Cervus elaphus, Linnaeus, 1758

Don E. Wilson & Russell A. Mittermeier, 2011, Cervidae, Handbook of the Mammals of the World – Volume 2 Hoofed Mammals, Barcelona: Lynx Edicions, pp. 350-443 : 422-423

25.

Western Red Deer

Cervus elaphus

French: Cerf élaphe / German: Rothirsch / Spanish: Ciervo

Other common names: Barbary Red Deer (barbarus), Bukhara Red Deer (bactrianus), Caspian Red Deer/Maral (maral), Tarim Red Deer (yarkandensis), Thyrrenian Red Deer (corsicanus)

Taxonomy. Cervus elaphus Linnaeus, 1758 ,

Sweden.

Red deer appeared in Europe in late Early Pleistocene, about 900,000 years ago. They were large-sized deer bearing five-tined antlers with a terminal fork, replaced in Middle Pleistocene by deer characterized by antlers with a terminal crown. For decades the red deer complex, including European and North African Red Deer ( C. elaphus ), Central Asian Red Deer ( C. wallichii ), and Eastern Asian and North American Wapiti (C. canadensis ), was considered as only one species. The general body shape, the conspicuous rump patch, and the occurrence of the bez tine in the antler are common traits. But the various forms differ in many characteristics (degree of sexual dimorphism, coat coloration, antler conformation, behavior postures, vocalizations) and have only partial interfertility. Several genetic studies have recently tried to clarify the taxonomy of this species complex, but their results are not always concordant. Two or three main clades are genetically distinguishable. The analysis of mtDNA revealed in the western part of the distribution range three main lineages: a western one that during the last glacial maximum (about 20,000 years ago) survived in an ice-free refugium in Spain and southern France, an eastern one subsisting in South-eastern Europe (Carpathians and Balkans) and a third lineage possibly living then in Italy and now represented only by “Thyrrenian Red Deer” (corsicanus) and “Barbary Red Deer” (barbarus). With the end of the last glaciacion most of Europe was recolonized from the western refugium. It is probable that the Thyrrenian Red Deer derives from introductions by man from mainland Italy into Sardinia about 8000 years ago. Barbary Red Deer of North Africa, could have been translocated there in ancient times. Here six subspecies of Western Red Deer are recognized.

Subspecies and Distribution.

C.e.elaphusLinnaeus,1758—GreatBritainandcontinentalEurope.

C.e.bactrianusLydekker,1900—WTurkestan(Kazakhstan,Uzbekistan,Tajikistan&NAfghanistan).

C.e.barbarusBennett,1833—AtlasMts(Algeria,Tunisia).

C.e.corsicanusErxleben,1777—Corsica(extinct,reintroducedin1985),Sardinia.

C.e.maralGray,1850—Anatolia,Caucasus,andNWIran.

C. e. yarkandensis Blanford, 1892 — E Turkestan (S Xinjiang, China).

It has been introduced into Chile, Argentina, Australia, and New Zealand.

Descriptive notes. Head-body 180-205 cm for males (stags) and 165-180 cm for females (hinds), tail 14-16 cm, shoulder height 105-130 cm (stags) and 95-115 cm (hinds); post-rutting weight of stags 110-220 kg and hinds 75-120 kg. The heaviest animals occur in the Carpathians and Bulgaria (with records of 350 kg lean weight for stags and 200 kg for hinds), the lightest in the Scottish Highlands, southern Spain, Sardinia, and Mesola Wood (Italian Peninsula). Adult stags are on average 50-70% heavier than hinds. Large-sized deer with large head and elongated snout. There are two main genetic groups: a western advanced one (typically with six-tined antlers with a terminal crown, from Europe to Iran) and an Asian primitive one (with five-tined antlers, from the Aral Sea to the Tarim Basin). The rump patch is relatively small, orange on the upper portion, whitish on the lower part, and bordered below in dark brown. The tail is orange and relatively short. The summer coat is reddish or reddishbrown, with grayish legs and whitish belly. The winter coat is grayish-brown; adult stags have a thick neck mane and the belly is heavily stained black. Molts in April-May and September—October. Newborn calves have a brown coat with scattered white spots on the sides, which are lost after 2-3 months. Preorbital, metatarsal, rear interdigital, and subcaudal glands are present. The preorbitals are particularly well developed and open in young calves and in rutting stags. Red deer have a permanent dentition of 34 teeth, with rudimentary upper canines. Permanent teeth erupt at 5-25 months of age. Antlers of adult stags are long, cylindrical, and typically well branched, with two basal tines (brow and bez) and a terminal cup-like “royal” crown or an upper fork. Pedicles begin to grow at 9-10 months of age, the first set of antlers at twelve months. Yearlings are typically “spikers,” with unbranched antlers 5-40 cm long. At 8-13 years the antlers of Western Red Deer attain their full size. In high quality populations antlers are on average 90-105 cm long, with a brutto weight (antler pair plus skull) of 6.6-5 kg, but there are records of 130-140 cm for antler beam length and of 18-21 kg for trophy mass. There are normally 5-7 tines per antler beam, with records of 16 in the wild and 24 in parks. In less productive habitats (Scottish Highlands, Sardinia, Mesola Wood, North Africa) antlers are on average only 60-70 cm in length, with 3-4 tines per beam. In adult stags velvet shedding occurs in July, antler casting in late February-March. Antler regression occurs at 15-16 years of age, sometimes earlier. Hooves are 9-11 cm long in adult stags and 7-8 cm in hinds, with toe pads occupying one third of the hoof.

