- Email: [email protected]
The maximum geographic extension of Late Pleistocene Mammuthus primigenius (Proboscidea, Mammalia) and its limiting factors
Quaternary International 379 (2015) 147e154
Contents lists available at ScienceDirect
Quaternary International journal homepage: www.elsevier.com/locate/quaint
The maximum geographic extension of Late Pleistocene Mammuthus primigenius (Proboscidea, Mammalia) and its limiting factors* Ralf-Dietrich Kahlke Senckenberg Research Institutes and Natural History Museums, Research Station of Quaternary Palaeontology, Am Jakobskirchhof 4, 99423 Weimar, Germany
a r t i c l e i n f o
a b s t r a c t
Article history: Available online 8 May 2015
The maximum geographic extension of fully developed woolly mammoth, Mammuthus primigenius, occurred during the last glacial period (MIS 5de2), i.e. during the 110e14 ka interval. The mapping of the species' widest distribution reveals principal barriers to its expansion that were acting independent of short term environmental fluctuations. The maximum spread of M. primigenius was primarily controlled by the following trans-regional factors: (A) The configuration of inland glaciers; (B) The configuration of high mountain chains; (C) The configuration of semi-deserts and deserts; (D) The configuration of marine shorelines at year-round open water surfaces; (E) The exposure of continental shelf regions; and (F) The replacement of tundra-steppe by extended grasslands. Some of the biogeographic barriers were climateindependent; others were determined by long- or medium-term climatic processes. The effect of Eurasian and North American intra-continental steppes as obstacles to migration has not yet been fully elucidated. The verifiable area of the Late Pleistocene distribution range of M. primigenius, based on geomorphological data, comprises approximately 33,301,000 km2 (surface area). When taking the geographical distribution of a species as a measure of its success, the Late Pleistocene woolly mammoth was one of the most successful large herbivores of the Holarctic Mammoth Fauna. © 2015 Elsevier Ltd and INQUA. All rights reserved.
Keywords: Mammuthus primigenius Maximum geographic extension Biogeographic barriers Late Pleistocene
1. Introduction The origin and distribution of individual species reflects, in a very clear way, the spatial and temporal variation of environmental r conditions (Ba anescu and Bos¸caiu, 1978; Lomolino and Heaney, 2004 and references therein). The barriers restricting the spread of a species are determined by their mode of life, as well as the physiological performance of single individuals in relation to the prevailing environmental conditions (Cox and Moore, 2010 and references therein). The determination of the limiting factors in the distribution of fossil species can be essential in understanding the dynamics of palaeoecosystems as well contributing to the elucidation of the causes underlying local or global extinction of individual species or whole faunas. The history of the development of cold adapted large mammal associations of the northern hemisphere during the Pleistocene
* This paper is dedicated to Larry D. Agenbroad (y), a colleague and friend. We miss him. E-mail addresses: [email protected], [email protected].
http://dx.doi.org/10.1016/j.quaint.2015.03.023 1040-6182/© 2015 Elsevier Ltd and INQUA. All rights reserved.
was, to a great extent, determined by the increasing aridity of significant parts of Eurasia, through rhythmical global cooling with extended and intensified cold phases, as well as increasing continentality (Kahlke, 2014). Between 2.6 Ma and 700 ka, the continental Asian steppe regions and the circumpolar tundra belt were biogeographically separated, each inhabited with its own, largely independent mammalian faunas (Kahlke, 1999, 2014). At about 900 ka, the transformation of the global climate cyclicity from a 41 ky to a clear 100 ky periodicity was completed (Lisiecki and Raymo, 2005). Correspondingly, prolonged climatic cycles, which appeared as extended glacial and interglacial phases, caused drastic changes in the faunal structure of the Northern Hemisphere (Kahlke et al., 2011). During the pronounced cold period of MIS 12, Arctic tundra species regularly expanded south- and southwestward into a newly formed type of biome, the so-called tundrasteppe. In parallel, species originating from the inner continental (Asian) steppe dispersed into new habitats north and northwest of their ancestral distribution areas. Such drastic faunal turnover led to the formation of the earliest Palaearctic Mammoth Fauna at around 460 ka (Kahlke and Lacombat, 2008; Kahlke, 2014). This new type of mammal assemblage, or its individual faunal elements,
148
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
expanded and contracted its geographic extension parallel to alternating environmental conditions of the subsequent late Middle to Late Pleistocene (Kahlke, 1999). During the last glacial period Mammuthus primigenius, the characteristic species of Pleistocene cold adapted faunas, reached its farthest spatial extent (Kahlke, 1999). With the help of a reconstruction of its distribution range based on the current state of knowledge, principal limiting factors of its extension can be detected and with them the maximum range of the species in the northern hemisphere clarified. 2. Holarctic mammoth faunas and the emergence of Mammuthus primigenius The aridisation of intra-continental regions and the duration and intensity of individual cold periods drove the evolution and dispersal of the genus Mammuthus. Mammuthus trogontherii as the proboscidean key species of the Palaearctic Mammoth Faunas, originated during the Early Pleistocene, between the Olduvai and Jaramillo subchrones, in continental areas north of the HimalayaneTibetan uplift (Wei et al., 2006). In the period around 1.2 Ma, a significant expansion event of M. trogontherii into the north of Asia is assumed to have occurred, as recorded by finds from the northeastern Siberian Early Olyorian (1.2e0.8 Ma; Lister and Sher, 2001). More or less at the same time, at the beginning of the Epivillafranchian, M. trogontherii also appeared in the Western Palaearctic for the first time. The species established itself there as a characteristic faunal element of cool and cold periods until the Middle Pleistocene (Lister et al., 2005; Kahlke et al., 2011; Kahlke, 2014). An even longer persistence of evolved trogontherii-populations until the Late Pleistocene is plausible for the Central Asian region, especially in northern China (Wei et al., 2010; Tong, 2010a, 2010b). Moreover, it cannot be excluded that Early Pleistocene forms on the level of M. trogontherii could have spread to the Nearctic and established the foundation of the Mammuthus columbi lineage (e.g. Lister and Bahn, 2007; Wei et al., 2010). The Columbian Mammoth (sensu lato, details in Agenbroad, 2005) dominated large areas of North America until the Late Pleistocene. Between 800 and 600 ka (Late Olyorian) in northeastern Eurasia, indigenous M. trogontherii started to transform into more advanced mammoth forms, mainly recognisable from the increase in the number of enamel lamellae in the dental elements (Lister et al., 2005). Continuous cold conditions, which probably fluctuated less on an annual and interannual basis than in inner continental regions of Eurasia, in combination with Far Northern vegetation phenology, caused the speciation of M. primigenius. The primigenius stage was already widespread in NE Asia from 400 ka onwards (Lister and Sher, 2001). In Europe, corresponding forms first occurred much later, during the late MIS 7 or at the beginning of MIS 6, 200e160 ka. The fossil record indicates an oscillating movement of synchronous western Palaearctic M. trogontherii and M. primigenius populations (Lister et al., 2005). During the Late Pleistocene, M. primigenius reached northern North America (Agenbroad, 2005), where already during the last (Sangamonian) interglacial unglaciated areas were successfully occupied (Harington, 2011, Tab. 1). Recorded finds show that occasionally woolly mammoths and Columbian mammoths co-occurred in one and the same site (e.g. Agenbroad, 1994), indicating temporal replacement or short term (perhaps seasonal) sympatry. 3. Maximum geographic extension of Late Pleistocene Mammuthus primigenius The maximum distribution of fully developed woolly mammoths occurred during the last glacial period (¼Devensian ¼
Weichselian ¼ Valdaian ¼ Dalian ¼ Wisconsinian) and spans the marine isotope stages (MIS) 5de2, i.e. approximately the interval between 110 and 14 ka (Lisiecki and Raymo, 2005). In addition to a number of overviews and summaries (Graham and Lundelius, 1994; Kahlke, 1999; Kuzmin et al., 2003, 2008; Agenbroad, 2005; AlvarezLao et al., 2009; Markova et al., 2013, 2014), the mapping of the last glacial maximum spread of M. primigenius is primarily based on regional studies. Particular emphasis is placed on data from areas close to the maximum distribution limits of the species, to derive principle limiting factors. Individual verifications of M. primigenius have significant regional variations in frequency. In addition to biogeographic conditions, these also arise from differences in regional research intensity. The Iberian Peninsula constituted the most southwesterly part of the last glacial Eurasian woolly mammoths' distribution range. Recent studies have established that the species was not only in the northern Spanish Basque country, but also in Portugal, in the Madrid region, and Granada (Alvarez-Lao, 2007; Alvarez-Lao et al., 2009; Alvarez-Lao and García, 2011). In the Pyrenees, areas of the northern flank were populated (Clot and Duranthon, 1990). Evidence of M. primigenius has also been found in almost all parts of France (e.g. Delpech, 1983; Fosse et al., 2014), in Ireland and Britain (Stuart, 1977, 1982, 1983; Stuart and Wijngaarden-Bakker, 1985; Lister, 2009), on Jersey (Scott et al., 2014), in the region of the Netherlands, as well as in the southern part of the North Sea (van Essen and Mol, 1996; Mol et al., 1999, 2006a, 2008; Kahlke, 2001). There are also numerous documented finds in southern Scandinavia, including Finland and in the Baltic states (AarisSørensen et al., 1990; Aaris-Sørensen, 1998, 2009; Ukkonen et al., 1999, 2011; Markova et al., 2013). In Central Europe, remains of M. primigenius are common occurrences in the fossil record (Kowalski, 1959; Musil, 1968, 1985; €ppes and Rabeder, 1997; Germonpre , 2003; Hünermann, 1985; Do Wojtal, 2007; Sommer and Benecke, 2009; Nadachowski et al., 2011; Ukkonen et al., 2011; Gaar and Preusser, 2012; Pawłowska, 2014). On the Apennine peninsula, the animals ventured into southern Italy (Sala et al., 1992; Biddittu and Celletti, 2001; Rustioni et al., 2003). To the southeast, the Pannonian Basin and the Balkan nossy and Vo € ro €s, 1979; Peninsula were occupied (Rakovec, 1975; Ja € ro € s, 1981; Malez, 1986; Konra d et al., 2010; Katona Koufos, 1981; Vo et al., 2012; Mauch Lenardi c, 2012). In Eastern Europe, the species ranged from the Russian Plain and Ukraine, up into the southern part of the Kola Peninsula (Strelkov, 1977), and to the north coast of the Black Sea (Markova et al., 2013). In the Crimea, the lowlands as well as northern foothills and low mountains of the Crimea mountain range were settled (Baryshnikov, 2003). The situation was similar in the margins of the northern foothills of the Greater Caucasus range (Baryshnikov, 2003). Occasionally, M. primigenius penetrated even into Transcaucasia, as sporadic evidence as far as Tbilisi has shown (Gabunia, 1952; Vereshchagin, 1959). To the east, the Caspian Sea as well as areas in north-west to north-eastern Kazakhstan were reached (Kozhamkulova, 1969; Zhylkibaev, 1975). The animals also expanded into the area between the Balkhash Lake and the Dzungarian Alatau, around present-day Almaty and the foothills of the Transili-Alatau. At higher latitudes, there is widespread evidence for their occupation in present-day Bashkortostan and the Urals (Dubrovo, 1966; Kuzmina, 1971; Danukalova and Yakovlev, 2001; Markova et al., 2013). Western Siberia is rich in Late Pleistocene mammoth evidence too (Alekseeva, 1980; Kuzmin et al., 2003, 2008; Markova et al., 2013). The species has also been found in the Xinjiang Uygur Region in westernmost China (Tong, 2004, p. 311) and in the northern part of the Mongol UIs (records in the collections of the Museum of Natural History Ulaanbaatar). The East Siberian region
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
