Roman Croitor
email: romancroitor@europe.com

A new form of wapiti Cervus canadensis Erxleben, 1777 (Cervidae, Mammalia) from the Late Pleistocene of France

Keywords: deer, antlers, Last Glacial Maximum, systematics, megafauna, refugium

A well-preserved antlered braincase of wapiti (Cervus canadensis) from the Late Pleistocene of Saint-Hippolyte (Puy de Dôme, France) is described herein. The specific morphology of antlers suggests peculiar adaptations to the open landscapes of periglacial tundra-steppe that permitted to identify a new fossil subspecies Cervus canadensis combrayicus n. ssp. The specimen attests the occurrence of Cervus canadensis in the paleontological record of Western Europe and helps to clarify the systematical position of some disputed findings of Cervus from Western Europe. The revised systematic position of some fossil and sub-fossil cervid findings reveals the paleobiogeographic story of wapiti in Western Europe during the Last Glacial Maximum and the postglacial time. The treeless Alpine altitudes and Sweden are proposed as the glacial refugia for the last European wapiti. The medium-sized deer from Capri Island is regarded as an insular dwarfed wapiti C. canadensis tyrrhenicus closely related to the continental form C. canadensis palmidactyloceros from the alpine refugium.

Introduction

The occurrence of another "elaphine" deer species close to modern Cervus canadensis in the paleontological record of Europe was a subject of long-lasting debates since [START_REF] Owen | A History of British Fossil Mammals, and Birds[END_REF] and then [START_REF] Pomel | Catalogue méthodique et descriptif des vertébrés fossiles découverts dans le bassin hydrographique de la Loire[END_REF] reported fossil remains of a large 2 deer similar to American wapiti. The reports of wapiti from Paleolithic sites of Western Europe were quite frequent over the next decades [START_REF] Gervais | Notes sur différentes espèces de vertébrés fossiles observés pour la plupart dans le midi de la France[END_REF][START_REF] Gervais | Rapport sur les découverts faites dans la Grotte de Loubeau, pres Melle[END_REF][START_REF] Rivière | Découverte d'un squelette humain de l'époque paléolithique dans les cavernes des Baoussé-Roussé dites Grottes de Menton[END_REF][START_REF] Rivière | Les parois gravées et peintes de la grotte de la Mouthe (Dordogne)[END_REF][START_REF] Belgrand | La Seine: I. Le bassin parisien aux âges antéhistoriques[END_REF][START_REF] De Rance | Notes on the Vale of Clwyd Caves[END_REF][START_REF] Martin | Le bassin du Pignon et sa faune: note sur la vallée du Buëch[END_REF].

Nonetheless, a conservative and more cautious approach to the systematics of the "elaphine" deer (the species complex of Cervus elaphus and Cervus canadensis) dominated in the zoological literature of the second half of 20th century, where American wapiti and Asian maral and izubr stags were considered as subspecies of common red deer Cervus elaphus [START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF][START_REF] Ellerman | Checklist of Palaearctic and Indian Mammals, 1758-1946[END_REF][START_REF] Flerov | Musk Deer and Deer. The Fauna of USSR[END_REF][START_REF] Sokolov | Hoofed Animals (Orders Perissodactyla and Artiodactyla). Fauna of the USSR[END_REF][START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF]. It is still possible to find this type of approach in some recent publications, which regard the subspecies of red deer and wapiti as informal groups "elaphoids" and "wapitoids" [START_REF] Meiri | Subspecies dynamics in space and time: A study of the red deer complex using ancient and modern DNA and morphology[END_REF]. The caution apparently was produced by lack of detailed morphological, biological, and ethological studies, while the body size -the only readily available feature distinguishing red deer and wapiti -varies greatly within the better known C. elaphus [START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF][START_REF] Flerov | Musk Deer and Deer. The Fauna of USSR[END_REF][START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF]. The Middle Pleistocene red deer subspecies C. elaphus acoronatus that has similar body size to modern wapiti is a good example of the challenge faced by researchers. Therefore, the exceptionally large remains of Cervus from the Late Pleistocene of Europe have often been regarded as an intraspecific variation of red deer [START_REF] Azzaroli | Il nanismo nei cervi insulari[END_REF]Lister, 1987a).

Although the presence of two distinct size forms of the "elaphine" deer in the Late Pleistocene of Western Europe has been recognized [START_REF] Prat | Remarques sur les cerfs contemporains des deux premiers stades würmiens[END_REF]Lister, 1987a;[START_REF] Guadelli | Les cerfs du Würm ancien en Aquitaine[END_REF], the adequate taxonomical interpretation based on this size difference was not possible due to lack of complete antlers preserving distal portions that provide the main diagnostic characters of "elaphine" deer species and subspecies [START_REF] Lydekker | On apparently new deer from North China, in the menagerie of the Duke of Bedford at Woburn Abbey[END_REF][START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF][START_REF] Flerov | Musk Deer and Deer. The Fauna of USSR[END_REF][START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF]. Therefore, the question on the occurrence of wapiti in the Late Pleistocene of Europe remained open for more than a century.

The new arguments for the reassessment of systematical criteria for "elaphine" deer were obtained much later when genetic studies revealed an important phylogenetic distance between European red deer and Asian and American wapitis [START_REF] Kuwayama | Phylogenetic relationships among European red deer, wapiti, and Sika deer inferred from mitochondrial DNA sequences[END_REF][START_REF] Polziehn | A phylogenetic comparison of red deer and wapiti using mitochondrial DNA[END_REF][START_REF] Ludt | Mitochondrial DNA phylogeography of red deer (Cervus elaphus)[END_REF][START_REF] Pitra | Evolution and phylogeny of old world deer[END_REF]. The molecular data permitted to reinstate the species status for Cervus canadensis [START_REF] Polziehn | A phylogenetic comparison of red deer and wapiti using mitochondrial DNA[END_REF], further supported by biological, ecological and ethological data [START_REF] Schonewald | Cervus canadensis and C. elaphus: North American subspecies and evaluation of clinal extremes[END_REF][START_REF] Croitor | On the presence of Late Pleistocene wapiti, Cervus canadensis Erxleben, 1777 (Cervidae, Mammalia) in the Palaeolithic site Climăuți II (Moldova)[END_REF]. of very large shed antler with brow and bez tines and an isolated large molar with strong entostyle. The same author [START_REF] Gaudry | L'existence des saïgas en France al époque Quaternaire[END_REF] reported the findings of wapiti from Laugerie-Basse (Dordogne) in association with abundant remains of a horse, as well as with remains of reindeer and saiga. [START_REF] Martin | Le bassin du Pignon et sa faune: note sur la vallée du Buëch[END_REF] described a basal fragment of antler of Cervus canadensis from the Bassin du Pignon (Hautes-Alpes) and gave some measurements of this specimen: the distance between the brow and bez tines is 23 mm, the maximal diameter of the beam between the bez and brow tines is 55 mm, the minimum diameter of the same part of beam is 40 mm. [START_REF] Belgrand | La Seine: I. Le bassin parisien aux âges antéhistoriques[END_REF] described a proximal part of antler with pedicle from Genelle in the Seine valley as Cervus (canadensis) (sic) noticing the exceptionally large size of the antler (the circumference of the pedicle is 200 mm; the antler beam circumference above the bez tine is 210 mm).

The discussion on the occurrence of the large-sized elaphine deer in France was reopened by [START_REF] Friant | Le Strongyloceros spelaeus Owen (Cerf géant des Cavernes)[END_REF][START_REF] Friant | Le Strongyloceros spelaeus Owen de Kent's Hole (Angleterre)[END_REF] who regarded the giant cave stag from Kent's Hole as an extremely specialized elaphine deer Strongyloceros spelaeus [START_REF] Owen | A History of British Fossil Mammals, and Birds[END_REF][START_REF] Friant | Le Strongyloceros spelaeus Owen de Kent's Hole (Angleterre)[END_REF] proposed to give Owen's Strongyloceros the full genus rank, but this suggestion was not supported (Lister, 1987a). [START_REF] Prat | Remarques sur les cerfs contemporains des deux premiers stades würmiens[END_REF] noticed an important and statistically meaningful difference in the dentition and third phalanx sizes recorded in the samples of the elaphine deer collected from different layers of the Paleolithic site Combe-Grenal (Dordogne).

