Evolution of the European regional large mammals assemblages in the end of the Middle Pleistocene – The first half of the Late Pleistocene (MIS 6–MIS 4)
Graphical abstract
Introduction
A large volume of data on the large mammals – inhabitants of Europe at the end of Middle and the first half of the Late Pleistocene – have been accumulated by the European palaeontologists over the recent decades. The information should be generalized and interpreted so as to get a notion of the general pattern and regional characteristics of the ecosystem evolution in the western Eurasia on the background of wide fluctuations of the global climate at the end of Middle and Late Pleistocene: Dnieper (= Saale, Wolston, Riss, Pechora) Glaciation – Mikulino (= Eem, Ipswich) Interglacial – Valdai (= Vistula, Weichsel, Würm, Devens) Glaciation (Glückert, 1974; Rose, 1987; Velichko et al., 2005, 2011; 2012; Litt et al., 2007; Fiebig and Preusser, 2008; Gibbard et al., 2009; Ehlers et al., 2013; Astakhov et al., 2016).
There are many published papers dealing with the mammal fauna at the time of the Last Glacial Maximum (MIS 2) and the transition to the Holocene Interglacial (MIS 1), the period most important also for understanding the evolution of human populations (Markova et al., 2019; Pavelková Řičánková et al., 2015; ; Álvarez-Lao and Méndez, 2016; Crees et al., 2016; Fernández-García et al., 2016; Danukalova et al., 2018; Puzachenko, 2019; Kovalchuk et al., 2020). At the same time, we have not yet a comprehensive knowledge of the general regularities, nor regional specific features of the large mammal fauna evolution at the boundary of MIS 6 (~200/191–140/130 ka BP) MIS 5e (~130–115/111 ka BP) (Shackleton et al., 2003; Lisiecki and Raymo, 2005; Molodkov and Bolikhovskaya, 2006; Otvos, 2015). The same is true, even to a greater extent, of the fauna evolution from Eemian interglacial to the Weishel (= Valdai) Glaciation (MIS 5d–MIS 4). The evolutionary changes in the fauna composition and the ranges of individual species at the Middle/Late Pleistocene boundary might have a pronounced effect on the range and quantity of hominids (and first of all, Neanderthals), which used many species of that group (including members of megafauna) as an important source of food (Defleur and Desclaux, 2019).
In this work we included species of Proboscidea, Artiodactyla (= Cetartiodactyla), Perissodactyla, Carnivora (including small-sized mustelids) and Rodentia (Hystrix, Castor) orders into the “large mammal” group. The environmental and feeding preferences of that species group are highly variable. Most of them are herbivorous, and the proportion of carnivorous is much smaller. Many herbivores are known for gregarious habits, some of them make seasonal migrations. Several species (mammoths, woolly rhino, reindeer, cave lion, and some others) form the main body of the “Mammuthus–Coelodonta” fauna that developed at the end of Middle – the early stage of the Late Pleistocene and reached its maximum at the second part of the Late Pleistocene in Northern Eurasia (Kahlke, 2014).
Unlike small mammals (Markova and Puzachenko, this volume), a great number of megafauna species became extinct, in particular at the end of the Late Pleistocene – beginning of the Holocene. Changes in the spatial distribution of certain species (straight-tusked elephant, mammoths, rhinoceros, reindeer, fallow deer, roe deer, elk, saiga, beaver, and porcupines) are reliable indicators of environmental changes. The large mammal fauna response to the sharp rise of the global temperature in MIS 6–MIS 5, and in MIS 5e phase especially, is of particular interest.
Large mammals are known for a high diversity of their morphology, various environmental preferences and ethological diversity; all those characteristics determine their leading position in the Pleistocene ecosystem functioning. It was the extinction of megafauna that resulted in a radical transformation of the terrestrial ecosystems not only in Europe, but also all over Northern Eurasia. Similar, though probably not quite identical, processes took place also in the South-East of Asia and in North America at the end of the Pleistocene (Grayson, 2007; Wan and Zhang, 2017).
