Dietary variation and overlap in Central and Northwest European Stephanorhinus kirchbergensis and S. hemitoechus (Rhinocerotidae, Mammalia) influenced by habitat diversity

To trace the dietary evolution of the two abundant Middle to Late Pleistocene rhinoceros species Stephanorhinus kirchbergensis and Stephanorhinus hemitoechus in Europe over several climatic cycles, we examined comprehensive material of stratigraphically well-defined alaeopopulations from different regions and interglacials. Using morphometrics and mesowear analysis, these reconstructions of Stephanorhinus diets indicate that habitat diversity and interspecific competition with closely related rhinoceros species induced variation in feeding behaviour. Although anatomical features of both species suggest significantly higher dietary specializations compared to the Early to early Middle Pleistocene Stephanorhinus hundsheimensis, their mesowear signals are characteristic of a mixed feeder diet, similar to that of extant mammal species in relatively open habitats. Both species retained a degree of dietary flexibility, enabling them to survive in a range of environments. Although each of these rhinoceroses preferred different habitats, species identity alone is not sufficient to establish the real dietary traits of a Stephanorhinus palaeopopulation. As a consequence, their occurrence in a faunal assemblage alone cannot be taken to indicate a specific habitat. S. kirchbergensis and S. hemitoechus were embedded in a dynamic process of temporo-spatial replacements and interspecific differentiation of rhinoceroses in the western Palaearctic. However, dietary specialization in these Middle to Late Pleistocene European rhinoceroses was not the result of a directed time-transgressive evolution. Rather, within the range of each species' ecological tolerance, it was controlled by environmental parameters, with habitat variability as the main factor

The woolly rhinoceros from Seweckenberge near Quedlinburg (Germany)

Bioarchaeolog

Skull Evolution Method and Analysis in The Rhinocerotidae: Phylogeny of Early Rhinocerotoids

After phylogeny is measurably disposed of, cranial elements utilized essentially for rumination ought to change most with hypsodonty (high-delegated cheek teeth). These structures should be least phylogenetically restricted. Corollary: structures with significant common ancestry will integrate more morphologically. All living rhinoceroses and many extinct European Plio-Pleistocene species We examined skull, mandible, and upper tooth row form in the dorsal, lateral, and occlusal perspectives using two-dimensional geometric morphometrics. Hypsodonty index was employed to represent eating behaviours. We divided form variation into function, phylogeny, and size using phylogenetically independent comparisons and variation partitioning. We used Escoufier's RV coefficient to evaluate morphological reconciliation. The mandible and upper tooth column covariate most with hypsodonty and least with phylogeny. Skull morphology corresponds least with hypsodonty and most with phylogeny. Low morphological joining between the top tooth line and different parts recommends it is the least phylogenetically restricted. As predicted, the chewing area is confined by function rather than phylogeny, unlike others

Les grands herbivores (Bovinés, Équidés, Rhinocérotidés, Proboscidiens) de la fin du Pléistocène Moyen : la couche 9 de Coudoulous II (Lot, Quercy, Sud-ouest France)

Les grands herbivores sont particulièrement bien représentés dans la couche 9 de la grotte de Coudoulous II (Quercy, Lot), en nombre d’espèces et nombre de restes. Les études morphologique et biométrique précisent les stades évolutifs de plusieurs taxons : Bos, Bison, Equus, Coelodonta, Stephanorhinus et Mammuthus. Les comparaisons effectuées, et l’analyse statistique, apportent des informations sur l’âge du dépôt, attribué à la toute fin du Pléistocène moyen. De plus, les données sur les tailles corporelles des taxons associées à celles sur la micro- et méso-usure dentaire précisent le paléoenvironnement climatique. L’ensemble permet de rendre compte d’une association faunique originale, marqueur d’une période de transition entre deux stades climatiques, entre le Pléistocène moyen (SIM 6) et supérieur (SIM 5), encore mal documentée dans le Quercy.The large Herbivores are particularly well represented in the level 9 of the Coudoulous II cave (Quercy, Lot) both in term of number of species and number of remains. The morphological and biometrical study precise the evolutionary degree of several taxa: Bos, Bison, Equus, Coelodonta, Stephanorhinus and Mammuthus. Comparative analyses, supported by statistical approach, allow to infer the chronology of the level, attributed to the extreme end of the Middle Pleistocene. Paleoenvironmental information’s are given thanks to complementary data such body-size of the species and micro-and meso-dental wear. The large ungulates of level 9 constitute an original faunal association which marks a transitional period between two climatic phases, between Middle (MIS 6) and Late (MIS 5) Pleistocene, period still poorly known in the Quercy region and beyond