Those layers correspond to Würm I and Würm II and have yielded different mammalian faunas that indicate alternating cold and warm climate conditions. The smaller elaphine form that comes from the "Würm I" phase is associated with typically forest fauna, while the large elaphine deer from the "Würm II" stage comes from the deposits characterized by more severe climate conditions. According to [START_REF] Prat | Remarques sur les cerfs contemporains des deux premiers stades würmiens[END_REF], this size difference may have a taxonomical significance at the subspecies level. [START_REF] Raynal | Le maar de Saint-Hippolyte (Puy de Dôme, France): datation par thermoluminescence, flores et faunes fossiles, présence humaine, climatochronologie et dynamique du système paléo-lacustre[END_REF] confirmed that the Late Pleistocene paleontological record of southwestern France has yielded the remains of two forms of "elaphine" deer distinguished by body size. According to [START_REF] Raynal | Le maar de Saint-Hippolyte (Puy de Dôme, France): datation par thermoluminescence, flores et faunes fossiles, présence humaine, climatochronologie et dynamique du système paléo-lacustre[END_REF], the relatively small red deer existed during the phase Wurm I and then was replaced by a larger elaphine deer form ("grand Elaphe") at the beginning of phase Wurm II. The antlered cervid braincase from Saint-Hippolyte belongs, according to [START_REF] Raynal | Le maar de Saint-Hippolyte (Puy de Dôme, France): datation par thermoluminescence, flores et faunes fossiles, présence humaine, climatochronologie et dynamique du système paléo-lacustre[END_REF], to the large "elaphine" form associated with "cold" fauna. [START_REF] Raynal | Le maar de Saint-Hippolyte (Puy de Dôme, France): datation par thermoluminescence, flores et faunes fossiles, présence humaine, climatochronologie et dynamique du système paléo-lacustre[END_REF] noticed the poor development of the distal antler crown in the deer from Saint-Hippolyte that differs from the multiaxial crown typical of the modern European red deer, therefore this specimen was arbitrarily described as C. elaphus acoronatus. According to [START_REF] Guadelli | Les cerfs du Würm ancien en Aquitaine[END_REF], the "larger elaphine" deer from Saint-Hippolyte is similar to the large cervid form C. elaphus ssp. from Combe-Grenale. [START_REF] Guadelli | Les cerfs du Würm ancien en Aquitaine[END_REF] reported some distinctions in dental morphology between "large elaphine" and "small elaphine" forms from Combe-Grenale, such as the relatively linguolabially broader upper molars and very frequent lingual groove in P 2 in the "large elaphine" deer. Those morphological differences are important since they indicate a stronger brachyodonty (a primitive morphological character) and a more advanced degree of molarization of P 2 (an advanced morphological character) in the "larger elaphine" deer. [START_REF] Guadelli | Les cerfs du Würm ancien en Aquitaine[END_REF] also indicated a stronger development of entostyle and a stronger developed lingual cingulum in molars of the "larger elaphine" deer, as well as a paraconid more frequently separated from parastylid in P2, a more frequent and higher degree of molarization of P4 (the metaconid and paraconid are connected and close the second dental valley) that distinguish the "large elaphine" deer from the "smaller" one. [START_REF] Guadelli | Les cerfs du Würm ancien en Aquitaine[END_REF] estimated the differences in dental morphology as taxonomically significant and proposed the species name Cervus simpicidens for the "smaller elaphine" deer from Combe-Grenale.

However, the species name Cervus simplicidens is already preoccupied by [START_REF] Lydekker | Indian Tertiary and post-Tertiary Vertebrata. Part II. Molar teeth and other remains of Mammalia[END_REF]. This is a case of primary homonymy that makes C. simplicidens

Guadelli, 1997 permanently invalid (ICZN: Article 52.2). Besides that, the selectedor, better to say, available -diagnostic morphological characters Guadelli's species possesses do not allow to distinguish it from other forms of red deer. The primitive stage of molarization of P4 indicated by [START_REF] Guadelli | Les cerfs du Würm ancien en Aquitaine[END_REF] is quite variable in C.

elaphus and is recorded in varying frequencies in different red deer samples.

According to [START_REF] Janis | The morphology of the lower fourth premolar as a taxonomic character in the Ruminantia (Mammalia; Artiodactyla), and the systematic position of Triceromeryx[END_REF], the simple unmolarized P4 is found in 8% of specimens of C. elaphus from the Middle Pleistocene of Mosbach (Germany) and in 47% of specimens of red deer from the early Holocene of England. A rather high frequency of unmolarized P4 is also recorded in the sample of indigenous Early Iron Age Crimean red deer (Croitor, 2016a).

The species status itself of the smaller elaphine deer from Combe-Grenale was a matter of doubts. [START_REF] Armand | La faune de la Grotte Bourgeois-Delaunay, Commune de la Chaise de Vouthon (Charente). Résultats préliminaires[END_REF], who reported the findings of Guadelli's 'C.

simplicidens' together with larger Cervus elaphus from the Mousterian site of Bourgeois-Delaunay, noticed that the morphology of P4 in the smaller deer form may be variable and occasionally shows a molarized condition. [START_REF] Steele | Variation in mortality profiles of red deer (Cervus elaphus) in middle Palaeolithic assemblages from Western Europe[END_REF] noticed that the arguments for the creation of a new species 'C. simplicidens' are not sufficient, while [START_REF] Monchot | Les chasseurs tardigravettiens de la Baume de Goulon (Salernes, Var, France)[END_REF] supposed that the small body size of this cervid form may be explained by the clinal body size variation of C. elaphus. [START_REF] Becker | Les dolines du Pléistocène supérieur de la Combe de «Vâ Tche Tchâ» (Ajoie, Suisse): un piège à restes de mammifères et artefacts lithiques[END_REF] regarded 'C. simplicidens' as a subspecies of C. elaphus.

Thus, the question on the taxonomical status and systematical position of the "large elaphine" deer from the Late Pleistocene of France remains unresolved. The present work proposes a new look at the cervid antlered braincase from the Late Pleistocene of Saint-Hippolyte (France) and other puzzling findings of elaphine deer from Europe. Special attention is paid to the morphological peculiarities of antlers of the deer from Saint-Hippolyte that reveal specific ecomorphological adaptations and thus provide arguments that permit the establishment of a new subspecies for the Late Pleistocene wapiti of Western Europe.

The site of Saint-Hippolyte

The volcano-sedimentary complex from Saint-Hippolyte is situated at the slope between the Combrailles elevation and the Limagne Plain in the northwestern part of the Massif Central (Puy-de-Dôme, France; Fig. 1) and is of particular interest, since it yields a well-preserved local record of the Würm pleniglacial stage [START_REF] Raynal | Paléoenvironnements et chronostratigraphie du Paléolithique moyen dans le Massif central français. Implications culturelles[END_REF].

The site of Saint-Hippolyte represents a sequence of palaeo-lake deposits that formed in a volcano crater erupted at the end of a temperate phase of the Early Würm, 94000 years B.P. The palaeo-lake deposits were accumulated during at least 30000 years [START_REF] Raynal | Le maar de Saint-Hippolyte (Puy de Dôme, France): datation par thermoluminescence, flores et faunes fossiles, présence humaine, climatochronologie et dynamique du système paléo-lacustre[END_REF][START_REF] Raynal | Evolution comparée de lacs de maars en Auvergne et Velay (France). Datation et contribution des climats Pléistocènes[END_REF]. The palynological analysis revealed a continuous climate evolution toward cold steppe during all the period of accumulation of paleolake deposits [START_REF] Raynal | Le maar de Saint-Hippolyte (Puy de Dôme, France): datation par thermoluminescence, flores et faunes fossiles, présence humaine, climatochronologie et dynamique du système paléo-lacustre[END_REF][START_REF] Raynal | Evolution comparée de lacs de maars en Auvergne et Velay (France). Datation et contribution des climats Pléistocènes[END_REF][START_REF] Raynal | Paléoenvironnements et chronostratigraphie du Paléolithique moyen dans le Massif central français. Implications culturelles[END_REF].

According to [START_REF] Raynal | Le maar de Saint-Hippolyte (Puy de Dôme, France): datation par thermoluminescence, flores et faunes fossiles, présence humaine, climatochronologie et dynamique du système paléo-lacustre[END_REF], the fauna of Saint-Hippolyte includes Deer remains were recovered from the uppermost layers 3 and 2 in association with remains of large bovids and traces of the presence of Mousterian hunters. Raynal (1987,1988) did not explicitly indicate the stratigraphic provenance of the skull stored in the Museum of Clermont-Ferrand, however they mentioned that a fragment of female skull of a similar large cervid came from layer 3. The climate conditions at the time of deposit accumulation were cold and dry and corresponded to the Würm Pleniglacial, MIS2 [START_REF] Raynal | Le maar de Saint-Hippolyte (Puy de Dôme, France): datation par thermoluminescence, flores et faunes fossiles, présence humaine, climatochronologie et dynamique du système paléo-lacustre[END_REF][START_REF] Raynal | Evolution comparée de lacs de maars en Auvergne et Velay (France). Datation et contribution des climats Pléistocènes[END_REF].

Material and methods

The The method of cranial measurements is described in detail elsewhere [START_REF] Croitor | Plio-Pleistocene Deer of Western Palearctic: Taxonomy, Systematics, Phylogeny[END_REF]. The measurements of antlers in most cases are taken according to [START_REF] Boeskorov | Taxonomic position of the red deer Cervus elaphus L. (Cervidae, Artiodactyla, Mammalia) from the Neopleistocene of Northwestern Asia[END_REF]. Some measurements (the circumferences of antler beam above burr, between brow and bez tines, and above bez tine) were taken according to the antler measurement methodology applied by [START_REF] Owen | A History of British Fossil Mammals, and Birds[END_REF] and Ward (1892). The diagnostic morphological characters of species-level taxa for cervids are selected according to the criteria established by Vislobokova (1990) and Croitor (2006aCroitor ( , 2006b)). The crown part of the antler was measured from the base of the first crown to the tip of the last (third) crown tine. Since the third crown tine is destroyed in the specimen from Saint-Hippolyte, the crown part of the antler was measured with approximation from the base of the first crown tine to the tip of the second crown tine. Locus typicus: Saint-Hippolyte (Puy-de-Dôme, Auvergne, France), 460 m of altitude [START_REF] Raynal | Paléoenvironnements et chronostratigraphie du Paléolithique moyen dans le Massif central français. Implications culturelles[END_REF].

Abbreviations

Stratum typicum: Late Pleistocene, "final Wurm I", MIS2 [START_REF] Raynal | Evolution comparée de lacs de maars en Auvergne et Velay (France). Datation et contribution des climats Pléistocènes[END_REF].

Description of the holotype:

The lambdoid and interfrontal sutures are not obliterated yet; the interparietal suture is completely obliterated (Fig. 2). It is not possible to estimate the stage of obliteration of the coronal suture between pedicles because this part of the skull was restored. However, according to [START_REF] Raynal | Le maar de Saint-Hippolyte (Puy de Dôme, France): datation par thermoluminescence, flores et faunes fossiles, présence humaine, climatochronologie et dynamique du système paléo-lacustre[END_REF] who studied the specimen before the restoration, this part of the coronal suture was already obliterated. Therefore, the whole stage of the cranial sutures obliteration corresponds to the age of four years in modern red deer [START_REF] Mystkowska | Morphological variability of the skull and body weight of the red deer[END_REF]. Therefore, the specimen under study belongs to a young mature individual.

