Old World Deer (Cervinae)

(Cervinae)

Class: Mammalia

Order: Artiodactyla

Family: Cervidae

Subfamily: Cervinae

Thumbnail description
Only proximal parts of metacarpal bones of the second and fourth fingers (Plesiometacarpalian deer) are retained; vomer is short and does not divide posterior nasal holes; only males have antlers

Size
Large- and medium-sized deer

Number of genera, species
4 genera; 14 species

Habitat
Tend to stay in forests, woodlands, forest-steppe, partly in forested mountains; can adapt to varieties of habitats from marshlands to alpine meadows

Conservation status
Extinct: 1 species; Critically Endangered: 1 species; Endangered: 3 species; Vulnerable 3 species; Data Deficient: 2 species

Distribution
Most of Old World, except Africa (only in northwest and introduced into South Africa); in North America, only one species (Cervus elaphus), which is a newcomer

Evolution and systematic

The most ancient deer forms, similar to the modern Axis deer, appeared in early Pleistocene, about two million years ago. A comparative study of extinct and existing forms revealed some trends in body construction and ecology, which helped distinguish between ancient and recent types of Cervinae. The most primitive of deer, they are distributed in India, China, and Indochina, and inhabit tropical forests, shrub lands, and grass thickets, often close to water basins. Species more advanced in evolution adapted to forests that interchanged with clearings so that their diet preferences are wider, and include leaves, tree and bush branches, forbs, and herbs. Some species such as Siberian maral (Cervus elaphus sibiricus) feed on grasses in high numbers, though they need a mixed diet that includes tree or bush leaves and twigs. None of the Cervinae became true grass and roughage eaters like sheep or cattle. A mixed diet facilitated the adaptation to a wide variety of habitats, including forest edges (with raids to neighboring grasslands), as did red deer, fallow deer, and barasingha. Teeth developed higher crowns and so are now adapted to partial feeding of herbal forages. Lower jaws grew longer to develop a significant gape between fangs and premolars. Incisors are still wide, with asymmetric outer edges, adapted to grazing softer vegetation and to browsing. Intestines in red deer and fallow deer are 15–17 times longer than body length, while in true grass and roughage eaters, the length of intestines exceeds body length by 25–30 times.

Deer that inhabited northern areas differed from southern ones by the type of their coat. They had a thick undercoat, and hair in winter become air-filled and wavy, serving as an impermeable insulate for the animals.

Pere David's deer (Elaphurus davidianus) and barasingha (Cervus duvaucelii), as inhabitants of marshlands, had long hooves. The species that occupied mountainous areas, such as white-lipped deer (Cervus albirostris) had shorter and higher hooves, more adapted to harder ground. As well, the heels of the feet were enlarged, adding to their resilience on hard surfaces. When they moved from a continent to archipelago island, deer usually diminished in size, such as the Axis deer of the Indonesian archipelago.

Most deer fawns develop white spots, probably for camouflage. Most primitive deer are often spotty in the adult state, though they manifest no color differences in winter or summer coats, or between sexes. Multicolored ornate coats and different colors in bulls and hinds distinguish the evolutionary advanced species. They also develop a bright, white rump patch (a mirror) around the tail. These color differences all relate to more complicated social behavior. Both mirror and white underparts of the tail enable the herd to more easily follow a leader when danger threatens.

Antlers have become more and more significant as weapons, with the development of one or two brow tines that are important in defending themselves in rival fights. Three-pronged deer with long tails (hog deer, Axis porcinus; spotted deer, Axis axis; barasingha, Cervus duvaucelii; and sambar, Cervus unicolor) are found in the tropics; four-pronged deer in the warm, temperate zone (sika deer, Cervus nippon; and fallow deer, Dama dama), five- and six-pronged deer in the cold and alpine zones (izubr in Siberia, Cervus elaphus xanthopygus, elk in northern America, C. e. canadensis; and white-lipped deer, Cervus albirostris).

Cervinae (in particular the genus Cervus), compared with Odocoileinae, have lower reproductive rates, though they better utilize feeding resources and are more resistant to helminth diseases. The high degree of polymorphism is good for selection. The ability of interspecies (and even intergenera) hybridism is an important evolutionary feature of Cervinae. They are well adapted for intensive husbandry.

Physical characteristics

Deer vary in size from small to very large, standing high at the shoulders from 25–59 in (63–150 cm). Body is elongated, while legs are short or medium in length. A large bald spot partly covers the nostrils. Ears vary in size from short to long. Mane is typical for some specialized forms; primitive forms have no mane. The tail varies from extremely short and hidden by hairs, to very long. Antlers are rounded, with no less than three tines, and the brow tine is always developed. Some species wear antlers palmate at the top. Incisors are wide, with high crowns, while middle incisors have an elongated outer edge. Some species lack upper fangs. Molars always have high crowns.