to Transbaikalia was also a part of the last glacial distribution area of the woolly mammoth (Ermolova, 1978; Imetkhenov and Kalmykov, 1988; Kuzmin et al., 2008; Alexeeva and Erbajeva, 2010; Markova et al., 2013). Northeast China, especially the provinces of Heilongjiang and Jilin, has provided numerous pieces of evidence for M. primigenius (Liu and Li, 1989; Jin and Kawamura, 1996). The southernmost distribution of woolly mammoths in East Asia extended to the coastal region of the Sea of Bohai (compilation in Alvarez-Lao et al., 2009, p. 66). A confirmed molar of M. primigenius has been recorded from Ji'nan at the Beidasha River (Shandong province) at 36 350 N. According to Takahashi et al. (2007), this find represents the world's most southerly record of the species. On the Korean peninsula evidence of M. primigenius has been found as far as the Ham Kyung province (Park, 1988) and in the Jungwon area in the northernmost part of South Korea at c. 37 500 N (Cho and Woo, 2004). Through the Russian Far East (e.g. Kuzmin et al., 2000), woolly mammoths reached as far as Sakhalin at 46 N (Matsumoto, 1937; Kuzmin et al., 2005) and to Hokkaido at 42 N (Dubrovo, 1981; Kamei, 1981; Takahashi et al., 2006). The detection of M. primigenius from the Sea of Japan off western Shimane Prefecture, 35 200 N, is taken to be a consequence of drifting from the Huang He area of mainland China (Kamei, 1990; Takahashi et al., 2006). The Siberian Arctic and the continental shelf of the Arctic Ocean, including today's New Siberian Islands were also part of the heartland of the last glacial mammoth distribution (e.g. Vereshchagin and Tikhonov, 1990; Sulerzhitsky and Romanenko, 1997; Kuzmina et al., 1999; MacPhee et al., 2002; Kahlke and Mol, 2005; Mol et al., 2006b; Kuznetsova, 2007; Kuzmin et al., 2008; Lazarev, 2008; Vartanyan et al., 2008; Markova et al., 2013). The Kamchatka peninsula was also reached (Kuzmin et al., 2000; Orlova et al., 2000). The distribution area expanded continuously as far as eastern Beringia and beyond (Agenbroad and Barton, 1991; Harington, 2003, 2011; Kuzmin et al., 2008). There are existing records of woolly mammoth occupation from the Pribilof Islands (Yesner et al., 2005) and from the western part of the Canadian Arctic Archipelago (MacPhee, 2007; Kuzmin et al., 2008). In addition to Alaska, numerous remains of M. primigenius are commonly found in northern Canada (Harington, 2003). In North America, the animals have been established in many sites in the Great Lakes region and across the Northern Plains (Agenbroad, 1985, 1989, 2005; Agenbroad et al., 1994; Burns, 1996; Kuzmin et al., 2008; Harington et al., 2011; Widga et al., 2014). Along the U.S. West Coast, they ventured to around 45 N (Fisher in Larmer, 2013). Easternmost evidence for M. primigenius is documented in Coastal Maine (Hoyle et al., 2004), Nova Scotia (Cooke et al., 1993), as well as on the Atlantic continental shelf of Virginia (Harington et al., 2011). 4. Principal biogeographic barriers The mapping of the last glacial maximum distribution of M. primigenius, inclusive of all territorial variations during this period documented by fossil records, reveals principal biogeographical barriers that are acting independently of short term environmental fluctuations. Thus, the configuration of the northern distribution limit of woolly mammoths very clearly reflects the position of the great inland glaciers (Fig. 1A, arrows). In particular, the Scandinavian (-West Siberian) ice sheet in Eurasia and the Laurentide and the Cordilleran ice sheets in North America (Ehlers and Gibbard, 2004a, 2004b, 2004c; Ehlers et al., 2011) were significant barriers to expansion. However, the two major New World glacial regions were only connected during the Last Glacial Maximum (LGM) (Dyke et al., 2002) and thus only briefly formed an effective single, continent-wide migration barrier. In most cases,
149
an ice-free corridor between the two ice sheets enabled the invasion of woolly mammoths to the southeast (see section 3). In other regions, advances of M. primigenius also trace deglaciated areas, as in the circum-Baltic region of Northern Europe (Ukkonen et al., 2011), on the Taimyr Peninsula of Arctic Siberia (Alexanderson et al., 2002; Kahlke and Mol, 2005, p. 50ff.), and in the western periphery of the Canadian Arctic Archipelago (MacPhee, 2007). High mountain chains with no passable valleys were another very effective barrier to migration (Fig. 1B, arrows). From the distribution of finds, this is particularly evident in the Pyrenees area, the Crimea mountain range and the Greater Caucasus. While M. primigenius could, in every case, colonise the (northern) flank of the mountain ranges facing the distribution areas, up to considerable altitudes, the ridges remained insurmountable. In different regions the high mountains were circumvented, as in the colonisation of the Iberian Peninsula and Transcaucasia. In Central Asia, a lowland corridor between the Altai and Dzungarian Alatau mountain ranges enabled the mammoths to advance from eastern Kazakhstan to China. The mountain ranges of the Alps and Rocky Mountains were only partially settled. Here, mammoths spread out mainly along broad valleys. Inside Eurasia, between Transcaucasia and northern China, the southern border of the last glacial distribution area of M. primigenius was very much shaped by the arrangement of semideserts and deserts (Fig. 1C, arrows). The mammoths recorded south of the Caucasus (see Section 3) are likely to have immigrated through the Kuban area (Krasnodar region), not via the arid areas of Azerbaijan. East of the Caspian Sea, the Kyzyl Kum desert and the semiarid area of the Betpak Dala (Hunger Steppe) limited the mammoths' range. The distribution patterns of Mongolian and northern Chinese M. primigenius also reflect the animals' response to extreme aridity. The desert regions of the Taklamakan and Gobi deserts were strictly avoided. Marine shorelines with year-round, ice-free water surfaces represented a more insurmountable obstacle to expansion for M. primigenius. In regions with near-coastal continental slope, the last glacial maximum distribution of the animals can be fairly accurately estimated (Fig. 1D, arrows). Thus, certainly the last glacial distribution area of the mammoths was directly limited by the majority of the French Mediterranean coast, the northern coast of the Black Sea, as well as some sections of the Pacific North West coast, especially in the Russian Far East at the Sea of Okhotsk and Kamchatka, and the west coast of North America. More difficult to assess is the mammoths' spread within present-day flooded continental shelf regions (Fig. 1E, arrows), which over the course of the Last Glacial eustatic sea level decrease of 120e135 m were exposed (Clark and Mix, 2002). In several regions the bathymetric situation opened up access routes to new expansion areas, as made evident by a M. primigenius migration route via the coastal part of the Cantabrian Sea into the Iberian Peninsula. Moreover, another temporary migration route is likely to have existed directly east of the Pyrenees. Here, a coastal part of the Mediterranean dried out during the extended coldest periods of the Late Pleistocene (Alvarez-Lao et al., 2009). The drop in sea level in the Mediterranean also caused large-scale drying up of northern parts of the Adriatic Sea (Gallini and Sala, 2001; Rustioni et al., 2003), so that an unbroken distribution area for M. primigenius developed between SE Europe and the Apennine Peninsula. Extensive finds-based knowledge accounting for mammoth distribution from the present-day flooded shelf areas exists only from the southern part of the North Sea between England and the Netherlands (see Section 3). Individual finds prove the presence of the animals on the shelf of the Arctic Ocean to the north of the New Siberian Islands and on the Atlantic shelf off the U.S. East Coast (see
150
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
Fig. 1. Localisation of principal barriers (arrows) of Late Pleistocene Mammuthus primigenius spread: A e ice sheets; B e high mountain chains; C e semi-deserts and deserts; D e year round open water surfaces; E e continental shelf regions; F e other types of grassland.
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
151
Fig. 1. (continued)
Section 3). A land bridge enabled the colonization of Sakhalin in the North Pacific (Kirillova and Tesakov, 2008). Evidence from the New Siberian Islands and Wrangel, as well as from the Pribilof Islands and the Canadian Arctic Archipelago (see Section 3) allow a conservative extrapolation of last glacial spread of M. primigenius in the region of today's Laptev- and East Siberian Sea and the Chuchi- and Bering Sea (Fig. 1E). New finds here could lead to significant increases in the areas which can be demonstrably shown to support woolly mammoth populations. In contrast to the aforementioned, comparatively welldocumented control factors (Fig. 1AeE), the effect of the intracontinental steppes of Eurasia and North America as migration barriers for M. primigenius has been little investigated. In recent times, it has become clear that sister taxa of several species involved in Eurasian Mammoth Faunas underwent separate evolution in Central Asia, thus indicating significant ecological differences between the Asian core steppe and Eurasian tundra-steppe habitats (Kahlke, 2014). Independent of the successively evolving M. primigenius of northern altitudes, M. trogontherii survived in Central Asia until the Late Pleistocene. A corresponding phenomenon of coexisting mammoth species, adapted to intra-continental steppe and tundra steppe habitats respectively, is known from North America. Here, M. columbi, which in many respects resembles M. trogontherii, occupied southerly extended grassland habitats, whereas M. primigenius synchronously spread into the mammoth steppe. It remains unclear whether the replacement of vegetation, the competition for food from the steppe mammoths (M. trogontherii and M. columbi respectively), or a combination of both prevented the successful spread of M. primigenius into intra-
continental steppe regions of Eurasia and North America (Fig. 1F, arrows). 5. Conclusions Based on the current fossil record, Fig. 2 shows the geographic extension of Late Pleistocene M. primigenius. The maximum spread of the species was primarily controlled by the following transregional factors: (A) (B) (C) (D)
The configuration of inland glaciers; The configuration of high mountain chains; The configuration of semi-deserts and deserts; The configuration of marine shorelines at year-round open water surfaces; (E) The exposure of continental shelf regions; (F) The replacement of tundra-steppe by extended grasslands (Central Asian core steppe, North American grasslands).
Locally acting factors could have modified the configuration of the distribution limits slightly here and there. The synopsis of the principal barriers to migration leads to an almost complete explanation of the Holarctic maximum spread of M. primigenius during the Late Pleistocene. AeD constituted more or less abrupt obstacles to expansion, which were not physiologically possible for individual mammoths to overcome. Some of the barriers mentioned were climate-independent (B, partly D), and others were determined by long- or medium-term climatic processes (A, C, E, F, partly D). In the case of F, the as yet still poorly
152
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
Fig. 2. Maximum extension of Mammuthus primigenius during the Late Pleistocene based on the current fossil record.
investigated replacement of tundra-steppe habitats by intracontinental grasslands would probably have had more of a gradual effect in limiting expansion. To date, the verifiable area of the Late Pleistocene distribution range of M. primigenius (Fig. 2) comprises approximately 33,089,000 km2 (status 09.20.2014; calculated by using Esri ArcGIS (C), surface volume tool). Taking into account the geomorphological data (based on Esri topographic map, raster resolution of 30 arc seconds) results in an actual mammoth-population surface area of approximately 33,301,000 km2, which is considered as the minimum of the actual distribution area. Gains in knowledge and thus increases in the known distribution area are to be expected, especially from the currently submerged shelf regions. To a lesser extent this is also possible on the mainland, especially in inaccessible or little studied regions. Despite the limiting factors mentioned, woolly mammoths enjoyed an enormous geographic extension during the Late Pleistocene. When we look at the geographical distribution of a species as a measure of its success, Late Pleistocene M. primigenius was one of the most successful large herbivores of the Holarctic Mammoth Fauna. Only the morphologically quite diverse Bison priscus group occupied a comparably large, and in many places identical, distribution area during the last glacial period (Kahlke, 1999, Fig. 20; Shapiro et al., 2004 and references therein). The woolly mammoth rightly proved itself as a palaeoecological characteristic species and a worthy namesake for the most cold-adapted large mammal assemblages in Earth's history, the Mammoth Fauna. Acknowledgements The study was conducted within the Senckenberg Research Institute's project group “Origin, dispersal and impoverishment of Eurasian cold adapted faunas” (RDK). Thanks go to E. Haase and G. Utschig (both Weimar) for their assistance in designing the graphics. For information and support on various topics we are grateful to B. Fiedler (Weimar; references), T. Hermsen (Offenbach; area calculations und graphics), C. Hertler (Frankfurt/M.; logistics), I. Kirillova (Moscow; literature), D. Mol (Hoofddorp; information on mammoth sites) and C. Nielsen-Marsh (Leipzig; language). The kind comments of E. Tsoukala (Thessaloniki) and an anonymous reviewer are greatly appreciated. References Aaris-Sørensen, K., 1998. Danmarks forhistoriske dyreverden. Om skovelefanter, næsehorn, bisoner, urokser, mammutter og kæmpehjorte. Gyldendal, Denmark.
