Cervus elaphus, Linnaeus, 1758

Croitor, Roman & Crégut-Bonnoure, Évelyne, 2025, From continents to islands: tracking the red deer (Cervus elaphus Linnaeus, 1758) and its “ troublesome cousin ”, the wapiti (Cervus canadensis Erxleben, 1777), in Europe, Anthropozoologica 60 (11), pp. 143-163 : 149-151

RED DEER CERVUS ELAPHUS

Cervus elaphus is a relatively old, highly successful, and adaptable species, primarily thriving in broadleaf forest biomes across the middle latitudes of western Eurasia ( Geist 1998; Di Stephano & Petronio 2021). Several distinctive cranial features distinguish this species. The facial portion of the skull in largest subspecies (as, for instance, in C. elaphus maral Ogilby, 1840 ), is elongated, primarily due to the lengthening of the orbitofrontal portion, with the anterior edge of the orbit projecting behind the posterior edge of the upper third molar, M3 ( Ogilby 1840). The lower mandible also displays elongated characteristics: its diastemal part is relatively long and attains from 63.0% to 82.5 % of the lower toothrow length, the angle between the horizontal and ascending ramuses is more open compared to other deer species, and the processus angularis is poorly pronounced. The lower fourth premolar typically exhibits molarization, although this trait can vary.

The earliest fossil records of red deer date back to the early Middle Pleistocene of Europe, around 900 000 years ago. The earliest red deer, described as subspecies C. elaphus acoronatus , is characterized by the development of a simple distal fork and a very strong and long bez tine ( Figs 3B, 4A). This subspecies is one of the largest forms of red deer, with an average weight estimated at c. 240 kg. It has been identified in England, Germany (including remains of young individuals reported as Cervus elaphoides Kahlke, 1960 , and Cervus reichenaui Kahlke, 1996 ), France, Italy, the Netherlands, and Moldova ( Beninde 1937; Lister 1990; Di Stefano & Petronio 1992). The early acoronate red deer evolved into several specialized endemic European subspecies, characterized by further complication of antler crown or specific evolutionary specializations in the case of Mediterranean dwarfed forms ( Beninde 1937; Azzaroli 1961; Di Stephano & Petronio 2021).

Between 600 000 and 500 000 years ago, European red deer began to evolve a multiaxial antler crown. The initial simple antler crown consisted of three tines. This evolutionary stage of red deer was identified as C. elaphus antiqui Pohlig, 1892 . In France, evidence of early crowned deer dates back to around 400 000 years ago at the site of La Caune de l’Arago (Pyrénées Orientales) ( Magniez et al. 2013).

The evolution of C. elaphus angulatus Beninde, 1937 (occurring at the end of the Middle Pleistocene in Germany, approximately 250 000 years ago) is marked by further complication of the distal portion of the antler. The primary evolutionary development of C. elaphus angulatus is the emergence of a third extremely hypermorphous posterior crown tine, in addition to a transversal fork ( Fig. 4B). This additional tine is directed backward and forms an angle with the antler beam, often flattens and bears additional digitations. This peculiar variant of antlers with flattened distal portion instead of multiaxial crown is still present in some modern red deer populations of Western Europe ( Johnston 1903).

Several endemic subspecies have been documented from the Pleistocene of Italy. C. elaphus rianensis Leonardi & Petronio, 1974 (also known as C. elaphus eostephanoceros Di Stefano & Petronio, 1993 ), from the end of the Middle Pleistocene of the Italian Peninsula, is distinguished by a reduction in body size (with an estimated body mass of approximately 100 kg) and a significant shortening of the distal portion of antlers terminating in a simple fork, similar to C. elaphus acoronatus .

C. elaphus siciliae Pohlig, 1893 (occurring at the end of the Middle Pleistocene and Upper Pleistocene of Sicily) was an endemic dwarfed insular red deer with an estimated body mass of approximately 60 kg ( Gliozzi et al. 1993). The dwarfing of body size resulted in disproportionately broad frontal bones, a relatively narrow occiput, and noticeably divergent antlers. Some specimens are characterized by a three-tined crown, suggesting that C. elaphus siciliae evolved from C. elaphus antiqui or a similar form.

C. elaphus jerseyensis Zeuner, 1940 represents a unique case of strong evolutionary specialization of red deer in the conditions of insular isolation on the northern island of Jersey in the English Channel ( Zeuner 1946; Lister 1989). This dwarfed form of red deer evolved during the last interglacial period (Eemian = MIS 5e) and underwent a rapid six-fold reduction in body size to approximately 36 kg ( Lister 1989). As a result, C. elaphus jerseyensis reached a size comparable to that of roe deer. It is the smallest form of red deer, with antlers that retained their trez (middle) tine and only one basal tine positioned at a certain distance from the burr ( Zeuner 1946).

In France, a small-sized deer with teeth exhibiting a more primitive structure compared to other fossil deer populations has been identified in several archaeological levels with Mousterian industry (Layers 54 to 50A; MIS 5) at the Combe-Grenal shelter (Dordogne, Western France). This deer is referred to as Cervus simplicidens Guadelli, 1997 . However, this taxon has been invalidated because the species name had already been used by Lydekker (1876). Furthermore, the particular morphological characteristics of the teeth, such as primitive unmolarized lower fourth premolar, are occasionally observed in other populations of C. elaphus ( Janis & Lister 1985) . The origin of small-sized red deer from Dordogne is most probably related to the Iberian glacial refugium, and further investigation is needed to determine its possible relationship with modern C. elaphus hispanicus Hilzheimer, 1909 (= C. elaphus bolivari Cabrera, 1914 ), a smaller red deer subspecies from the Iberian Peninsula ( Cabrera 1914).

Today, the red deer is represented by several subspecies and forms. The hypothesized area of its origin is the Tarim region in China ( Ludt et al. 2004). Among the modern subspecies, the Bactrian deer ( C. elaphus bactrianus ) is one of the forms that retains the most primitive characteristics, inhabiting riparian forests of Central Asia. This subspecies is characterized by antlers that feature a bez tine and terminate in a simple transverse distal fork, typically exhibiting a total of six antler tines. According to some reports, primitive white spotting on their backs is retained even in some adults ( Flerov 1952).

Taxonomic interpretations based on mitochondrial cytochrome b and control region marker analyses of primitive red deer subspecies ( Lorenzini & Garofalo 2015), merit closer examination. Lorenzini & Garofalo (2015) concluded that Central Asian red deer subspecies are genetically distinct from their Western counterparts, forming a regional monophyletic group. They proposed classifying this group as a separate species, Cervus hanglu Wagner, 1844 , comprising the subspecies hanglu , bactrianus, and yarkandensis.

However, this genetic study, which sought to redefine the taxonomic status of red deer subspecies, notably did not include type specimens in its analysis. Additionally, Lorenzini & Garofalo (2015) acknowledged that their findings do not align with the morphology-based subspecific taxonomy within C. elaphus . Despite this discrepancy, they failed to provide revised diagnoses for the taxa whose statuses were altered. This tripartite classification of elaphines ( C. elaphus , C. hanglu , and C. canadensis ) has since been adopted by other researchers (e.g., Meiri et al. 2018). In our opinion, the elevation of C. elaphus hanglu to species rank appears to have been driven more by conservation objectives ( Lorenzini & Garofalo 2015; Narayan et al. 2024) than by strict adherence to established taxonomic principles and criteria.

Lorenzini & Garofalo (2015) classified Central Asian red deer as a distinct species, Cervus hanglu , citing genetic distances from Western red deer subspecies comparable to those between C. canadensis and Cervus nippon . However, to date this classification still has some important gaps. Their interpretations do not align with the biological species concept ( Mayr 1942), which emphasizes genetic isolation, nor with the ecological species concept ( Van Valen 1976). According to Van Valen, a species is defined by its occupation of a distinct adaptive zone and its evolutionary separation from other lineages. The classification proposed by Lorenzini and Garofalo lacks consideration of whether Cervus hanglu occupies a unique ecological niche or exhibits diagnostic ecological adaptations, both of which are crucial to supporting its status as a distinct species under the ecological framework. Divergence time alone is not a sufficient criterion for taxonomic classification. Subspecies can remain isolated for extended glaciation periods, yet if environmental conditions remain stable and evolutionary rates are slow –such as in glacial refugia– they tend to retain conservative traits and remain subspecies. This type of natural selection in glacial refugia is actually an example of stabilizing selection. Modern Central Asian red deer exhibit traits more consistent with the oldest known red deer subspecies, C. elaphus acoronatus , rather than displaying unique adaptations to new ecological niches.

In contrast, the relatively recent divergence between C. nippon and C. canadensis illustrates rapid evolution within a new adaptive zone, driven by directional selection under extreme ecological and climatic pressures. This divergence resulted in

A B

C

distinct morphological and biological adaptations to their respective niches, justifying their classification as separate species.

Based on these considerations, we still consider C. elaphus bactrianus , C. elaphus yarkandensis Blanford, 1892 , and C. elaphus hanglu as a group of primitive subspecies within C. elaphus .

The Caspian red deer, C. elaphus maral , ranges from North Iran to the Carpathian region in Europe. This subspecies is the largest among red deer, characterized by several primitive features, including the frequent absence of the bez tine ( Fig. 4C) and rudimentary white spots on the back, particularly notable in populations from the Caucasian area ( Heptner & Zalkin 1947; Flerov 1952). The crown part of the antler exhibits a distinct pattern of complexity, which can be considered metameric: it consists of an axial tine with one or two to three transversal forks, resulting in a potential increase in the number of crown tines up to seven ( Fig. 4C). This subspecies has been documented in the Balkan area since the Late Pleistocene and is likely closely related to the Late Pleistocene red deer subspecies C. elaphus aretinus Azzaroli, 1961 from the Italian Peninsula ( Croitor & Cojocaru 2016).

The modern West European red deer, C. elaphus elaphus L., is distinguished by its highly evolved multiaxial cup-shaped crown and the consistent presence of the bez tine ( Smith 1881; Lydekker 1898; Heptner & Zalkin 1947; Flerov 1952; Sokolov 1959; Geist 1998). The most basic form of an antler crown consists of an initial terminal transversal fork with an additional third tine emerging from the posterior side of the bifurcation base. As antler development progresses, it undergoes hypermorphosis characterized by the multiplication of terminal forks ( Fig. 3C). In the most extreme cases observed in adult males, the terminal growth forms a cup-like structure composed of multiple tines in the crown ( Smith 1881; Lydekker 1898; Johnston 1903; Beninde 1937; Heptner & Zalkin 1947; Flerov 1952). White spots on the back of adults have never been reported. This subspecies of red deer is considered the most evolved, with its origin traced back to the Iberian glacial refugium. Its current distribution expanded following the retreat of glaciers during the Holocene ( Ludt et al. 2004; Meiri et al. 2013).

Other forms of red deer found in Western Europe have more restricted distributions and are often considered synonyms of the nominotypical subspecies C. elaphus elaphus . These include the Iberian red deer ( C. elaphus hispanicus ), the British red deer ( C. elaphus scoticus Lönnberg, 1906 ), and the Norwegian red deer ( C. elaphus atlanticus Lönnberg, 1906 ). These subspecies are characterized by smaller body size and simplified antlers, features that may represent clinal variation ( Lönnberg 1906; Heptner & Zalkin 1947).