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AccueilNumérosvol.33/1The bears of the European steppe:...

The bears of the European steppe: a review

Les ours de la steppe européenne : un état de l’art
Johann Galdies
p. 47-62

Résumé

Bears exhibit marked evolution for Pleistocene Europe. Both lineages are thought to have arisen from etruscan bear U. etruscus in the Early Pleistocene, however their high degree of polymorphism has prevented the establishment of an accepted evolutionary scenario. Isotopic analysis and tooth morphology of fossil brown bear U. arctos suggests that it was an omnivorous opportunist. The deningeri bear U. deningeri represents the spelaean bear of the Middle Pleistocene, sharing certain morphological affinities with brown bear U. arctos (frontal bulge and face; occlusal surface of jugular teeth). Within U. deningeri, several subspecies have been distinguished as evolutionary stages leading to the speciation of the cave bear U. spelaeus, the typical spelaean bear of the Late Pleistocene, which dominates cave fossil deposits. The speloïd lineage might serve as a good chronological marker for Pleistocene stratigraphic levels. There are several morphologically distinct lineages within U. spelaeus “sensu lato”, of controversial taxonomic status. Herbivorous feeding habits for U. spelaeus “s.l.” have been inferred from morphology (tooth, skull, jaw), demographics, and stable isotope analysis. This dietary difference between brown bears and cave bears shows that ecological competition was probably limited between both types. Paleo-genetic studies suggest that cave bears gradually lowered their reproductive rate (between 52,800 and 27,800 y BP) which led to their extinction at the onset of the last glacial maximum. Climatic changes are the main suggested causes responsible for the extinction of U. spelaeus.

Les ours présentent une évolution marquée pour l’Europe du Pléistocène. On pense que les deux lignées sont issues de l’ours étrusque U. etruscus au Pléistocène inférieur, mais leur degré élevé de polymorphisme a empêché l’établissement d’un scénario évolutif accepté. L’analyse isotopique et la morphologie des dents de l’ours brun fossile U. arctos suggèrent qu’il s’agissait d’un omnivore opportuniste. L’ours de Deninger U.deningeri représente l’ours spéléen du Pléistocène moyen, partageant certaines affinités morphologiques avec l’ours brun U. arctos (renflement frontal et face; surface occlusale des dents jugales). Au sein d’Ursus deningeri, plusieurs sous-espèces ont été distinguées comme des stades évolutifs conduisant à la spéciation de l’ours des cavernes U. spelaeus, l’ours spéléen typique du Pléistocène supérieur, qui domine les dépôts fossiles des cavernes. La lignée spéloïde pourrait servir de bon marqueur chronologique pour les niveaux stratigraphiques du Pléistocène. Il existe plusieurs lignées morphologiquement distinctes au sein de U. spelaeus «sensu lato», de statut taxonomique controversé. Des habitudes alimentaires herbivores de l’U. spelaeus «s.l.» ont été déduits par la morphologie (dent, crâne, mâchoire), la démographie et l’analyse des isotopes stables. cette différence alimentaire entre les ours bruns et les ours des cavernes montre que la concurrence écologique était probablement limitée entre les deux types. Des études paléogénétiques suggèrent que les ours des cavernes ont progressivement abaissé leur taux de reproduction (entre 52800 et 27800 ans BP), ce qui a conduit à leur extinction au début du dernier maximum glaciaire. Il est suggéré que les changements climatiques sont les causes principales de l’extinction de l’U. spelaeus.

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The author would like to thank Clara Galdies for translating all the necessary text to French, and Robin Liesens for allowing his palaeoart to be included in this publication. The author is indebted to the anonymous referees and journal editors for improving the quality of this manuscript through their valid suggestions.

1 - Introduction

1The family Ursidae (Ursidae Fischer, 1817) belongs to the order Carnivora, class Mammalia (commonly known as bears). Eight species of bears are extant; these species are widespread and appear in a wide variety of habitats throughout the Northern Hemisphere and partially in the Southern Hemisphere. Common characteristics of modern bears include large bodies with stocky legs, long snouts, small rounded ears, shaggy hair, plantigrade paws with five non-retractile claws, and short tails. The  story of bears started 38 Ma BP – the oldest known bear, of the subfamily Amphicyondontinae, is Parictis from Late Eocene North America (Kemp, 2005) – and spans all the world’s contents (except Australia and Antarctica). It deals with animals which have adapted to a variety of habitats ranging from oceanic ice flows to tropical cloud forests, and which have occupied different ecological niches like marine hypercarnivores and strict low-metabolic monotypic herbivores. However, this work only summarises the history of bears in Europe during the Pleistocene Epoch (fig. 1).

2Within Europe, bears underwent a very complex and interesting biological evolution during the Pleistocene, and their remains represent an important and substantial part of the fossil record in the Middle and Upper Pleistocene deposits in Europe, with the cave bear (Ursus spelaeus “sensu lato”) having one of the most extensive fossil records amongst the megafaunal species of the Pleistocene (Kurtén, 1976 ; Musil, 1980 ; Baryshnikov, 1998, 2007 ; Rabeder, 1999 ; Rabeder et al., 2000), with a high degree of polymorphism (Grandal-D’Anglade & LópezGonzález, 2005) to boot. This is not only the result of a very pronounced sexual dimorphism (although Withalm (2001) and Athen et al. (2005) contest this) and a marked inter-individual morphological variability amongst bears (Koby, 1949 ; Kurtén, 1955, 1976 ; Auguste, 1995 ;

3Grandal-D’Anglade & Vidal Romaní, 1997 ; GrandalD’Anglade & López-González, 2005 ; Toškan, 2006 ; Goubel, 2011 ; Baryshnikov & Puzachenko, 2017), but also because of their high diversity throughout the Pleistocene (tab. 1).

4The earliest Pleistocene ursid to appear in the fossil record of Europe is Ursus etruscus Cuvier, 1823; which is characteristic of the Late Pliocene and Early Pleistocene Europe and North Africa. It is hypothesised that the U. etruscus lineage then split into two branches (Argant, 2009 ; Barlow et al., 2018 ; Krause et al., 2008).

  1. The arctoid lineage, with Ursus arctos Linnaeus, 1758; commonly known as the brown bear. It is still present in Asia, Europe and North America.

  2. The speloïd lineage, with Ursus deningeri Von Reichenau, 1904, of the Middle Pleistocene. It was present in Europe and Asia. Ursus deningeri evolved into Ursus spelaeus Rosenmüller and Heinroth, 1794, at the end of the Middle Pleistocene; but there are no clear separations between these two species. U. spelaeus is commonly known as the cave bear. Today extinct, many are the speculations concerning the causes of its extinction (e.g. Barnosky et al., 2004 ; Cooper et al., 2015 ; Mondanaro et al., 2019).