The frontal bones are concave in the lateral view. The occipital part of the skull is broadened and forms an almost right angle with the skull roof. The shape of the lambdoid suture is trapezoid, a characteristic feature of the genus Cervus. The proportions of braincase do not show any important particularities and fit with values of other species of the genus Cervus involved in the comparison. However, the braincase behind pedicles in the specimen from Saint-Hippolyte is somewhat broader than in the Asian forms (Fig. 3; Table 1). The frontal bones breadth falls within the variation of the frontal breadth of the fossil wapiti from Siberia and northeast Asia described by [START_REF] Boeskorov | Taxonomic position of the red deer Cervus elaphus L. (Cervidae, Artiodactyla, Mammalia) from the Neopleistocene of Northwestern Asia[END_REF] (Fig. 3B).

The pedicles are moderately long (their length exceeds the half of pedicle diameter) that corresponds to the young adult age of the deer. The circumference of pedicles is quite large exceeding the maximal value of the modern sample of Asian wapiti reported by [START_REF] Boeskorov | Taxonomic position of the red deer Cervus elaphus L. (Cervidae, Artiodactyla, Mammalia) from the Neopleistocene of Northwestern Asia[END_REF] and approaches the maximal value of the fossil Asian wapiti C. canadensis cf. 'sibiricus' (Fig. 4A). Taking into account the individual age and the robustness of the pedicles of the deer from Saint-Hippolyte, one can assume that its antlers attained the stage of full development but not reached the potential maximum size according to the available data on the ontogenetic development of antlers in modern red deer and wapiti [START_REF] Sokolov | Hoofed Animals (Orders Perissodactyla and Artiodactyla). Fauna of the USSR[END_REF][START_REF] Danilkin | Deer (Cervidae)[END_REF][START_REF] Bender | Body mass and antler development patterns of Rocky Mountain elk (Cervus elaphus nelsoni) in Michigan[END_REF].

The antlers maintain the typical for wapiti bauplan with six tines: two proximal tines (brow and bez tines), a middle tine (trez tine), and three crown tines (Figs. 5,6).

The distal part of the third crown tine is not preserved. The antler measurements are generally similar to measurements of Asian fossil wapiti (Table 2). The total length of the left antler (measured from the antler base to the tip of the second crown tine)

amounts to 114 cm. The antler length of the French specimen equals the largest antlers of C. elaphus maral measured by Ward (1892), however, it is much more robust than the antlers of modern red deer. The length of the antler from Saint-Hippolyte is rather moderate if compared to the large sample of antlers of modern American wapiti measured by Ward ( 1892), but the antler beam is generally very robust and corresponds to the stoutest specimens of modern wapiti (Fig. 7A). The almost horizontal (Fig. 5). In the area of middle tine, the antler beam turns upward and the distal fork of the antler crown is set almost vertically and oriented in the parasagittal plane. The antler beam becomes remarkably flattened in the areas of the trez tine and the crown tines ramifications as it is seen in wapitis [START_REF] Croitor | On the presence of Late Pleistocene wapiti, Cervus canadensis Erxleben, 1777 (Cervidae, Mammalia) in the Palaeolithic site Climăuți II (Moldova)[END_REF]. The crown part of the beam is significantly compressed from the sides.

The wapiti from Saint-Hippolyte is distinguished from all modern and fossil forms of wapiti involved in the present study by its relatively and absolutely longer distance between the bez and the trez tines (43% of the antler length; Fig. 4B). The crown part attains 30.3% of the antler length. The antler proportions of the wapiti from Saint-Hippolyte are very distinct from those of C. canadensis mongoliae (= C. canadensis fossilis). Unlike French wapiti, Chinese form is characterized by the relatively short distance between bez and trez tines (21.6% of the antler length), while the distal crown part is very long and attains 55.4% of the antler length. The type specimen of C. canadensis cherskii may be regarded as a transitional form between the French and Chinese wapitis: the beam segment between bez and trez tines attains 32%, while the crown part makes 32% of the antler length (Fig. 8).

The bez-tine is longer than the brow-tine and is situated at a certain distance from it. This character distinguishes the French wapiti from C. canadensis canadensis, C. canadensis mongoliae, and C. canadensis cherskii, which are characterized by the greatest length of the first crown tine. The distance between the brow and the bez tines amounts to 89 mm on the left antler and 83 mm on the right antler.

The trez tine (or the middle tine) in the deer from Saint-Hippolyte is inserted on the dorsal (lower) side of the beam and is pointed sideward and downward. The apical part of the trez-tine of the left antler is turned toward the posterior. This is one of the most peculiar morphological features of C. canadensis combrayicus.

The antler crown is uniaxial and is formed by three tines: the longer and stronger first crown tine and a distal fork. The first crown tine is directed sideward, upward and toward the anterior. The distal crown fork is composed of the second and the third crown tines situated in the parasagittal plane. The distance between the first and the second crown ramifications is larger than the length of the second crown tine. The angle between the trez-tine ramification plane and the crown plane amounts to ca.

45°. The angle between the plane of the first crown tine ramification and the plane of the distal fork is approximatively the same.

The antler surface is clearly pearled in their proximal part but becomes smoother in the distal portion of the preserved left antler. The surface of the crown part is not pearled. The longitudinal furrows of blood vessels are well-expressed.

Discussion

Red deer versus wapiti

The biogeographic, genetic, biological and paleontological data suggest that C. elaphus and C. canadensis represent a typical case of vicarious species. The sharp geographical and genetic division between red deer and wapiti established at least from the pre-LGM times [START_REF] Meiri | Subspecies dynamics in space and time: A study of the red deer complex using ancient and modern DNA and morphology[END_REF]. The eastern border of the former species areal and the western border of the latter one moved repeatedly eastwards and westwards with climate changes, therefore the dispersals of C. canadensis in Europe could occur several times [START_REF] Stankovic | First ancient DNA sequences of the Late Pleistocene red deer (Cervus elaphus) from the Crimea, Ukraine[END_REF][START_REF] Meiri | Subspecies dynamics in space and time: A study of the red deer complex using ancient and modern DNA and morphology[END_REF]. The vicarious character of distribution in modern faunas and in the paleontological record is mostly due to the ecological adaptations and ethological distinctions between C. elaphus and C. canadensis [START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF]. The morphological distinctions between wapiti and red deer are the main challenge. Very little has been done on the comparative morphological study of C. elaphus and C. canadensis, apparently, because these species were regarded for a long time as subspecies groups within the single species C. elaphus. The attention of researchers was focused mostly upon the shape of antlers and linear cranial measurements that were regarded as taxonomic criteria at the subspecies level [START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF][START_REF] Danilkin | Deer (Cervidae)[END_REF]. Therefore, the available data on the morphological distinction between those two species are scant and incomplete, while the size distinction may be an insufficient criterion (Fig. 3).

Actually, the detailed comparative study of the cranial and dental morphology of wapiti and red deer was never carried out. The report by [START_REF] Guadelli | Les cerfs du Würm ancien en Aquitaine[END_REF] regarding the differences in molar morphology between "small" and "large" elaphine deer is the only study that brings some data on comparative dental morphology of C. elaphus and C. canadensis. The relatively linguolabially broader upper molars and the more frequent lingual groove in P 2 of the "large elaphine" deer observed by [START_REF] Guadelli | Les cerfs du Würm ancien en Aquitaine[END_REF] are important dental characters since they indicate the stronger brachyodonty (a primitive morphological character according to Vislobokova, 1990) and the more advanced degree of molarization of P 2 (an advanced morphological character according to Vislobokova, 1990) in the "larger elaphine" deer if compared to European C. elaphus. [START_REF] Guadelli | Les cerfs du Würm ancien en Aquitaine[END_REF] observations suggest that the dental morphology characters may be an effective diagnostic tool and require special attention and verification on a larger fossil and modern osteological material.

We know very little about morphological distinctions in cranial morphology between red deer and wapiti. [START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF] reported just some general differences in cranial proportions of modern C. canadensis and C. elaphus: the skull in wapiti is generally broader and more robust with a somewhat shorter facial part.

The ratio of the face length measured from the anterior part of orbit rim to the tips of premaxillary bones (prosthion) to the condylobasal length normally is below 60%, while in red deer this ratio is usually above 60%, especially in the largest red deer subspecies C. elaphus maral, although the sample values are overlapping [START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF].

The distinctive features in antler morphology are better known. The antlers of C. canadensis are distinguished from C. elaphus by their generally poor development of crown that normally consists of three tines arranged in the parasagittal plane [START_REF] Lydekker | On apparently new deer from North China, in the menagerie of the Duke of Bedford at Woburn Abbey[END_REF][START_REF] Lydekker | Artiodactyla, Families Cervidae (Deer), Tragulidae (Chevrotains), Camelidae (Camels and Llamas), Suidae (Pigs and Peccaries), and Hippopotamidae (Hippopotamuses). Trustees of the British Museum[END_REF][START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF]. The antler crown in wapiti rarely evolves additional tines, more or less compressed from the sides, and often tends to develop palmations in optimal environmental conditions [START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF].