Distribution

They occur in Europe and Asia, excluding areas to the north of 60°N, to northwest Africa. In North America, they occur from 60°N to Mexico. They are farmed in New Zealand, Australia, and the South African Republic.

Habitat

Deer inhabit predominantly ecotone habitats such as forest edges, and tend to move to more open habitats like alpine meadows, steppes, and farmed fields, rather than deep into forests. Some species inhabit tall shrub and grass thickets or marshlands.

Behavior

Most species live in small to moderately sized groups. In places of abundant forage, aggregations of tens and hundreds of deer gather. Bulls in these aggregations behave reasonably well, though they immediately establish hierarchy, which causes young males to stay at the periphery. In many species, bulls gather and defend harems of several does during the rut, trying to keep them on their breeding patches, and defend territories against rivals. Deer that inhabited thick groves, jungles, tall grass thickets in river or lake banks behave more like the Capriolinae: they live a solitary life, often are nocturnal, and are strictly linked to their home ranges.

Feeding ecology and diet

The type of feeding in these deer is mixed—they consume both concentrated forage (leaves, soft forest herbs, and fruits) and meadow grasses. This feature facilitates the farming and park breeding of them as they find the food supply (hay) to be acceptable.

Reproductive biology

The reproductive rate of Cervinae is lower than in Odocoileinae. Does give birth to one fawn, and start mating at one and half years. Stags start mating at the age of five years, when they are in full physical maturity and are strong and heavy enough to dominate rivals. Most species are polygamous, with bulls gathering and defending a harem of several does.

Conservation status

For a long time, deer of these subgenera have been game animals, some since ancient times. As a result, many species have become threatened or scarce. Schomburgk's deer (Cervus schomburgki) may be Extinct. Pere David's deer (Elaphurus davidianus) is Critically Endangered; Calamian deer (Axis calamianensis), Bawean hog deer (Axis kuhlii), and Philippines spotted deer (Cervus alfredi) are Endangered. Many are considered

Vulnerable species, including barasingha (Cervus duvaucelii) and Eld's deer subspecies (C. eldi eldi from Manipur state in India and C. e. siamensis from Thailand, Vietnam, Kampuchea, Laos, and Hainan Island).

Significance to humans

Cervinae includes important game and farming deer due to venison (recently, it has become more highly esteemed due to its low-fat content) and skins (its raw material makes for the best suede). Many parts are valuable for Asian medicine (their prices have exploded since 1960), including velvet antlers, hard antlers, tail, bones, penis, heart, liver, sinews, placenta, and blood. The annual import of velvet antlers to Taiwan reaches 12 tons (11 tonnes) (including a minor percentage of reindeer velvet antlers); Korea also imports 12 tons (11 tonnes) annually and Thailand imports 1.1–3.3 tons (1–3 tonnes). The main antler manufacturer is China; its output is 44–55 tons (40–50 tonnes) per year (mostly from sika deer, and some from red deer). In Russia, which exports 13.2–15.4 tons (12–14 tonnes) per year, there are farms of red deer and sika deer. Antlers are also considered valuable trophies. Since medieval times, castles were adorned with deer heads with magnificent antlers.

Species accounts

Resources

Baskin, Leonid, and Kjell Danell. Ecology of Ungulates. A Handbook of Species in Eastern Europe, Northern and Central Asia. Heidelberg: Springer Verlag, 2003.

Bedi, Ramesh. Wildlife of India. New Delhi: Bridgebasi Printers Private Ltd., 1984.

Flerov, Konstantin K. Musk Deer and Deer. Moscow: Izdatelstvo Akademii Nauk SSSR, 1952.

Geist, Valerius. Elk Country. Minoqua, WI: North Word Press, 1991. ——. Deer of the World: Their Evolution, Behavior, and Ecology. Mechanicsburg, PA: Stackpole Books, 1998.

Hudson, Robert J., Karl R. Drew, and Leonid M. Baskin. Wildlife Production Systems. Economic Utilization of Wild Ungulates. Cambridge: Cambridge University Press, 1989.

Harrington, R. "Evolution and Distribution of the Cervidae." In Biology of Deer Production, edited by P. F. Fennessy and K. R. Drew. Wellington: The Royal Society of New Zealand, 1985.

Hofmann, R. R. "Evolution Digestive Physiology of the Deer—Their Morphophysiological Specialization and Adaptation." In Biology of Deer Production, edited by P. F. Fennessy and K. R. Drew. Wellington: The Royal Society of New Zealand, 1985.

Schaller, George B. Wildlife of the Tibetan Steppe. Chicago: University of Chicago Press, 1998.

Sheng, Helin, and Lu Houji. The Mammalian of China. Beijing, China Forestry Publishing, 1999.

[Article by: Leonid Baskin, PhD]