New World deer

(Capriolinae)

Class Mammalia

Order Artiodactyla

Family Cervidae

Subfamily Capriolinae

Thumbnail description
New World deer have preserved the distal rudiments of the lateral metacarpal bones; the middle parts of the lateral metacarpae are reduced; and posterior portion of the nasal cavity is divided into two chambers by the vomer, which feature is retained in South American deer, but is lacking in moose and roe deer

Size
Vary in size from very small (pudu) to the largest among Cervidae (moose)

Number of genera, species
9 genera; 27 species

Habitat
Woodlands and shrublands, often forest edges; many populations of reindeer inhabit tundra or arctic desert yearlong, others migrate to openness of subarctic tundra for summer

Conservation status
Endangered: 1 species; Vulnerable: 2 species; Lower Risk/Near Threatened: 3 species; Data Deficient: 7 species

Distribution
North America, South America, Europe, and northern Asia

Evolution and systematics

Recent paleontological research in the high Arctic suggest that as late as mid-Pliocene times the Arctic portions of North America and Asia were covered by temperate forests. Together with the zoogeography and fossil record of New World deer species, this suggests that their origin lay in these northern land masses in late Tertiary times. From here they dispersed southward as climates cooled into periodic continental glaciations. The fossil record of New World deer is limited by the facts that huge continental glaciers destroyed Tertiary fossil deposits over enormous areas, while in unglaciated regions where rich plant fossil deposits are found, the acidic conditions favoring these dissolve bones. We have thus no idea what early New World deer looked like. When they first appear in areas south of the continental glaciations in North America or Eurasia, they are already well differentiated and close to modern genera. After they enter South America with the onset of major glaciations some 2 million years ago (mya), they evolved rapidly into a large array of diverse species, only some of which survived to the present. Here they evolved dwarfs, such as pudu (Pudu pudu, P. mephistophiles) and brocket deer (Mazama spp.); mountain climbing, short-legged specialists such as the huemul (Hippocamelus antisensis, H. bisculus); sophisticated swamp dwellers such as the marsh deer (Blastocerus dichotomus); gregarious plains dwellers such as pampas deer (Ozotocerus bezoarticus); as well as massively antlered, large-bodied steppe deer that are now extinct (Morenelaphus). In the north they evolved giants such as moose (Alces); herd forming reindeer or caribou (Rangifer tarandus); a large-bodied extinct form Torontoceros; forest and swamp dwellers such as the ancient white-tailed deer (Odocoileus virginianus), the oldest deer species in the world; rainforest and mountain dwellers such as black-tailed and mule deer (O. hemionus); as well as a large, short-legged, cliff adapted Rocky Mountain deer, Navahoceros, which vanished at the end of the Pleistocene. Closely related to these North Americans are the roe deer (Capreolus capreolus, C. pygargus) of Eurasia, which is a cold-climate specialist and the Chinese water deer (Hydropotes inermis). A sparse fossil record suggests that there were a few more species present in late Pliocene times. Compared to Old World deer, New World deer are more differentiated from and less related to on another. They thrived in the ecological turmoil of the Pleistocene and have often taken advantage of humanmade landscape changes in the Recent. They also suffered fewer extinctions than other groups of large mammals.

A characteristic of New World deer is that they appear in the fossil record in order of adaptation to cold climates. The

species most tolerant of heat, and which therefore disperses into and colonizes tropical South America, is the white-tailed deer. A species very similar, if not identical, to it shows up in southern North America at the end of the Blancan period almost 4 mya. Moose appear about 2.6 mya. Roe deer appear about 2 mya, and reindeer, the most cold-adapted genus, appears about 1 mya. This genus is the sister genus to Navahoceros, suggesting that competition segregated these originally montane lineages, with reindeer exploiting the alpine and subalpine slopes while Navahoceros sought refuge in cliffs. Similar ecological division is seen in extant Asiatic goat (Capra) and sheep (Ovis).