Aaris-Sørensen, K., 2009. Diversity and dynamics of the mammalian fauna in Denmark throughout the last glacial-interglacial cycle, 115-0 kyr BP. Fossils and Strata 57, 1e59. Aaris-Sørensen, K., Petersen, K.S., Tuber, H., 1990. Danish finds of Mammoth (Mammuthus primigenius (Blumenbach)). Stratigraphical position, dating and evidence of Late Pleistocene environment. Danmarks Geologiske Undersøgelse, DGU Serie B 14, 1e44. Agenbroad, L.D., 1985. The distribution and chronology of Mammoth in the New World. Acta Zoologica Fennica 170, 221e224. Agenbroad, L.D., 1989. New World mammoth distribution. In: Martin, P.S., Klein, R.G. (Eds.), Quaternary Extinctions. A Prehistoric Revolution. University of Arizona Press, Tucson, pp. 90e108. Agenbroad, L.D., 1994. Geology, hydrology, and excavation of the site. In: Agenbroad, L.D., Mead, J.I. (Eds.), The Hot Springs Mammoth Site: A Decade of Field and Laboratory Research in Paleontology, Geology, and Paleoecology. Fenske, Rapid City, SD, pp. 15e27. Agenbroad, L.D., 2005. North American Proboscideans: Mammoths: The state of Knowledge, 2003. Quaternary International 126e128, 73e92. Agenbroad, L.D., Barton, B.R., 1991. North American mammoths: An annotated bibliography 1940-1990. Mammoth Site of Hot Springs Scientific Papers 2, 1e118. Agenbroad, L.D., Lister, A.M., Mol, D., Roth, V.L., 1994. Mammuthus primigenius remains from the Mammoth Site of Hot Springs, South Dakota. In: Agenbroad, L.D., Mead, J.I. (Eds.), The Hot Springs Mammoth Site: A Decade of Field and Laboratory Research in Paleontology, Geology, and Paleoecology. Fenske, Rapid City, pp. 269e281. Alekseeva, E.V., 1980. Mlekopitayushchie plejstocena yugo-vostoka Zapadnoj Sibiri. Nauka, Moskva (in Russian). € ller, P., Antonov, O., Eriksson, S., Alexanderson, H., Adrielsson, L., Hjort, C., Mo Pavlov, M., 2002. Depositional history of the North Taymyr ice-marginal zone, Siberia e a landsystem approach. Journal of Quaternary Science 17, 361e382. Alexeeva, N., Erbajeva, M., 2010. Habitat and distribution area of woolly mammoth and some core mammal species in Transbaikalia during the Late Pleistocene. In: Lacombat, F., Mol, D. (Eds.), The Vth International Conference on Mammoths rie 3, 177e178. and Their Relatives. Quaternaire, Hors-se n paleontolo gica de los macromamíferos indicadores Alvarez-Lao, D.J., 2007. Revisio rica (Ph.D. thesis). Universidad de Oviedo, de clima frío en la Península Ibe Departamento de Geología, Oviedo, Spain. Alvarez-Lao, D.J., García, N., 2011. Geographical distribution of Pleistocene coldadapted large mammal faunas in the Iberian Peninsula. Quaternary International 233, 159e170. Alvarez-Lao, D.J., Kahlke, R.-D., García, N., Mol, D., 2009. The Padul mammoth finds e On the southernmost record of Mammuthus primigenius in Europe and its southern spread during the Late Pleistocene. Palaeogeography, Palaeoclimatology, Palaeoecology 278, 57e70. ra nescu, P., Bos¸caiu, N., 1978. Biogeographie. Fauna und Flora der Erde und ihre Ba geschichtliche Entwicklung. Gustav Fischer, Jena. Baryshnikov, G., 2003. Mammuthus primigenius from the Crimea and the Caucasus. In: Reumer, J.W.F., de Vos, J., Mol, D. (Eds.), Advances in Mammoth Research. Proceedings of the Second International Mammoth Conference, Rotterdam, 1620 May 1999. Deinsea 9, 41e56. Biddittu, I., Celletti, P., 2001. Plio-Pleistocene Proboscidea and Lower Palaeolithic bone industry of southern Latium (Italy). In: Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), The World of Elephants. Proceedings of the 1st International Congress, Roma, 16-20 Ottobre 2001. Consiglio Nazionale delle Ricerche, Roma, pp. 91e96. Burns, J.A., 1996. Vertebrate paleontology and the alleged ice-free corridor: the meat of the matter. Quaternary International 32, 107e112. historique et la subsistence de la re gion Cho, T.-S., Woo, J.-Y., 2004. La faune pre e. In: Lee, Y.-J., Cho, T.-S. (Eds.), Paleoenvironnement et culture Jungwon en Core
R.-D. Kahlke / Quaternary International 379 (2015) 147e154 e. Institute for Jungwon Culture, paleolithique de la region Jungwon, Core Chungbuk National University, Seoul, pp. 89e99. Clark, P.U., Mix, A.C., 2002. Ice sheets and sea level of the Last Glacial Maximum. Quaternary Science Reviews 21, 1e7. res fossiles du Quaternaire dans les Clot, A., Duranthon, F., 1990. Les mammife ne es. Museum d’Histoire Naturelle de Toulouse, Toulouse. Pyre Cooke, H.B.S., Harington, C.R., Sollows, J.D., 1993. Undescribed mammoth (Mammuthus) teeth from Georges Bank and Nova Scotia. Proceedings of the Nova Scotian Institute of Science 40, 19e28. Cox, C.B., Moore, P.D., 2010. Biogeography. An Ecological and Evolutionary Approach. Wiley, Hoboken. Danukalova, G.A., Yakovlev, A.G., 2001. Finds of Proboscidean remains in the territory of the Southern Urals region. In: Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), The World of Elephants. Proceedings of the 1st International Congress, Roma, 16-20 Ottobre 2001. Consiglio Nazionale delle Ricerche, Roma, pp. 201e204. olithique supe rieur dans le Sud-Ouest de la Delpech, F., 1983. Les faunes du Pale France. Cahiers du Quaternaire 6. Editions du C.N.R.S., Paris. € €ppes, D., Rabeder, G. (Eds.), 1997. Plioz€ €ne Faunen Osterreichs. Do ane und pleistoza Ein Katalog der wichtigsten Fossilfundstellen und ihrer Faunen (Endbericht €rderung der wissendes Forschungsprojektes Nr. 9320 des “Fonds zur Fo schaftlichen Forschung”). Mitteilungen der Kommission für Quart€ arforschung € der Osterreichischen. Akademie der Wissenschaften Band 10. Verlag der € Osterreichischen Akademie der Wissenschaften, Wien. Dubrovo, I.A., 1966. Plejstocenovye mlekopitayushchie Sredgeo i Yushogo Urala i ikh stratigrafichekoye znachenie. Nauchnye Trudy Politechnicheskogo Instituta, Sbornik 20, 12e135 (in Russian). €ontologie 4, 49e84. Dubrovo, I.A., 1981. Die fossilen Elefanten Japans. Quart€ arpala Dyke, A.S., Andrews, J.T., Clark, P.U., England, J.H., Miller, G.H., Shaw, J., Veillette, J.J., 2002. The Laurentide and Innuitian ice sheets during the Last Glacial Maximum. Quaternary Science Reviews 21, 9e31. Ehlers, J., Gibbard, P.L. (Eds.), 2004a. Quaternary Glaciations e Extent and Chronology. Part I: Europe. Developments in Quaternary Science, vol. 2. Elsevier, Amsterdam. Ehlers, J., Gibbard, P.L. (Eds.), 2004b. Quaternary Glaciations e Extent and Chronology. Part II: North America. Developments in Quaternary Science, vol. 2. Elsevier, Amsterdam. Ehlers, J., Gibbard, P.L. (Eds.), 2004c. Quaternary Glaciations e Extent and Chronology. Part III: South America, Asia, Africa, Australasia, Antarctica. Developments in Quaternary Science, vol. 2. Elsevier, Amsterdam. Ehlers, J., Gibbard, P.L., Hughes, P.D. (Eds.), 2011. Quaternary Glaciations e Extent and Chronology. A Closer Look. Developments in Quaternary Science, vol. 15. Elsevier, Amsterdam. Ermolova, N.M., 1978. Teriofauna doliny Angary v pozdnem antropogene. Izdatelstvo Nauka, Sibirskoe otdelenie, Novosibirsk (in Russian). van Essen, H., Mol, D., 1996. Plio-Pleistocene proboscideans from the southern bight of the North Sea and the Eastern Scheldt, The Netherlands. In: Shoshani, J., Tassy, P. (Eds.), The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives. Oxford University Press, Oxford, New York, Tokyo, pp. 214e224. gut-Bonnoure, E., Fleury, G., Fourvel, J.B., Madeleine, S., Fosse, P., Fritz, C., Cre re, L., Rivero, O., Tosello, G., 2014. Woolly mammoths (MamSauvet, G., Aurie muthus primigenius) in southern France during the Late Palaeolithic: a geochronological assessment based on the palaeontological, rock art and portable art records. In: Kostopoulos, D.S., Vlachos, E., Tsoukala, E. (Eds.), Abstract Book of the VIth International Conference on Mammoths and their Relatives. Scientific Annals of the School of Geology, Aristotle University of Thessaloniki Special volume 102, 62. Gaar, D., Preusser, F., 2012. Luminescence dating of mammoth remains from northern Switzerland. Quaternary Geochronology 10, 257e263. Gabunia, L.K., 1952. O zube “Karlikovogo” mamonta iz Gori. Soobshcheniya Akademii Nauk Gruzinskoj SSR 13, 153e154 (in Russian). Gallini, V., Sala, B., 2001. Settepolesini di Bondeno (Ferrara e Eastern Po Valley): the first example of mammoth steppe in Italy. In: Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), The World of Elephants. Proceedings of the 1st International Congress, Roma, 16-20 Ottobre 2001. Consiglio Nazionale delle Ricerche, Roma, pp. 272e275. , M., 2003. Mammoth taphonomy of two fluvial sites from the Flemish Germonpre Valley, Belgium. In: Reumer, J.W.F., de Vos, J., Mol, D. (Eds.), Advances in Mammoth Research, Proceedings of the Second International Mammoth Conference, Rotterdam, May 16-20 1999. Deinsea 9, 171e183. Graham, R.W., Lundelius Jr., E.L., Faunmap Working Group, 1994. Faunmap: A Database Documenting Late Quaternary Distributions of Mammal Species in the United States. Illinois State Museum Scientific Papers 25(1), viii þ 1-287, 25 (2), vii þ 289e690. Harington, C.R. (Ed.), 2003. Annotated Bibliography of Quaternary Vertebrates of Northern North America e With Radiocarbon Dates. Toronto University Press, Toronto, Buffalo, London. Harington, C.R., 2011. Pleistocene vertebrates of the Yukon Territory. Quaternary Science Reviews 30, 2341e2354. Harington, C.R., Mol, D., van der Plicht, J., 2011. The Muirkirk Mammoth: A Late Pleistocene woolly mammoth (Mammuthus primigenius) skeleton from southern Ontario, Canada. Quaternary International 255, 106e113. Hoyle, B.G., Fisher, D.C., Borns Jr., H.W., Churchill-Dickson, L.L., Dorion, C.C., Weddle, T.K., 2004. Late Pleistocene mammoth remains from Coastal Maine, USA. Quaternary Research 61, 277e288.
153
€ugetiere aus dem Kanton Zürich. Hünermann, K.A., 1985. Eiszeit-Sa Vierteljahresschrift der Naturforschenden Gesellschaft in Zürich 130, 229e250. Imetkhenov, A.B., Kalmykov, N.P., 1988. Fauna i flora kajnozoya Buryatii. Buryatskij filial SO AN SSSR, Ulan-Ude (in Russian). nossy, D., Vo € ro €s, I., 1979. Gross€ Ja auger-Streufunde aus dem Pleistoz€ an Ungarns. Fragmenta Mineralogica et Palaeontologica 9, 21e60. Jin, C.-Z., Kawamura, Y., 1996. Late Pleistocene mammal fauna in Northeast China: Mammal fauna including woolly mammoth and woolly rhinoceros in association with Paleolithic tools. Earth Science (Chikyu Kagaku) 50, 315e330. Kahlke, R.-D., 1999. The History of the Origin, Evolution and Dispersal of the Late Pleistocene Mammuthus-Coelodonta Faunal Complex in Eurasia (Large Mammals). Fenske Companies, Rapid City. Kahlke, R.-D., 2001. Ein Meer voller Knochen? Pleistoz€ ane Wirbeltierreste aus der Scheldemündung und vom Nordseeboden. Natur und Museum 131, 417e432. Kahlke, R.-D., 2014. The origin of Eurasian Mammoth Faunas (Mammuthus-Coelodonta faunal complex). Quaternary Science Reviews 96, 32e49. Kahlke, R.-D., Lacombat, F., 2008. The earliest immigration of woolly rhinoceros (Coelodonta tologoijensis, Rhinocerotidae, Mammalia) into Europe and its adaptive evolution in Palaearctic cold stage mammal faunas. Quaternary Science Reviews 27, 1951e1961. €ugetiere der Sibirischen Arktis. Die Kahlke, R.-D., Mol, D., 2005. Eiszeitliche Großsa Cerpolex/Mammuthus-Expeditionen auf Tajmyr. Schweizerbart, Stuttgart. Kahlke, R.-D., García, N., Kostopoulos, D.S., Lacombat, F., Lister, A.M., Mazza, P.P.A., Spassov, N., Titov, V.V., 2011. Western Palaearctic palaeoenvironmental conditions during the Early and early Middle Pleistocene inferred from large mammal communities, and implications for hominin dispersal in Europe. Quaternary Science Reviews 30, 1368e1395. Kamei, T., 1981. Faunal succession of Pleistocene Mammals in the Japanese Islands: €rpala €ontologie 4, 165e174. An aspect. Quarta Kamei, T., 1990. The Japan Sea and Elephants. The Quaternary Research 29, 163e172 (in Japanese, with English summary). g mamKatona, L., Kov acs, J., Kordos, L., Szappanos, B., Linkai, I., 2012. The Csaja moths (Mammuthus primigenius): Late Pleniglacial finds from Hungary and their chronological significance. Quaternary International 255, 130e138. Kirillova, I.A., Tesakov, A., 2008. New mammalian elements of the Ice Age assemblage of the Sakhalin Island. Mammal Study 33, 87e92. cs, J., Hal Konr ad, G., Kova asz, A., Sebe, K., P alffy, H., 2010. Late Quaternary woolly mammoth (Mammuthus primigenius Blum) remains from southern Transdanubia, Hungary. Comptes Rendus Palevol 9, 47e54. Koufos, G., 1981. A New Late Pleistocene (Würmian) Mammal Locality from the Basin of Drama (Northern Greece). Scientific Annals of the Faculty of Physics and Mathematics, University of Thessaloniki 21, 129e148. w plejstocenu Polski. Panstwowe Wydawnictwo Kowalski, K., 1959. Katalog ssako Nauk. Warszawa, Wrocław. Kozhamkulova, B.S., 1969. Antropogenovaya iskopaemaya teriofauna Kazakhstana. Nauka, Alma-Ata (in Russian). Kuzmin, Y.V., Orlova, L.A., Zol'nikov, I.D., Igol'nikov, A.E., 2000. The history of mammoth (Mammuthus primigenius Blum.) population in Siberia and adjacent areas (according to radiocarbon data). Russian Geology and Geophysics 41, 723e730. Kuzmin, Y.V., Orlova, L.A., Zolnikov, I.D., 2003. Dynamics of the mammoth (Mammuthus primigenius) population in Northern Asia: radiocarbon evidence. In: Reumer, J.W.F., Vos, J., de, Mol, D. (Eds.), Advances in Mammoth Research, Proceedings of the Second International Mammoth Conference, Rotterdam, May 16-20 1999. Deinsea 9, 221e237. Kuzmin, Y.V., Gorbunov, S.V., Orlova, L.A., Vasilevsky, A.A., Alekseeva, E.V., Tikhonov, A.N., Kirillova, I.V., Burr, G.S., 2005. 14C Dating of the Late-Pleistocene Faunal Remains from Sakhalin Island (Russian Far East). Current Research in the Pleistocene 22, 78e80. Kuzmin, Y.V., Orlova, L.A., Dementiev, V.N., 2008. Dynamics of Mammoth (Mammuthus primigenius Blum.) Populations of Asia and North America and Its Correlation with Climatic Changes in the Late Neopleistocene (45,000e9700 Years BP). Doklady Earth Sciences 421, 978e982. Kuzmina, I.E., 1971. Formirovanie teriofauny severnogo Urala v pozdnem antropogene. Trudy Zoologicheskogo Instituta AN SSSR 49, 44e122 (in Russian). Kuzmina, S., Kuznetsova, T., Sher, A., 1999. Paleontological research on the Bykovsky Peninsula. Reports on Polar Research 315, 179e190. Kuznetsova, T., 2007. Paleontological collection of the “Mammoth” fauna from the museum of the Lena Delta Reserve. Reports on Polar and Marine Research 550, 135e140. Larmer, B., 2013. Of Mammoths and Man. National Geographic 223, 44e63. Lazarev, P.A., 2008. Kupnye mlekopitayushchie antropogena Yakutii. Nauka, Novosibirsk. Lisiecki, L.E., Raymo, M.E., 2005. A Pliocene-Pleistocene stack of 57 globally distributed benthic d18O records. Paleoceanography 20, PA1003. http:// dx.doi.org/10.1029/2004PA001071. Lister, A.M., 2009. Late-glacial mammoth skeletons (Mammuthus primigenius) from Condover (Shropshire, UK): anatomy, pathology, taphonomy and chronological significance. Geological Journal 44, 447e479. Lister, A.M., Bahn, P., 2007. Mammoths. Giants of the Ice Age. University of California Press, Berkeley, Los Angeles. Lister, A.M., Sher, A.V., 2001. The Origin and Evolution of the Woolly Mammoth. Science 294, 1094e1097. Lister, A.M., Sher, A.V., van Essen, H., Wei, G., 2005. The pattern and process of mammoth evolution in Eurasia. Quaternary International 126e128, 49e64.
