply that high-latitude climates dominated the area long before the EOT, probably related to incipient ice-volume fluctuations. Furthermore, our results reveal two paleoenvironmental deterioration steps preceding the Eocene–Oligocene Transition. The first step occurs in the top of chron C17n.1n at ~ 36.6 Ma. This age closely corresponds to (1) the high-altitude pollen appearance in chron C16.2r at ~ 36.4 Ma in the same section, (2) the recently dated final retreat of the Tarim Sea in western China, and (3) a shift from precession to obliquity dominance in the Atlantic Ocean. This near co-occurrence suggests global change at this time. We hypothesize this change is related to an increase in incipient ice sheet volume leading to passing threshold conditions for the high-altitude pollen appearance and Tarim Sea retreat, finally leading to decreased moisture availability in the Xining Basin. At the second step, in the base of chron C13r at ~ 34.7 Ma, a substantial increase in clastic sedimentation rates is observed. This might relate to increased climate variability preceding the greenhouse to icehouse transition at the EOT that prevented landscapes to attain equilibrium configurations.     

Holocene and Late Pleistocene climate in the sub-Mediterranean continental environment: A speleothem record from Poleva Cave (Southern Carpathians,Romania)

The PP10 stalagmite from Poleva Cave provides a Late Pleistocene and Holocene isotopic record characteristic for the SW of Romania, a sub-Mediterranean climatic region. The speleothem was dated by eight TIMS and one alpha U-series dates which showed that it was precipitated between ∼ 75 ka and ∼ 2 ka with at least two hiatuses. The basal sector of the stalagmite showed a slow-growing regime of ∼ 0.26 cm/ka, while the upper one grew relatively fast with about 5 cm/ka. The temporal resolution for the isotopic sampling is thus ∼ 2 ka/sample for the lower sector, and ∼ 150 years/sample for the upper one. The relationship between δ¹⁸O and temperature was found positive. The isotopic record of the lower sector shows two marked cold intervals during ∼ 67 and 58 ka and ∼ 40–35 ka, respectively, which correlate well with the Villars and Soreq records. The upper sector record is so far the most detailed Holocene isotopic record in Romania and the only one available for the regions located at the exterior of the Carpathians Range. The signal shows a gradual warming after the GS1 event punctuated by several cold events at ∼ 8, 7.2 and 4.2 ka and also by warm oscillations centered at about 5.2 and 3.3 ka. The results seem to indicate that if the North-Atlantic first-order signals may extend well to the south-eastern Europe, their amplitude and general trend may be diminished by the interferences with the Mediterranean circulation.     

Geopalaeontological setting,chronology and palaeoenvironmental evolution of the Baccinello-Cinigiano Basin continental successions (Late Miocene,Italy)

The Latest Miocene succession of the Baccinello-Cinigiano Basin in southern Tuscany (Italy) recorded a faunal turnover documenting the extinction of an older, insular, endemic faunal complex characterised by the extinct ape Oreopithecus bambolii and the setting of a new, continental, European faunal complex including the colobine monkey Mesopithecus. A similar turnover pattern (Late Miocene ape/Latest Miocene Cercopithecidae) is generally observed in Late Miocene continental successions of Eurasia, from Spain to central Europe, Southwest Europe, the near East, and Southwest Asia. Abundant literature reports that the Late Miocene Eurasian hominoid primate distribution closely tracks the climatic/environmental changes occurring during the 12–9 Ma interval, until their extinction in western Europe. In the primate record, the dispersion of Cercopithecidae and the contraction of hominids is interpreted as an event depicting a pattern of “continentalisation” in the Old World. The sedimentary succession of the Baccinello-Cinigiano basin, one of the longest continuous vertebrate-bearing continental successions in the Neogene Italian record, contributes to the debate on this hypothesis. This paper provides an overview of the main characteristics of the sedimentary succession, the chronological constraints (biochronology, radiometric datings, magnetostratigraphy), and the palaeoenvironmental evolution as derived from palaeobiological approaches and from the study of stable carbon and oxygen isotope contents along the entire sedimentary succession. The 2 myr geological history of the Baccinello Cinigiano Basin, which documents the evolutionary history of Oreopithecus and associated faunas, does not have a direct relation with the event of the Messinian Salinity Crisis. The evolutionary history of Baccinello-Cinigiano Basin and its palaeontological record have been mainly driven by the regional tectonism and palaeogeographic changes that affected the northern Tyrrhenian regions in Late Miocene (Latest Tortonian–Messinian) times.     