5The origin of the speloïd lineage and its relation with the arctoid lineage are still unclear. The high degree of polymorphism observed within the arctoid and speloïd bear specimens has prevented the establishment of a single, universally accepted evolutionary scenario. Many authors share the idea of an etruscan origin of the speloïd lineage (Erdbrink, 1953 ; Kurtén, 1976 ; Argant, 2009); while Mazza & Rustioni (1994) have suggested that the speloïd lineage might have diverged from an archaic arctoid stock. Spelaean bears are an exclusively Eurasian lineage of ursids (Baryshnikov, 2008 ; Rabeder et al., 2004, 2010) that had become established as a distinct lineage by the start of the Middle Pleistocene (Kurtén, 1976 ; Moullé, 1992 ; MadurellMalapeira et al., 2009) following their supposed separation from the arctoid lineage (Kurtén, 1976 ; Torres Pérez-Hidalgo, 1992 ; Rabeder et al., 2010). Krause et al. (2008) have placed this divergence at 2.8 Ma BP; however, according to molecular data, the arctoid and speloïd lineages may have diverged later from their common ancestor, roughly between 1.2 and 1.6 Ma BP (Hänni et al., 1994 ; Loreille et al., 2001 ; Bon et al., 2008). U. deningeri, the archaic spelaean bear, led rise to the Late Pleistocene spelaean taxon U. spelaeus (Orlando et al., 2002 ; Hofreiter et al., 2002, 2004 ; Rabeder & Hofreiter, 2004 ; Knapp et al., 2009) toward the end of Middle Pleistocene (Torres Pérez-Hidalgo, 1992 ; Rabeder et al., 2000). U. spelaeus persisted until the end of the Late Pleistocene c. 27 ka BP (Pacher & Stuart, 2009 ; Bocherens et al., 2014 ; Martini et al., 2014 ; Baca et al., 2016 ; Terlato et al., 2018).

6Throughout the Pleistocene Epoch, the abrupt succession of climatic events which occurred in Europe inevitably resulted in profound changes in the floral landscape. Alternations between open landscapes on one hand and forest environments on the other reflected the highly volatile climatic context of Europe during the Pleistocene (many continental sedimentary sequences rich in pollen make it possible to precisely trace the development of these floral communities, e.g. Woillard, 1978). It is thought that mammalian populations, particularly herbivores, were directly influenced by these fluctuations (Kahlke, 1999) – and as a result of these palaeoenvironmental changes, ursids displayed morphological, physiological and ethological adaptations which mainly affected their body dimensions and proportions (Kahlke, 1999). Large-scale migration phenomena, involving significant faunal turnover, also occurred (Kahlke, 1999). Indeed, climatic fluctuations resulted in the contractions and extensions of habitable areas, with each taxon being limited by its own biogeography at a specific time-period (Azzaroli et al.,1988 ; Kahlke,1999 ; Pushkina, 2007).

7Mainly during the glacial episode of the Late Pleistocene, when atmospheric parameters were generally arid, large non-permanently-glaciated swathes of Eurasia and Northern Yukon in North America consisted of a palaeo-realm which has been termed the “Mammoth Steppe” (Guthrie, 2001 ; Bocherens, 2015), described by Guthrie (1982, 1984, 1985, 1990, 2001) as a homogeneous ecosystem which extended from the westernmost European regions (French Brittany, Spain, England) to the easternmost Asian region (Siberia) and across the Bering Strait and beyond into North America, constituting “Beringia” (Harington, 2005). The “Mammoth Steppe” was characterised by a vegetal mosaic or patchwork cover dominated by grasses, mixed with temperate and boreal tree communities in the form of scrubs, groves and forests, and their associated mammalian megafauna (Guthrie, 1968, 1984, 1990 ; Zazula et al., 2002, 2003 ; Van Geel et al., 2008 ; Zimov et al., 2012). Guthrie (2001) stated that the nature of the vegetation was indicative of intensive browsing by large herbivores: the soils of the “Mammoth Steppe” were characterised by their high productivity and high nutrient content. Indeed, several Late Pleistocene deposits from Europe display a faunal biodiversity far richer than the one present in today’s northern latitudes (Sher, 1971 ; Matthews, 1982 ; Auguste, 1995 ; Kahlke, 1999 ; Mol et al., 2001 ; Van Geel et al., 2008 ; Marot, 2009): the characteristic “Mammoth Steppe” fauna was characterised by its high diversity. Truly, in terms of mammalian megafaunal diversity, Bocherens (2015) likened this ecosystem with today’s African savannah rather than to the boreal or arctic terrestrial ecosystems of today – pointing out the one major difference between the savannah and the “Mammoth Steppe” faunas being the presence of ursids.

8Among the taxa typical of the “Mammoth Steppe” of the Upper Pleistocene, Auguste (1995) cited the following exhaustive list: the woolly mammoth Mammuthus primigenius Blumenbach, 1827; the woolly rhinoceros Coelodonta antiquitatis Blumenbach, 1799; the steppe bison Bison priscus Bojanus, 1827; horse Equus sp. Linnaeus, 1758; the reindeer Rangifer tarandus Linnaeus, 1758; the elk Alces alces Linnaeus, 1758; the saiga Saiga tatarica Linnaeus, 1758; the musk ox Ovibos moschatus Zimmermann, 1780; the cave bear Ursus spelaeus Rosenmüller and Heinroth, 1794; the wolverine Gulo gulo Linnaeus, 1758; the wolf Canis lupus Linnaeus, 1758; the cave lion Panthera spelaea Goldfuss, 1810; the lynx Lynx lynx Linnaeus, 1758; the cave hyena Crocuta crocuta spelaea Goldfuss, 1832; the arctic fox Vulpes lagopus Linnaeus, 1758; etc. Several species or types of hominids were also present in some areas at some times of the “Mammoth Steppe” realm (Ripple & Van Valkenburgh, 2010 ; Turner II et al., 2013 ; Stuart & Lister, 2014), namely Homo neanderthalensis in Europe and western Asia until 40 ka BP, being replaced by anatomically modern humans (H. sapiens) around 35 ka BP (Higham et al., 2014).

9The end of the Late Pleistocene marked the almost complete disappearance of the “Mammoth Steppe”, causing the mass extinction of many large mammals (Stuart & Lister, 2007), including the speloïd lineage of bears. Numerous discussions have highlighted the several explanations employed to explain the cause for the disappearance of the “Mammoth Steppe” (Martin, 1967 ; Martin & Klein, 1984 ; Barnosky et al., 2004 ; Hofreiter & Stewart, 2009) and many (but not all) of its associated megafauna.

Fig. 1: A representation of the evolutionary origin and history during the Middle/Late Pleistocene of the arctoid-speloïd lineage.

Fig. 1: A representation of the evolutionary origin and history during the Middle/Late Pleistocene of the arctoid-speloïd lineage.