The type specimen of C. canadensis merriami from Arizona, for instance, represents such a variant of the flattened distal portion of antler [START_REF] Nelson | A new species of elk from Arizona[END_REF]. According to [START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF], the strongest first crown tine in wapitis is a continuation of the antler beam, while the second and third crown tines should be regarded as a bifurcated crown tine situated on the posterior side of the beam. The Geist's viewpoint is confirmed by the paleontological evidence: some of the early ontogenetic stages of antler development in Tibetan wapiti C. canadensis macneilli (Geist, 1998, fig. 8.6c) strikingly remind the antler shape of Cervus magnus (= Pseudaxis magnus Zdansky, 1925) from the late Tertiary of China (Fig. 9A,B). The antlers of C. magnus are characterized by the presence of a single basal tine (brow tine), a trez tine, and a small crown tine situated on the posterior side of the distal part of the main beam, which apparently is a homologue of the first crown tine in wapitis of modern type (Fig. 9A).

According to [START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF] [START_REF] Lydekker | Artiodactyla, Families Cervidae (Deer), Tragulidae (Chevrotains), Camelidae (Camels and Llamas), Suidae (Pigs and Peccaries), and Hippopotamidae (Hippopotamuses). Trustees of the British Museum[END_REF][START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF][START_REF] Flerov | Musk Deer and Deer. The Fauna of USSR[END_REF][START_REF] Sokolov | Hoofed Animals (Orders Perissodactyla and Artiodactyla). Fauna of the USSR[END_REF][START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF]. However, this morphological feature appeared very early in the evolution of the genus Cervus. For example, Cervus nestii [START_REF] Azzaroli | I cervi fossili della Toscana[END_REF]) (= Cervus abesalomi [START_REF] Kahlke | Neufunde fon Cerviden-resten aus dem Unterpleistozän von Untermassfeld. Das Pleistozän von Untermassfeld bei Meningen (Thüringen), Teil 2[END_REF] from the Early Pleistocene of Italy and Georgia already developed the transversal elaphus-like crown fork but possessed only one basal (brow) tine (Croitor, 2006a). Therefore, the difference in construction and development of crown part in C. elaphus and C. canadensis indicates an independent evolutionary development of the antler crown in Cervus starting from the three-pointed antler stage that we can see today in Rusa unicolor and Rusa timorensis. Such an early evolutionary split of the magnus-canadensis and nestiielaphus lineages raises a question on how red deer and wapiti acquired the second basal tine: the bez tine, which is a less constant element of antler in both taxa. The most plausible explanation is that there was a genetic exchange between those two lineages, a phenomenon that is often recorded in the animal world [START_REF] Arnold | Divergence with Genetic Exchange[END_REF].

Multiple cases of such interspecific gene exchange within the Cervus lineage were recently reported by [START_REF] Hu | Genome-wide study on genetic diversity and phylogeny of five species in the genus Cervus[END_REF].

The position of trez tine is a good diagnostic character that distinguishes wapiti from red deer. In C. canadensis, the trez tine is inserted on the lateral side of the beam and the plane of trez ramification is clearly situated in a different plane with respect to the parasagittal plane of the crown tines and the brow and bez tines that are set on the anterior side of the beam.

The smooth surface of the distal portion of antler represents another specific morphological feature of wapiti distinguishing it from the completely pearled antlers of the red deer C. elaphus [START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF]. Partially pearled antlers are also reported for modern C. canadensis from Yakutia, Russia, that evolve surface pearling only in the area of brow and bez tines (Stepanova and Argunov, 2016). The knobby appearance of the red deer antler surface derives from the periosteum of growing antlers that is responsible for the development of the surface pearling or "perlation" [START_REF] Goss | Deer Antlers: Regeneration, Function and Evolution[END_REF].

The pearled antler surface is a specific morphological feature of C. elaphus and some related forms that sometimes are also included in Cervus, such as Rusa unicolor and Rusa timorensis.

Fossil subspecies of Eurasian wapiti

The past diversity of wapiti is little known. 1925) is characterized by a rather moderate length, however, its maximum diameter above the burr (74.0 mm) indicates that this specimen is as robust as other Eurasian fossil wapiti antlers (Fig. 7A). The crown part of the antler is welldeveloped and consists of three large crown tines that lie in the same plane (Fig. 9D).

The crown tines are very strong and clearly more robust than the bez and trez tines.

The distal crown tine is compressed laterally and terminated with two small prongs.

The crown part of the Chinese wapiti attains more than half of the antler length (55%)

and represents the main distinctive morphological feature of this subspecies (Fig. 8).

The distance between the bez tine and the trez tine in the Zdansky's specimen, however, is remarkably short and the distance between the bez tine and the first crown tine (ca. 485 mm) is significantly shorter than that in the type specimen of C.

mongoliae. Zdansky (1925) also ascribed to the fossil Chinese wapiti extremely pachyostotic mandibles, but those specimens actually belong to Sinomegaceros.

Cervus canadensis spelaeus Owen, 1846

Owen (1846) described and figured a basal fragment of an extremely large shed antler from the Late Pleistocene deposits of Kent's Cavern (England) that is distinguished from the modern European red deer by its extraordinarily large size, approaching to the size of the largest modern North American wapiti specimens.

Owen (1846) reported this deer as Cervus (Strongyloceros) spelaeus. According to [START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF], the cave stag from Kent's Cave was most likely a wapiti. The antler of C. canadensis spelaeus bears two basal tines (brow and bez) separated by a certain distance; the circumference of the beam between the brow and the bez tines, according to [START_REF] Owen | A History of British Fossil Mammals, and Birds[END_REF], measures 381 mm and significantly exceeds the maximal value for modern American wapiti (330.2 mm) reported by Ward (1892) (Fig. 7B).

This value exceeds by ca. 30% the circumferences of other fossil forms of wapiti. The circumference above the burr attains 375 mm, thus greatly exceeds the analogous measurements of modern European red deer Cervus elaphus Linnaeus, which range between 220 and 230 mm [START_REF] Friant | Le Strongyloceros spelaeus Owen de Kent's Hole (Angleterre)[END_REF]. According to [START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF], the basal beam circumference indicates a predicted condylobasal length of 528 mm and a body weight of at least 550 kg lean. The circumferences above the burr of other antlers found in Kent's Cavern range from 250 to 300 mm (Lister, 1987a) and correspond to the values of the largest modern American wapitis [START_REF] Geist | Deer of the World: Their Evolution, Behavior and Ecology[END_REF]. Besides the exceptionally large size, not much could be said about the cave stag. The distal fragments of wapiti antlers described from the Late Paleolithic site Climauți II are characterized by exceptionally large size and possibly represent a wapiti similar to cave stag. The length of the segment between the trez and the first crown tines amounts to 410 mm and exceeds by ca. 38% the analogous measurement of other known fossil forms of wapiti [START_REF] Croitor | On the presence of Late Pleistocene wapiti, Cervus canadensis Erxleben, 1777 (Cervidae, Mammalia) in the Palaeolithic site Climăuți II (Moldova)[END_REF]. Most of Kent's Cavern fauna from the main cave deposits has a pre-LMG age and is dated back to 50000-25000 years before present [START_REF] Higham | The earliest evidence for anatomically modern humans in northwestern Europe[END_REF].

Possibly, the mandible 2003-4-420-Sol of a large-sized deer from the Pleistocene of Soleilhac (France) that was originally described as Megaceros (Megaceroides)

solilhacus by Azzaroli (1979, pl. 3, fig. 2) also belongs to C. canadensis. Besides the extremely large size, this specimen from Soleilhac is characterized by a very long diastema. The length ratio between diastema and lower cheek tooth row is 82.6%, approaching the mandible 2003-4-420-Sol to modern red deer and distinguishing it from other Pleistocene large-sized deer [START_REF] Croitor | Origin and evolution of the late Pleistocene island deer Praemegaceros (Nesoleipoceros) cazioti (Depéret) from Corsica and Sardinia[END_REF]. The mandible in question comes from the old paleontological collections of Musée Crozatier (Le Puyel-Velay, Haute-Loire, France), so its exact stratigraphic context is unknown.

Cervus canadensis cherskii Boeskorov, 2005

The type specimen of this subspecies comes from the Late Pleistocene of the lower part of Kolyma valley in Yakutia, Russia. According to [START_REF] Boeskorov | Taxonomic position of the red deer Cervus elaphus L. (Cervidae, Artiodactyla, Mammalia) from the Neopleistocene of Northwestern Asia[END_REF], this subspecies is distinguished from the modern C. canadensis sibirica Severtsov, 1873 (= C. canadensis canadensis according to [START_REF] Flerov | Musk Deer and Deer. The Fauna of USSR[END_REF] by its larger size, more robust antlers and by the longer distance between the bez and trez tines (Fig. 9E). The antlers of C. canadensis cherskii correspond to the most robust specimens of modern American wapiti measured by Ward (1892), however, the antler length falls within the mean values of Canadian wapiti (Fig. 7). The crown part is relatively shorter than in C. canadensis mongoliae and attains 36% of the antler length (Fig. 8). The distal part of the third crown tine in the type specimen is terminated by a small fork as in C.

canadensis mongoliae. The first crown tine is the strongest one as in modern C. canadensis canadensis. The bez tine may be almost of the same length as the brow tine or shorter. The trez tine is well-developed and clearly longer than the brow and bez tines. The area of distribution of this wapiti in northwestern Asia ranged from the Lena River to Western Chukotka. The age of findings of C. canadensis cherskii corresponds to the Kargin Interglacial stage (55-25 ky BC) [START_REF] Boeskorov | Taxonomic position of the red deer Cervus elaphus L. (Cervidae, Artiodactyla, Mammalia) from the Neopleistocene of Northwestern Asia[END_REF] characterized by the maximal distribution of forests and bogs in northeastern Asia (Ukraintseva, 1979).