Moose appear in the Pliocene fossil record first in western Eurasia. They are then large, plains-dwelling, long-legged runners (Alces [Libralces] gallicus), about the size of a red deer (Cervus elaphus), with a normal deer face and extraordinary long spoon-shaped antlers. There appear to be several species. Moose are next seen in mid-Pleistocene Eurasia during the major glaciations. It has grown into a massive giant (Alces [Cervalces] latifrons). The huge antlers are more palmate and have shorter beams than the Pliocene specimen. The skull is intermediate between that of a normal deer and contemporary moose. Such moose cross into North America, where they grow into long-legged trotters with complex large tri-lobed antlers, but retain the primitive deer-like skull (Cervalces scotti). This American stag-moose is narrowly associated with huge Pleistocene pro-glacial lakes, which were apparently its escape terrain from the many large predator species found in North America. In Siberia moose continue to evolve into the modern moose. These have even larger palms and much shorter antler beams, the muzzle became adapted to feeding on underwater vegetation, while the body changed from that of a cursor (runner) to that of a trotter. After post-Pleistocene megafaunal extinction in North America, Siberian moose colonize the northern half of that continent beginning about 10,000 years ago, where they are now widely distributed. Two fairly distinct modern moose evolved, one the west Siberian-European form, and the other the east Siberian-American form. Despite great physical differences, moose share diagnostic behaviors with white-tailed deer, mule deer, and caribou. They twin readily, and moose remained in cold climates throughout.

The Odocoileinae comprise four groups of species: North American Odocoileus deer and South American deer, reindeer, moose, and roe deer. They all branched early from an ancestral stem into different evolutionary radiations, but retain common features in body plan, behavior, and ecology. The Odocoileus and South American deer comprise one tribe, according

to G. G. Simpson. These deer may be found from the Arctic Circle in Canada south to the glaciers of southern Chile, but the majority of species live in warm climates. Moose, reindeer, and roe deer are each placed into separate tribes. These deer live in the cold northern environments, including the high arctic, and have never colonized southern latitudes.

All Odocoileinae retain distal rudiments of the lateral metacarpal bones II and V, but which still retain hoof functions. Another feature is the division of the posterior portion of the nasal cavity into two chambers by the vomer. However, this feature is missing in moose and roe deer. Males have a pendular penis. Antlers are found in all genera. The dwarf deer of South America may be secondarily dwarfed, as they are exceedingly closely related to the older and more ubiquitous white-tailed deer. This species ranges from near the Arctic Circle in Canada to 18° south of the equator in South America. The short dagger-like antlers of the dwarf deer may be related to territorial defense. Reindeer and caribou have the largest antlers relative to body size among deer, while their

females also carry antlers except in some woodland populations. They also are the most cursorial and gregarious deer alive.

Physical characteristics

All Capriolinae retain distal rudiments of the lateral metacarpal bones II and V, though they are important to the functioning of lateral toes. Another feature, the posterior portion of the nasal cavity that is divided into two chambers by the vomer, is retained in South American deer, but is lacking in moose and roe deer. There are different varieties of antler structure in Odocoileinae: simple spiked (Mazama, Pudu), bifurcate (Hippocamelus), dichotomous (Blastocerus), or branched (Odocoileus, Rangifer). Moose often sport wide spade-like antlers. Normally, only males wear antlers, though reindeer females grow antlers as well.

Distribution

Six genera, including Odocoileus, Ozotocerus, Blastocerus, Hippocamelus, Mazama, and Pudu, live only in the New World. One genus, Capreolus, is known in Eurasia only. Alces and Rangifer inhabit both North America and Eurasia.

Habitat

Deer belonging to Capriolinae adapt to diverse habitats. Dwarf forms of deer with short antlers and long tails (brocket deer, Mazama) are inhabitants of tropical latitudes, while large

deer with dichotomous antlers (marsh deer, Blastocerus dichotomus) inhabit tropical and subtropical marshlands. Tropical savanna is the favored habitat for pampas deer (Blastocerus campestris). Subarctic dwellers are roe deer, white-tailed deer, black-tailed deer (Odocoileus hemionus), and moose and reindeer inhabit cold temperate subarctic forests, alpine, and subarctic tundra.