Habitat. Originally associated with the interface between woodland and grassland, but highly adaptable. It also lives in Mediterranean maquis scrub, in alpine summit meadows, and in the treeless moorlands of Scotland. It prefers broadleaved woodlands interspersed by large meadows. Up to 2800 m above sea level on the Alps. In west-central Asia it occurs in woody and shrubby thickets along riverbanks in desert areas.

Food and Feeding. As an intermediate feeder with a relatively large rumen,it tends to eat a mixed diet of about 30% grass and sedges and about 65% browse, without significant differences among seasons. Fruits and seeds are important items in autumn. It is a very versatile species, consuming up to 300 different species of plants. The two sexes tend to differ in diet composition, hinds eating higher quality food. Stags eat on average 14-16 kg and females around 9-10 kg offresh vegetation per day. Actual food intake follows appetite cycles related to photoperiod and hormones, which are highest in March—June and lowest in late September-December. Lactating hinds more than double their energy requirements.

Breeding. In good quality habitats most females attain puberty at 16 months of age. In less productive habitats sexual maturity may be postponed by at least one year. Stags reach puberty at about 16 months, but can actually begin to reproduce at five years of age. Rutting season is in September—October. Females are polyestrous, with cycles of 18 days. During estrus they are receptive for only 6-24 hours. Conceptions are well synchronized, occurring for 80% of females within 3-4 weeks. Stags leave their summer quarters and move to their traditional rutting stands, where they try to collect and defend a harem against other stags. Competition among stags is high, with some failing completely to reproduce and others able to father up to a dozen or more calves in a season. Stags are involved in roaring contests, dominance displays, and overt fights, with a real risk of serious injury, or more rarely, death. During rut stags lose as much as 15-20% of their pre-rut weight and up to 80% of body fat. The mean length of gestation is 235 days. Before giving birth, hinds leave their group and move away from their usual core areas, looking for better cover. Calving season peaks in May-June. Hinds almost invariably give birth to one calf, which weighs on average 8-9 kg (about 6-5 kg in Scottish moorlands). Male calves are about 5% heavier than females. For the first 7-10 days calves are kept hidden, usually lying in tall vegetation, then begin to follow their mother. During the first few days calves are fed ten times a day. From five days to 90 days they are nursed eight times, and after that, four times a day. Lactating hinds produce 2-4 liters of milk per day, with a peak 50 days after parturition and a decline after three months. By 15-18 days of age calves have doubled their birth weight. They begin to ingest their first vegetation at ten days, and at one month they feed more regularly on plants. Weaning occurs at 6-9 months (November—February). Stags can typically breed for 3-5 years, peaking in Scotland at 7-10 years of age (possibly at 9-12 years in continental Europe). Hinds have a much longer reproductive life span, of up to 12-16 years. The sexes have also a differential pattern of body growth; stags do not reach their full body weight before 8-9 years of age, whereas hinds attain adult size at 3-6 years. Normally Western Red Deerlive to a maximum of 17-18 years, with records of 22 years for stags and 26 for hinds. Main predators are Gray Wolves (Canis lupus) and Eurasian Lynxes (Lynx lynx).

Activity patterns. It is active all the day, with peaks at dawn and in early evening, alternating 6-8 periods of foraging, ruminating, resting, and moving from place to place. It dedicates 7-12 hours to feeding. Human disturbance increases nocturnal activity.