Asian distribution is represented with blue, European distribution with yellow, and both European and Asian distribution in green (Argant, 2009 ; Athen & Pfretzschner, 2005 ; Baca et al., 2016 ; Barlow et al., 2018 ; Baryshnikov, 2008 ; Baryshnikov & Foronova, 2001 ; Baryshnikov & Kalmykov, 2005 ; Baryshnikov & Puzachenko, 2011, 2019 ; Bocherens et al., 2011b, 2014 ; Bon et al., 2008, 2011 ; Curry, 2010 ; Erdbrink, 1953 ; Hofreiter et al., 2002, 2004 ; Knapp et al., 2009 ; Kohn et al., 1995 ; Krause et al., 2008 ; Kurtén, 1976 ; Loreille et al., 2001 ; Madurell-Malapeira et al., 2009 ; Martini et al., 2014 ; Moullé, 1992 ; Münzel et al., 2011 ; Orlando et al., 2002 ; Pacher & Stuart, 2009 ; Rabeder et al., 2000, 2004, 2010 ; Rabeder & Hofreiter, 2004 ; Sabol et al., 2013 ; Stiller et al., 2009, 2013 ; Taberlet & Bouvet, 1994 ; Tamako et al., 2001 ; Tchernov & Tsoukala, 1997 ; Terlato et al., 2018 ; Torres Pérez-Hidalgo, 1992 ; Zdansky, 1928). Several alternative scenarios have been postulated, namely: (A) U. etruscus is the ancestor of the arctoid lineage (Argant, 1991 ; Ehrenberg, 1929 ; Erdbrink, 1953 ; Kurtén, 1968) – alternatively (B) U. etruscus became extinct without direct descent and the arctoid-speloïd lineage derives from cf. U. minimus (Mazza & Rustioni 1994). (C) U. dolinensis is nested within U. arctos (Argant, 2009 ; Goubel, 2011). (D) Western and eastern lineages of U. arctos diverged around 850 ka BP (Loreille et al., 2001), or more recently around 550 ka BP (Bon et al., 2008). (E) Origin of speloïd lineage is etruscan (Argant, 2009 ; Erdbrink, 1953 ; Kurtén, 1976) – alternatively (F) speloïd lineage diverged from archaic U. arctos (Mazza & Rustioni, 1994). (H) Mitochondrial studies position U. rossicus closest to U. ingressus (Pacher et al., 2009 ; Stiller et al., 2013) which in turn is synonymous with U. kanivetz in this scenario (Baryshnikov & Puzachenko, 2017) – alternatively whole-genomic studies position U. rossicus as first to diverge from U. spelaeus “s.l.” (Barlow et al., 2021). (I) Barlow et al. (2021) present evidence for distinguishing between U. ingressus and U. kanivetz, positioning U. ingressus as closer to U. spelaeus “s.s.” than to U. kanivetz. (J) The following taxa are recognised by most researchers (Baryshnikov & Puzachenko, 2011 ; Bocherens et al., 2011b ; Curry, 2010 ; Hofreiter et al., 2002, 2004 ; Orlando et al., 2002 ; Rabeder & Hofreiter, 2004 ; Rabeder et al., 2004 ; Sabol et al., 2013; Stiller et al., 2009, 2013 ; Torres Pérez-Hidalgo et al., 1991) as belonging to the same species U. spelaeus: U. spelaeus ladinicus, U. spelaeus eremus, U. spelaeus spelaeus, and U. spelaeus parvilatipedisalternatively, Barlow et al. (2021) presents wholegenomic evidence for classifying U. eremus as a distinct species from U. spelaeus. (K) inter-breeding between European U. spelaeus and U. arctos (Barlow et al., 2018). Palaeoclimatological data from: Böse et al. (2012), Gibbard & Head (2009), Gibbard et al. (2005, modified 2007), John (1997), Litt et al. (2007), McMillan (2005) and Zagwijn (1960).

Tab. 1: With Pleistocene bear remains being so extensive throughout Eurasia, it is difficult to compile an exhaustive list of all the records to date. This table lists some of the more studied finds, as identified by 1: Argant (1995), 2: Auguste (2012), 3: Baryshnikov & Foronova (2001), 4: Baryshnikov et al. (2003), 5: Bocherens et al. (2011b), 6: De Carlis et al. (2005), 7: Enloe et al. (2000), 8: Hofreiter et al. (2004), 9: Knapp et al. (2009), 10: Nagel et al. (2005), 11: Rabeder et al. (2004), 12: Sabol et al. (2013), 13: Stiller et al. (2013) and 14: Tsoukala et al. (2001).

Tab. 1: With Pleistocene bear remains being so extensive throughout Eurasia, it is difficult to compile an exhaustive list of all the records to date. This table lists some of the more studied finds, as identified by 1: Argant (1995), 2: Auguste (2012), 3: Baryshnikov & Foronova (2001), 4: Baryshnikov et al. (2003), 5: Bocherens et al. (2011b), 6: De Carlis et al. (2005), 7: Enloe et al. (2000), 8: Hofreiter et al. (2004), 9: Knapp et al. (2009), 10: Nagel et al. (2005), 11: Rabeder et al. (2004), 12: Sabol et al. (2013), 13: Stiller et al. (2013) and 14: Tsoukala et al. (2001).

2 - Ursus arctos

10As discussed above, the main candidate as the common ancestor of the arctoid-speloïd complex is U. etruscus (Ehrenberg, 1929 ; Erdbrink, 1953 ; Kurtén, 1968 ; Argant, 1991). Mazza & Rustioni (1994) rejected this hypothesis and suggested that U. etruscus, found in Europe, became extinct without direct descent; suggesting that the arctoid-speloïd complex (which eventually replaced the U. etruscus group in Europe) derives from an Asian stock of black bears (cf. U. minimus). Regardless, the earliest occurrence of U. arctos dates from the Middle Pleistocene in China (Zdansky, 1928). This archaic form, sometimes distinguished from the present forms as a separate subspecies U. arctos fossilis Goldfuss, 1821, is only definitively identified in Europe from the Middle Pleistocene (Erdbrink, 1953). However, the earlier European form U. dolinensis Garcia and Arsuaga, 2001 from the Early Pleistocene (Argant, 2009) might actually be nested within U. arctos (Goubel, 2011). The information available on the diet of the fossil representatives of brown bear suggests that it was probably an omnivorous opportunist just like the extant representatives (Herrero, 1985 ; GarcíaVázquez et al., 2018).