Behavior

The dwarf and mountain deer of South and Central America are classical territory defenders and hiders. Capreolus males defend large territories and bond females that live within their territory. This species is a classical saltor (jumper) that relies on short runs over obstacles and then hides. Odocoileus may defend fawning territories, but otherwise related females form clans so that they move over a shared home range. Males form unstable fraternal groups but disperse and compete individually over females during the rut. These deer rely on sprinting or specialized locomotion to escape predators as well as on hiding. They may form a large selfish herd in open grasslands. Pampas deer also rely on large selfish herds to escape predators, while swamp deer use extensive wetlands for that purpose. South American deer, unlike North American species, are exceedingly sensitive to predation by feral dogs as there are no native wolf-sized canids in South America. Moose live dispersed much of the time, but bulls may form unstable fraternal groups in spring and after the rut in fall. Female moose are likely to join bulls socially only if they are neither pregnant nor have a calf at heel. Moose escape predators primarily by running over obstacles that are low relative

to the moose, but high relative to the pursuing predators, who then must expend great amounts of energy to follow the fleeing moose. In deep snow moose turn and fight predators using both front and hind legs. Reindeer and caribou form highly gregarious selfish herds and excel at sustained high speed running to escape predators. In sedentary populations females disperse and give birth in hiding. In migratory populations they move northward in spring onto open terrain where they congregate into birthing herds. This overloads pursuing wolves and grizzly bears with calves. The calves are highly developed at birth and soon follow the female, who produces the richest milk among all deer, ensuring rapid growth of the calf to a survivable size. This is the most migratory species of large terrestrial mammal. Bulls advertise with antlers during the rut. Females use antlers to ward off young bulls in winter, who might otherwise parasitize the female's work of digging craters in deep snow to reach lichens. Both Rangifer and Alces are large Ice Age giants compared to other species in their family.

Feeding ecology and diet

The Capriolinae diet comprises highly nutritive forages of low-fiber content: forbs, flowers, and leaves, but rarely

grass. The type of diet determines speciation of the digestive system: a large mouth enables the browsing of branches, while a long sensitive tongue helps to choose among forbs and foliage. Some species have a relatively small rumen, large salivary glands, and rapid digestion. Intestines are rather short—12 to 15 times longer than body. Consequently, interchanging periods of grazing and ruminating are short. If these deer, by necessity, feed on rough fibrous forage, there are fewer interchanging of grazing and ruminating periods during their diurnal activity (only five to six, rather than the usual eight to 12).

Reproductive biology

Capriolinae are polygynous, though the overall reproductive strategy of Capriolinae differs from that of Cervinae. They have a higher reproductive rate and mature sexually earlier; most genera produce two fawns per birth. Consequently, Capriolinae species have higher population densities, including mule deer, caribou, and moose. At the same time, the species are reproductively isolated: for instance, male hybrids between the European and Siberian roe deer as well as between white-tailed deer and mule deer are sterile. Hybridization between white-tailed and black-tailed deer is restricted. Capriolinae poorly resist diseases and parasites, and poorly adapt to new conditions.

Conservation status

Deer that inhabit Central and South America (pampas deer, Ozotoceros bezoarticus; marsh deer, Blastocerus dichotomus; huemul and pudu, Pudu spp.; brocket deer, Mazama spp.); and

some subspecies from the southern part of North America have experienced severe pressure from hunters and have been on the brink of extinction. These species would greatly benefit from preservation and conservation efforts. Pampas deer, for instance, were harvested by the millions in the nineteenth century, and currently is Lower Risk/Near Threatened. The subfamily Capriolinae also includes the most numerous species on Earth, including white-tailed deer, black-tailed deer, reindeer, roe deer, and moose.

Most threatened are the Capriolinae in South America, where poor residents use subsistence hunting and do not distinguish between rare species that need to be preserved and flourishing species that need regular game management. In many South American countries, there is a fierce competition between deer and livestock for pastures. Also, a drastic decrease of habitats occurs due to drainage, farm development of grasslands, and forest cut. The Chilean huemul (Hippocamelus bisulcus) is considered Endangered. Marsh deer (Blastocerus dichotomus) and Chilean pudu (Pudu pudu) are considered Vulnerable. Less studied, Data Deficient are: Peruvian huemul (Hippocamelus antisensis), and six species (of seven known) of brocket deer (Mazama spp.).