Numerous and diversified findings of large mammals make it possible to trace the changes in the species assemblages both in time (at a scale of the oxygen isotope stages – MIS) and in space – over large areas and individual regions of Europe. The abundance of data permits to use statistical analysis of the data for describing regularities of evolution, including its regional characteristics. In the present paper the statistical methods were applied to estimating changes in some parameters of the biological diversity of the large mammals at the end of Middle Pleistocene – the first half of the Late Pleistocene, the models of faunal assemblage evolution being developed for some regions. In this work we leaned on our previous experience in statistics application to the analysis of large massifs of palaeontological data available on the second half of the Late Pleistocene and the Holocene in Europe (Puzachenko and Markova, 2014, 2016; 2020; Puzachenko, 2019).
Our investigations were primarily aimed at estimating of general regularities in temporal changes in the large mammal fauna in Europe beginning from individual species and up to regional faunal assemblage. Multivariate analysis of a large array of palaeontological data was used to obtain descriptive models of evolution of regional mammal assemblages. We also studied changes of species richness and spatial (within regions) diversity of species distribution for the time intervals corresponding to MIS 6, MIS 5e, MIS 5d–MIS 5a, and MIS 4. We discussed the problem of the stability/instability of mammalian faunal complexes at the Middle Pleistocene – Early Late Pleistocene and at the end of the Late Pleistocene – Holocene, and, in conclusion, we gave the evidence of the hypothesis about relative temporal stability of European zoogeographic units (bioregions).
Section snippets
European bioregions
The European biogeographical zones used in this studies as in the previous ones (Markova and Puzachenko, 2018; Puzachenko and Markova, 2019; Puzachenko, 2019) are based on modern terrestrial mammal ranges (Spatial Data, IUCN Red List of Threatened Species: https://www.iucnredlist.org/resources/spatial-data-download). We used regular spatial grid cells (150 × 150 km) with lists of species and classified them that have been described in literature (Heikinheimo et al., 2007; Kreft and Jetz, 2010).
South European Mediterranean Iberian region
Two bioregions have been identified on the Iberian Peninsula, divided by a boundary running from east to west (Fig. 1A (I, II); Table 1). Here we combine the data on both regions for the purpose of constructing a better substantiated model of the faunal assemblage evolution over the entire MIS 6–MIS 4 interval. With this in mind, we have chosen 32 localities (Table S1). A general idea on the regional fauna diversity may be obtained from local faunas recovered from localities Ambrona, Bolomor,
Discussion
The paper presents the first attempt at the analysis of the evolution of the European regional faunal assemblages of mammals at the transition from the Middle Pleistocene to the Late Pleistocene (MIS 6–MIS4) (MLPT: Middle–Late Pleistocene Transition). We succeeded in constructing descriptive models of the evolution and finding general regularities in variations of relative frequency of occurrence of localities of species and species richness, reflecting changes in the European environments
Conclusions
For the first time, we examined a big array of palaeontological data on large mammals from a numerous number of sites and localities for the whole of Europe dated by the Dnieper (= Saale, Wolston, Riss, Pechora) glaciation (MIS 6), Mikulino (= Eem, Ipswich) interglacial (MIS 5e) and Early – Middle Valdai (= Weichsel, Würm, Devens) glaciation MIS 5d–MIS 4). Despite the different methods of dating and possible errors in dating of some sites and localities as well as differences in their
Declaration of competing interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Acknowledgements
This study financially supported by the Russian Foundation for Basic Research (grant numbers 18-05-00076-a (A. Puzachenko andV. Titiv) and 18-04-00982-a (P. Kosintsev)), and in part by the Federal theme of the Laboratory of Biogeography of Institute of Geography of RAS “Evaluation of physical and geographical, hydrological, and biotic environmental changes and their effects for the base of sustainable environmental management” (theme number 0148-2019-0007, АААА-А19-119021990093-8).
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