11Today, the brown bear is currently the most widely distributed bear in the world (fig. 2), present in Eurasia and North America (and was only extirpated from North Africa in historical times). Its large spatial distribution has resulted in several geographically isolated forms, ranked by most researchers as subspecies. This variability is mainly expressed in their body dimensions and coat hue. All of these haplotypes fit into two main mitochondrial lineages: the western lineage and the eastern lineage (Taberlet & Bouvet, 1994 ; Kohn et al., 1995 ; Loreille et al., 2001 ; Tamako et al., 2001 ; Bon et al., 2008). These two lines are supposed to have diverged during the Early Pleistocene around 850 ka BP (Loreille et al., 2001) or more recently around 550 ka BP (Bon et al., 2008). Beyond the European stock, the world population of brown bears is divided into at least five major phylogenetic lines (Tamako et al., 2001). A latitudinal morphometric gradient is observed with an increase in size from south to north (Bergmann’s Rule) and a longitudinal gradient in Eurasia where the increase of dimensions is from west to east (Kurt, 1990). Morphological variability is also expressed at the individual level and by a particularly pronounced sexual dimorphism (Mazza & Rustioni, 1994). Extant brown bears do not have a restricted diet but a diverse one, including: leaves, fruits, berries, fungi, honey, insects, fish (especially salmon), micromammals, birds, and sometimes large mammals such as deer or bison (Kurt, 1990). It has been observed that its diet is dependent on the kind of food that is available and also on the energy requirements needed at that time (Welch et al., 1997 ; Rode and Robbins, 2000). Omnivory can be inferred for Pleistocene U. arctos (dated around 30,000 14C BP to 50,000 14C BP), with δ15N values of Pleistocene U. arctos from Austria being higher than for the sympatric U. spelaeus, hypothesised to be a strict herbivore, but lower than for the cave lion, a hypercarnivore (Bocherens et al., 1997, 2011a ; Bocherens & Drucker, 2006 ; Döppes et al., 2008 ; Münzel et al., 2008). However, the same pronounced discrepancy in δ15N was not observed between sympatric U. arctos and U. spelaeus from Pleistocene and Holocene (for U. arctos only of course) deposits in Cantabria, Spain (García-Vázquez et al., 2018), which highlights the adaptability of the U. arctos with regards to its feeding behaviour.

12Second to the polar bear which can reach the weight of a tonne in winter, the largest extant bears are the Kodiak brown bears of Alaska, weighing about 780 kg (Kurt, 1990).

Fig. 2: Geographic distribution of U. arctos.Red = current range, Yellow = historical range.

Fig. 2: Geographic distribution of U. arctos.Red = current range, Yellow = historical range.

I have included the range of U. maritimus as it is generally considered to be phylogenetically nested within U. arctos.

3 - Ursus deningeri

13Ursus deningeri first occurred at the Early Pleistocene – Middle Pleistocene boundary (Moullé, 1992) and represents the typical spelaean bear of the Middle Pleistocene. Its anatomy is less robust than that of the later species U. spelaeus. It shares certain morphological affinities with  U. arctos, namely the accentuation of its frontal bulge and face (Moullé, 1992 ; Mazza & Rustioni, 1994); and the occlusal surface of the jugular teeth is much less complex than that of the later U. spelaeus and more similar to that of U. arctos (Goubel, 2011). Within U. deningeri, several subspecies have been distinguished as evolutionary stages leading to the speciation of the chrono-species U. spelaeus, including: U. deningeri deningeri Von Reichnenau, 1904; U. deningeri savini Andrews, 1922; U. deningeri suessenbornensis Soergel, 1926; U. deningeri suevicus Koy, 1951; U. deningeri romeviensis Prat, 1976; U. deningeri hercynicus Rode, 1935 and U. kudarensis Baryshnikov, 1985. Stiller et al. (2009, 2013) stated that, based on genetic data, the earliest cave bear lineage to diverge from the rest of the speloïd stock was U. kudarensis, whose fossils have been found in the Caucasus region of Eurasia; this is further confirmed by whole-genomic analysis (Barlow et al., 2018 ; 2021).

4 - Ursus spelaeus “sensu lato”

14Usually dominating faunal associations of cave deposits (Bocherens et al., 2011b), U. spelaeus remains often amount to large bone accumulations. It is thought that this is the result of U. spelaeus being more dependent on caves for hibernation than other ursid species (Kurtén, 1976 ; Fortes et al., 2016 ; Pérez-Rama et al., 2011a), and thus Late Pleistocene caves of northern Eurasia yield so many remains of U. spelaeus that probably died during their hibernation periods (Kurtén, 1976 ; Pacher & Stuart, 2009 ; Grandal D’Anglade et al., 2019).

15U. spelaeus is thought to have evolved from U. deningeri (Athen & Pfretzschner, 2005) around the Middle Pleistocene - Late Pleistocene transition (Knapp et al., 2009): the appearance of U. spelaeus seems to have occurred at the beginning of the last Eemian interglacial, at the Middle Pleistocene - Upper Pleistocene boundary (Rabeder et al., 2000), i.e. around 100 ka BP; however the date to which the biological event of U. spelaeus arising from U. deningeri is assigned is still not fully determined, but varies according to the type of data used, whether palaeontological or biomolecular (Argant, 2009). Argant (1991) proposed using the intermediate taxon U. spelaeus deningeroides Mottl, 1964 for the earliest forms. The problematic taxonomic attribution of these transitional specimens has led to the use of open nomenclature U. deningeri sub. sp. or U. spelaeus sub. sp. (Goubel, 2011), especially in cases when the age of the deposits may be difficult to establish (Virion, 2016). As mentioned above, it is generally accepted (Rabal-Garcés & Sauqué, 2015) that both U. deningeri and U. spelaeus are a single chronospecies, i.e. arbitrary divisions of a single evolutionary lineage defined on the basis of morphological change, thus making any precise differentiation between both taxa problematic (Mazza & Rustioni, 1994 ; Argant, 2001 ; Grandal-D’Anglade & López-González, 2004 ; Santi & Rossi, 2014). It can be argued that the speloïd lineage evolution might serve as a good chronological marker for Pleistocene stratigraphic levels in which these bones are found, since strong phylogenetic and ecological forcing was at play during the Quaternary. All typical spelaean characteristics appear to have been acquired by the beginning of the Eemian interglacial (PérezRamos et al., 2020; Rabeder et al., 2000).

16The fossil distribution range of U. spelaeus “sensu lato” extends from northwest Spain to the Urals and the Altai and from Belgium and the Harz region of Germany to Italy and Greece and from Crimea to North-eastern Siberia (Nordmann, 1858 – 1860 ; Grandal D’Anglade, 1993 ; Minieri et al., 1995 ; Grandal D’Anglade & Vidal Romaní, 1997 ; Enloe et al., 2000 ; Baryshnikov, 2007 ; Kosintsev, 2007 ; Knapp et al., 2009 ; Kostopoulos, 2006 ; Sher et al., 2011 ; Rabal-Garcés & Sauqué, 2015). In the large area occupied by speloïd bears, elevation, biotopes and vegetation varied widely (Rabeder et al., 2006 ; Knapp et al., 2009 ; Pacher & Stuart, 2009). There would have been an east-west gradient from continental to coastal climates, which is associated with flowering time of plants (Lewandowska-Sabat et al., 2012) and types of plants (Löffler, 2003). There would also have been a northsouth gradient, which is associated with temperature and seasonality (Legates & Willmott, 1990) and plant response to climatic changes (Peñuelas et al., 2004).