Significance to humans

Roe deer, moose, and white-tailed deer benefit from successful, extremely productive game husbandry, though farming proved to be unsuccessful. Consequently, their future lies in game management.

Reindeer is the only deer species to become domesticated. It supports local cultures throughout northern Eurasia in this capacity. Wild migratory reindeer and caribou are also significant in northern economies. While moose can be tamed and used for riding, as beasts of burden, and for milking, their fickle feeding habits and susceptibility to livestock diseases makes them difficult to keep. They are a productive, highly appreciated meat source throughout their range. Whitetailed, black-tailed, and mule deer, after recovery from severe depletion at the end of the nineteenth century, support today a rich hunting economy in North America. Roe deer fulfill a similar role in Europe.

Species accounts

List of Species

European roe deer

Capreolus capreolus

taxonomy

Capreolus capreolus (Linnaeus, 1758), Sweden.

other common names

French: Chevreuil; German: Reh; Spanish: El corzo.

physical characteristics

Small animal. Shoulder height: 24–35 in (60–90 cm); body length: males 39–54 in (100–137 cm), females 37–54 in (94–136 cm); weight: males 46–75 lb (21–34 kg), females 42–71 lb (19–32 kg). Very slender, with long, slim legs and nearly invisible tail. Preorbital gland is rudimentary. There is large bald spot on the end of the muzzle between and around nostrils. Antlers are small, usually forked in three at the end, with a lack of brow tine. Small protuberances develop along beams. Coat is monotonously colored, gray in winter (with brownish or reddish tint) and red with lighter belly in summer. Rump patch is white. Fawns develop spots.

distribution

Throughout Europe, including Britain and Sicily (except in Corsica and Sardinia); none in close coniferous forests of Scandinavia and northern Russia.

habitat

Inhabit dry plains and mountains, in varieties of landscapes where forest islands interchange with steppe and meadows. Mature broadleaved forests also attract them, as well as shrubs and tall grass, which are used for shelter. In many countries of Central and Western Europe, they spend most time on farmed fields.

behavior

Live solitarily or in small family groups (doe, fawn, yearling), except in rut season. Aggregations to tens of animals without social

bonds appear in areas with abundant forage. Once disturbed, animals scatter. Males defend their territories against invaders: they mark it by horning trees, as well as leaving secretions from scent glands. Does keep to their home ranges and defend it from other does. Strong bonds to home ranges are a distinguishing feature of roe deer. A male home range about 990–2,100 ac (400–850 ha) usually overlap some ranges of does (490–700 ac [200–700 ha]).

Combats between bulls are frequent, even out of rut season; males even fight females. Good swimmers and jumpers. Diurnal activity is highest at dawn and dusk. Extremely wary; looking around takes up to 50% of active time. When disturbed, they make a typical bark, strike the ground with the front leg hooves, run away showing bright white tail mirror, and making signal jumps.

feeding ecology and diet

Feed mostly on herbs, less on leaves, buds, fruits, cereals, and sedges. In winter, twigs are preferable, as well as dry grasses, fallen leaves, mosses, and tree lichens; in summer, they browse green soft shoots of cereals and sedges. Fungi are eaten year-round. To reach upper branches or highly hanging fruits, they rise on hind legs. Animals even dig at snow or soil to get to roots in farmed fields. Can stay without water, using moisture in vegetation, and snow in winter.

reproductive biology

Polygynous. First signs of rut appear in July into the beginning of August, or sometimes in May–June; peak of rut lasts from mid-July to the end of September (depending on latitudes). There are no harems; mating takes place when females stay in male's home range. Does are in heat for 4–5 days, gestation period lasts about 240 days. Calving takes place in herb or shrub thickets; one to two fawns per birth; fawns stay hidden for the first 6–8 days, rising only to be nursed; they later follow mothers, start feeding on herbs and leaves at one month, and are weaned at two months. Life expectancy is 11–12 years.

conservation status

Not threatened.

significance to humans

Subject of game hunting, both venison and antlers are main attractions.