Cervus canadensis palmidactyloceros De Stefano, 1911

Numerous remains of a peculiar deer come from the Late Pleistocene and Early Holocene of Northern Italy and the peat bogs of Switzerland [START_REF] Rütimeyer | Die Fauna der Pfahlbauten der Schweiz[END_REF][START_REF] De Stefano | I mammiferi preistorici dell'Imolese[END_REF]. The rather heavy antler beams are directed sideward and terminated with a strongly flattened crown segment that bears four tines (Figs. 9C,10B). This strong divergence of antlers may be seen also in the antlered skull from Settecannelle, Italy (Abbazzi, 1995, figs. 5, 7). De Stefano (1911) reported the remarkably large size of skeletal remains of this deer and the development of a distal palmation that varies in breadth from 142 to 190 mm. All palmations figured by De Stefano (1911) and [START_REF] Abbazzi | Occurrence of palmated Cervus elaphus from Italian late Pleistocene localities[END_REF] generally show a constant shape pattern: three crown tines that terminate palmation are situated in the same parasagittal plane; the first crown tine is the strongest one and in all cases is bifurcated. Sometimes this bifurcation is set in the transversal plane or looks like a small accessory prong of the stronger first crown tine [START_REF] De Stefano | I mammiferi preistorici dell'Imolese[END_REF]. The antler tines are comparatively short and the bez tine is often missing [START_REF] De Stefano | I mammiferi preistorici dell'Imolese[END_REF][START_REF] Abbazzi | Occurrence of palmated Cervus elaphus from Italian late Pleistocene localities[END_REF] Balkakra, Scania (Sweden) (Ahlen, 1965, fig. 9) shows a great affinity with Alpine wapiti. The antlers of Swedish wapiti are characterized by a pronounced compression from the sides above the trez tine and the typical for wapiti parasagittal orientation of the crown plane (Fig. 10A). The trez tine is the strongest and the longest one. Unlike the typical form of C. canadensis palmidactyloceros, the wapiti individual from Sweden lacks the bifurcation of the first crown tine. [START_REF] Azzaroli | Il nanismo nei cervi insulari[END_REF][START_REF] Azzaroli | Il nanismo nei cervi insulari[END_REF] divergent and characterized by the presence of short brow and trez tines (Fig. 10C).

Cervus canadensis tyrrhenicus

The bez tine is always missing, while the distal part of the antler is extended into a small palmation terminated with three tines situated in the parasagittal plane: the stronger anterior crown tine that represents a continuation of the antler beam, and two smaller tines that have a posterior position on the main antler axis. The parasagittal orientation of crown tines in the dwarfed deer from Capri is remarkable and apparently was noticed by [START_REF] Azzaroli | Il nanismo nei cervi insulari[END_REF] who dedicated a part of his discussion to morphological features of wapiti. Nonetheless, according to the state of knowledge at that time, the occurrence of a dwarfed wapiti on a Mediterranean island was incredible, so [START_REF] Azzaroli | Il nanismo nei cervi insulari[END_REF] assumed that the dwarfed deer from Capri was phylogenetically related to C. elaphus aretinus from the Late Pleistocene of Italy.

However, C. elaphus aretinus is characterized by a multiaxial crown and may be very close to, or even synonymous with, C. elaphus maral [START_REF] Croitor | An antlered skull of a subfossil red deer, Cervus elaphus L., 1758 (Mammalia: Cervidae), from Eastern Romania[END_REF]. This extinct form of wapiti is generally as large as C. canadensis cherskii and modern C. canadensis sibiricus and by 11-15% larger than C. canadensis palmidactyloceros (Fig. 3). The estimated body mass of C. canadensis combrayicus based on the occipital height is ca. 364 kg according to the methodology proposed by [START_REF] Janis | Correlation of cranial and dental variables with body size in ungulates and macropodoids[END_REF]. The comparatively smaller antlers of the French wapiti may reflect the rather young individual age of the type specimen (Fig. 7A). Unlike C. canadensis mongoliae, the wapiti from Saint-Hippolyte is characterized by relatively shorter crown part and the relatively and absolutely longer proximal portion of the antler between the bez and the trez tines. C. canadensis cherskii has rather an intermediate position between the extreme forms of antler shape seen in C. canadensis mongoliae from one hand, and in C. canadensis combrayicus from another hand (Fig. 8). Unlike both C. canadensis mongoliae and C. canadensis cherskii, the French wapiti is distinguished by the relatively small size of the first crown tine that is deviated sideward and therefore is not situated in the same parasagittal plane with other two crown tines (Fig. 9F). The trez tine in C. canadensis combrayicus is much smaller and is directed downward and represents a peculiar feature of this fossil form.

Functional morphology of antlers and paleoecology of the wapiti from Saint-

Hippolyte

The morphological peculiarities of antlers of C. canadensis combrayicus represent interesting evolutionary specializations that require a special discussion. The main paleoecologically meaningful antler specializations are the strong divergence of antlers, the relatively long distance between the bez and the trez tines, the short trez tine directed downward, and the sideward deviation of the first crown tine. Here I propose a discussion of the functional significance of morphological specializations of the antlers of the wapiti from Saint-Hippolyte.

The antler spread in the stag from Saint-Hippolyte attained about 205 cm, while the antler spread in modern American wapitis hardly exceeds 130 cm (Ward, 1892).

According to Lister (1987b) According to [START_REF] Davitashvili | The Theory of Sexual Selection[END_REF], the branched and complicated antlers in the advanced cervids represent a specific adaptation that "softens" the sexual selection and diminish the excessive male mortality during the intraspecific combats. The most proximal antler tine (brow tine) appears very early in the cervid evolution and has a function of the locally very restricted protection of eyes and face during combats (Lister, 1987b). The second proximal tine, the bez tine, is specific only for C. elaphus and C. canadensis and appears in the evolution of Cervus during the early Middle Pleistocene simultaneously with a significant body size increase. The estimated body mass of primitive small-sized Cervus nestii was about 60 kg [START_REF] Croitor | Plio-Pleistocene Deer of Western Palearctic: Taxonomy, Systematics, Phylogeny[END_REF], while the body mass of the most ancient subspecies of red deer C. elaphus acoronatus attained ca. 237 kg (the body mass is estimated on the basis of occipital height; the sample includes the cranial material from the Tiraspolian gravel and specimens studied by Di Stefano and Petronio, 1992; N total =14). Therefore, a ca. four-time body size increase is observed in the C. nestii-C. elaphus lineage during the Early Pleistoceneearly Middle Pleistocene transition. [START_REF] Petrie | Intraspecific variation in structures that display competitive ability: large animals invest relatively more[END_REF] assumed that the costs of intraspecific contests may be relatively greater for larger animals that may have a greater capacity to injure each other. This, probably, may explain why red deer (and, apparently, wapiti too) acquired the second basal tine enhancing the face protection during combats. It is interesting to notice that the bez tine has a trend to be diminished in European red deer during Late Pleistocene and Holocene simultaneously with body size reduction (Lister, 1987b). Nonetheless, the direct relationship between the body size and the strength of the bez tine in red deer cannot be considered as proved, since the bez tine is quite weak in the largest modern red deer subspecies, C. elaphus maral [START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF][START_REF] Flerov | Musk Deer and Deer. The Fauna of USSR[END_REF]. Lister assumed that the reduction of bez tine in red deer is correlated with the development of richly branched antler crown, but, again, the example of C. elaphus maral does not support this hypothesis.

Nonetheless, the bez tine in C. canadensis combrayicus is particularly strong and long, a feature that distinguished the stag from Saint-Hippolyte from Late Pleistocene European red deer.

In turn, the trez (or middle) tine appears in the evolution of the subfamily Cervinae quite early as an additional second ramification and coincides with the increase of body size and the reduction of upper canines when male antlers became the only dangerous weapon against conspecific rivals [START_REF] Geist | The relation of social evolution and dispersal in ungulates during the Pleistocene, with emphasis on the old world deer and the genus Bison[END_REF]. Apparently, the development of this tine in the early stages of cervid evolution also must increase the safety of both combatting stags that attained a certain body mass threshold that increases the risk of lethal wounding. The protection function of the trez tine against the rival's antlers was suggested by [START_REF] Kitchener | Fighting behaviour of the extinct Irish elk[END_REF] also for the cervids that bear highly evolved complicated antlers, such as red deer and giant deer. [START_REF] Davitashvili | The Theory of Sexual Selection[END_REF] demonstrated the importance of the additional antler tines in red deer stags by the examples of old bucks with degenerated and simplified antlers that can easily wound and kill their conspecific rivals. A forwardly pointed tine situated in the middle of the antler beam appeared several times independently in the evolution of cervines. This is the case of the middle tine of Praemegaceros verticornis from the Middle Pleistocene of Europe, that appeared quite late in the Praemegaceros lineage, as it is completely absent in P. pliotarandoides from the late Early Pleistocene (Croitor, 2006b). The middle tine of P. verticornis is analogous to the trez tine of C. elaphus and C. canadensis and, apparently, it had the same function of locking of rival's antlers, but is not homologous, since its evolutionary story is different. The evolutionary importance of the middle tine may be confirmed by the fact that many insular dwarfed deer with diminished and simplified antlers maintain the middle tine, which may be the only remained additional branch in the extreme cases of antler simplification [START_REF] Azzaroli | Il nanismo nei cervi insulari[END_REF]van der Geer, 2018).

It is important to note, that the trez tine or its analogues normally evolved in cervids from the middle latitudes and is not present in southern lineages with large antlers, such as Rucervus and Panolia. A possible explanation is that the male survival is exposed to a stronger pressure in cervids from the temperate latitudes with cold winters and a seasonal drop of forage. The pressure of seasonal male mortality (especially of young and prim-adult age) is important [START_REF] Clutton-Brock | Red Deer: Behavior and Ecology of Two Sexes[END_REF], so the additional security adaptations in antler shape diminish the excessive loss of males during the combats and therefore compensate the seasonal male mortality. Therefore, one can assume the direct relationship between the harder environmental conditions with a seasonal drop of temperatures and forage availability and richly branched antlers. The consequences of seasonality in the southern latitudes are less costly for deer stags, and apparently, this may explain the smaller number of ramifications and the absence of the "second-level protection" tine (the trez tine or its analogues) in large South Asian deer species.