17Several morphologically-distinct forms have been identified and it has been suggested that there are several different lineages within  U. spelaeus “sensu lato” (Torres Pérez-Hidalgo et al., 1991 ; Baryshnikov, 1998, 2006 ; Orlando et al., 2002 ; Hofreiter et al., 2002, 2004 ; Rabeder & Hofreiter, 2004 ; Rabeder et al., 2004, 2008, 2010 ; Hofreiter, 2005 ; Nagel et al., 2005 ; Knapp et al., 2009 ; Stiller et al., 2009, 2013 ; Baryshnikov & Puzachenko, 2011 ; Münzel et al., 2011 ; Baca et al., 2012 ; Cvetković & Dimitrijević, 2014): U. kanivetz Vereshchagin, 1973; U. ingressus Rabeder et al., 2004; U. rossicus Borissiak, 1930; U. spelaeus ladinicus Rabeder et al., 2004; U. spelaeus eremus Rabeder et al., 2004; U. spelaeus spelaeus Rosenmüller, 1794 and U. spelaeus parvilatipedis Torres, 1991 (fig. 3). The taxonomic status of these morphologically distinct groups is controversial and it where previously classified as a distinct sister clade to U. spelaeus “s.s.”]; (b) placing U. rossicus from Asia as the most divergent cave bear; and (c) classifying the European large-bodied taxa, U. spelaeus, U. ingressus and U. kanivetz, as a sister clade to the smaller bodied cave bear U. eremus thus warranting species level (Barlow et al., 2021).

18U. spelaeus spelaeus (fig. 4 A) remains have been found in West Europe while U. ingressus (fig. 4 B) had a more East European distribution, with an overlap in range between the two taxa in central Europe; the U. spelaeus ladinicus/eremus complex was confined to the Alps (Stiller et al., 2013). In some cases (e.g. Austria), the two cave bear lineages U. eremus and U. ingressus lived sympatrically for 15,000 years (Hofreiter et al., 2004). Remains of small cave bears, morphologically identified as U. rossicus (fig. 4 C), have been found in western Siberia (Baryshnikov & Foronova, 2001), a lineage which Stiller et al. (2013) suggested formed a sister group to U. ingressus. However, nuclear genetic studies have once again challenged this classification, with Barlow et al. (2021) positioning U. rossicus as the most divergent from the U. spelaeus“s.l.” stock. With morphological and genetic variation being so high within relatively small geographic regions, homogeneity in diet, habitat and life-ways seems unlikely (Bocherens et al., 2011b; Seetah et al., 2012). In common with the extant brown bear (U. arctos), U. spelaeus “sensu lato” shows high intraspecific variability, and it can be argued that the similar high variable intraspecific genetic structure of the brown bear U. arctos (Hirata et al., 2013) warrants caution when identifying putative species status for distinct cave bearhaplogroups (Stiller et al., 2013). This considerable and evident morphological variation between bear populations and the controversial nature of taxonomic affinities in cave bear groups is most clearly illustrated by the diminutive ‘Alpine’ spelaean form (Grandal D’anglade, 1993; Rossi & Santi, 2001; Hofreiter et al., 2004; De Carlis et al., 2005; Withalm, 2005; Rabeder et al., 2008; Baryshnikov & Puzachenko, 2011; Seetah et al., 2012). Typical U. spelaeus jugal teeth are large in comparison to those of U. arctos, and the dental occlusal surface is particularly complex (Goubel, 2011). The roundness has been argued that they may represent different species (Hofreiter et al., 2004 ; Rabeder & Hofreiter, 2004), but also that they are not even distinct enough to warrant subspecies status (Baryshnikov & Puzachenko, 2011). The genetic data are consistent with the principal classification above (Stiller et al., 2013) – several authors (Hofreiter et al., 2004 ; Rabeder & Hofreiter, 2004 ; Stiller et al., 2009 ; Curry, 2010 ; Sabol et al., 2013) have argued that there were two main lineages of Late Pleistocene cave bears, to the extent that they may be classified as distinct species (Bocherens et al., 2011b): these being U. spelaeus “sensu stricto” and U. ingressus, with Hofreiter et al. (2002) and Orlando et al. (2002) concluding that there are four different haplogroups: U. ingressus, U. spelaeus spelaeus, U. spelaeus eremus and U. spelaeus ladinicus (Rabeder & Hofreiter, 2004 ; Rabeder et al., 2004); meanwhile, Stiller et al. (2013) cited five different species or subspecies of U. spelaeus “sensu lato” based on morphological differences: U. spelaeus spelaeus, U. spelaeus eremus, U. spelaeus ladinicus, U. ingressus, and U. rossicus; and further noted that the distinctness between U. ingressus, U. rossicus, U. kudarensis, and U. spelaeus “s.s.” is also well supported by mitochondrial genetic data. However, the same mitochondrial genetic distinctness is not supported between U. spelaeus ladinicus and U. spelaeus eremus which appear not to vary when looking at their mitochondrial DNA. Nuclear genetics however paint a slightly different picture, mainly (a) distinguishing between U. ingressus and U. kanivetz [which, based on the earlier mitochondrial studies (Stiller et al., 2013), Austria), the two cave bear lineages U. eremus and U. ingressus lived sympatrically for 15,000 years (Hofreiter et al., 2004). Remains of small cave bears, morphologically identified as  U. rossicus (fig. 4 C), have been found in western Siberia (Baryshnikov & Foronova, 2001), a lineage which Stiller et al. (2013) suggested formed a sister group to U. ingressus. However, nuclear genetic studies have once again challenged this classification, with Barlow  etal. (2021) positioning U. rossicus as the most divergent from the U. spelaeus “s.l.” stock. With morphological and genetic variation being so high within relatively small geographic regions, homogeneity in diet, habitat and life-ways seems unlikely (Bocherens et al., 2011b; Seetah et al., 2012).

19In common with the extant brown bear (U. arctos), U. spelaeus “sensu lato” shows high intraspecific variability, and it can be argued that the similar high variable intraspecific genetic structure of the brown bear U. arctos (Hirata et al., 2013) warrants caution when identifying putative species status for distinct cave bear haplogroups (Stiller et al., 2013). This considerable and evident morphological variation between bear populations and the controversial nature of taxonomic affinities in cave bear groups is most clearly illustrated by the diminutive ‘Alpine’ spelaean form (Grandal D’anglade, 1993; Rossi & Santi, 2001; Hofreiter et al., 2004; De Carlis et al., 2005; Withalm, 2005; Rabeder et al., 2008; Baryshnikov & Puzachenko, 2011; Seetah et al., 2012).