Paleoenvironmental reconstruction of the Early Pleistocene site of Quibas (SE Spain) using a rodent assemblage

In this paper we analyze the rodent assemblage from the Early Pleistocene site of Gruta1 (Quibas karstic complex, Murcia, SE Spain), providing taphonomic comments and paleoecological and paleoclimatic data. The studied assemblage includes 209 identified rodent teeth corresponding to at least four taxa. The karstic source of the site and the low presence of digested remains suggest that the accumulation is scatological in origin, with the influence of predators such as owls; the accumulation also shows certain evidence of slight hydrodynamic sorting. For the area around Quibas/Gruta1 the paleoecological study indicates a predominance of rocky areas (31.7%), forested environments (31.7%) and open dry meadows or shrublands (29.7%), which would indicate a mosaic forest environment, and to a lesser extent the presence of open humid land (5.4%) and areas along streams or ponds (1.5%). The distribution of the bioclimatic spectra yields the highest percentage for a Mediterranean climate (37.5%), while the climatic parameters calculated (MAT = 15 °C, MTW = 25.7 °C, MTC = 4.3 °C, MAP = 390 mm) suggest colder and slightly more humid conditions than today at the time of the deposition of the remains. The data also point to a relative decrease in temperature within the phase in which the site was produced, which possibly occurred during a relative cold period between MIS 36 and MIS 40 according to the age.     

The open-air Paleolithic site of Mirak,northern edge of the Iranian Central Desert (Semnan,Iran): Evidence of repeated human occupations during the late Pleistocene

The northern edge of the Iranian Central Desert has provided valuable evidence of terminal Pleistocene human settlements. Mirak constitutes one of the largest open-air lithic scatters in the region, consisting of eight natural mounds. Fieldwork was initiated in 2015 by the joint Iranian-French program at Mirak 8. Preliminary results have demonstrated at least three successive phases of human occupation during the MIS3: an upper layer with clear Upper Paleolithic affinities and a maximum age of 28 ky, a lower layer with clear Middle Paleolithic affinities that dates around 47 ky, and an intermediate layer with mixed characteristics that can be seen as an intermediate Paleolithic phase which dates between 28 ± 2 and 38 ± 2 ky. At the time when Upper Paleolithic cultures originated in the Zagros Mountains, cultures with clear Middle Paleolithic affinities persisted nearby along the northern edge of the Iranian Central Plateau.     

Early human occupations at the westernmost tip of Eurasia: The lithic industries from Menez–Dregan I (Plouhinec,Finistère,France)

The stratigraphical complex preserved at the site of Menez–Dregan I (Brittany, France) displays an alternating sequence of 17 occupation levels and of 4 marine deposits, between ca. 450 and 150,000 years (stratigraphical units 9a and 3b). The lithic industry retrieved at the site is extremely abundant, with more than 153,000 artefacts over 3 cm in length, as well as millions of knapping debris, and corresponds to a regional facies of the Acheulean, with heavy-duty tools essentially represented by choppers, as well as a few handaxes and cleavers. This paper presents a first synthesis of the data produced for almost 30 years on the site of Menez–Dregan I. The typological study of the entire lithic collection is now complete and allows us to assess the different human occupations that occurred at this site during the Lower Palaeolithic.     