Movements, Home range and Social organization. Western Red Deer have home ranges of variable size, according to sex and habitat; sometimes they occupy only 2-5 km?, sometimes 8-40 km” and more. Adult stags tend to be more mobile and to have separate seasonal ranges. In mountains part of the population can be sedentary, part can make short distance movements, and part can be migratory, moving 10-60 km to reach lower pastures in the winter and high feeding areas in summer. Ranges are normally undefended. Red deer are social animals. The basic social unit is the matrilineal family group, with a dominant old hind associated with her daughters, granddaughters, and their dependent offspring. Stags segregate from hinds for most of the year and form small, less stable, bachelor bands. In woodlands, female family groups can split into smaller units; in open habitat they may unite with neighboring hind groups. During winter and early spring several groups can form temporary large aggregations in open ground.

Status and Conservation. “Bukhara Red Deer” (bactrianus) CITES Annex I and Barbary Red Deer CITES Annex III. Classified as Least Concern on The IUCN Red List (under C. elaphus complex), but it can be considered as not currently at risk and globally increasing. During the 18" and the 19" century overhunting with modern weapons resulted in a drastic contraction of the distribution range and abundance in most parts of Europe. Strict rules for culling, the enforcement of laws with precise hunting seasons, and reintroductions combined to produce an increase in range and numbers. This has become particularly evident since the 1950s-1960s. At the beginning of the 20" century in the Italian Peninsula only a small population in the Po River delta (Mesola Wood) survived. Thanks to the natural recolonization from neighboring countries of the Alpine region and releases, in 1970 deer numbered 3000 individuals. The number grew to 41,000 in 2000 and 57,000 in 2005. In all of Europe, excluding Russia, Western Red Deer numbered 1-25 million individuals in 1985 and 2-4 million in 2005. The annual harvest grew in the same period from 270,000 to 500,000 individuals. Densities are normally 1-5 ind/km?, sometimes up to 15 ind/km?*; higher numbers often imply supplementary feeding. Densities are artificially raised to unnaturallevels (up to 100 ind/km?) to increase revenues from hunting. The Western Red Deeris extinct in Albania and in the Near East ( Lebanon, Israel, Jordan, Syria). Four subspecies are still rare: the Thyrrenian Red Deer, the Barbary Red Deer, the Bukhara Red Deer, and the “Tarim Red Deer” (yarkandensis). Thyrrenian Red Deer faced a dramatic decline in the 20™ century, disappearing from Corsica in 1969 and decreasing to a minimum population size of 100 animals in Sardinia in 1970. Deforestation,fires, and poaching with guns and snares were the main causes. Red Deer from Sardinia have recovered slowly, numbering 6000 animals in 2005. In 1985 a reintroduction program was started in Corsica. Barbary Red Deer became extinct in Morocco by 1932, and in early 1960 only a few dozen animals survived in forests in the Atlas Mountains between Algeria and Tunisia. More effective protection allowed a recovery in Algeria and Tunisia, and in 1994 a reintroduction was begun in Morocco. The Bukhara Red Deer originally lived in the basins of the Amu Darya and Syr Darya rivers. Tree felling, cattle grazing, and overhunting led to a rapid decline during the 19" century. In early 1900 it disappeared from Syr Darya. In the mid-1960s the population reached a minimum of 350-400 animals; in the early 1980s it had grown to 1100. The Tarim Red Deer of Chinese Xinjiang had a declining population of 4000-5000 in 1991, scattered along the Tarim and Karakash rivers.

Bibliography. Apollonio et al. (2010), Butzler (1986), Carnevali et al. (2009), Clutton-Brock & Albon (1989), Clutton-Brock et al. (1982), Danilkin (1999), Gebert & Verheyden (2008), Gill (1990), Mattioli, Fico et al. (2003), Mattioli, Meneguz et al. (2001), von Raesfeld & Reulecke (1988), Skog et al. (2009), Sommeret al. (2008), Staines et al. (2008), Wagenknecht (1986), Zachos & Hartl (2011).

Figures

23. Sika Deer (Cervus nippon), 24. Central Asian Red Deer (Cervus wallichii), 25. Western Red Deer (Cervus elaphus), 26. Wapiti (Cervus canadensis)

Subspecies and Distribution. C.e.elaphusLinnaeus,1758—GreatBritainandcontinentalEurope. C.e.bactrianusLydekker,1900—WTurkestan(Kazakhstan,Uzbekistan,Tajikistan&NAfghanistan). C.e.barbarusBennett,1833—AtlasMts(Algeria,Tunisia). C.e.corsicanusErxleben,1777—Corsica(extinct,reintroducedin1985),Sardinia. C.e.maralGray,1850—Anatolia,Caucasus,andNWIran. C. e. yarkandensis Blanford, 1892 — E Turkestan (S Xinjiang, China ). It has been introduced into Chile , Argentina , Australia , and New Zealand .