20Typical U. spelaeus jugal teeth are large in comparison to those of U. arctos, and the dental occlusal surface is particularly complex (Goubel, 2011). The roundness of the main cusps and the multiplication of secondary cuspids and wrinkles (Rabeder, 1999) mean that occlusal surface greatly differs from the dental pattern typically encountered for carnivores (Kurtén, 1976). But the dental elements of U. spelaeus are also characterised by such a large intra-specific morphological variability that traditional morphometry cannot quantify precisely; thus, within the same population of U. spelaeus, there are differences in shape profoundly altering the outline of the teeth (Goubel, 2011). A north-south geographic separation was noted following morphometric analysis of cheek teeth from Belgium, Poland and Russia (northern assemblages) and Spain, France and Ukraine (southern sites) (Baryshnikov et al., 2003). The problem with such geographically separate assemblages is that it is not clear whether the variation has a genetic, regional, chronological or climatic basis (Seetah et al., 2012). Similar studies using assemblages on an east-to-west cline have shown variation in cheek teeth morphology between distinct geographic groups, without the possibility of confidently assigning taxonomic affinity (Baryshnikov, 2006). Research on skull and post-cranial bear elements from Italy have suggested size variation linked to chronology, with older assemblages correlated to smaller individual animals. This has been interpreted as a response to climatic change where animals increased in relative size in response to climate cooling (De Carlis et al., 2005). Size changes related to climate change have also been shown for sites in Croatia (Miracle, 2011) and Slovenia (Toškan, 2007).

21The metapodia of U. spelaeus are much more robust or plump than those of U. arctos, and, in this regard, the metapodia of U. deningeri can be described as having intermediate features, being more robust than those of U. arctos but less than those of U. spelaeus (RabalGarcés & Sauqué, 2015). According to Baryshnikov & Puzachenko (2017), the biometrical difference between brown and spelaean bears is more pronounced for the metacarpals than for the metatarsals, and they suggest that the latter are thus more ‘conservative’ and less variable over time; leading them to hypothesise that a very rapid modification of metacarpal bones occurred during an early stage of the evolutionary emergence of the speloïd lineage, perhaps a result of their occupying a distinct ecological niche (Martín-Serra et al., 2014). Spelaean bears have metapodia that show a dorso-palmar and dorsoplantar flattening (Rabal-Garcés & Sauqué, 2015), a phenomenon unobserved for U. arctos. Withalm (2001), Athen & Pfretzschner (2005) and Athen (2006) found significant morphometrical differences between metapodia belonging to the two species U. deningeri and U. spelaeus, unlike dental material (Grandal D’Anglade & López Gonzalez, 2004). Baryshnikov & Puzachenko (2017) found that metapodial bones of U. spelaeus “sensu lato” are easily distinguished from those of U. etruscus and U. arctos. However, Münzel & Athen (2009) performed a metapodial biometry study to separate the two spelaean bear types, U. spelaeus and U. ingressus, with no satisfactory results. U. spelaeus also shows robust femora and tibiae (Martín-Serra et al., 2014), an adaptation to resist axial and bending stresses (Anyonge, 1996).

22Herbivorous feeding habits for U. spelaeus “sensu lato” have been inferred from tooth, skull and jaw morphology (Kurtén, 1958, 1976 ; Richards et al., 2008 ; Figueirido et al., 2009 ; Grandal-d’Anglade, 2010 ;

23Van Heteren et al., 2009, 2013, 2016 ; PérezRamos et al., 2020) - this dietary difference between brown bears and U. spelaeus “s.l.” bears shows that ecological competition was probably limited between both types of ursids (Bocherens et al., 2011b) – and at the heart of these many debates, the ideas on the food spectrum of the diet of U. spelaeus have continued to evolve with new discoveries and technological advances. Its bone and dental anatomy have long been interpreted as an evolutionary adaptation to an increasingly herbivorous diet, until reaching an exclusive herbivorous specialisation with the most evolved forms at the end of the Late Pleistocene. These morphological features include, but are not limited to, frontonasal glabellar bulging, massive zygomatic arches, the loss of earlier pre-molars (creation of a diastema), large dimensions, conformation of juvenile teeth (Kurtén, 1976) and progressive complication of the tooth surface (Santi & Rossi, 2014). Geometric morphometry outline analysis provided information on the morphological affinities of the dental elements between current and fossil ursids. However, morphometric analysis and comparison of teeth carried out by Goubel (2011) concluded that the dental features of U. spelaeus do not confirm that it was hyper-herbivorous, but rather that it might have been slightly more omnivorous than commonly suggested, while archaic U. arctos was probably more herbivorous than extant variants. That said, Pérez-Ramos et al. (2020) have observed an adaptation for applying pressure on molars during biting scenarios (rather than on the canines and fourth premolars as is typical with more carnivorous bears) in U. spelaeus “s.l.”. In addition, U. spelaeus was generally larger than U. deningeri, suggesting increased herbivory; with weight-supporting elements being strengthened, especially the anterior extremities, suggesting they would have been increasingly used by U. spelaeus to dig for roots and other food (Athen, 2006 ; Santi & Rossi, 2014).

24Stiner et al. (1998) claimed that the sex ratio (female/ male) in cave bears is less pronounced than those of other extant bears (Van Heteren et al., 2009), and that this might be indicative of a strong dependence upon highly seasonal food resources. Bocherens et al. (2011b) argued that the high proportion of males in hibernating populations of cave bears might be evidence for a depletion of plant food availability during winter – something which would not be observed for male carnivorous bears which do not need to hibernate, as opposed to females who, independent of their diet, give birth only during hibernation (Stiner, 1999). Hibernation is a seasonal decrease of metabolic activity and food intake during the winter due to seasonal food shortage, a behaviour widely spread among extant ursids and was also even more pronounced in the case of cave bears (Kurtén, 1976; Stiner, 1998).

25The stable isotopic compositions of bulk collagen in U. spelaeus “s.l.” from a large number of sites support the hypothesis of strict herbivory (Bocherens et al., 1994, 1997, 2001, 2006, 2011b, 2013, 2014 ; Bocherens, 2000, 2008,

262015, 2018; Bocherens & Drucker, 2003 ; FernandezMosquera, 1998 ; Stiner, 1999 ; Fox-Dobbs et al., 2007 ; Barnett et al., 2009 ; Sabol et al., 2013 ; Münzel et al., 2014 ; Krajcarz et al., 2016 ; Naito et al., 2020). This is in total contrast with modern brown bears as well as fossil U. arctos (Naito et al., 2016), with isotopic evidence indicating that, whenever the two species co-occurred, Pleistocene brown bears occupied the opposite (i.e. high animal protein) end of the dietary spectrum to that of U. spelaeus “sensu lato” (Bocherens & Drucker, 2006 ; Münzel et al., 2008 ; Bocherens et al., 2011b). However, despite the evidence that all spelaean bears were (relatively) herbivorous (i.e. that they consumed less animal protein than U. arctos), firm evidence shows an ecological difference between  U. eremus and U. ingressus, with both spelaean types feeding on different plant types and/or plants from different habitats (Bocherens et al., 2011b). To date, direct 14C dates are not precise enough to sort out whether U. eremus and U. ingressus were chronologically isolated or not. Contrastingly, isolated reports of high δ15N levels have also been used to demonstrate that cave bears were omnivorous or even carnivorous (Hilderbrand et al., 1996 ; Richards et al., 2008 ; Naito et al., 2016). However, specimens with high δ15N values have low stable carbon isotopic (δ13C) values that are not found in coeval predators but rather approximate the values observed in Late Pleistocene woolly mammoth and horse (Bocherens, 2008; Bocherens et al., 2011b; Drucker et al., 2015, 2017; Naito et al., 2016; Wissing et al., 2019). In addition, the high δ15N may also be a consequence of urea recycling during hibernation (Fernández Mosquera et al., 2001 ; Bocherens, 2004 ; Grandal D’Anglade & Fernández Mosquera, 2008 ; Pérez-Rama et al., 2011b ; GrandalD’Anglade et al., 2019) or the consumption of δ15 Nrich plants as a response to the lack of competition by mammoths in such locations (Naito et al., 2020).