Pulleniatina Minimum Events in the Andaman Sea (NE Indian Ocean): Implications for winter monsoon and thermocline changes

The late Quaternary record of Pulleniatina obliquiloculata was investigated from two well dated sediment cores from the Andaman Sea (NE Indian Ocean) to examine its paleoceanographic significance and the presence of the Pulleniatina Minimum Events (PME) in the western North Pacific. As in the Pacific, our study shows that PMEs exist in the Indian Ocean albeit with a lower intensity. The Holocene PME occurs between 4.5 and 3.0 cal ka BP with a considerable reduction in P. obliquiloculata abundance, and which matches well with the Pacific records influenced by the Kuroshio Current. Additionally, two significant minimum events of P. obliquiloculata are also seen during the Younger Dryas (YD) and late Last Glacial Maximum (LGM, 20–18 cal ka BP). Overall, the PMEs of the Andaman Sea are not current driven events like in the western Pacific margin either by the weakening of the Kuroshio Current or reduced winter SSTs. The PMEs of the Andaman Sea are characterised by lower abundances of thermocline species indicating the increased depth of the thermocline (DOT) and reduced winter SSTs mainly during the minimum events of the YD and late LGM. The high SSTs during the Holocene PME make this event a mystery. However, the presence of PMEs in the Andaman Sea suggests that these events are not confined to areas influenced by the Kuroshio Current but may be responding to a broad scale oceanographic–climatic process or mechanism which needs to be explored with a detailed study.     

Using climatic suitability thresholds to identify past,present and future population viability

Often climatic niche models predict that any change in climatic conditions will impact species abundance or distribution. However, the accuracy of models that just incorporate climatic information to predict future species habitat use is widely debated. Alternatively, environmental conditions may simply need to be above some minimum threshold of climatic suitability, at which point, other factors drive population size. Using the example of nesting sites of loggerhead sea turtles (Caretta caretta) in the Mediterranean (n = 105), we developed climatic niche models to examine whether a climatic suitability threshold could be identified as a climatic indicator in order for large populations of a widespread species to exist. We then assessed the climatic suitability of sites above and below this threshold in the past (∼1900) and future (∼2100). Most large sites that are currently above the climatic threshold were above the threshold in the past and future, particularly when future nesting seasonality shifted to start 1–2 months earlier. Our analyses highlight the importance of future phenological shifts for maintaining suitability. Our results provide a positive outlook for sea turtle conservation, suggesting that climatic conditions may remain suitable in the future at sites that currently support large nesting populations. Our study also provides an alternative way of interpreting the outputs of climatic niche models, by generating a threshold as an index of a minimum climatic suitability required to sustain large populations. This type of approach offers the possibility to benefit from information provided by climate-driven models, while reducing their inherent uncertainties.     

Miocene bristlemouths (Teleostei: Stomiiformes: Gonostomatidae) from the Makrilia Formation,Ierapetra, Crete

Bristlemouths of the genus Cyclothone are currently regarded as the most abundant vertebrates on Earth. The fossil record seems to suggest that these fishes diversified during the Miocene in the Pacific Ocean, but there is no evidence of their presence in the Miocene of the Atlantic Ocean and Mediterranean basin. A new bristlemouth, Cyclothone gaudanti sp. nov. (Teleostei, Stomiiformes, Gonostomatidae), is described herein based on 16 specimens from the Upper Miocene Makrilia Formation (late Tortonian of Crete, Greece). The small sized species is characterized by light pigmentation, 30–31 (14–15 + 15–16) vertebrae, dorsal fin with 10–13 rays, anal fin with 10–14 rays, premaxilla bearing seven closely spaced teeth, maxilla with 42–55 teeth, epipleurals, and autogenous parhypural. The presence of epipleurals appears to be unique of this Miocene species, and the re-establishment of this ancestral character state may be possibly interpreted as related to a phylogenetic character reversal. Morphological and paleoecological considerations suggest that this species possibly inhabited the upper mesopelagic layer, at depths ranging from 2–300 and 500 meters.     