154
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
Liu, T.-S., Li, X.-G., 1989. Mammoths in China. In: Martin, P.S., Klein, R.G. (Eds.), Quaternary Extinctions. A Prehistoric Revolution. University of Arizona Press, Tucson, pp. 517e527. Lomolino, M.V., Heaney, L.R. (Eds.), 2004. Frontiers of Biogeography. New Directions in the Geography of Nature. Sinauer, Sunderland. MacPhee, R.D.E., 2007. Mammoths in the insular Nearctic? Some constrains on the existence of a Pleistocene megafaunal refugium in the Canadian Arctic Archipelago. Quaternary International 169e170, 29e38. MacPhee, R.D.E., Tikhonov, A.N., Mol, D., de Marliave, C., van der Plicht, H., Greenwood, A.D., Flemming, C., Agenbroad, L., 2002. Radiocarbon Chronologies and Extinction Dynamics of the Late Quaternary Mammalian Megafauna of the Taimyr Peninsula, Russian Federation. Journal of Archaeological Science 29, 1017e1042. €ren Vertebraten-Faunen in der SFR Jugoslawien. Malez, M., 1986. Die quarta €ontologie 6, 101e117. Quart€ arpala Markova, A.K., Puzachenko, A. Yu, van Kolfschoten, T., van der Plicht, J., Ponomarev, D.V., 2013. New data on changes in the European distribution of the mammoth and the woolly rhinoceros during the second half of the Late Pleistocene and the early Holocene. Quaternary International 292, 4e14. Markova, A., Puzachenko, A., van Kolfschoten, T., van der Plicht, J., Kuznezova, T., Tikhonov, A., Ponomarev, D., 2014. The Eurasian mammoth distribution during the second half of the Late Pleistocene and the Holocene. In: Kostopoulos, D.S., Vlachos, E., Tsoukala, E. (Eds.), Abstract Book of the VIth International Conference on Mammoths and Their Relatives. Scientific Annals of the School of Geology, Aristotle University of Thessaloniki Special Volume 102, 118e119. Matsumoto, H., 1937. Two Species of Mammoths from off Odomari and Vicinity, Karafuto (Saghalin). Zoological Magazine 49, 9e11 (in Japanese, with English summary). Mauch Lenardi c, J., 2012. Miocene to Late Pleistocene proboscideans of Croatia. Quaternary International 276e277, 120e128. Mol, D., van den Bergh, G.D., de Vos, J., 1999. Fossil proboscideans from The Netherlands, the North Sea and the Oosterschelde Estuary. In: Haynes, G., Klimovicz, J., Reumer, J.W.F. (Eds.), Mammoths and the Mammoth Fauna: Studies of an Extinct Ecosystem. Deinsea 6, 119e145. Mol, D., Post, K., Reumer, J.W.F., van der Plicht, J., de Vos, J., van Geel, B., van Reenen, G., Pals, J.P., Glimmerveen, J., 2006a. The Eurogeul e first report of the palaeontological, palynological and archaeological investigations of this part of the North Sea. Quaternary International 142e142, 178e185. Mol, D., Tikhonov, A., van der Plicht, J., Kahlke, R.-D., Debruyne, R., van Geel, B., van Reenen, G., Pals, J.P., de Marliave, C., Reumer, J.W.F., 2006b. Results of the CERPOLEX/Mammuthus expeditions on the Taimyr Peninsula, Arctic Siberia, Russian Federation. Quaternary International 142e143, 186e202. Mol, D., de Vos, J., Bakker, R., van Geel, B., Glimmerveen, J., van der Plicht, H., Post, K., 2008. Kleine encyclopedie van het leven in het Pleistoceen. Mammoeten, neushoorns en andere dieren van de Noordzeebodem. Veen Magazines B.V., Diemen. €ontologische Musil, R., 1968. Die Mammutmolaren von Predmostí (CSSR). Pala Abhandlungen, A 3, 1e192. Musil, R., 1985. Paleobiography of terrestrial communities in Europe during the Last rodniho Muzea v Praze, B 41, 1e83. Glacial. Sbornik Na Nadachowski, A., Lipecki, G., Wojtal, P., Mie˛ kina, B., 2011. Radiocarbon chronology of woolly mammoth (Mammuthus primigenius) from Poland. Quaternary International 245, 186e192. Orlova, L.A., Kuzmin, Y.V., Zolnikov, I.D., 2000. Time-space systematics for mammoths (Mammuthus primigenius Blum.) and prehistoric humans in Siberia (on the basis of radiocarbon dating). Archaeology, Ethnology & Anthropology of Eurasia 3, 31e41. Park, S.J., 1988. The Palaeoenvironmental Changes and Macromammal Evolution during the Pleistocene in East Asia. The Korean Journal of Quaternary Research 2, 51e85. Pawłowska, K., 2014. Elephantids from the Pleistocene of Poland: state of knowledge. In: Kostopoulos, D.S., Vlachos, E., Tsoukala, E. (Eds.), Abstract Book of the VIth International Conference on Mammoths and Their Relatives. Scientific Annals of the School of Geology, Aristotle University of Thessaloniki Special volume 102, 152. Rakovec, I., 1975. Razvoj kvartarne sesalske favne Sloveniji. Slovenska Akademija Znanosti in Umetnosti, Razred za Zgodovinske in Druzbene Vede, Cl. I, Hist. Sociol. 24, pp. 225e270. Rustioni, M., Ferretti, M.P., Mazza, P., Pavia, M., Varola, A., 2003. The vertebrate fauna from Cardamone (Apulia, southern Italy): an example of Mediterranean mammoth fauna. In: Reumer, J.W.F., de Vos, J., Mol, D. (Eds.), Advances in Mammoth Research. Proceedings of the Second International Mammoth Conference, Rotterdam, 16-20 May 1999. Deinsea 9, 395e403. Sala, B., Masini, F., Ficcarelli, G., Rook, L., Torre, S., 1992. Mammal dispersal events in the Middle and Late Pleistocene of Italy and Western Europe. In: von Koenigswald, W., Werdelin, L. (Eds.), Mammalian Migration and Dispersal Events in the European Quaternary. Courier Forschungsinstitut Senckenberg 153, 59e68. Scott, B., Bates, M., Bates, R., Conneller, C., Pope, M., Shaw, A., Smith, G., 2014. A new view from La Cotte de St Brelade, Jersey. Antiquity 88, 13e29. Shapiro, B., Drummond, A.J., Rambaut, A., Wilson, M.C., Matheus, P.E., Sher, A.V., Pybus, O.G., Gilbert, M.T.P., Barnes, I., Binladen, J., Willerslev, E., Hansen, A.J.,
Baryshnikov, G.F., Burns, J.A., Davydov, S., Driver, J.C., Froese, D.G., Harington, C.R., Keddie, G., Kosintsev, P., Kunz, M.L., Martin, L.D., Stephenson, R.O., Storer, J., Tedford, R., Zimov, S., Cooper, A., 2004. Rise and fall of the Beringian steppe bison. Science 306, 1561e1565. Sommer, R.S., Benecke, N., 2009. First radiocarbon dates on woolly mammoth (Mammuthus primigenius) from northern Germany. Journal of Quaternary Science 24, 902e905. Strelkov, S.A., 1977. Pervaya nakhodka mamonta v podmorennykh sloyakh na Kol'skom poluostove. Trudy Zoologicheskogo Instituta AN SSSR 73, 36e39 (in Russian). Stuart, A.J., 1977. The vertebrates of the Last Cold Stage in Britain and Ireland. Philosophical Transactions of the Royal Society B: Biological Sciences 280, 295e312. Stuart, A.J., 1982. Pleistocene vertebrates in the British Isles. Longman, London. Stuart, A.J., 1983. Pleistocene Bone Caves in Britain and Ireland. Studies in Speleology 4, 9e36. Stuart, A.J., van Wijngaarden-Bakker, L.H., 1985. Quaternary vertebrates. In: Edwards, K.J., Warren, W.P. (Eds.), The Quaternary History of Ireland. Academic Press, London, pp. 221e249. Sulerzhitsky, L.D., Romanenko, F.A., 1997. Vozrast i rasselenie “mamontovoj” fauny aziatskogo zapolyar'ya (po radiouglerodnym dannym). Kriosfera Zemli 1, 12e19 (in Russian). Takahashi, K., Soeda, Y., Izuho, M., Yamada, G., Akamatsu, M., Chang, C.-H., 2006. The chronological record of the woolly mammoth (Mammuthus primigenius) in Japan, and its temporary replacement by Palaeoloxodon naumanni during MIS 3 in Hokkaido (northern Japan). Palaeogeography, Palaeoclimatology, Palaeoecology 233, 1e10. Takahashi, K., Wei, G., Uno, H., Yoneda, M., Jin, C., Sun, C., Zhang, S., Zhong, B., 2007. AMS 14C chronology of the world's southernmost woolly mammoth (Mammuthus primigenius Blum.). Quaternary Science Reviews 26, 954e957. Tong, H.-W., 2004. Paleoenvironmental significance of Coelodonta in different fossil assemblages. Acta Anthropologica Sinica 23 (Suppl.), 306e314 (in Chinese, with English summary). Tong, H., 2010a. Proboscidean fossil records in Nihewan Basin, North China. In: Lacombat, F., Mol, D. (Eds.), The Vth International Conference on Mammoths rie vol. 3, 173e174. and Their Relatives. Quaternaire, Hors-se Tong, H., 2010b. New materials of Mammuthus trogontherii (Proboscidea, Mammalia) of Late Pleistocene age from Yuxian, Hebei. Quaternary Sciences 30, 307e318 (in Chinese, with English abstract). Ukkonen, P., Lunkka, J.P., Jungner, H., Donner, J., 1999. New radiocarbon dates from Finnish mammoths indicating large ice-free areas in Fennoscandia during the Middle Weichselian. Journal of Quaternary Science 14, 711e714. Ukkonen, P., Aaris-Sørensen, K., Arppe, L., Clark, P.U., Daugnora, L., Lister, A.M., ~ ugas, L., Seppa €, H., Sommer, R.S., Stuart, A.J., Wojtal, P., Zupins, I., 2011. Woolly Lo mammoth (Mammuthus primigenius Blum.) and its environment in northern Europe during the last glaciation. Quaternary Science Reviews 30, 693e712. Vartanyan, S.L., Arslanov, K.A., Karhu, J.A., Possnert, G., Sulerzhitsky, L.D., 2008. Collection of radiocarbon dates on the mammoths (Mammuthus primigenius) and other genera of Wrangel Island, northeast Siberia, Russia. Quaternary Research 70, 51e59. Vereshchagin, N.K., 1959. Mlekopitayushchie Kavkaza. Istoriya formirovaniya fauny. Izdatelstvo Akademii Nauk SSSR, Moskva, Leningrad (in Russian). Vereshchagin, N.K., Tikhonov, A.N., 1990. Ekster'er mamonta. Akademiya Nauk SSSR, Sibirskoye otdelenie, Yakutskaya respublikanskaya kommissiya po izucheniyu mamontovoj fauny, Yakutsk. € ro € s, I., 1981. The skulls of Mammoth in Hungary. Fragmenta Mineralogica et Vo Palaeontologica 10, 97e106. Wei, G.B., Taruno, H., Kawamura, Y., Jin, C.Z., 2006. Pliocene and Early Pleistocene Primitive Mammoths of Northern China: Their revised taxonomy, biostratigraphy and evolution. Journal of Geosciences, Osaka City University 49, 59e101. Wei, G.B., Hu, S.M., Yu, K.F., Hou, Y.M., Li, X., Jin, C.Z., Wang, Y., Zhao, J.X., Wang, W.H., 2010. New materials of the steppe mammoth, Mammuthus trogontherii, with discussion on the origin and evolutionary patterns of mammoths. Science China, Earth Science 53, 956e963. Widga, C., Lengyel, S., Saunders, J., Hodgins, G., Walker, J.D., 2014. Proboscidean biogeography in midwestern North America, with a comment on the extinction of Mammut americanum. In: Kostopoulos, D.S., Vlachos, E., Tsoukala, E. (Eds.), Abstract Book of the VIth International Conference on Mammoths and Their Relatives. Scientific Annals of the School of Geology, Aristotle University of Thessaloniki Special Volume 102, 221. Wojtal, P., 2007. Zooarchaeological studies of the Late Pleistocene sites in Poland. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, w. Krako Yesner, D.R., Veltre, D.W., Crossen, K.J., Graham, R.W., 2005. 5,700-year-old Mammoth Remains from Qagnax Cave, Pribilof Islands, Alaska. In: Agenbroad, L.D., Symington, R.L. (Eds.), 2nd World of Elephants Congress. Short Papers and Abstracts. Mammoth Site of Hot Springs, South Dakota, pp. 200e204. Zhylkibaev, K. Zh, 1975. Drevnie slony Kazakhstana. Izdatelstova Nauka Kazakhskoj SSR, Alma-Ata (in Russian).