27Yet strict herbivory for cave bears is not supported by dental micro-surface analyses (Pinto Llona, 2006 ; Peigné et al., 2009), but rather suggest that U. spelaeus had a more carnivorous diet than is otherwise proposed. Furthermore, puncture marks found on cave bear bones have been shown to be made by other cave bears, which may point to bone gnawing for micronutrients (Pinto Llona, 2006 ; Quilès et al., 2006 ; RabalGarcés et al., 2012).

28Mitochondrial DNA studies by Knapp et al. (2009) and by Stiller et al. (2010, 2013) suggested that a decline in the cave bear population occurred between 52,800 and 27,800 BP, which gradually lowered its reproductive rate and led to its extinction. Radiocarbon dating suggests that the extinction happened c. 27,800 BP, before the end of the Weichselian or Würm glaciation (Pacher & Stuart, 2009; Münzel et al., 2011 ; Bocherens et al., 2014). Cave bear populations may have declined from east to west (Bon et al., 2011 ; Stiller et al., 2013). Cave bears became totally extinct around 25,000 BP (Pacher & Stuart, 2009; Bocherens et al., 2014).

29Sandom et al. (2014) described West Europe at the end the Weichselian/Würm glacial as an extinction hotspot. Climatic cooling and inferred decreased vegetation productivity at the end of the Late Pleistocene were suggested causes responsible for the extinction of U. spelaeus and the other extinct Eurasian megafauna (Lascu & Puscas, 2002 ; Orlando et al., 2002 ; Barnosky et al., 2004 ; Pacher & Stuart, 2009 ; NoguésBravo et al., 2010). Many have argued that it is unlikely that their extinction was brought about by anthropogenic causes (Pacher & Stuart, 2009 ; Curry, 2010). However, other authors argued that climatic changes alone cannot explain the extinction of the spelaean bears, and that anthropogenic factors also contributed to their extinction (Grayson & Delpech, 2003 ; Münzel & Conard, 2004 ; Germonpré & Hämäläinen, 2007 ; Stiller et al., 2010 ; Münzel et al., 2011 ; Sandom et al., 2014) – either a direct result from hunting, or as a result of competition for caves (which are necessary for female cave bears). In fact, several cave bear remains show signs of hunting by humans (Münzel et al., 2011), which supports the hypothesis that humans and cave bears were in competition for resources and that the cave bear decline was caused by anthropogenic factors (Münzel & Conard, 2004). Based on genetic discoveries from sites in North Spain, which have shown that remains of cave bears (but significantly not of brown bears) from specific caves often belonged to unique and exclusive lineages of closely related individuals, Fortes et al. (2016) suggest that cave bears might have exhibited homing behaviour, i.e. having bears return to their caves of birth to hibernate. If this was indeed the case, having access to their caves of birth hindered by any cause, whether climatic or anthropological or both, would have made cave bears more susceptible to extinction than brown bears, and disturbance in the accessibility of ancestral caves might be one of the reasons which led to cave bear, but not brown bear, extinction.

30Yet in some sense, the biological legacy of cave bears endures. Genetic analysis (Barlow et al., 2018) has also revealed that inter-breeding between cave bears and bbrown bears occurred in the Pleistocene, with gene flow being observed in both directions. Brown bears have alleles derived from cave bears, with this introgression being highest for brown bears from Late Pleistocene Austria which were contemporaneous with U. spelaeus (c. 2.4%); while among extant brown bears, the spelaean bear admixture is highest in the Georgian brown bear (≥1.8%), intermediate in Western European brown bears (1.3 - 1.4%), and lowest in the Russian and American brown bears (0.9 - 1.0%). This is expected since spelaean bears were never present in North America. Brown bear alleles are also reported from European spelaean bear genomes, but not from U. kudarensis from the Caucasus region, suggesting that the first instance of interbreeding between brown and spelaean bears post-dates the divergence of the Caucasian U. kudarensis from the rest of the Eurasian speloïd stock.

Fig. 3: Palaeoartistic reconstructions of several Pleistocene bear taxa, from left to right: U. arctos (silhouette), U. dolinensis, U. deningeri, U. kudarensis, U. ingressus, U. rossicus, U. spelaeus ladinicus, U. (spelaeus) eremus, U. spelaeus spelaeus. Produced by Robin Liesens ©, and reproduced here with his permission.

Fig. 3: Palaeoartistic reconstructions of several Pleistocene bear taxa, from left to right: U. arctos (silhouette), U. dolinensis, U. deningeri, U. kudarensis, U. ingressus, U. rossicus, U. spelaeus ladinicus, U. (spelaeus) eremus, U. spelaeus spelaeus. Produced by Robin Liesens ©, and reproduced here with his permission.

Fig. 4: Geographic distribution of (A) U. spelaeus s.s., (B) U. ingressus, and (C) U. rossicus In the case of figure 4 A, solid black colour represents U. spelaeus spelaeus and U. spelaeus ladinicus while U. (spelaeus) eremus (whose classification as a subspecies of U. spelaeus is argued against by Barlow et al., 2021) is represented by horizontal bars. The diagonal bars in figure 4 A indicate cave bear remains which have not been identified to subspecies level but are generally considered to fall within the U. spelaeus taxon. Records are listed in table 1.