Siberian roe deer

Capreolus pygargus

taxonomy

Capreolus pygargus (Pallas, 1771), Volga area, Russia.

other common names

French: Chevreuil de Siberie; German: Reh von Sibirien; Spanish: Corzo siberiano.

physical characteristics

Middle sized deer. Body length: males 47–61 in (120–156 cm), females 46–59 in (116–150 cm); weight: males 66–132 lb (30–60 kg), females 55–121 lb (28–55 kg). Coat in winter is grayish to brownish on back, and creamy at belly and inside legs. Rump patch is white or cream. Summer coat is red at head and body. Fawns develop distinct spots arranged in four or five rows. Antlers to 16 in (40 cm) and long, pockmarked with bumps, some of which transform to protuberances and tines.

distribution

From the Volga to Russian Far East and northern China via northern Kazakhstan and north of Middle Asia.

habitat

Inhabit both plains and mountains to altitudes of 6,900 ft (2,100 m). The species adapt to deep snow to 20 in (50 cm) and to harsh winters, surviving in Yakutia and Transbaikal; also inhabit pine forests and mature coniferous-deciduous forests.

behavior

In winter, they form groups of four to six, live solitary (does with fawns). Daily home range of 99 ac (40 ha) and annual range about 990 ac (500 ha). In Amur basin, home range in winter is to 34,500 ac (14,000 ha). Migrations (to avoid deep snow) to distances 62–250 mi (100–400 km) are frequent. In mountain ranges of the Caucasus, Altai, and the Urals, they make short migrations between altitudes 1,600–3,300 ft (500–1,000 m).

feeding ecology and diet

In winter, feed on tree and shrub branches, dry herbs, fallen leaves, mosses; in summer, mostly on sedges and grasses.

reproductive biology

Polygynous. Rut peak is in August to the beginning of September, calving is May–June. Gestation period lasts 264–318 days. Does give birth to one, twins, or triples. Lifespan is about seven years; average age in wild is two and a half years. Strong hunting pressure and predators cause high mortality.

conservation status

Not threatened.

significance to humans

Important game animal; in Russia, annual harvest is 5,000–10,000 deer, mostly in the Urals and in Amur valley.

White-tailed deer

Odocoileus virginianus

taxonomy

Odocoileus virginianus (Zimmermann, 1780), Virginia, United States.

other common names

French: Cerf de Virginie; German: Weisswedelhirsch; Spanish: Ciervo de Virginia.

physical characteristics

Small. Shoulder height: males 39 in (100 cm), females 35 in (90 cm); body length: males 77 in (195 cm), females 67 in (170 cm); tail length: 11 in (27 cm); weight: males 128–300 lb (58–136 kg), females 110–175 lb (50–79 kg). Newborns weight 4–9 lb (1.8–4 kg); reach 55–86 lb (25–39 kg) by six months. In summer, coat is a foxy-red color, changing in autumn to tawny gray, with longer and thicker hairs. Newborns have reddish coat sprinkled with light spots. Under part of tail is bright white. Usually only bucks wear antlers. Glands producing strong scent are well developed on all four hooves.

distribution

Distributed from Atlantic shore to Pacific shore of North America; from central Canada to Bolivia, Guiana, and northern Brazil.

habitat

Recently, they have come to inhabit edges of forest clearings: burned areas, logged sites, fields, and meadows. Clumps of broadleaved and coniferous forests in the middle of fields are their favored habitats. Also inhabit rugged river valleys, sandy hills covered by grass and trees, and sometimes stay in large forests. In summer, habitats are more diverse, including fields and meadows. In winter, deer keep to forests, especially coniferous stands, where they find shelter from harsh elements. Home range is from 0.1–1.5 mi² (0.2–4 km²). Snow deeper than 17.7 in (45 cm), lasting more than two months, harsh weather, and ice crust are important elements of habitat. Use thick forest for cover from predators and hunters, as well as in tall grasses and bushes in prairies and riparian habitats.