Contents lists available at ScienceDirect
Quaternary International journal homepage: www.elsevier.com/locate/quaint
The maximum geographic extension of Late Pleistocene Mammuthus primigenius (Proboscidea, Mammalia) and its limiting factors* Ralf-Dietrich Kahlke Senckenberg Research Institutes and Natural History Museums, Research Station of Quaternary Palaeontology, Am Jakobskirchhof 4, 99423 Weimar, Germany
a r t i c l e i n f o
a b s t r a c t
Article history: Available online 8 May 2015
The maximum geographic extension of fully developed woolly mammoth, Mammuthus primigenius, occurred during the last glacial period (MIS 5de2), i.e. during the 110e14 ka interval. The mapping of the species' widest distribution reveals principal barriers to its expansion that were acting independent of short term environmental fluctuations. The maximum spread of M. primigenius was primarily controlled by the following trans-regional factors: (A) The configuration of inland glaciers; (B) The configuration of high mountain chains; (C) The configuration of semi-deserts and deserts; (D) The configuration of marine shorelines at year-round open water surfaces; (E) The exposure of continental shelf regions; and (F) The replacement of tundra-steppe by extended grasslands. Some of the biogeographic barriers were climateindependent; others were determined by long- or medium-term climatic processes. The effect of Eurasian and North American intra-continental steppes as obstacles to migration has not yet been fully elucidated. The verifiable area of the Late Pleistocene distribution range of M. primigenius, based on geomorphological data, comprises approximately 33,301,000 km2 (surface area). When taking the geographical distribution of a species as a measure of its success, the Late Pleistocene woolly mammoth was one of the most successful large herbivores of the Holarctic Mammoth Fauna. © 2015 Elsevier Ltd and INQUA. All rights reserved.
Keywords: Mammuthus primigenius Maximum geographic extension Biogeographic barriers Late Pleistocene
1. Introduction The origin and distribution of individual species reflects, in a very clear way, the spatial and temporal variation of environmental r conditions (Ba anescu and Bos¸caiu, 1978; Lomolino and Heaney, 2004 and references therein). The barriers restricting the spread of a species are determined by their mode of life, as well as the physiological performance of single individuals in relation to the prevailing environmental conditions (Cox and Moore, 2010 and references therein). The determination of the limiting factors in the distribution of fossil species can be essential in understanding the dynamics of palaeoecosystems as well contributing to the elucidation of the causes underlying local or global extinction of individual species or whole faunas. The history of the development of cold adapted large mammal associations of the northern hemisphere during the Pleistocene
* This paper is dedicated to Larry D. Agenbroad (y), a colleague and friend. We miss him. E-mail addresses: [email protected], [email protected].
http://dx.doi.org/10.1016/j.quaint.2015.03.023 1040-6182/© 2015 Elsevier Ltd and INQUA. All rights reserved.
was, to a great extent, determined by the increasing aridity of significant parts of Eurasia, through rhythmical global cooling with extended and intensified cold phases, as well as increasing continentality (Kahlke, 2014). Between 2.6 Ma and 700 ka, the continental Asian steppe regions and the circumpolar tundra belt were biogeographically separated, each inhabited with its own, largely independent mammalian faunas (Kahlke, 1999, 2014). At about 900 ka, the transformation of the global climate cyclicity from a 41 ky to a clear 100 ky periodicity was completed (Lisiecki and Raymo, 2005). Correspondingly, prolonged climatic cycles, which appeared as extended glacial and interglacial phases, caused drastic changes in the faunal structure of the Northern Hemisphere (Kahlke et al., 2011). During the pronounced cold period of MIS 12, Arctic tundra species regularly expanded south- and southwestward into a newly formed type of biome, the so-called tundrasteppe. In parallel, species originating from the inner continental (Asian) steppe dispersed into new habitats north and northwest of their ancestral distribution areas. Such drastic faunal turnover led to the formation of the earliest Palaearctic Mammoth Fauna at around 460 ka (Kahlke and Lacombat, 2008; Kahlke, 2014). This new type of mammal assemblage, or its individual faunal elements,
148
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
expanded and contracted its geographic extension parallel to alternating environmental conditions of the subsequent late Middle to Late Pleistocene (Kahlke, 1999). During the last glacial period Mammuthus primigenius, the characteristic species of Pleistocene cold adapted faunas, reached its farthest spatial extent (Kahlke, 1999). With the help of a reconstruction of its distribution range based on the current state of knowledge, principal limiting factors of its extension can be detected and with them the maximum range of the species in the northern hemisphere clarified. 2. Holarctic mammoth faunas and the emergence of Mammuthus primigenius The aridisation of intra-continental regions and the duration and intensity of individual cold periods drove the evolution and dispersal of the genus Mammuthus. Mammuthus trogontherii as the proboscidean key species of the Palaearctic Mammoth Faunas, originated during the Early Pleistocene, between the Olduvai and Jaramillo subchrones, in continental areas north of the HimalayaneTibetan uplift (Wei et al., 2006). In the period around 1.2 Ma, a significant expansion event of M. trogontherii into the north of Asia is assumed to have occurred, as recorded by finds from the northeastern Siberian Early Olyorian (1.2e0.8 Ma; Lister and Sher, 2001). More or less at the same time, at the beginning of the Epivillafranchian, M. trogontherii also appeared in the Western Palaearctic for the first time. The species established itself there as a characteristic faunal element of cool and cold periods until the Middle Pleistocene (Lister et al., 2005; Kahlke et al., 2011; Kahlke, 2014). An even longer persistence of evolved trogontherii-populations until the Late Pleistocene is plausible for the Central Asian region, especially in northern China (Wei et al., 2010; Tong, 2010a, 2010b). Moreover, it cannot be excluded that Early Pleistocene forms on the level of M. trogontherii could have spread to the Nearctic and established the foundation of the Mammuthus columbi lineage (e.g. Lister and Bahn, 2007; Wei et al., 2010). The Columbian Mammoth (sensu lato, details in Agenbroad, 2005) dominated large areas of North America until the Late Pleistocene. Between 800 and 600 ka (Late Olyorian) in northeastern Eurasia, indigenous M. trogontherii started to transform into more advanced mammoth forms, mainly recognisable from the increase in the number of enamel lamellae in the dental elements (Lister et al., 2005). Continuous cold conditions, which probably fluctuated less on an annual and interannual basis than in inner continental regions of Eurasia, in combination with Far Northern vegetation phenology, caused the speciation of M. primigenius. The primigenius stage was already widespread in NE Asia from 400 ka onwards (Lister and Sher, 2001). In Europe, corresponding forms first occurred much later, during the late MIS 7 or at the beginning of MIS 6, 200e160 ka. The fossil record indicates an oscillating movement of synchronous western Palaearctic M. trogontherii and M. primigenius populations (Lister et al., 2005). During the Late Pleistocene, M. primigenius reached northern North America (Agenbroad, 2005), where already during the last (Sangamonian) interglacial unglaciated areas were successfully occupied (Harington, 2011, Tab. 1). Recorded finds show that occasionally woolly mammoths and Columbian mammoths co-occurred in one and the same site (e.g. Agenbroad, 1994), indicating temporal replacement or short term (perhaps seasonal) sympatry. 3. Maximum geographic extension of Late Pleistocene Mammuthus primigenius The maximum distribution of fully developed woolly mammoths occurred during the last glacial period (¼Devensian ¼
Weichselian ¼ Valdaian ¼ Dalian ¼ Wisconsinian) and spans the marine isotope stages (MIS) 5de2, i.e. approximately the interval between 110 and 14 ka (Lisiecki and Raymo, 2005). In addition to a number of overviews and summaries (Graham and Lundelius, 1994; Kahlke, 1999; Kuzmin et al., 2003, 2008; Agenbroad, 2005; AlvarezLao et al., 2009; Markova et al., 2013, 2014), the mapping of the last glacial maximum spread of M. primigenius is primarily based on regional studies. Particular emphasis is placed on data from areas close to the maximum distribution limits of the species, to derive principle limiting factors. Individual verifications of M. primigenius have significant regional variations in frequency. In addition to biogeographic conditions, these also arise from differences in regional research intensity. The Iberian Peninsula constituted the most southwesterly part of the last glacial Eurasian woolly mammoths' distribution range. Recent studies have established that the species was not only in the northern Spanish Basque country, but also in Portugal, in the Madrid region, and Granada (Alvarez-Lao, 2007; Alvarez-Lao et al., 2009; Alvarez-Lao and García, 2011). In the Pyrenees, areas of the northern flank were populated (Clot and Duranthon, 1990). Evidence of M. primigenius has also been found in almost all parts of France (e.g. Delpech, 1983; Fosse et al., 2014), in Ireland and Britain (Stuart, 1977, 1982, 1983; Stuart and Wijngaarden-Bakker, 1985; Lister, 2009), on Jersey (Scott et al., 2014), in the region of the Netherlands, as well as in the southern part of the North Sea (van Essen and Mol, 1996; Mol et al., 1999, 2006a, 2008; Kahlke, 2001). There are also numerous documented finds in southern Scandinavia, including Finland and in the Baltic states (AarisSørensen et al., 1990; Aaris-Sørensen, 1998, 2009; Ukkonen et al., 1999, 2011; Markova et al., 2013). In Central Europe, remains of M. primigenius are common occurrences in the fossil record (Kowalski, 1959; Musil, 1968, 1985; €ppes and Rabeder, 1997; Germonpre , 2003; Hünermann, 1985; Do Wojtal, 2007; Sommer and Benecke, 2009; Nadachowski et al., 2011; Ukkonen et al., 2011; Gaar and Preusser, 2012; Pawłowska, 2014). On the Apennine peninsula, the animals ventured into southern Italy (Sala et al., 1992; Biddittu and Celletti, 2001; Rustioni et al., 2003). To the southeast, the Pannonian Basin and the Balkan nossy and Vo € ro €s, 1979; Peninsula were occupied (Rakovec, 1975; Ja € ro € s, 1981; Malez, 1986; Konra d et al., 2010; Katona Koufos, 1981; Vo et al., 2012; Mauch Lenardi c, 2012). In Eastern Europe, the species ranged from the Russian Plain and Ukraine, up into the southern part of the Kola Peninsula (Strelkov, 1977), and to the north coast of the Black Sea (Markova et al., 2013). In the Crimea, the lowlands as well as northern foothills and low mountains of the Crimea mountain range were settled (Baryshnikov, 2003). The situation was similar in the margins of the northern foothills of the Greater Caucasus range (Baryshnikov, 2003). Occasionally, M. primigenius penetrated even into Transcaucasia, as sporadic evidence as far as Tbilisi has shown (Gabunia, 1952; Vereshchagin, 1959). To the east, the Caspian Sea as well as areas in north-west to north-eastern Kazakhstan were reached (Kozhamkulova, 1969; Zhylkibaev, 1975). The animals also expanded into the area between the Balkhash Lake and the Dzungarian Alatau, around present-day Almaty and the foothills of the Transili-Alatau. At higher latitudes, there is widespread evidence for their occupation in present-day Bashkortostan and the Urals (Dubrovo, 1966; Kuzmina, 1971; Danukalova and Yakovlev, 2001; Markova et al., 2013). Western Siberia is rich in Late Pleistocene mammoth evidence too (Alekseeva, 1980; Kuzmin et al., 2003, 2008; Markova et al., 2013). The species has also been found in the Xinjiang Uygur Region in westernmost China (Tong, 2004, p. 311) and in the northern part of the Mongol UIs (records in the collections of the Museum of Natural History Ulaanbaatar). The East Siberian region
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