A multi-proxy reconstruction and comparison of Holocene palaeoenvironmental changes in the Alvor and Alcantarilha estuaries (southern Portugal)

The present study investigates the history of two estuaries on the Algarve coast, southern Portugal, through a multi-proxy approach that links sedimentology, geochemistry, palynology, microfaunal analyses and radiocarbon dating. These analyses provide an overview of the development of these estuaries over the last ～7500 years. Palynological data reveal climate-driven vegetational shifts over the whole period. Microfauna, which is composed predominantly of foraminifers and ostracods, provides evidence for periods of marine and brackish water conditions, observed in both estuaries. Whereas the sediment record from the Alvor estuary shows a clear, nearly continuous silting-up sequence, the development of the Alcantarilha estuary is characterised by fluvial sedimentation, replaced by a marine environment at about 7300 yr cal BP. Marine transgression is indicated by marine-brackish foraminifera and ostracod assemblages, while upper-marsh pollen decreased and wetland communities increased at the same time. A continuous shallowing of the estuaries followed between ～6700 yr cal BP and ～5700 yr cal BP in Alvor and 4500 yr cal BP in Alcantarilha. This was interrupted by a distinct high-energy event, possibly a storm or tsunami, between 6400 and 5800 yr cal BP in Alvor; the event was not recorded in the Alcantarilha estuary. Afterwards, the silting-up restarted with clear similarities to the pre-event situation and continued until ～3500/2000 yr cal BP. This process was influenced by the build-up of sand barriers in front of the estuaries. This is also documented by a higher percentage of upper-marsh vegetation. The core from Alcantarilha estuary shows another layer of high-energy deposits, with mainly marine foraminifers and ostracods after 4500 yr cal BP. The youngest parts of the two records are characterised by decreasing tidal influence and increasing anthropogenic impact, indicated by increasing values of cultivated plants and other human-induced changes of vegetation cover.     

Differentiating between isolates of Vibrio vulnificus with monoclonal antibodies

Monoclonal antibodies (MAbs) against Vibrio vulnificus (isolate I, VVC and isolate II, VVB) were raised using heat-killed and heat-killed plus SDS–mercaptoethanol treated forms of VVC and VVB for immunizing Swiss mice. Twenty three hybridomas producing MAbs against V. vulnificus were selected and divided into five groups according to their specificities to different V. vulnificus isolates and apparent protein antigens which ranged from ∼ 3–50 kDa. Four groups were specific to V. vulnificus without cross reactivity to either other Vibrio spp. or other bacterial species. In dot blot based assays, one group of MAbs were specific to VVC, with a sensitivity of ∼ 1.6 × 10⁷ CFU ml^− 1 (∼ 1.6 × 10⁴ cells spot^− 1), and bound to proteins of ∼ 50 and ∼ 39 kDa. Other MAbs, binding to proteins ranging from ∼ 3–14 and ∼ 40 kDa, detected VVB (but not VVC) with high sensitivity at ∼ 1.6 × 10⁵ and 4 × 10⁶ CFU ml^− 1 (∼ 1.6 × 10² and 4 × 10³ cells spot^− 1), respectively. In addition, certain MAbs were able to recognize V. vulnificus in tissues by means of immunohistochemistry. The remaining groups demonstrated cross reactivity to Vibrio fluvialis. MAbs from this study can, therefore, detect the difference between some isolates of V. vulnificus and in addition to pathogen detection may, with further antibodies, form the basis of serovar typing isolates in the future.     

Holocene paleoclimate change in the Antarctic Peninsula: evidence from the diatom,sedimentary and geochemical record

Holocene, marine deposition in Lallemand Fjord, Antarctic Peninsula, is reinterpreted using statistical analyses (cluster analysis, analysis of variance, nonmetric multidimensional scaling and multiple regression) to compare diatom assemblages and the primary sedimentological proxies. The assemblages have been deposited in a variable sea ice zone over the last ca. 10,500 yr BP in response to a climate change. In the Late Pleistocene/early Holocene (10,580–7890 yr BP), a sea ice diatom assemblage was deposited in the presence of a retreating ice shelf at the head of the fjord. In the mid Holocene (7890–3850 yr BP), an open water assemblage was deposited and sea ice cover was at a minimum. We associate the assemblage with climatic warming, which characterizes much of the Antarctic Peninsula during this time. A second sea ice assemblage, different from that deposited in the early Holocene, has been deposited in Lallemand Fjord since the late Holocene (<3850 yr BP). The assemblage reflects Neoglacial cooling, an increase in sea ice extent and/or an advance of the Müller Ice Shelf.     