Ancient and Modern Genomes Unravel the Evolutionary History of the Rhinoceros Family

Only five species of the once-diverse Rhinocerotidae remain, making the reconstruction of their evolutionary history a challenge to biologists since Darwin. We sequenced genomes from five rhinoceros species (three extinct and two living), which we compared to existing data from the remaining three living species and a range of outgroups. We identify an early divergence between extant African and Eurasian lineages, resolving a key debate regarding the phylogeny of extant rhinoceroses. This early Miocene (∼16 million years ago [mya]) split post-dates the land bridge formation between the Afro-Arabian and Eurasian landmasses. Our analyses also show that while rhinoceros genomes in general exhibit low levels of genome-wide diversity, heterozygosity is lowest and inbreeding is highest in the modern species. These results suggest that while low genetic diversity is a long-term feature of the family, it has been particularly exacerbated recently, likely reflecting recent anthropogenic-driven population declines. © 2021 The Authors.The authors acknowledge support from the Science for Life Laboratory, the Garvan Institute of Medical Research, the Knut and Alice Wallenberg Foundation, and the National Genomics Infrastructure funded by the Swedish Research Council and Uppsala Multidisciplinary Center for Advanced Computational Science for assistance with massively parallel sequencing and access to the UPPMAX computational infrastructure. We thank the Natural History Museum at the University of Oslo for providing the Javan rhinoceros sample. We thank the Museum of the Institute of Plant and Animal Ecology (UB RAS, Ekaterinburg) for providing the sample of Siberian unicorn. M.T.P.G. was supported by European Research Council (ERC) Consolidator grant 681396 (Extinction Genomics). E.D.L. was supported by Independent Research Fund Denmark grant 8021-00218B . A.C. was supported by an Australian Research Council Laureate Fellowship ( FL140100260 ). T.M.B. is supported by funding from the ERC under the European Union’s Horizon 2020 research and innovation program (grant agreement 864203 ), grant BFU2017-86471-P ( MINECO /FEDER, UE), “Unidad de Excelencia María de Maeztu” funded by the AEI ( CEX2018-000792-M ), Howard Hughes International Early Career, and Secretaria d’Universitats i Recerca and CERCA Programme del Departament d’Economia i Coneixement de la Generalitat de Catalunya ( GRC 2017 SGR 880 ). L.D. was supported by the Swedish Research Council ( 2017-04647 ) and Formas ( 2018-01640 ). We thank Dmitry Bogdanov and Roger Hall for giving us permission to use their rhinoceros artwork

Middle Pleistocene fauna and palaeoenvironment in the south of Eastern Europe: A case study of the Medzhybizh 1 locality (MIS 11, Ukraine)

The Middle Pleistocene was a period of dynamic changes in Europe. During MIS 11, a number of modern mammal taxa appeared and environmental conditions remained warm and favourable for a relatively long time. The Medzhybizh 1 locality of Ukraine dated to this very period comprises alluvial deposits with rich animal remains, which allow not only to reconstruct the fauna composition, but also to highlight the environmental conditions that dominated at this locality. A revision of the fauna of Medzhybizh 1 locality based on remains of all vertebrate groups revealed a taxonomically diverse fish community (16 species of 11 genera) dominated by cyprinids common for lacustrine or riverine assemblages. Amphibians are represented by 11 species, while the number of reptile and bird remains are less significant. Mammals are the most represented group at the locality, including small mammals (30 species), carnivorans (2 species), and ungulates (5 taxa), the latter dominated by C. elaphus. The taxonomic composition of terrestrial groups indicates temperate climate with boreal-type forests and meadows similar to cold steppe, as well as low wet areas and riparian habitats inhabited by amphibians, reptiles, insectivores, beavers, and various voles. The fish assemblage indicates a partially overgrown but well-aerated water body (lake or slow-flowing river) with sandy-silty bottom. Lithic artefacts found at the Medzhybizh 1 locality contribute to a better understanding of relationships between ancient hominins and faunas during the Middle Pleistocene of Eastern Europe.Archaeological field works at Medzhybizh were supported by the NASU state research projects 0105U001383 (2005–2009), 0109U008921 (2010–2014) and partly funded by the State Fund of Fundamental Research of Ukraine grant 0118U001457 (F77/50–2018). This research was also supported by the grant 0201/2048/18 ‘Life and death of extinct rhino (Stephanorhinus sp.) from Western Poland: a multi-proxy palaeoenvironmental approach’ financed from the funds of the National Science Centre, Poland. JvdM received support from the Spanish Ministerio de Ciencia, Inovación y Universidades (current grant numbers PGC2018-093925-B-C31 and PGC2018-095489-B-I00).Peer reviewe