to Transbaikalia was also a part of the last glacial distribution area of the woolly mammoth (Ermolova, 1978; Imetkhenov and Kalmykov, 1988; Kuzmin et al., 2008; Alexeeva and Erbajeva, 2010; Markova et al., 2013). Northeast China, especially the provinces of Heilongjiang and Jilin, has provided numerous pieces of evidence for M. primigenius (Liu and Li, 1989; Jin and Kawamura, 1996). The southernmost distribution of woolly mammoths in East Asia extended to the coastal region of the Sea of Bohai (compilation in Alvarez-Lao et al., 2009, p. 66). A confirmed molar of M. primigenius has been recorded from Ji'nan at the Beidasha River (Shandong province) at 36 350 N. According to Takahashi et al. (2007), this find represents the world's most southerly record of the species. On the Korean peninsula evidence of M. primigenius has been found as far as the Ham Kyung province (Park, 1988) and in the Jungwon area in the northernmost part of South Korea at c. 37 500 N (Cho and Woo, 2004). Through the Russian Far East (e.g. Kuzmin et al., 2000), woolly mammoths reached as far as Sakhalin at 46 N (Matsumoto, 1937; Kuzmin et al., 2005) and to Hokkaido at 42 N (Dubrovo, 1981; Kamei, 1981; Takahashi et al., 2006). The detection of M. primigenius from the Sea of Japan off western Shimane Prefecture, 35 200 N, is taken to be a consequence of drifting from the Huang He area of mainland China (Kamei, 1990; Takahashi et al., 2006). The Siberian Arctic and the continental shelf of the Arctic Ocean, including today's New Siberian Islands were also part of the heartland of the last glacial mammoth distribution (e.g. Vereshchagin and Tikhonov, 1990; Sulerzhitsky and Romanenko, 1997; Kuzmina et al., 1999; MacPhee et al., 2002; Kahlke and Mol, 2005; Mol et al., 2006b; Kuznetsova, 2007; Kuzmin et al., 2008; Lazarev, 2008; Vartanyan et al., 2008; Markova et al., 2013). The Kamchatka peninsula was also reached (Kuzmin et al., 2000; Orlova et al., 2000). The distribution area expanded continuously as far as eastern Beringia and beyond (Agenbroad and Barton, 1991; Harington, 2003, 2011; Kuzmin et al., 2008). There are existing records of woolly mammoth occupation from the Pribilof Islands (Yesner et al., 2005) and from the western part of the Canadian Arctic Archipelago (MacPhee, 2007; Kuzmin et al., 2008). In addition to Alaska, numerous remains of M. primigenius are commonly found in northern Canada (Harington, 2003). In North America, the animals have been established in many sites in the Great Lakes region and across the Northern Plains (Agenbroad, 1985, 1989, 2005; Agenbroad et al., 1994; Burns, 1996; Kuzmin et al., 2008; Harington et al., 2011; Widga et al., 2014). Along the U.S. West Coast, they ventured to around 45 N (Fisher in Larmer, 2013). Easternmost evidence for M. primigenius is documented in Coastal Maine (Hoyle et al., 2004), Nova Scotia (Cooke et al., 1993), as well as on the Atlantic continental shelf of Virginia (Harington et al., 2011). 4. Principal biogeographic barriers The mapping of the last glacial maximum distribution of M. primigenius, inclusive of all territorial variations during this period documented by fossil records, reveals principal biogeographical barriers that are acting independently of short term environmental fluctuations. Thus, the configuration of the northern distribution limit of woolly mammoths very clearly reflects the position of the great inland glaciers (Fig. 1A, arrows). In particular, the Scandinavian (-West Siberian) ice sheet in Eurasia and the Laurentide and the Cordilleran ice sheets in North America (Ehlers and Gibbard, 2004a, 2004b, 2004c; Ehlers et al., 2011) were significant barriers to expansion. However, the two major New World glacial regions were only connected during the Last Glacial Maximum (LGM) (Dyke et al., 2002) and thus only briefly formed an effective single, continent-wide migration barrier. In most cases,
149
an ice-free corridor between the two ice sheets enabled the invasion of woolly mammoths to the southeast (see section 3). In other regions, advances of M. primigenius also trace deglaciated areas, as in the circum-Baltic region of Northern Europe (Ukkonen et al., 2011), on the Taimyr Peninsula of Arctic Siberia (Alexanderson et al., 2002; Kahlke and Mol, 2005, p. 50ff.), and in the western periphery of the Canadian Arctic Archipelago (MacPhee, 2007). High mountain chains with no passable valleys were another very effective barrier to migration (Fig. 1B, arrows). From the distribution of finds, this is particularly evident in the Pyrenees area, the Crimea mountain range and the Greater Caucasus. While M. primigenius could, in every case, colonise the (northern) flank of the mountain ranges facing the distribution areas, up to considerable altitudes, the ridges remained insurmountable. In different regions the high mountains were circumvented, as in the colonisation of the Iberian Peninsula and Transcaucasia. In Central Asia, a lowland corridor between the Altai and Dzungarian Alatau mountain ranges enabled the mammoths to advance from eastern Kazakhstan to China. The mountain ranges of the Alps and Rocky Mountains were only partially settled. Here, mammoths spread out mainly along broad valleys. Inside Eurasia, between Transcaucasia and northern China, the southern border of the last glacial distribution area of M. primigenius was very much shaped by the arrangement of semideserts and deserts (Fig. 1C, arrows). The mammoths recorded south of the Caucasus (see Section 3) are likely to have immigrated through the Kuban area (Krasnodar region), not via the arid areas of Azerbaijan. East of the Caspian Sea, the Kyzyl Kum desert and the semiarid area of the Betpak Dala (Hunger Steppe) limited the mammoths' range. The distribution patterns of Mongolian and northern Chinese M. primigenius also reflect the animals' response to extreme aridity. The desert regions of the Taklamakan and Gobi deserts were strictly avoided. Marine shorelines with year-round, ice-free water surfaces represented a more insurmountable obstacle to expansion for M. primigenius. In regions with near-coastal continental slope, the last glacial maximum distribution of the animals can be fairly accurately estimated (Fig. 1D, arrows). Thus, certainly the last glacial distribution area of the mammoths was directly limited by the majority of the French Mediterranean coast, the northern coast of the Black Sea, as well as some sections of the Pacific North West coast, especially in the Russian Far East at the Sea of Okhotsk and Kamchatka, and the west coast of North America. More difficult to assess is the mammoths' spread within present-day flooded continental shelf regions (Fig. 1E, arrows), which over the course of the Last Glacial eustatic sea level decrease of 120e135 m were exposed (Clark and Mix, 2002). In several regions the bathymetric situation opened up access routes to new expansion areas, as made evident by a M. primigenius migration route via the coastal part of the Cantabrian Sea into the Iberian Peninsula. Moreover, another temporary migration route is likely to have existed directly east of the Pyrenees. Here, a coastal part of the Mediterranean dried out during the extended coldest periods of the Late Pleistocene (Alvarez-Lao et al., 2009). The drop in sea level in the Mediterranean also caused large-scale drying up of northern parts of the Adriatic Sea (Gallini and Sala, 2001; Rustioni et al., 2003), so that an unbroken distribution area for M. primigenius developed between SE Europe and the Apennine Peninsula. Extensive finds-based knowledge accounting for mammoth distribution from the present-day flooded shelf areas exists only from the southern part of the North Sea between England and the Netherlands (see Section 3). Individual finds prove the presence of the animals on the shelf of the Arctic Ocean to the north of the New Siberian Islands and on the Atlantic shelf off the U.S. East Coast (see
150
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
Fig. 1. Localisation of principal barriers (arrows) of Late Pleistocene Mammuthus primigenius spread: A e ice sheets; B e high mountain chains; C e semi-deserts and deserts; D e year round open water surfaces; E e continental shelf regions; F e other types of grassland.
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
151
Fig. 1. (continued)
Section 3). A land bridge enabled the colonization of Sakhalin in the North Pacific (Kirillova and Tesakov, 2008). Evidence from the New Siberian Islands and Wrangel, as well as from the Pribilof Islands and the Canadian Arctic Archipelago (see Section 3) allow a conservative extrapolation of last glacial spread of M. primigenius in the region of today's Laptev- and East Siberian Sea and the Chuchi- and Bering Sea (Fig. 1E). New finds here could lead to significant increases in the areas which can be demonstrably shown to support woolly mammoth populations. In contrast to the aforementioned, comparatively welldocumented control factors (Fig. 1AeE), the effect of the intracontinental steppes of Eurasia and North America as migration barriers for M. primigenius has been little investigated. In recent times, it has become clear that sister taxa of several species involved in Eurasian Mammoth Faunas underwent separate evolution in Central Asia, thus indicating significant ecological differences between the Asian core steppe and Eurasian tundra-steppe habitats (Kahlke, 2014). Independent of the successively evolving M. primigenius of northern altitudes, M. trogontherii survived in Central Asia until the Late Pleistocene. A corresponding phenomenon of coexisting mammoth species, adapted to intra-continental steppe and tundra steppe habitats respectively, is known from North America. Here, M. columbi, which in many respects resembles M. trogontherii, occupied southerly extended grassland habitats, whereas M. primigenius synchronously spread into the mammoth steppe. It remains unclear whether the replacement of vegetation, the competition for food from the steppe mammoths (M. trogontherii and M. columbi respectively), or a combination of both prevented the successful spread of M. primigenius into intra-
continental steppe regions of Eurasia and North America (Fig. 1F, arrows). 5. Conclusions Based on the current fossil record, Fig. 2 shows the geographic extension of Late Pleistocene M. primigenius. The maximum spread of the species was primarily controlled by the following transregional factors: (A) (B) (C) (D)
The configuration of inland glaciers; The configuration of high mountain chains; The configuration of semi-deserts and deserts; The configuration of marine shorelines at year-round open water surfaces; (E) The exposure of continental shelf regions; (F) The replacement of tundra-steppe by extended grasslands (Central Asian core steppe, North American grasslands).
Locally acting factors could have modified the configuration of the distribution limits slightly here and there. The synopsis of the principal barriers to migration leads to an almost complete explanation of the Holarctic maximum spread of M. primigenius during the Late Pleistocene. AeD constituted more or less abrupt obstacles to expansion, which were not physiologically possible for individual mammoths to overcome. Some of the barriers mentioned were climate-independent (B, partly D), and others were determined by long- or medium-term climatic processes (A, C, E, F, partly D). In the case of F, the as yet still poorly
152
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
Fig. 2. Maximum extension of Mammuthus primigenius during the Late Pleistocene based on the current fossil record.
investigated replacement of tundra-steppe habitats by intracontinental grasslands would probably have had more of a gradual effect in limiting expansion. To date, the verifiable area of the Late Pleistocene distribution range of M. primigenius (Fig. 2) comprises approximately 33,089,000 km2 (status 09.20.2014; calculated by using Esri ArcGIS (C), surface volume tool). Taking into account the geomorphological data (based on Esri topographic map, raster resolution of 30 arc seconds) results in an actual mammoth-population surface area of approximately 33,301,000 km2, which is considered as the minimum of the actual distribution area. Gains in knowledge and thus increases in the known distribution area are to be expected, especially from the currently submerged shelf regions. To a lesser extent this is also possible on the mainland, especially in inaccessible or little studied regions. Despite the limiting factors mentioned, woolly mammoths enjoyed an enormous geographic extension during the Late Pleistocene. When we look at the geographical distribution of a species as a measure of its success, Late Pleistocene M. primigenius was one of the most successful large herbivores of the Holarctic Mammoth Fauna. Only the morphologically quite diverse Bison priscus group occupied a comparably large, and in many places identical, distribution area during the last glacial period (Kahlke, 1999, Fig. 20; Shapiro et al., 2004 and references therein). The woolly mammoth rightly proved itself as a palaeoecological characteristic species and a worthy namesake for the most cold-adapted large mammal assemblages in Earth's history, the Mammoth Fauna. Acknowledgements The study was conducted within the Senckenberg Research Institute's project group “Origin, dispersal and impoverishment of Eurasian cold adapted faunas” (RDK). Thanks go to E. Haase and G. Utschig (both Weimar) for their assistance in designing the graphics. For information and support on various topics we are grateful to B. Fiedler (Weimar; references), T. Hermsen (Offenbach; area calculations und graphics), C. Hertler (Frankfurt/M.; logistics), I. Kirillova (Moscow; literature), D. Mol (Hoofddorp; information on mammoth sites) and C. Nielsen-Marsh (Leipzig; language). The kind comments of E. Tsoukala (Thessaloniki) and an anonymous reviewer are greatly appreciated. References Aaris-Sørensen, K., 1998. Danmarks forhistoriske dyreverden. Om skovelefanter, næsehorn, bisoner, urokser, mammutter og kæmpehjorte. Gyldendal, Denmark.