Concerning the wapiti from Saint-Hippolyte, there are doubts if its short and downward-directed trez tine was an effective protection against antlers of stag rivals:

in the combat posture of antlers, the trez tines in C. canadensis combrayicus are directed backwards. Such a specific position of the trez tine may be explained by its newly acquired function. Apparently, the majority of deer adapted to more or less open landscapes evolve the so-called posterior tine situated on the back side of the beam and positioned below the crown part of the antler [START_REF] Croitor | Plio-Pleistocene Deer of Western Palearctic: Taxonomy, Systematics, Phylogeny[END_REF]. Usually, antlers are more or less sharply bent in the area of the posterior tine, which is situated on the convex side of the antler bow. The posterior tine appears independently in several cervid lineages of the both Cervinae and Capreolinae subfamilies (genera Rangifer, Megaloceros, Praemegaceros, Praedama, Sinomegaceros, some species of the genus Dama). In some cases, the lowermost posterior crown tine may have a position similar to that of the posterior tine seen in the Pleistocene fallow deer: this tine in Dama geiselana and Dama clactoniana is normally very large, strong and detached from the palmation [START_REF] Leonardi | The fallow deer of European Pleistocene[END_REF][START_REF] Pfeiffer | Die fossilen Damhirsche von Neumark-Nord (Sachsen-Anhalt) D. dama geiselana n. ssp[END_REF]. The occurrence of a posterior tine and its analogues apparently does not depend on body size (it is developed in the giant continental Praemegaceros and Megaloceros, in some small-sized insular dwarfs, and in medium-sized Rangifer) and is not correlated decreases the behavioural and immune defense of males against ticks, which distract them from the rutting behaviour and bring damages to their physical and health state [START_REF] Hart | Effects of hormones on behavioral defenses against parasites[END_REF][START_REF] Mooring | The evolution of parasite-defense grooming in ungulates[END_REF]. The priority of the investment into the rutting activity over the antiparasite defense has a negative consequence for males, which are exposed to a greater parasite load. Rutting impala males, for example, which self-oral groomed at one-third the rate of females, were found to carry about six times more adult ticks than females [START_REF] Hart | Effects of hormones on behavioral defenses against parasites[END_REF]. Observations on male white-tailed deer reveal they had about 3 times as many adult ticks as females [START_REF] Main | Ixodes dammini (Acari: Ixodidae) on white-tailed deer Odocoileus virginianus in Connecticut, USA[END_REF]. [START_REF] Mooring | The evolution of parasite-defense grooming in ungulates[END_REF] suppose that the elevated testosterone level in rutting stags suppresses the grooming behaviour in order to increase male's vigilance, so the increased ectoparasite burden is a frequent side effect in rutting males. It is interesting to notice that the scratch grooming in males of the observed species was not affected by the rutting behaviour [START_REF] Hart | Effects of hormones on behavioral defenses against parasites[END_REF]. One can assume that the posterior tine in cervids is a specialized adaptation for scratch grooming in rutting males that eventually indirectly increase their combat capacities (Croitor, 2016b). Apparently, the evolutionary acquisition of the scratch grooming tine (the posterior tine) indirectly influenced mating success by diminishing of the ectoparasite burden and improving the vigilance and physical state of the rutting stags. According to the available bibliographic data, the disturbing effect of ectoparasites during the rutting period may be important. The observations upon modern North American wapiti revealed that September and October constitute the peak time for feeding by larvae of winter tick Dermacentor albipictus [START_REF] Mooring | Tick-removal grooming by elk (Cervus elaphus): testing the principles of the programmed-grooming hypothesis[END_REF]. This stage of the parasite biological cycle coincides with the rutting season in elaphine deer [START_REF] Flerov | Musk Deer and Deer. The Fauna of USSR[END_REF][START_REF] Sokolov | Hoofed Animals (Orders Perissodactyla and Artiodactyla). Fauna of the USSR[END_REF] and therefore has an important negative influence over the physical state and the rutting success of stags. The use of antlers for scratch grooming is rarely reported for modern cervids, which are mostly adapted to wooded and ecotone habitats [START_REF] Mooring | The evolution of parasite-defense grooming in ungulates[END_REF], but it is recorded in Rangifer tarandus tarandus, the only modern deer adapted to open environments [START_REF] Kynkäänniemi | Acute impacts of the deer ked (Lipoptena cervi) infestation on reindeer (Rangifer tarandus tarandus) behavior[END_REF]

Paleobiogeography and evolution of wapiti in Eurasia

The paleogenetic studies revealed a close relationship of the modern Asian wapiti with the Late Pleistocene C. canadensis cherskii and the pre-Last Glacial Maximum specimens from Romania and Crimea [START_REF] Stankovic | First ancient DNA sequences of the Late Pleistocene red deer (Cervus elaphus) from the Crimea, Ukraine[END_REF][START_REF] Meiri | Subspecies dynamics in space and time: A study of the red deer complex using ancient and modern DNA and morphology[END_REF]. The area of distribution of Asian wapiti Cervus canadensis sibirica [START_REF] Severtzoff | Altitudinal and latitudinal distribution of Turkestan animals[END_REF] (including the darker coloured C. canadensis songarica [START_REF] Severtzoff | Altitudinal and latitudinal distribution of Turkestan animals[END_REF] today is limited to the mountains that range from Tian-Shan and Altai to Sayan and southern Transbaikal Area [START_REF] Heptner | Deer of the USSR (systematics and zoogeography). Transactions on Study of Fauna and Flora of the USSR[END_REF][START_REF] Flerov | Musk Deer and Deer. The Fauna of USSR[END_REF][START_REF] Danilkin | Deer (Cervidae)[END_REF]. 12). Most probably, dense broad-leafed forests were the main limiting factor for C. canadensis in Europe.

Conclusions

The new look at the fine antlered stag skull from Saint-Hippolyte permits to close (1995). (1902). The circumference of C. canadensis spelaeus is adapted from [START_REF] Owen | A History of British Fossil Mammals, and Birds[END_REF]. Cavern [START_REF] Higham | The earliest evidence for anatomically modern humans in northwestern Europe[END_REF]. The time span of wapiti presence in Sweden is hypothetical.

Figure captions

1

  antlered braincase from Saint-Hippolyte (catalogue number MHLCLFE-R-M118) is stored in the Natural History Museum Henri-Lecoq, Clermont-Ferrand (France). The material herein studied is compared with craniological material stored in the osteological and paleontological collections of the Natural History Museum of London; the National Museum of Natural History, Paris; the Museum of Zoology and Natural History "La Specola" of Florence; the Institute of Zoology, Chișinău, Moldova; and the National Museum of Ethnography and Natural History of Moldova.

  length and stoutness of the antlers generally correspond to those of C. canadensis cherskii from the Late Pleistocene of northeast Asia and C. canadensis mongoliae from the Late Quaternary of China. Nonetheless, the antlers of wapiti from Saint-Hippolyte are less massive if compared to C. canadensis ssp. from Climauti II (Moldova) and are much smaller than the type specimen of Strongyloceros spelaeus from Kent's Cavern (Fig. 7B). The antler beams of the specimen from Saint-Hippolyte are remarkably divergent. They are more divergent than antlers of modern C. canadensis sibirica and C. canadensis canadensis, so their middle portions (below the middle tines) are

  The fossil and subfossil remains of C. elaphus maral from Italy were reported by De[START_REF] De Stefano | I mammiferi preistorici dell'Imolese[END_REF], therefore, the synonymy of C. elaphus aretinus and C. elaphus maral should not be excluded. The shape of the flattened distal portion of antlers of C. canadensis tyrrhenicus shows a clear affinity with antler morphology of C. canadensis palmidactyloceros: the strong anterior crown tine corresponds to the strongest first crown tine in continental wapiti that represents a continuation of the antler beam; the second smaller crown tine is actually a homologue of the accessory prong or bifurcation of the first crown tine in C. canadensis palmidactyloceros (Fig.10B). The lowermost posterior crown tine in C. canadensis tyrrhenicus is a simplified homologue of the distalmost part of antler in continental wapiti (second and third crown tines). This part of antler in C. canadensis tyrrhenicus is reduced in size and has lost the bifurcation usually reported as the second and the third crown tines in continental wapitis. The isolation of the island population of Italian wapiti, apparently, occurred during the post-Glacial sea level rise. 5.2.6. Cervus canadensis combrayicus n. ssp.

  Unlike modern C. canadensis canadedsis and closely related C. canadensis sibirica, the French fossil wapiti is characterized by a much stronger divergence of antlers, the longer distance between the bez and trez tines, the downward direction of the trez tine, which is comparatively much shorter, and the weak development of the first crown tine, which is pointed more sideward and therefore deviates from the plane of other two crown tines. The difference from modern C. canadensis xantopygus is much deeper: besides the all above-mentioned distinctions from other modern wapitis, C. canadensis combrayicus is distinguished from C. canadensis xantopygus by curved antler beams between the bez and trez and above the trez tine. Certain parallelism may be seen in the weak development of the crown part of antler in both C. canadensis combrayicus and C. canadensis xantopygus. The strongly divergent antlers of the French wapiti remind the C. canadensis palmidactyloceros, however, the deer from Saint-Hippolyte is distinguished from the Alpine wapiti by the lack of distal palmation, the downward direction of the trez tine, and the unbranched first crown tine.