behavior

When deer run from danger, the tail bounces loosely from side to side, with white hairs erected; it serves as a signal of danger to all surrounding deer. It also helps a fawn following after the mother. They can run as fast as 30 mph (50 km/h), jump to 10 ft (3 m) high and to 30 ft (9 m) long.

feeding ecology and diet

Up to 70% of diet consists of tree and shrub leaves and twig ends; 19% is forbs; 11% is grasses. Agricultural crops accounted for 3% of yearlong diet. An adult needs 5–11 lb (2.5–5 kg) of forage daily. In winter, they can survive on two lb (1 kg)

of food daily. If succulent plants are available, they can manage without water for a long time, though watering places often are center of home range.

Preferable winter and autumn food: blackberry (Rubus spp.), dogwood (Cornus spp.), snowberry (Symphoricarpos spp.), chokecherry (Prunus virginiana), rose (Rosa spp.), Oregon grape (Berberis repens), ponderosa pine (Pinus ponderosa), elms (Ulmus spp.), and maples (Acer spp.). Nuts of oaks, hickories, beech, and walnuts and fungi are important.

reproductive biology

Polygynous. Bucks shed antlers annually in winter; re-growth starts at the end of April to the beginning of May. Rut begins in October, to reach peak in mid November and end in December. At the beginning of rut, bucks get a swollen neck and they clean antlers of velvet; establish dominance hierarchy in October, displaying scraping activity: pawing the ground until leaf or grass litter is removed and bare ground is exposed. The animal then urinates on the bare spot. Does are in heat only one day, those that do not breed go back into heat in 28 days. Bulls keep no harems; stay with a doe a day or two, and after breeding, they search for another one, thus having 6–8 does mated during rut. Gestation period is 188–222 days, fawns (usually twins, rarely triplets) appeared in May to the beginning of June. Within the first hours, they can suckle and follow mother; nevertheless, stay hidden in bushes or tall grass first days of their life. In 10 days, they start nibbling green shoots. Calves participate in rut from their first year. There are 40–75% of pregnant does among very young ones. High productivity goes with high mortality in this species. Annual rate of mortality in the species is 30–50%. Life expectancy is six years; some individuals in wild live to 14 years, in captivity to 20 years.

conservation status

Not threatened.

significance to humans

Important game species; the most numerous of big game animal in the world. Of 15 million white-tailed deer in North America, annual harvest is about three million animals.

Black-tailed deer

Odocoileus hemionus

taxonomy

Odocoileus hemionus (Rafinesque, 1817), South Dakota, United States.

other common names

English: Mule deer; French: Cerf-mulet; German: Schwarzwedelhirsch; Spanish: Ciervo mulo.

physical characteristics

Medium size. Shoulder height: 37–39 in (95–100 cm); bulls weigh 220 lb (100 kg) with body length to 77 in (195 cm); does weigh 143 lb (65 kg) with body length to 62 in (160 cm); tail: 7 in (18 cm). Head is narrow, elongated, with big nasal bald patch. Hairs are longer at the top of neck, no mane. Adult have a mono-color coat, mostly black tail surrounded by a smaller white rump patch. Develop long ears, big antlers, and white tail with a black end. Fawns are reddish, lighter colored at under parts, with very long tail. Only bulls wear antlers, varying from simple spikes to forked, depending on bull's age.

distribution

Western North America from the southernmost Yukon to Mexico.

habitat

Interspersion of food and cover is the major factor in habitats. Animals gather at cut clearings, rich in forage, and keep no farther than 330 ft (100 m) from edge of forest to escape when disturbed. In areas with deep snow, prefer southern slopes at altitudes less than 3,300 ft (1,000 m). Rich communities of herbs, ferns, and shrubs make the best spring ranges, usually at low elevations of less than 2,600 ft (800 m), southerly aspects, and moderate to steep slopes. Good spring range sometimes extends onto lower, gentler slopes where rich, moist soil produces more herbs and snowmelt earlier. Burned clear-cut sites often support abundant herbs—especially fireweed. Dense thickets of young cedar and hemlock provide security cover when they retain live branches within 7 ft (2 m) over the ground. Deep snow is critical for survival; though they move even through incrusted snow as deep as 20–25 in (50–60 cm), it takes too much energy.