Aaris-Sørensen, K., 2009. Diversity and dynamics of the mammalian fauna in Denmark throughout the last glacial-interglacial cycle, 115-0 kyr BP. Fossils and Strata 57, 1e59. Aaris-Sørensen, K., Petersen, K.S., Tuber, H., 1990. Danish finds of Mammoth (Mammuthus primigenius (Blumenbach)). Stratigraphical position, dating and evidence of Late Pleistocene environment. Danmarks Geologiske Undersøgelse, DGU Serie B 14, 1e44. Agenbroad, L.D., 1985. The distribution and chronology of Mammoth in the New World. Acta Zoologica Fennica 170, 221e224. Agenbroad, L.D., 1989. New World mammoth distribution. In: Martin, P.S., Klein, R.G. (Eds.), Quaternary Extinctions. A Prehistoric Revolution. University of Arizona Press, Tucson, pp. 90e108. Agenbroad, L.D., 1994. Geology, hydrology, and excavation of the site. In: Agenbroad, L.D., Mead, J.I. (Eds.), The Hot Springs Mammoth Site: A Decade of Field and Laboratory Research in Paleontology, Geology, and Paleoecology. Fenske, Rapid City, SD, pp. 15e27. Agenbroad, L.D., 2005. North American Proboscideans: Mammoths: The state of Knowledge, 2003. Quaternary International 126e128, 73e92. Agenbroad, L.D., Barton, B.R., 1991. North American mammoths: An annotated bibliography 1940-1990. Mammoth Site of Hot Springs Scientific Papers 2, 1e118. Agenbroad, L.D., Lister, A.M., Mol, D., Roth, V.L., 1994. Mammuthus primigenius remains from the Mammoth Site of Hot Springs, South Dakota. In: Agenbroad, L.D., Mead, J.I. (Eds.), The Hot Springs Mammoth Site: A Decade of Field and Laboratory Research in Paleontology, Geology, and Paleoecology. Fenske, Rapid City, pp. 269e281. Alekseeva, E.V., 1980. Mlekopitayushchie plejstocena yugo-vostoka Zapadnoj Sibiri. Nauka, Moskva (in Russian). € ller, P., Antonov, O., Eriksson, S., Alexanderson, H., Adrielsson, L., Hjort, C., Mo Pavlov, M., 2002. Depositional history of the North Taymyr ice-marginal zone, Siberia e a landsystem approach. Journal of Quaternary Science 17, 361e382. Alexeeva, N., Erbajeva, M., 2010. Habitat and distribution area of woolly mammoth and some core mammal species in Transbaikalia during the Late Pleistocene. In: Lacombat, F., Mol, D. (Eds.), The Vth International Conference on Mammoths rie 3, 177e178. and Their Relatives. Quaternaire, Hors-se n paleontolo gica de los macromamíferos indicadores Alvarez-Lao, D.J., 2007. Revisio rica (Ph.D. thesis). Universidad de Oviedo, de clima frío en la Península Ibe Departamento de Geología, Oviedo, Spain. Alvarez-Lao, D.J., García, N., 2011. Geographical distribution of Pleistocene coldadapted large mammal faunas in the Iberian Peninsula. Quaternary International 233, 159e170. Alvarez-Lao, D.J., Kahlke, R.-D., García, N., Mol, D., 2009. The Padul mammoth finds e On the southernmost record of Mammuthus primigenius in Europe and its southern spread during the Late Pleistocene. Palaeogeography, Palaeoclimatology, Palaeoecology 278, 57e70. ra nescu, P., Bos¸caiu, N., 1978. Biogeographie. Fauna und Flora der Erde und ihre Ba geschichtliche Entwicklung. Gustav Fischer, Jena. Baryshnikov, G., 2003. Mammuthus primigenius from the Crimea and the Caucasus. In: Reumer, J.W.F., de Vos, J., Mol, D. (Eds.), Advances in Mammoth Research. Proceedings of the Second International Mammoth Conference, Rotterdam, 1620 May 1999. Deinsea 9, 41e56. Biddittu, I., Celletti, P., 2001. Plio-Pleistocene Proboscidea and Lower Palaeolithic bone industry of southern Latium (Italy). In: Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), The World of Elephants. Proceedings of the 1st International Congress, Roma, 16-20 Ottobre 2001. Consiglio Nazionale delle Ricerche, Roma, pp. 91e96. Burns, J.A., 1996. Vertebrate paleontology and the alleged ice-free corridor: the meat of the matter. Quaternary International 32, 107e112. historique et la subsistence de la re gion Cho, T.-S., Woo, J.-Y., 2004. La faune pre e. In: Lee, Y.-J., Cho, T.-S. (Eds.), Paleoenvironnement et culture Jungwon en Core
R.-D. Kahlke / Quaternary International 379 (2015) 147e154 e. Institute for Jungwon Culture, paleolithique de la region Jungwon, Core Chungbuk National University, Seoul, pp. 89e99. Clark, P.U., Mix, A.C., 2002. Ice sheets and sea level of the Last Glacial Maximum. Quaternary Science Reviews 21, 1e7. res fossiles du Quaternaire dans les Clot, A., Duranthon, F., 1990. Les mammife ne es. Museum d’Histoire Naturelle de Toulouse, Toulouse. Pyre Cooke, H.B.S., Harington, C.R., Sollows, J.D., 1993. Undescribed mammoth (Mammuthus) teeth from Georges Bank and Nova Scotia. Proceedings of the Nova Scotian Institute of Science 40, 19e28. Cox, C.B., Moore, P.D., 2010. Biogeography. An Ecological and Evolutionary Approach. Wiley, Hoboken. Danukalova, G.A., Yakovlev, A.G., 2001. Finds of Proboscidean remains in the territory of the Southern Urals region. In: Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), The World of Elephants. Proceedings of the 1st International Congress, Roma, 16-20 Ottobre 2001. Consiglio Nazionale delle Ricerche, Roma, pp. 201e204. olithique supe rieur dans le Sud-Ouest de la Delpech, F., 1983. Les faunes du Pale France. Cahiers du Quaternaire 6. Editions du C.N.R.S., Paris. € €ppes, D., Rabeder, G. (Eds.), 1997. Plioz€ €ne Faunen Osterreichs. Do ane und pleistoza Ein Katalog der wichtigsten Fossilfundstellen und ihrer Faunen (Endbericht €rderung der wissendes Forschungsprojektes Nr. 9320 des “Fonds zur Fo schaftlichen Forschung”). Mitteilungen der Kommission für Quart€ arforschung € der Osterreichischen. Akademie der Wissenschaften Band 10. Verlag der € Osterreichischen Akademie der Wissenschaften, Wien. Dubrovo, I.A., 1966. Plejstocenovye mlekopitayushchie Sredgeo i Yushogo Urala i ikh stratigrafichekoye znachenie. Nauchnye Trudy Politechnicheskogo Instituta, Sbornik 20, 12e135 (in Russian). €ontologie 4, 49e84. Dubrovo, I.A., 1981. Die fossilen Elefanten Japans. Quart€ arpala Dyke, A.S., Andrews, J.T., Clark, P.U., England, J.H., Miller, G.H., Shaw, J., Veillette, J.J., 2002. The Laurentide and Innuitian ice sheets during the Last Glacial Maximum. Quaternary Science Reviews 21, 9e31. Ehlers, J., Gibbard, P.L. (Eds.), 2004a. Quaternary Glaciations e Extent and Chronology. Part I: Europe. Developments in Quaternary Science, vol. 2. Elsevier, Amsterdam. Ehlers, J., Gibbard, P.L. (Eds.), 2004b. Quaternary Glaciations e Extent and Chronology. Part II: North America. Developments in Quaternary Science, vol. 2. Elsevier, Amsterdam. Ehlers, J., Gibbard, P.L. (Eds.), 2004c. Quaternary Glaciations e Extent and Chronology. Part III: South America, Asia, Africa, Australasia, Antarctica. Developments in Quaternary Science, vol. 2. Elsevier, Amsterdam. Ehlers, J., Gibbard, P.L., Hughes, P.D. (Eds.), 2011. Quaternary Glaciations e Extent and Chronology. A Closer Look. Developments in Quaternary Science, vol. 15. Elsevier, Amsterdam. Ermolova, N.M., 1978. Teriofauna doliny Angary v pozdnem antropogene. Izdatelstvo Nauka, Sibirskoe otdelenie, Novosibirsk (in Russian). van Essen, H., Mol, D., 1996. Plio-Pleistocene proboscideans from the southern bight of the North Sea and the Eastern Scheldt, The Netherlands. In: Shoshani, J., Tassy, P. (Eds.), The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives. Oxford University Press, Oxford, New York, Tokyo, pp. 214e224. gut-Bonnoure, E., Fleury, G., Fourvel, J.B., Madeleine, S., Fosse, P., Fritz, C., Cre re, L., Rivero, O., Tosello, G., 2014. Woolly mammoths (MamSauvet, G., Aurie muthus primigenius) in southern France during the Late Palaeolithic: a geochronological assessment based on the palaeontological, rock art and portable art records. In: Kostopoulos, D.S., Vlachos, E., Tsoukala, E. (Eds.), Abstract Book of the VIth International Conference on Mammoths and their Relatives. Scientific Annals of the School of Geology, Aristotle University of Thessaloniki Special volume 102, 62. Gaar, D., Preusser, F., 2012. Luminescence dating of mammoth remains from northern Switzerland. Quaternary Geochronology 10, 257e263. Gabunia, L.K., 1952. O zube “Karlikovogo” mamonta iz Gori. Soobshcheniya Akademii Nauk Gruzinskoj SSR 13, 153e154 (in Russian). Gallini, V., Sala, B., 2001. Settepolesini di Bondeno (Ferrara e Eastern Po Valley): the first example of mammoth steppe in Italy. In: Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), The World of Elephants. Proceedings of the 1st International Congress, Roma, 16-20 Ottobre 2001. Consiglio Nazionale delle Ricerche, Roma, pp. 272e275. , M., 2003. Mammoth taphonomy of two fluvial sites from the Flemish Germonpre Valley, Belgium. In: Reumer, J.W.F., de Vos, J., Mol, D. (Eds.), Advances in Mammoth Research, Proceedings of the Second International Mammoth Conference, Rotterdam, May 16-20 1999. Deinsea 9, 171e183. Graham, R.W., Lundelius Jr., E.L., Faunmap Working Group, 1994. Faunmap: A Database Documenting Late Quaternary Distributions of Mammal Species in the United States. Illinois State Museum Scientific Papers 25(1), viii þ 1-287, 25 (2), vii þ 289e690. Harington, C.R. (Ed.), 2003. Annotated Bibliography of Quaternary Vertebrates of Northern North America e With Radiocarbon Dates. Toronto University Press, Toronto, Buffalo, London. Harington, C.R., 2011. Pleistocene vertebrates of the Yukon Territory. Quaternary Science Reviews 30, 2341e2354. Harington, C.R., Mol, D., van der Plicht, J., 2011. The Muirkirk Mammoth: A Late Pleistocene woolly mammoth (Mammuthus primigenius) skeleton from southern Ontario, Canada. Quaternary International 255, 106e113. Hoyle, B.G., Fisher, D.C., Borns Jr., H.W., Churchill-Dickson, L.L., Dorion, C.C., Weddle, T.K., 2004. Late Pleistocene mammoth remains from Coastal Maine, USA. Quaternary Research 61, 277e288.
153
€ugetiere aus dem Kanton Zürich. Hünermann, K.A., 1985. Eiszeit-Sa Vierteljahresschrift der Naturforschenden Gesellschaft in Zürich 130, 229e250. Imetkhenov, A.B., Kalmykov, N.P., 1988. Fauna i flora kajnozoya Buryatii. Buryatskij filial SO AN SSSR, Ulan-Ude (in Russian). nossy, D., Vo € ro €s, I., 1979. Gross€ Ja auger-Streufunde aus dem Pleistoz€ an Ungarns. Fragmenta Mineralogica et Palaeontologica 9, 21e60. Jin, C.-Z., Kawamura, Y., 1996. Late Pleistocene mammal fauna in Northeast China: Mammal fauna including woolly mammoth and woolly rhinoceros in association with Paleolithic tools. Earth Science (Chikyu Kagaku) 50, 315e330. Kahlke, R.-D., 1999. The History of the Origin, Evolution and Dispersal of the Late Pleistocene Mammuthus-Coelodonta Faunal Complex in Eurasia (Large Mammals). Fenske Companies, Rapid City. Kahlke, R.-D., 2001. Ein Meer voller Knochen? Pleistoz€ ane Wirbeltierreste aus der Scheldemündung und vom Nordseeboden. Natur und Museum 131, 417e432. Kahlke, R.-D., 2014. The origin of Eurasian Mammoth Faunas (Mammuthus-Coelodonta faunal complex). Quaternary Science Reviews 96, 32e49. Kahlke, R.-D., Lacombat, F., 2008. The earliest immigration of woolly rhinoceros (Coelodonta tologoijensis, Rhinocerotidae, Mammalia) into Europe and its adaptive evolution in Palaearctic cold stage mammal faunas. Quaternary Science Reviews 27, 1951e1961. €ugetiere der Sibirischen Arktis. Die Kahlke, R.-D., Mol, D., 2005. Eiszeitliche Großsa Cerpolex/Mammuthus-Expeditionen auf Tajmyr. Schweizerbart, Stuttgart. Kahlke, R.-D., García, N., Kostopoulos, D.S., Lacombat, F., Lister, A.M., Mazza, P.P.A., Spassov, N., Titov, V.V., 2011. Western Palaearctic palaeoenvironmental conditions during the Early and early Middle Pleistocene inferred from large mammal communities, and implications for hominin dispersal in Europe. Quaternary Science Reviews 30, 1368e1395. Kamei, T., 1981. Faunal succession of Pleistocene Mammals in the Japanese Islands: €rpala €ontologie 4, 165e174. An aspect. Quarta Kamei, T., 1990. The Japan Sea and Elephants. The Quaternary Research 29, 163e172 (in Japanese, with English summary). g mamKatona, L., Kov acs, J., Kordos, L., Szappanos, B., Linkai, I., 2012. The Csaja moths (Mammuthus primigenius): Late Pleniglacial finds from Hungary and their chronological significance. Quaternary International 255, 130e138. Kirillova, I.A., Tesakov, A., 2008. New mammalian elements of the Ice Age assemblage of the Sakhalin Island. Mammal Study 33, 87e92. cs, J., Hal Konr ad, G., Kova asz, A., Sebe, K., P alffy, H., 2010. Late Quaternary woolly mammoth (Mammuthus primigenius Blum) remains from southern Transdanubia, Hungary. Comptes Rendus Palevol 9, 47e54. Koufos, G., 1981. A New Late Pleistocene (Würmian) Mammal Locality from the Basin of Drama (Northern Greece). Scientific Annals of the Faculty of Physics and Mathematics, University of Thessaloniki 21, 129e148. w plejstocenu Polski. Panstwowe Wydawnictwo Kowalski, K., 1959. Katalog ssako Nauk. Warszawa, Wrocław. Kozhamkulova, B.S., 1969. Antropogenovaya iskopaemaya teriofauna Kazakhstana. Nauka, Alma-Ata (in Russian). Kuzmin, Y.V., Orlova, L.A., Zol'nikov, I.D., Igol'nikov, A.E., 2000. The history of mammoth (Mammuthus primigenius Blum.) population in Siberia and adjacent areas (according to radiocarbon data). Russian Geology and Geophysics 41, 723e730. Kuzmin, Y.V., Orlova, L.A., Zolnikov, I.D., 2003. Dynamics of the mammoth (Mammuthus primigenius) population in Northern Asia: radiocarbon evidence. In: Reumer, J.W.F., Vos, J., de, Mol, D. (Eds.), Advances in Mammoth Research, Proceedings of the Second International Mammoth Conference, Rotterdam, May 16-20 1999. Deinsea 9, 221e237. Kuzmin, Y.V., Gorbunov, S.V., Orlova, L.A., Vasilevsky, A.A., Alekseeva, E.V., Tikhonov, A.N., Kirillova, I.V., Burr, G.S., 2005. 14C Dating of the Late-Pleistocene Faunal Remains from Sakhalin Island (Russian Far East). Current Research in the Pleistocene 22, 78e80. Kuzmin, Y.V., Orlova, L.A., Dementiev, V.N., 2008. Dynamics of Mammoth (Mammuthus primigenius Blum.) Populations of Asia and North America and Its Correlation with Climatic Changes in the Late Neopleistocene (45,000e9700 Years BP). Doklady Earth Sciences 421, 978e982. Kuzmina, I.E., 1971. Formirovanie teriofauny severnogo Urala v pozdnem antropogene. Trudy Zoologicheskogo Instituta AN SSSR 49, 44e122 (in Russian). Kuzmina, S., Kuznetsova, T., Sher, A., 1999. Paleontological research on the Bykovsky Peninsula. Reports on Polar Research 315, 179e190. Kuznetsova, T., 2007. Paleontological collection of the “Mammoth” fauna from the museum of the Lena Delta Reserve. Reports on Polar and Marine Research 550, 135e140. Larmer, B., 2013. Of Mammoths and Man. National Geographic 223, 44e63. Lazarev, P.A., 2008. Kupnye mlekopitayushchie antropogena Yakutii. Nauka, Novosibirsk. Lisiecki, L.E., Raymo, M.E., 2005. A Pliocene-Pleistocene stack of 57 globally distributed benthic d18O records. Paleoceanography 20, PA1003. http:// dx.doi.org/10.1029/2004PA001071. Lister, A.M., 2009. Late-glacial mammoth skeletons (Mammuthus primigenius) from Condover (Shropshire, UK): anatomy, pathology, taphonomy and chronological significance. Geological Journal 44, 447e479. Lister, A.M., Bahn, P., 2007. Mammoths. Giants of the Ice Age. University of California Press, Berkeley, Los Angeles. Lister, A.M., Sher, A.V., 2001. The Origin and Evolution of the Woolly Mammoth. Science 294, 1094e1097. Lister, A.M., Sher, A.V., van Essen, H., Wei, G., 2005. The pattern and process of mammoth evolution in Eurasia. Quaternary International 126e128, 49e64.