  , outstretched antlers are indicative of adaptation of cervids to open landscape, since in forest they would impede movement. Therefore, C. canadensis combrayicus shows a more pronounced adaptation to open landscapes than modern wapitis. The specific shape of antler tines in the wapiti from Saint-Hippolyte, however, cannot be explained directly by the adaptation to open landscape and requires a broader discussion of their functional significance in Cervidae and in the deer from Saint-Hippolyte in particular.

  with the development of palmations or other parts of antlers. The stability of development of posterior tine within the individual variation range (for instance, in the series of antlers of Megaloceros giganteus figured by[START_REF] Reynolds | A Monograph on the British Pleistocene Mammalia. Vol. III; Part III. The Giant Deer[END_REF] suggests an important evolutionary selection in favour of this character, which appeared repeatedly and independently in several lineages of the family Cervidae. Therefore, one can assume a certain biological importance of this part of antlers. The adaptation to more or less open environments is the only circumstance that unites the deer with posterior antler tine. The most plausible explanation of the iterative development of the posterior tine during the evolution of cervids is its specific function of the defense against ectoparasites in rutting males. The study of rutting males of North American deer and territorial African antelopes revealed that the involvement of males in rutting

  . The downward-directed trez tine in C. canadensis combrayicus, apparently, got the function of removal of flying parasites and ticks analogous to that of the posterior tine in other Pleistocene giants and in modern reindeer. The reduced size of first crown tine in C. canadensis combrayicus is an advanced evolutionary specialization. The first crown is deflected sideward and a little toward the anterior and spatially is detached from the second and third tines that are directed toward the posterior. A most plausible explanation of the modifications in the crown part of the antler is the acquired clutch function by the first crown tine that becomes in C. canadensis combrayicus a functional analogue of the trez tine in other subspecies of wapiti and in red deer. Therefore, the wapiti from Saint-Hippolyte shows the most advanced adaptations to the open landscape within the genus Cervus and the evolutionary specialization of its antlers involved the changes in shape and function of the trez and the first crown tine.

  Therefore, the area of distribution of the lineage of Asian wapitis during the Glaciation epoch was extremely vast and ranged from Chukotka in Northeast Asia to Western Europe (Fig.11). From the biogeographic point of view, it means that the modern Siberian wapiti should be regarded as a remnant of the successful Ice Age megafaunal species, which survived the Holocene climate warming in the mountains of Central Asia that act as a glacial refugium. According to[START_REF] Stankovic | First ancient DNA sequences of the Late Pleistocene red deer (Cervus elaphus) from the Crimea, Ukraine[END_REF], the modern Altai environmental conditions, where Asian wapiti survived, represent the recent analogue of the environment of the full-glacial period of Central Europe. This assumption was confirmed by Pavelková Řičánková et al. (2014) who reported a marked similarity in ecological structure between Recent eastern Altai-Sayan mammalian assemblages and Pleistocene faunas. The Last Glacial and recent eastern Altai-Sayan faunal assemblages are peculiar in the co-occurrence of large herbivore and predator species associated with steppe, desert and alpine biomes. According to Pavelková Řičánková et al. (2014), the relic glacial fauna seems to persist up to present in the Eastern part of the Altai-Sayan region where, for instance, reindeer and saiga antelope still live in sympatry. The occurrence of the cold-adapted Cervus canadensis in Europe during the Last Glacial Maximum explains the "paradoxical" record of elaphine deer in the southern extension of permafrost areas in Eastern Europe, namely in the Late Paleolithic sites Rașcov 7 and Cosouți from Moldova[START_REF] Sommer | Glacial refugia of mammals in Europe: evidence from fossil records[END_REF][START_REF] Sommer | Late Quaternary distribution dynamics and phylogeography of the red deer (Cervus elaphus) in Europe[END_REF].[START_REF] Banks | Reconstructing ecological niches and geographic distributions of caribou (Rangifer tarandus) and red deer (Cervus elaphus) during the Last Glacial Maximum[END_REF] elaborated the ecological niche modelling of red deer during the Last Glacial Maximum that does not support the hypothesis of the "East Carpathian Last Glacial Maximum refugium" of Cervus elaphus. The elaphine remains from the Late Paleolithic sites of Moldova are very scanty and represented by a few poor and isolated teeth and bone fragments[START_REF] David | Theriofauna of the Pleistocene of Moldova[END_REF]. The recently described antler fragments of C. canadensis from the Moldavian Late Paleolithic site of Climăuți[START_REF] Croitor | On the presence of Late Pleistocene wapiti, Cervus canadensis Erxleben, 1777 (Cervidae, Mammalia) in the Palaeolithic site Climăuți II (Moldova)[END_REF] urges a reconsideration of the elaphine remains from the Late Glacial Maximum of the East Carpathian area.Wapitis became extinct postglacially in the vast North Eurasian area as LatePleistocene horses and steppe bison did. However, the relatively abundant subfossil remains of C. canadensis palmidactyloceros from Switzerland and Italy, as well as the articulated skeleton with antlers from southern Sweden indicate that the Alpine altitudes and Scandinavia, apparently, served as postglacial refugia for wapiti in Western Europe. The retraction of European wapiti after the post-Pleistocene climate warming into the cool treeless alpine altitudes and its ecological and biological isolation from European red deer may explain the puzzling results obtained by[START_REF] Drucker | Evolution of habitat and environment of red deer (Cervus elaphus) during the Late-glacial and early Holocene in eastern France (French Jura and the western Alps) using multi-isotope analysis (δ 13 C, δ 15 N, δ 18 O, δ 34 S) of archaeological remains[END_REF]. The multi-isotope analysis of bone collagen revealed a meaningful difference in paleoecology and paleoenvironment of the postglacial elaphine deer samples from Jura and western Alps[START_REF] Drucker | Evolution of habitat and environment of red deer (Cervus elaphus) during the Late-glacial and early Holocene in eastern France (French Jura and the western Alps) using multi-isotope analysis (δ 13 C, δ 15 N, δ 18 O, δ 34 S) of archaeological remains[END_REF]. According to the obtained results, C. elaphus from French Jura inhabited the dense forests in warmer and ecologically more productive ecosystems, while the "red deer" from the Alps persisted in the open landscape and rather cool conditions with low soil productivity by dwelling at higher altitudes than before the climate warming. Drucker et al. (2011) explained this difference in response to the postglacial climate change as remarkable ecological flexibility of red deer and, possibly, as a specific selection by prehistoric human hunters who preferred to hunt red deer in open areas for technical and/or cultural reasons. Most probably, the postglacial elaphine remains from the high Alpine altitudes belong to the remnant population of Cervus canadensis that survived during the early stages of Holocene in the Alpine refugium. Therefore, C. canadensis palmidactyloceros from Switzerland, Italy, and possibly French Alps, is a relict population of wapiti that survived in Europe to the Holocene climate warming (Fig.

  the long-lasting debate of occurrence of wapiti Cervus canadensis in the paleontological record of Western Europe. The specific features of antler morphology (the very strong divergence of antler beams, the relatively long antler segment between the bez and the trez tine; the relatively short crown tine of antler; the short downward-directed trez tine and the reduced size of the first crown tine) indicate the advanced adaptations of the wapiti from Saint-Hippolyte to open environments of the Late Pleistocene periglacial ecosystems. The revealed specific evolutionary adaptations are taxonomically meaningful and distinguish the deer from Saint-Hippolyte from all fossil and modern subspecies of wapiti, therefore a subspecies name Cervus canadensis combrayicus n. ssp. is proposed for this cervid form. The taxonomical status and exact diagnosis of the extremely large C. canadensis spelaeus Owen, 1846 from Kent's Cavern, England, is not clear yet due to the incomplete paleontological material. The confirmed occurrence of wapiti in the Late Pleistocene of Western Europe answers some questions on evolution and paleoecology of the so-called "elaphine" deer, including the different ecological reaction of lowland red deer and highland Alpine populations of Cervus to the postglacial climate change and the conflicting hypothesis on the Last Glacial Maximum refugium for red deer in the East Carpathian Area. The specific adaptations of Cervus canadensis to the cold periglacial tundrasteppes explain perfectly the above-mentioned discrepancies.

Fig. 1 .Fig. 2 .

 12 Fig. 1. The geographical location of the paleolake and fossiliferous site Saint-Hippolyte.

Fig. 3 .

 3 Fig. 3. The cranial measurements of Cervus canadensis combrayicus n. ssp. compared to the modern and fossil forms of wapiti and red deer. (A) Braincase breadth; (B) frontal breadth; (C) occipital breadth. The data on C. elaphus priscus/aretinus, C. canadensis canadensis, and a part of data on C. elaphus acoronatus and C. elaphus elaphus are adapted from Di Stefano and Petronio (1992); the data on C. canadensis sibirica and C. canadensis cherskii are adapted from[START_REF] Boeskorov | Taxonomic position of the red deer Cervus elaphus L. (Cervidae, Artiodactyla, Mammalia) from the Neopleistocene of Northwestern Asia[END_REF]; the data on C. canadensis palmidactyloceros are adapted from[START_REF] Rütimeyer | Die Fauna der Pfahlbauten der Schweiz[END_REF] and[START_REF] Abbazzi | Occurrence of palmated Cervus elaphus from Italian late Pleistocene localities[END_REF].

Fig. 4 .

 4 Fig. 4. The pedicle and antler size of Cervus canadensis combrayicus n. ssp. compared with modern and fossil wapitis. (A) Pedicle circumference in the correlation with the antler circumference above burr; (B) length of bez-trez antler segment in the correlation with the antler circumference above burr. The data on C. canadensis cf. sibirica and C. canadensis cherskii from the Late Pleistocene of Siberia and modern C. canadensis sibirica and C. canadensis canadensis are adapted from Boeskorov (2005); the data on C. canadensis from the Seine Basin (France) are adapted from Belgrand (1883).

Fig. 5 .