Reframing the mammoth steppe: Insights from analysis of isotopic niches

Woolly mammoth (Mammuthus primigenius), horse (Equus spp.) and bison (Bison spp.) coexisted with a variety of mammalian megafauna across the Pleistocene mammoth steppe – a megacontinental ecosystem that spanned northern Eurasia and northwestern North America. Previous research has suggested that highly conserved niches with minimal niche overlap allowed high levels of species diversity on the mammoth steppe. Here we evaluate previously published and some new collagen carbon and nitrogen isotope data (δ13C, δ15N) for mammoth steppe megaherbivores using Stable Isotope Bayesian Ellipses in R (SIBER) and linear regression models to determine isotopic niches for individual species during broad time intervals (pre-, during and post-Last Glacial Maximum (LGM)) at multiple geographic regions across the mammoth steppe. Individual species maintained relatively consistent isotopic niche positions at different geographic locations and during different times. Diet and habitat niches for any given species appear to have been similar across the mammoth steppe. Between some regions and times, however, species' isotopic niches changed, suggesting adaptation to local climatic conditions and/or changes in the nitrogen isotope patterns at the base of the food web. Isotopic niche overlap, including at the level of core niche overlap (>60% overlap), was observed in at least one time and region for most species. This overlap suggests high levels of functional redundancy in the ecosystem, whereby one species could fulfil another's ecological role in the latter's absence. Despite spatial and temporal environmental variation, species' adaptability and functional redundancy within the ecosystem would have made the mammoth steppe a highly resilient ecosystem

The Great Fossil Mine of the southern North Sea: exploring the potential of submerged Palaeolithic archaeology

This research explores the potential of the submerged Palaeolithic archaeology of the southern North Sea for answering questions about how hominins occupied and adapted within their environments in these northerly latitudes throughout the Pleistocene. Recent coastal discoveries in East Anglia have demonstrated occupation as far back as ~1 million years, and yet our appreciation of the how, why and who of this occupation is missing a crucial piece of its puzzle; excluding these now-submerged landscapes is an active bias on our understanding, truncating the archaeological record. Having been subjected to repeated glaciations, trans- and regressions, the very processes that led to the terrestrial exposure of these areas have subsequently led to their neglect: the assumption that pre-LGM deposits will have been eroded or re-worked has prevailed. Recent work, however, has demonstrated the inaccuracy of this assumption, with evidence for extant Pleistocene-age deposits, landscape features and archaeology. Unlocking the clear potential of these submerged landscapes now relies on the approaches that we take to their investigation as, to-date, all archaeological finds have been entirely by chance. In order to move beyond this reactive style of archaeology, methodologies must be developed which tackle these areas in a more focused and reasoned way. The research undertaken throughout this PhD makes steps towards this. Starting from no baseline understanding of the nature of the existing resource, this work located, collated and analysed a prolific collection of 1,019 faunal specimens. Recovered by the 19th and 20th Century UK trawling industry, the development of historical methods has elucidated their locations and conditions of collection. Combining this locational information with species taxonomic evolution, the emergent spatio-temporal patterns provide a fresh understanding of the integrity of the extant deposits and unique opportunities for locating them on the seabed. These results are presented at a range of scales: • First, a broad-scale understanding of offshore regions across the southern North Sea which have demonstrated a dominance of cold-stage species from MIS 8-MIS 2. • Secondly, a local scale: linking faunal remains with seabed features in the near shore area off Happisburgh, identifying Early and early Middle Pleistocene assemblages related to exposures of the CFbF. • Finally, a discrete, high resolution area of seabed off the coast of Clacton has been identified. Through the collection of swath bathymetry, this area has shown the exciting correlation of Pleistocene seabed deposits and faunal remains. This research presents a significant move towards a proactive approach to these submerged landscapes and represents a step-change in our ability to understand, locate and engage with this undervalued archaeological resource. <br/