154
R.-D. Kahlke / Quaternary International 379 (2015) 147e154
Liu, T.-S., Li, X.-G., 1989. Mammoths in China. In: Martin, P.S., Klein, R.G. (Eds.), Quaternary Extinctions. A Prehistoric Revolution. University of Arizona Press, Tucson, pp. 517e527. Lomolino, M.V., Heaney, L.R. (Eds.), 2004. Frontiers of Biogeography. New Directions in the Geography of Nature. Sinauer, Sunderland. MacPhee, R.D.E., 2007. Mammoths in the insular Nearctic? Some constrains on the existence of a Pleistocene megafaunal refugium in the Canadian Arctic Archipelago. Quaternary International 169e170, 29e38. MacPhee, R.D.E., Tikhonov, A.N., Mol, D., de Marliave, C., van der Plicht, H., Greenwood, A.D., Flemming, C., Agenbroad, L., 2002. Radiocarbon Chronologies and Extinction Dynamics of the Late Quaternary Mammalian Megafauna of the Taimyr Peninsula, Russian Federation. Journal of Archaeological Science 29, 1017e1042. €ren Vertebraten-Faunen in der SFR Jugoslawien. Malez, M., 1986. Die quarta €ontologie 6, 101e117. Quart€ arpala Markova, A.K., Puzachenko, A. Yu, van Kolfschoten, T., van der Plicht, J., Ponomarev, D.V., 2013. New data on changes in the European distribution of the mammoth and the woolly rhinoceros during the second half of the Late Pleistocene and the early Holocene. Quaternary International 292, 4e14. Markova, A., Puzachenko, A., van Kolfschoten, T., van der Plicht, J., Kuznezova, T., Tikhonov, A., Ponomarev, D., 2014. The Eurasian mammoth distribution during the second half of the Late Pleistocene and the Holocene. In: Kostopoulos, D.S., Vlachos, E., Tsoukala, E. (Eds.), Abstract Book of the VIth International Conference on Mammoths and Their Relatives. Scientific Annals of the School of Geology, Aristotle University of Thessaloniki Special Volume 102, 118e119. Matsumoto, H., 1937. Two Species of Mammoths from off Odomari and Vicinity, Karafuto (Saghalin). Zoological Magazine 49, 9e11 (in Japanese, with English summary). Mauch Lenardi c, J., 2012. Miocene to Late Pleistocene proboscideans of Croatia. Quaternary International 276e277, 120e128. Mol, D., van den Bergh, G.D., de Vos, J., 1999. Fossil proboscideans from The Netherlands, the North Sea and the Oosterschelde Estuary. In: Haynes, G., Klimovicz, J., Reumer, J.W.F. (Eds.), Mammoths and the Mammoth Fauna: Studies of an Extinct Ecosystem. Deinsea 6, 119e145. Mol, D., Post, K., Reumer, J.W.F., van der Plicht, J., de Vos, J., van Geel, B., van Reenen, G., Pals, J.P., Glimmerveen, J., 2006a. The Eurogeul e first report of the palaeontological, palynological and archaeological investigations of this part of the North Sea. Quaternary International 142e142, 178e185. Mol, D., Tikhonov, A., van der Plicht, J., Kahlke, R.-D., Debruyne, R., van Geel, B., van Reenen, G., Pals, J.P., de Marliave, C., Reumer, J.W.F., 2006b. Results of the CERPOLEX/Mammuthus expeditions on the Taimyr Peninsula, Arctic Siberia, Russian Federation. Quaternary International 142e143, 186e202. Mol, D., de Vos, J., Bakker, R., van Geel, B., Glimmerveen, J., van der Plicht, H., Post, K., 2008. Kleine encyclopedie van het leven in het Pleistoceen. Mammoeten, neushoorns en andere dieren van de Noordzeebodem. Veen Magazines B.V., Diemen. €ontologische Musil, R., 1968. Die Mammutmolaren von Predmostí (CSSR). Pala Abhandlungen, A 3, 1e192. Musil, R., 1985. Paleobiography of terrestrial communities in Europe during the Last rodniho Muzea v Praze, B 41, 1e83. Glacial. Sbornik Na Nadachowski, A., Lipecki, G., Wojtal, P., Mie˛ kina, B., 2011. Radiocarbon chronology of woolly mammoth (Mammuthus primigenius) from Poland. Quaternary International 245, 186e192. Orlova, L.A., Kuzmin, Y.V., Zolnikov, I.D., 2000. Time-space systematics for mammoths (Mammuthus primigenius Blum.) and prehistoric humans in Siberia (on the basis of radiocarbon dating). Archaeology, Ethnology & Anthropology of Eurasia 3, 31e41. Park, S.J., 1988. The Palaeoenvironmental Changes and Macromammal Evolution during the Pleistocene in East Asia. The Korean Journal of Quaternary Research 2, 51e85. Pawłowska, K., 2014. Elephantids from the Pleistocene of Poland: state of knowledge. In: Kostopoulos, D.S., Vlachos, E., Tsoukala, E. (Eds.), Abstract Book of the VIth International Conference on Mammoths and Their Relatives. Scientific Annals of the School of Geology, Aristotle University of Thessaloniki Special volume 102, 152. Rakovec, I., 1975. Razvoj kvartarne sesalske favne Sloveniji. Slovenska Akademija Znanosti in Umetnosti, Razred za Zgodovinske in Druzbene Vede, Cl. I, Hist. Sociol. 24, pp. 225e270. Rustioni, M., Ferretti, M.P., Mazza, P., Pavia, M., Varola, A., 2003. The vertebrate fauna from Cardamone (Apulia, southern Italy): an example of Mediterranean mammoth fauna. In: Reumer, J.W.F., de Vos, J., Mol, D. (Eds.), Advances in Mammoth Research. Proceedings of the Second International Mammoth Conference, Rotterdam, 16-20 May 1999. Deinsea 9, 395e403. Sala, B., Masini, F., Ficcarelli, G., Rook, L., Torre, S., 1992. Mammal dispersal events in the Middle and Late Pleistocene of Italy and Western Europe. In: von Koenigswald, W., Werdelin, L. (Eds.), Mammalian Migration and Dispersal Events in the European Quaternary. Courier Forschungsinstitut Senckenberg 153, 59e68. Scott, B., Bates, M., Bates, R., Conneller, C., Pope, M., Shaw, A., Smith, G., 2014. A new view from La Cotte de St Brelade, Jersey. Antiquity 88, 13e29. Shapiro, B., Drummond, A.J., Rambaut, A., Wilson, M.C., Matheus, P.E., Sher, A.V., Pybus, O.G., Gilbert, M.T.P., Barnes, I., Binladen, J., Willerslev, E., Hansen, A.J.,
Baryshnikov, G.F., Burns, J.A., Davydov, S., Driver, J.C., Froese, D.G., Harington, C.R., Keddie, G., Kosintsev, P., Kunz, M.L., Martin, L.D., Stephenson, R.O., Storer, J., Tedford, R., Zimov, S., Cooper, A., 2004. Rise and fall of the Beringian steppe bison. Science 306, 1561e1565. Sommer, R.S., Benecke, N., 2009. First radiocarbon dates on woolly mammoth (Mammuthus primigenius) from northern Germany. Journal of Quaternary Science 24, 902e905. Strelkov, S.A., 1977. Pervaya nakhodka mamonta v podmorennykh sloyakh na Kol'skom poluostove. Trudy Zoologicheskogo Instituta AN SSSR 73, 36e39 (in Russian). Stuart, A.J., 1977. The vertebrates of the Last Cold Stage in Britain and Ireland. Philosophical Transactions of the Royal Society B: Biological Sciences 280, 295e312. Stuart, A.J., 1982. Pleistocene vertebrates in the British Isles. Longman, London. Stuart, A.J., 1983. Pleistocene Bone Caves in Britain and Ireland. Studies in Speleology 4, 9e36. Stuart, A.J., van Wijngaarden-Bakker, L.H., 1985. Quaternary vertebrates. In: Edwards, K.J., Warren, W.P. (Eds.), The Quaternary History of Ireland. Academic Press, London, pp. 221e249. Sulerzhitsky, L.D., Romanenko, F.A., 1997. Vozrast i rasselenie “mamontovoj” fauny aziatskogo zapolyar'ya (po radiouglerodnym dannym). Kriosfera Zemli 1, 12e19 (in Russian). Takahashi, K., Soeda, Y., Izuho, M., Yamada, G., Akamatsu, M., Chang, C.-H., 2006. The chronological record of the woolly mammoth (Mammuthus primigenius) in Japan, and its temporary replacement by Palaeoloxodon naumanni during MIS 3 in Hokkaido (northern Japan). Palaeogeography, Palaeoclimatology, Palaeoecology 233, 1e10. Takahashi, K., Wei, G., Uno, H., Yoneda, M., Jin, C., Sun, C., Zhang, S., Zhong, B., 2007. AMS 14C chronology of the world's southernmost woolly mammoth (Mammuthus primigenius Blum.). Quaternary Science Reviews 26, 954e957. Tong, H.-W., 2004. Paleoenvironmental significance of Coelodonta in different fossil assemblages. Acta Anthropologica Sinica 23 (Suppl.), 306e314 (in Chinese, with English summary). Tong, H., 2010a. Proboscidean fossil records in Nihewan Basin, North China. In: Lacombat, F., Mol, D. (Eds.), The Vth International Conference on Mammoths rie vol. 3, 173e174. and Their Relatives. Quaternaire, Hors-se Tong, H., 2010b. New materials of Mammuthus trogontherii (Proboscidea, Mammalia) of Late Pleistocene age from Yuxian, Hebei. Quaternary Sciences 30, 307e318 (in Chinese, with English abstract). Ukkonen, P., Lunkka, J.P., Jungner, H., Donner, J., 1999. New radiocarbon dates from Finnish mammoths indicating large ice-free areas in Fennoscandia during the Middle Weichselian. Journal of Quaternary Science 14, 711e714. Ukkonen, P., Aaris-Sørensen, K., Arppe, L., Clark, P.U., Daugnora, L., Lister, A.M., ~ ugas, L., Seppa €, H., Sommer, R.S., Stuart, A.J., Wojtal, P., Zupins, I., 2011. Woolly Lo mammoth (Mammuthus primigenius Blum.) and its environment in northern Europe during the last glaciation. Quaternary Science Reviews 30, 693e712. Vartanyan, S.L., Arslanov, K.A., Karhu, J.A., Possnert, G., Sulerzhitsky, L.D., 2008. Collection of radiocarbon dates on the mammoths (Mammuthus primigenius) and other genera of Wrangel Island, northeast Siberia, Russia. Quaternary Research 70, 51e59. Vereshchagin, N.K., 1959. Mlekopitayushchie Kavkaza. Istoriya formirovaniya fauny. Izdatelstvo Akademii Nauk SSSR, Moskva, Leningrad (in Russian). Vereshchagin, N.K., Tikhonov, A.N., 1990. Ekster'er mamonta. Akademiya Nauk SSSR, Sibirskoye otdelenie, Yakutskaya respublikanskaya kommissiya po izucheniyu mamontovoj fauny, Yakutsk. € ro € s, I., 1981. The skulls of Mammoth in Hungary. Fragmenta Mineralogica et Vo Palaeontologica 10, 97e106. Wei, G.B., Taruno, H., Kawamura, Y., Jin, C.Z., 2006. Pliocene and Early Pleistocene Primitive Mammoths of Northern China: Their revised taxonomy, biostratigraphy and evolution. Journal of Geosciences, Osaka City University 49, 59e101. Wei, G.B., Hu, S.M., Yu, K.F., Hou, Y.M., Li, X., Jin, C.Z., Wang, Y., Zhao, J.X., Wang, W.H., 2010. New materials of the steppe mammoth, Mammuthus trogontherii, with discussion on the origin and evolutionary patterns of mammoths. Science China, Earth Science 53, 956e963. Widga, C., Lengyel, S., Saunders, J., Hodgins, G., Walker, J.D., 2014. Proboscidean biogeography in midwestern North America, with a comment on the extinction of Mammut americanum. In: Kostopoulos, D.S., Vlachos, E., Tsoukala, E. (Eds.), Abstract Book of the VIth International Conference on Mammoths and Their Relatives. Scientific Annals of the School of Geology, Aristotle University of Thessaloniki Special Volume 102, 221. Wojtal, P., 2007. Zooarchaeological studies of the Late Pleistocene sites in Poland. Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, w. Krako Yesner, D.R., Veltre, D.W., Crossen, K.J., Graham, R.W., 2005. 5,700-year-old Mammoth Remains from Qagnax Cave, Pribilof Islands, Alaska. In: Agenbroad, L.D., Symington, R.L. (Eds.), 2nd World of Elephants Congress. Short Papers and Abstracts. Mammoth Site of Hot Springs, South Dakota, pp. 200e204. Zhylkibaev, K. Zh, 1975. Drevnie slony Kazakhstana. Izdatelstova Nauka Kazakhskoj SSR, Alma-Ata (in Russian).