Fig. 4: Geographic distribution of (A) U. spelaeus s.s., (B) U. ingressus, and (C) U. rossicus In the case of figure 4 A, solid black colour represents U. spelaeus spelaeus and U. spelaeus ladinicus while U. (spelaeus) eremus (whose classification as a subspecies of U. spelaeus is argued against by Barlow et al., 2021) is represented by horizontal bars. The diagonal bars in figure 4 A indicate cave bear remains which have not been identified to subspecies level but are generally considered to fall within the U. spelaeus taxon. Records are listed in table 1.
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Table des illustrations

Titre Fig. 1: A representation of the evolutionary origin and history during the Middle/Late Pleistocene of the arctoid-speloïd lineage.
Légende Asian distribution is represented with blue, European distribution with yellow, and both European and Asian distribution in green (Argant, 2009 ; Athen & Pfretzschner, 2005 ; Baca et al., 2016 ; Barlow et al., 2018 ; Baryshnikov, 2008 ; Baryshnikov & Foronova, 2001 ; Baryshnikov & Kalmykov, 2005 ; Baryshnikov & Puzachenko, 2011, 2019 ; Bocherens et al., 2011b, 2014 ; Bon et al., 2008, 2011 ; Curry, 2010 ; Erdbrink, 1953 ; Hofreiter et al., 2002, 2004 ; Knapp et al., 2009 ; Kohn et al., 1995 ; Krause et al., 2008 ; Kurtén, 1976 ; Loreille et al., 2001 ; Madurell-Malapeira et al., 2009 ; Martini et al., 2014 ; Moullé, 1992 ; Münzel et al., 2011 ; Orlando et al., 2002 ; Pacher & Stuart, 2009 ; Rabeder et al., 2000, 2004, 2010 ; Rabeder & Hofreiter, 2004 ; Sabol et al., 2013 ; Stiller et al., 2009, 2013 ; Taberlet & Bouvet, 1994 ; Tamako et al., 2001 ; Tchernov & Tsoukala, 1997 ; Terlato et al., 2018 ; Torres Pérez-Hidalgo, 1992 ; Zdansky, 1928). Several alternative scenarios have been postulated, namely: (A) U. etruscus is the ancestor of the arctoid lineage (Argant, 1991 ; Ehrenberg, 1929 ; Erdbrink, 1953 ; Kurtén, 1968) – alternatively (B) U. etruscus became extinct without direct descent and the arctoid-speloïd lineage derives from cf. U. minimus (Mazza & Rustioni 1994). (C) U. dolinensis is nested within U. arctos (Argant, 2009 ; Goubel, 2011). (D) Western and eastern lineages of U. arctos diverged around 850 ka BP (Loreille et al., 2001), or more recently around 550 ka BP (Bon et al., 2008). (E) Origin of speloïd lineage is etruscan (Argant, 2009 ; Erdbrink, 1953 ; Kurtén, 1976) – alternatively (F) speloïd lineage diverged from archaic U. arctos (Mazza & Rustioni, 1994). (H) Mitochondrial studies position U. rossicus closest to U. ingressus (Pacher et al., 2009 ; Stiller et al., 2013) which in turn is synonymous with U. kanivetz in this scenario (Baryshnikov & Puzachenko, 2017) – alternatively whole-genomic studies position U. rossicus as first to diverge from U. spelaeus “s.l.” (Barlow et al., 2021). (I) Barlow et al. (2021) present evidence for distinguishing between U. ingressus and U. kanivetz, positioning U. ingressus as closer to U. spelaeus “s.s.” than to U. kanivetz. (J) The following taxa are recognised by most researchers (Baryshnikov & Puzachenko, 2011 ; Bocherens et al., 2011b ; Curry, 2010 ; Hofreiter et al., 2002, 2004 ; Orlando et al., 2002 ; Rabeder & Hofreiter, 2004 ; Rabeder et al., 2004 ; Sabol et al., 2013; Stiller et al., 2009, 2013 ; Torres Pérez-Hidalgo et al., 1991) as belonging to the same species U. spelaeus: U. spelaeus ladinicus, U. spelaeus eremus, U. spelaeus spelaeus, and U. spelaeus parvilatipedisalternatively, Barlow et al. (2021) presents wholegenomic evidence for classifying U. eremus as a distinct species from U. spelaeus. (K) inter-breeding between European U. spelaeus and U. arctos (Barlow et al., 2018). Palaeoclimatological data from: Böse et al. (2012), Gibbard & Head (2009), Gibbard et al. (2005, modified 2007), John (1997), Litt et al. (2007), McMillan (2005) and Zagwijn (1960).
URL http://journals.openedition.org/quaternaire/docannexe/image/16605/img-1.jpg
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Titre Tab. 1: With Pleistocene bear remains being so extensive throughout Eurasia, it is difficult to compile an exhaustive list of all the records to date. This table lists some of the more studied finds, as identified by 1: Argant (1995), 2: Auguste (2012), 3: Baryshnikov & Foronova (2001), 4: Baryshnikov et al. (2003), 5: Bocherens et al. (2011b), 6: De Carlis et al. (2005), 7: Enloe et al. (2000), 8: Hofreiter et al. (2004), 9: Knapp et al. (2009), 10: Nagel et al. (2005), 11: Rabeder et al. (2004), 12: Sabol et al. (2013), 13: Stiller et al. (2013) and 14: Tsoukala et al. (2001).
URL http://journals.openedition.org/quaternaire/docannexe/image/16605/img-2.jpg
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Titre Fig. 2: Geographic distribution of U. arctos.Red = current range, Yellow = historical range.
Légende I have included the range of U. maritimus as it is generally considered to be phylogenetically nested within U. arctos.
URL http://journals.openedition.org/quaternaire/docannexe/image/16605/img-3.jpg
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Titre Fig. 3: Palaeoartistic reconstructions of several Pleistocene bear taxa, from left to right: U. arctos (silhouette), U. dolinensis, U. deningeri, U. kudarensis, U. ingressus, U. rossicus, U. spelaeus ladinicus, U. (spelaeus) eremus, U. spelaeus spelaeus. Produced by Robin Liesens ©, and reproduced here with his permission.
URL http://journals.openedition.org/quaternaire/docannexe/image/16605/img-4.jpg
Fichier image/jpeg, 27k
Titre Fig. 4: Geographic distribution of (A) U. spelaeus s.s., (B) U. ingressus, and (C) U. rossicus In the case of figure 4 A, solid black colour represents U. spelaeus spelaeus and U. spelaeus ladinicus while U. (spelaeus) eremus (whose classification as a subspecies of U. spelaeus is argued against by Barlow et al., 2021) is represented by horizontal bars. The diagonal bars in figure 4 A indicate cave bear remains which have not been identified to subspecies level but are generally considered to fall within the U. spelaeus taxon. Records are listed in table 1.
URL http://journals.openedition.org/quaternaire/docannexe/image/16605/img-5.jpg
Fichier image/jpeg, 46k
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Johann Galdies, « The bears of the European steppe: a review »Quaternaire, vol.33/1 | 2022, 47-62.

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Johann Galdies, « The bears of the European steppe: a review »Quaternaire [En ligne], vol.33/1 | 2022, mis en ligne le 01 mars 2024, consulté le 15 février 2025. URL : http://journals.openedition.org/quaternaire/16605 ; DOI : https://doi.org/10.4000/quaternaire.16605

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Johann Galdies

Unité Evo-Eco-Paléo (EEP) - UMR 8198, CNRS / Université de Lille - Sciences et Technologies, Batiment SN5, Cité Scientifique, FR-59655 VILLENEUVE D’ASCQ ; Department of Geosciences, Room 211B, Maths & Physics Building, University of Malta, Msida, MT-MALTA. Email: johann.galdies@um.edu.mt

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