Vegetation history of the savanna corridor between the Guinean and the Congolian rain forest during the last 150,000 years

Pollen and spores from a deep-sea core located west of the Niger Delta record an uninterrupted area of lowland rain forest in West Africa from Guinea to Cameroon during the last Interglacial and the early Holocene. During other periods of the last 150 ka, a savanna corridor between the western — Guinean — and the eastern — Congolian — part of the African lowland rain forest existed. This so-called Dahomey Gap had its largest extension during Glacial Stages 6, 4, 3, and 2. Reduced surface salinity in the eastern Gulf of Guinea as recorded by dinoflagellate cysts indicates sufficient precipitation for extensive forest growth during Stages 5 and 1. The large modern extension of dry forest and savanna in West Africa cannot be solely explained by climatic factors. Mangrove expansion in and west of the Niger Delta was largest during the phases of sea-level rise of Stages 5 and 1. During Stages 6, 4, 3, and 2, shelf areas were exposed and the area of the mangrove swamps was minimal.     

Distribution of large Emiliania huxleyi in the Central and Northeast Atlantic as a tracer of surface ocean dynamics during the last 25,000 years

The coccolithophore species Emiliania huxleyi is characterized by a wide range of sizes, which can be easily distinguished in the light microscope. In this study we have quantified the abundance of large (coccoliths > 4 µm in maximum length) E. huxleyi specimens during the last 25 kyr in sedimentary records from eleven cores and drill sites in the NE Atlantic and W Mediterranean Sea, to prove its usefulness in the reconstruction of water mass dynamics and biostratigraphic potential.During the Last Glacial Maximum this large form, a cold-water indicator, was common in the NE Atlantic and Mediterranean, and its regional variation in abundance indicates a displacement of the climatic zones southwards in agreement with the development of ice sheets and sea ice in the Northern Hemisphere during this period. On the other hand, the gradient between northern and southern surface water masses in the Subtropical Gyre appears to have been more pronounced than at present, while the Portugal and Canary Currents were more intense. In the western Mediterranean basin temperatures were cooler than in the adjacent Atlantic, provoking a quasi-endemism of these specimens until the end of Heinrich Event 1. This may have been due to a restriction in the communication between the Atlantic and Mediterranean through the Strait of Gibraltar, the arrival of cold surface water and the amplification of cooling after the development of ice sheets in the Northern Hemisphere.During the deglaciation, large E. huxleyi specimens decreased in abundance at medium and low latitudes, but were still numerous close to the Subarctic region during the Holocene. In transitional waters this decrease to present day abundances occurred after Termination Ib.The abrupt change in abundance of this large E. huxleyi form is proposed as a new biostratigraphic event to characterize the Holocene in mid- to low-latitude water masses in the North Atlantic, although this horizon seems to be diachronous by 5 kyr from tropical to subarctic regions, in agreement with the gradual onset of warm conditions.     

A new whale barnacle from the early Pleistocene of Italy suggests an ancient right whale breeding ground in the Mediterranean

The fragmentary fossil history of whale barnacles (Cirripedia: Coronulidae) is mainly constituted by remains of Coronula spp. in Plio-Pleistocene deposits found along extant humpback whale migration routes, and especially in occurrence of breeding areas. Here we report the presence of a single compartment of Cetopirus along with remains of Coronula bifida in Lower Pleistocene open shelf deposits of Salento (South Italy). This is the first occurrence of the genus Cetopirus prior to the Late Glacial period (ca. 15–10 ky before Present), and the specimen here studied is designated as the holotype of the new fossil species Cetopirus fragilis. Since Cetopirus is currently found exclusively inhabiting the skin of the right whales (Eubalaena spp.), the fossil material here studied suggests the persistence of Balaenidae in the Mediterranean Basin during the Early Pleistocene and supports the existence of a baleen whale migratory route active between the central Mediterranean and the North Atlantic during the Plio-Pleistocene.