 5 Fig. 5. The antlers of Cervus canadensis combrayicus n. ssp. (holotype; A, frontal view; B, upper view of the distal part) compared to modern Cervus canadensis sibirica (C; adapted from Stepanova and Argunov, 2016).

Fig. 6 .

 6 Fig. 6. The side view of the holotype of Cervus canadensis combrayicus n. ssp.

Fig. 7 .

 7 Fig. 7. The antler size in modern and fossil red deer and wapitis. (A) Antler length plotted against the beam circumference measured between the brow and bez tines; (B) the comparison of antler circumference between brow and bez tines including the incomplete antlers. The data on modern C. elaphus and C. canadensis are adapted from Ward (1892); the circumference of C. canadensis mongoliae is estimated from the maximum diameter of antler above the burr reported by Zdansky (1925). The measurements of C. canadensis merriami from Arizona are adapted from Nelson

Fig. 8 .

 8 Fig. 8. The antler segments of Cervus canadensis combrayicus compared to C. canadensis cherskii and C. canadensis mongoliae.

Fig. 9 .

 9 Fig. 9. The antler bauplan and specific morphological peculiarities of fossil Eurasian wapiti and wapiti-like deer. (A) Cervus magnus (adapted from Zdansky, 1925; inverted image). (B) Antler of an extant MacNeill's stag Cervus canadensis macneilli (adapted from Geist, 1998; inverted image). (C) Cervus canadensis palmidactyloceros (adapted from De Stefano, 1911). (D) Cervus canadensis mongoliae (adapted from

Fig. 11 .

 11 Fig. 11. The maximum area of distribution of wapiti in Eurasia and the actual area of distribution of Cervus canadensis sibirica in the Tian-Shan/Altai/Sayan postglacial refugium.

  The Late Pleistocene and Early Holocene paleontological record of Eurasia has yielded several forms of C. canadensis, however, some of them were misunderstood and here are included in C.[START_REF] Gaudry | Sur les ossements d'animaux quaternaires que M. l'abbé David a recueillis en Chine[END_REF] described a series of antler fragments from the late Quaternary deposits of Suen Hoa Fou, northeast China as a new species Cervus mongoliae. The best-preserved specimen is a shed antler characterized by the rather straight beam that bears the brow and bez tines, while the trez tine is missing. The specimen is broken at the first crown ramification where antler becomes remarkably flattened as in modern wapiti. The total length of the antler fragment attains 82 cm, while the approximate distance between the bez and crown ramifications (calculated from the figure provided by[START_REF] Gaudry | Sur les ossements d'animaux quaternaires que M. l'abbé David a recueillis en Chine[END_REF] attains ca. 75.7 cm. According to Zdansky (1925), C. mongoliae is based on an abnormal variant of wapiti antler. Zdansky (1925) reported the frequent occurrence in the Late Quaternary of northeast China of deer remains that show a great affinity with modern C. canadensis canadensis and therefore extended the past geographical distribution of Asian wapiti southwards, as far as Henan Province in China. Zdansky (1925) described the fossil Chinese form of wapiti as C.

	canadensis for the first time.

5.2.1. Cervus canadensis mongoliae

[START_REF] Gaudry | Sur les ossements d'animaux quaternaires que M. l'abbé David a recueillis en Chine[END_REF] (= Cervus canadensis fossilis Zdansky, 1925)

canadensis fossilis, however, this subspecies name is preoccupied with Cervus elaphus fossilis

[START_REF] Goldfuss | Osteologische Beiträge zur Kwnntniss Verschiedener Säugthiere der Vormelt: Ueber den Edelhirsch der Vorzeit (Cervus elaphus fossilis)[END_REF][START_REF] Goldfuss | Osteologische Beiträge zur Kwnntniss Verschiedener Säugthiere der Vormelt: Ueber den Edelhirsch der Vorzeit (Cervus elaphus fossilis)[END_REF]

. Therefore the name proposed by Zdansky (

1925

) is invalid. Cervus canadensis mongoliae Gaudry, 1872 should be applied to the fossil wapiti from China. The almost complete antler described and figured by Zdansky (

  . The condylobasal length and the upper tooth row length of the skull from Settecannelle (Central Italy) figured by Abbazzi (1995; CBL measured from the figure is ca. 430 mm) correspond to the largest individuals of modern C. canadensis sibirica (according to the data of Heptner and

	Zalkin, 1947). The facial part of the skull from Settecannelle is relatively short
	(59.5% of the CBL length) as in modern wapiti. Some remains of this endemic wapiti
	form are reported from the Holocene of southern Italy, however, the osteological
	remains ascribed to C. canadensis palmidactyloceros require a thorough revision
	since the second coeval elaphine deer C. elaphus maral Ogilby, 1840 (probably is a
	senior synonym of C. elaphus aretinus Azzaroli, 1961) from Italy developed rather
	robust crown tines that sometimes look like small palmations (De Stefano, 1911).

This morphological parallelism was interpreted by

[START_REF] Abbazzi | Occurrence of palmated Cervus elaphus from Italian late Pleistocene localities[END_REF][START_REF] Di Stefano | The morphometry and the occurrence of Cervus elaphus (Mammalia, Cervidae) from the Late Pleistocene of the Italian Peninsula[END_REF]

as an intraspecific variation of C. elaphus. However, there are some important morphological distinctions between C. elaphus maral/aretinus and C. canadensis palmidactyloceros: the crown part in the Italian maral is multiaxial and richly branched, while in the Alpine wapiti the crown part always consists of three main crown tines aligned in the parasagittal plane. The strongest first crown tine normally is bifurcated and this feature is rather constant in C. canadensis palmidactyloceros from Switzerland and Italy. The almost complete (splanchnocranium is missing) subfossil articulated skeleton of an adult male from

  The third lobe of M3 is often reduced; P4 is molarized; the lower premolar series is relatively short and attains 58% of the molar series length; the protocone in upper molars is supplemented with an additional enamel fold(Azzaroli, 1961, table 7, fig.1) that was never recorded in C. elaphus. The antlers are less

described this dwarfed insular deer from the cave deposits of Capri (Italy) as C. elaphus tyrrhenicus. The body size is close to that of C. elaphus corsicanus and C. elaphus barbarus, however, the teeth are relatively larger (the lower tooth series exceeds in length the largest specimens of modern C. elaphus from France), suggesting a rapid and significant body size reduction. Unlike C. elaphus corsicanus, the metacarpal bone in C. canadensis tyrrhenicus is significantly shorter than the radius.

  Stepanova, V.V., Argunov, A.V., 2016. Morphometry of antlers of red deer (Cervus elaphus L., 1758) of Yakutia. Bulletin of Udmurd University, Series Biology,

	Earth Sciences 26 (1), 126-132 (in Russian).
	Ukraintseva, V.V., 1979. Vegetation of warm late Pleistocene intervals and the
	extinction of some large herbivorous mammals. Botanical Journal 3, 318-330 (in
	Russian).
	van der Geer, A.A.E., 2018. Uniformity in variety: Antler morphology and evolution
	in a predator-free environment. Palaeontologia Electronica, Article number
	21.1.9A, doi: 10.26879/834.
	Vislobokova, I.A., 1990. Fossil deer of Eurasia. Transactions of Paleontological
	Institute 240, 1-208 (in Russian).
	Ward, R., 1892. Horn Measurements and Weights of the Great Game of the World.
	Rowland Ward & Co., London, 264 pp.
	Zdansky, O., 1925. Fossile Hirsche Chinas. Palaeontologia Sinica, Series C, 2 (3), 1-
	94.

Table 1 .

 1 Cervus canadensis combrayicus n. ssp., cranial measurements (mm) of the holotype compared to fossil wapiti C. canadensis cherskii from northeast Asia (adapted from[START_REF] Boeskorov | Taxonomic position of the red deer Cervus elaphus L. (Cervidae, Artiodactyla, Mammalia) from the Neopleistocene of Northwestern Asia[END_REF].

Table 2 .

 2 Cervus canadensis combrayicus n. ssp., measurements (mm) of pedicles and antlers of the holotype compared to C. canadensis mongoliae from the Quaternary of China (adapted fromZdansky, 1925) and C. canadensis cherskii from the Late Pleistocene of northeast Asia (adapted from[START_REF] Boeskorov | Taxonomic position of the red deer Cervus elaphus L. (Cervidae, Artiodactyla, Mammalia) from the Neopleistocene of Northwestern Asia[END_REF].

	Measurement	C. canadensis combrayicus C. canadensis	C. canadensis
		right	left	mongoliae	cherskii
					(hopotype)
	L antler		1140	1090	1235
	H pedicle	34.5	36.8
	DAP pedicle	62.1	61.2
	DLM pedicle	60.7	61.8
	DAP burr	80.0	80.3	81.0
	DLM burr	76.7	78.2
	CF pedicle	188.0	190.0
	DLM above burr	54.7	59.8
	DAP above burr	70.3	73.2	74.0
	CF above burr	217.0	223.0		255
	H brow tine	96.5	92.0	72.0
	ramification
	L brow tine	ca. 260	ca. 260		440
	DLM brow tine base 45.5	41.0
	DAP brow tine base 31.4	29.8
	CF brow tine base	124.0	115.0
	H bez tine	147.7	195.3
	ramification
	L bez tine	ca. 315	ca. 390	> 162
	DLM bez tine base	40.1	44.3
	DAP bez tine base	37.2	37.3

Acknowledgements

I am very thankful to Michael Lebras and Charles Lemarchand (Museum of Natural History Henri-Lecoq, Clermont-Ferrand) for the granted access to the fossil specimen that is under their care and for the kind assistance during my work. I thank Prof. Wei Dong and an anonymous reviewer for their comments and suggestions that improved the manuscript. This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors.