Origin and zoology of the Deer
Systematics
The noble stag, the "crowned king" and the second largest Cervid in Europe, is scientifically designated by a Latin binomial comprising the name of the genus and that of the species, each followed by the name of the first describer and the date of description: genus Cervus Linné, 1758 - species Cervus elaphus Linné, 1758. It is known by several vernacular names: cerf noble, cerf rouge and cerf élaphe. This last name is a pleonasm because Cervus (in Latin) and elaphos (in Greek, ἔλαφος) both mean "stag"! This animal is an elegant, high-legged ungulate with a short tail. Its head has long ears and two deciduous, more or less branching, bony expansions called antlers, the prerogative of males only.
From a systematics point of view, the noble stag is part of the order Cetartiodactyla, the super-family Cervoidea and the family Cervidae, whose classification is complex. There are around 17 genera and 47 species in two subfamilies: Capreolinae and Cervinae. The Capreolinae have two tribes, Odocoileini and Caprolini, while the Cervinae have a single tribe, Cervini. The noble stag is classified in the latter two categories.
In addition to the noble stag, the subfamily Cervinae includes the North American and Asian elk, the Philippine stag, the sika stag (all in the genus Cervus), the Eld stag (or Thamin or Thameng; genus Panolia), the barasinga (or barasinghaou or Duvaucel's stag; genus Rucervus), the white-nosed stag (or Thorold's stag ; genus Przewalskium), fallow deer (genus Dama), axis deer, hog deer, Bawean deer and Calamian deer (or Calamian hog deer) (all of the genus Axis), father David's deer (or elaphurus ; genus Elaphurus), the muntjac (genus Muntiacus), the elaphid (or crested deer; genus Elaphodus), the rusa deer (or Java deer) and the sambar deer (both of the genus Rusa).
Red stag. Cervus elaphus, web source
Wapiti. Cervus canadensis, web source
Sika deer. Cervus nippon, web source
Eld deer Cervus (Rucervus) eldi, web source
Barasingha. Cervus (Rucervus) duvaucelli, web source
Morphology
As in all ketartiodactyls, the noble stag has an even number of digits, with the axis of the limb passing between radii n° III and n° IV, i.e. between metacarpals/metatarsals n° III and n° IV which become dominant and between the corresponding digits, and a talus (or talus) with a double pulley. Certain bones in the limbs were fused: in the forearm, the radius and ulna, which prevented any rotation of the hand; in the wrist (called the "knee" in ungulates), the capitate (or large bone) and the trapezoid (capitatotrapezoid); in the leg, the tibia and fibula, of which only the lower part remains, forming an isolated bone: the malleolar bone; at the ankle (called the "hock" in ungulates), the cuboid and navicular which form the cubonavicular, the receptacle for the lower pulley of the talus and the base of the calcaneus.
The inclusion of the noble stag in the Cervidae family is due to the presence of antlers covered by a vascularised skin, the "velvet", which covers the bone itself, the fusion of metacarpals/metatarsals n° III and n° IV and the presence of metacarpals/metatarsals n° II and n° V: it therefore has 4 toes per leg, two developed central ones and 2 smaller lateral ones.
The noble stag is a herbivorous ruminant. It therefore has dentition adapted to eating plants and a stomach with 4 pouches. After pre-digestion in the rumen, the food is regurgitated and chewed a second time before being swallowed again to pass into the other gastric reservoirs. It has 34 teeth. There are no incisors on the upper dental arch and the upper canines (or spits) are reduced. The lower canine resembles an incisor. Its premolars (three per side; the first premolar has disappeared) and molars (three per side) have a structure that allows food to be torn into flaps: the constituent denticles are stretched from front to back and differentiated into ridges (selenodont type). The crown is low (brachyodont type).
Distribution
Cervids are found in a variety of habitats throughout Eurasia, Malaysia, North Africa and the Americas.
The noble deer is adapted to deciduous and coniferous forests, but also to open spaces such as meadows. It is capable of colonising high mountains. Several subspecies have been identified, due to its wide geographical distribution, which extends from Western to Central Europe. Genetic analyses have made it possible to clarify the systematic position of some of them, which are now placed in different species (Markiewicz et al. 2022).
In Western and Central Europe, five subspecies of noble stag are recognised: the noble stag stricto sensu (C. e. hippelaphus Erxleben, 1777; France, Germany, Switzerland, Austria, Denmark, western Carpathians); the Spanish stag (C. e. hispanicus Hilzheimer, 1909), the British stag (C. e. scoticus Lönnberg, 1906), the Norwegian stag (C. e. atlanticus Lönnberg, 1906) and the Corsican stag (C. e. corsicanus Erxleben, 1777; Corsica and Sardinia). In the Caucasus, Anatolia and Crimea the species is represented by the maral (C. e. maral Ogilby, 1840). North Africa is home to the Atlas deer (C. e. barbarus Bennett, 1833).
Origin and diversity
Cervids have been known since the Cenozoic, more specifically in the Miocene, with the genera Procervulus, Dicrocerus, Acteocemas, Euprox, Amphiprox and Pliocervus. Around 5.3 Ma (early Pliocene), Croizetoceros (1 m high at the withers, around 60 kg), with its "modern" appearance, came to prominence (Heintz 1970). It was around the middle and end of this period, at around 2.6 Ma, that the genus Cervus appeared in China with C. magnus (= Pseudaxis magnus Zdansky, 1925), which could be the basis of the elk lineage (Croitor 2020). At around 2 Ma, C. nestii (Azzaroli, 1947) is the oldest European representative of the elaphus group (Croitor 2006, 2011, 2018). It occurs in Georgia (= C. abesalomi Kahlke, 2001; Dmanisi deposit: 1.8 Ma) and in Italy (Val d'Arno, Olivola). Its average weight is estimated at 60 kg.
The first traces of the noble stag are known from around 900,000 years ago (early Middle Pleistocene) with a subspecies whose antlers have no crown at the top: this is the acoronate stag, C. e. acoronatus Beninde, 1937, with an estimated average weight of 237 kg, which corresponds to the size of a modern elk. It has been identified in Germany (= C. elaphoides Kahlke, 1960 = C. reichenaui Kahlke, 1996), France, Italy, Spain, the Netherlands and England. Between 600,000 and 500,000 years ago, the antlers began to become more complex. In France, the crowned stag type is known from around 400,000 years ago at the Caune de l'Arago site (Pyrénées Orientales) (Magniez et al. 2013).
The species' great ecological plasticity and its ability to colonise a variety of environments led to the distinction of several subspecies in the Middle and Upper Pleistocene (Croitor 2018). In Germany, C. e. angulatus Beninde, 1937 (late Middle Pleistocene, 250,000 years old) is known; it is thought to be the direct descendant of C. e. acoronatus. In Italy, three subspecies have been defined. These are C. e rianensis Leonardi & Petronio, 1974 (= C. e. eostephanoceros Di Stefano & Petronio, 1993), from the late Middle Pleistocene, which is thought to be an endemic descendant of C. e. acoronatus, C. e. siciliae Pohlig, 1893 (late Middle Pleistocene, Late Pleistocene) found in Sicily and C. e. aretinus Azzaroli, 1961 (Late Pleistocene; possibly synonymous with C. e. maral Lydekker, 1898).
In France, a small stag, with teeth of a more primitive structure than in other fossil stag populations, has been identified in several archaeological levels of Mousterian industry (Layers 54 to 50A; MIS 5) from the Combe-Grenal shelter (Dordogne): it is C. simplicidens Guadelli, 1997. However, this taxon was invalidated because the name of the species had already been used by R. Lydekker (1876) and the particular morphological criteria of the teeth are occasionally observed in populations of C. elaphus (for more details see Croitor 2018). Recently, several fossil subspecies attributed to the noble deer have been redefined and are considered to be elk, C. canadensis,
Evelyne Crégut-Bonnoure, Honorary Curator, Requien Museum, Avignon & UMR 5608 TRACES, Toulouse, France
Quote this text (English version): Crégut-Bonnoure É.,2025. Deer origin and zoology in: Averbouh A., Feruglio F. & Plassard F. Dir. The Jean-Clottes database, Animal representation in Prehistory (BJC), "Deer File", English version online on 12/032025.
Quote this text (French version) : Crégut-Bonnoure É.,2023 (version française). Origine et zoologie du cerf in : Averbouh A., Feruglio F. & Plassard F. Dir. Base Jean-Clottes - Animal Representation, Les représentations animales depuis la Préhistoire, "Deer File“, on line 18/10/2023
References used
Croitor R. & Crégut-Bonnoure É (sous presse). From Continents to Islands: Tracking the Red Deer (Cervus elaphus) and its "Troublesome Cousin", the Wapiti (Cervus canadensis) in Europe. In Averbouh A. & Mashkour M. dir. Anthropozoologie du cerf élaphe, Anthropozoologica.
Croitor R. 2006. Early Pleistocene small-sized deer of Europe. Hellenic Journal of Geosciences, vol. 41, 89-117
Croitor R. 2011. Early Pleistocene deer Cervus nestii (Cervidae, Mammalia) and the earliest dispersal of the genus Cervus (sensu stricto) in Western Eurasia. In Actual problems of protection and sustainable use of animal world diversity, International Conference of zoologists, Chisinau: 203-204
Croitor R. 2018. Plio-Pleistocene deer of Western Palearctic: Taxonomy, Systematics, phylogeny. Institute of Zoology of the Academy of Sciences of Moldova: 140 p.
Croitor R., Obada Th., 2018. On the presence of Late Pleistocene wapiti, Cervus canadensis Erxleben, 1777 (Cervidae, Mammalia) in the Palaeolithic site Climăuți II (Moldova). Contributions to Zoology, 87 (1) 1-10
Di Stephano G., Petronio C., 2021. Importance of the morphological plasticity of Cervus elaphus in the biochronology of the Middle and the Late Pleistocene of the Italian peninsula. The Science of nature, 108 (5): 1-12 https://doi.org/10.1007/s00114-021-01753-x
Guadelli, J.L., 1997. Les cerfs du Würm ancien en Aquitaine. Paleo 8, 99-108.
Heintz, E. 1970. Les Cervidés villafranchiens de France et d'Espagne, Mémoires du Muséum national d'Histoire naturelle, 22: 1–303.
Magniez P., Moigne A.-M., Testu A., Lumley H. de, 2013. Biochronologie des mammifères Quaternaires. Apport des Cervidae du site Pléistocène moyen de la Caune de l’Arago (Tautavel, Pyrénées-Orientales, France). Quaternaire, 24(4), 477-502
Mackiewicz, P., Matosiuk, M., Świsłocka, M. et al. 2022. Phylogeny and evolution of the genus Cervus (Cervidae, Mammalia) as revealed by complete mitochondrial genomes. Sciences Report 12, 16381. https://doi.org/10.1038/s41598-022-20763-x