behavior

Live solitarily or in small groups (a mother and her young of the current and previous years) most of the year. Gather at places with abundant food or in shelters against harsh weather; no social bonds between individuals. Distance between summer and winter home range rarely exceeds 8 ft (2–3 m). Summer home range is 0.6–2.3 mi² (1.5–6 km²); winter home range is 1.4–2.3 mi² (3.5–6 km²). In winter and spring, they live in groups comprised of both sexes and any age. As spring approaches, does wean yearlings and soon leave herd. In summer, does live isolated, nursing their fawns. Bulls in summer live in separate groups; young form temporary groups, easily scattered, and gather in new ones. At the approach of rut, in September–October, bulls become intolerant of each other. Does return to their winter groups, especially for feeding time. At the peak of rut, in November–December, bulls antagonistic toward each other, start antler fighting (mostly frequent among two-year-old males). Does in heat are driven from their groups by dominating bulls. At the end of rut, the estrous does rejoin their clans after breeding. Dominating bulls lose interest in does and search for good feeding places to graze in solitude. Later bulls return to their clans.

feeding ecology and diet

In winter, western red cedar, Douglas fir, western hemlock, blueberry, deer fern, bunchberry, salal, and the arboreal beard lichens are the most important forages. In spring, diet is Douglas fir, different species of Rubus (salmonberry, blackberry, thimbleberry, raspberry, bramble), salal, willows, bracken, Pteridium acquilinum, fireweed, horsetail, and pearly everlasting. There is a significant seasonal difference in nutritive value and digestibility of forage. In winter, they lose 20–25% of their autumn weight. During spring, most of the important nutrients in newly grown material are readily digestible.

reproductive biology

Polygynous. Bucks shed antlers in January and re-grow during summer. Rut occurs from mid November to the beginning of December. Does are in heat repeatedly 22–29 days, with most does conceiving during their second ovulation. Three to four days before heat, a bull starts following a doe and stays with her three to four days after mating; during a rut, a bull can only mate three or four does. Gestation period lasts 200 days; fawns appear at the first half of June. Sometimes yearlings mate at their first year, and 45–80% become pregnant, while 90–95% of does older than two years become pregnant. By end of year, each 100 does bring 41–78 fawns. Along with high rate of productivity, display high mortality. Lifespan is six years. One-third of population perishes each year, especially after a harsh winter.

conservation status

Not threatened.

significance to humans

Important game species. In North America, population numbers of black-tailed deer is about 1.5 million and annual game harvest about 0.5 million.

Marsh deer

Blastocerus dichotomus

taxonomy

Blastocerus dichotomus (Illiger, 1815), Lake Ypoa, Paraguay.

other common names

French: Cerf des marais, cerf de marecages; Spanish: Ciervo del los pantanos.

physical characteristics

Largest of Cervidae in South America. Shoulder height: 3.6–4 ft (110–120 cm); head and body length: 70-80 in (180–200 cm); tail length: 4–6 in (10–15 cm); weight: 154–242 lb (70–110 kg), maximum 330 lb (50 kg). Male antlers: 24 in (60 cm) in length, dark yellow, doubly forked, each with four or sometimes five tines; weight of antlers 5.5 lb (2.5 kg). Harsh and longhaired coat is reddish brown to chestnut, lighter in lower parts, with

black muzzle, lips, and legs, and white orbital rounds. Large ears adorned by fluffy white hair. Tail is yellowish red above and dark brown to black underside. Fawns develop no spots, unlike other deer species. Hooves are well adapted to boggy habitats: with developed dewclaws, long hooves to 2.8–3.2 in (7–8 cm), with widely splayed middle "fingers."

distribution

Keeps to upper reaches of Rio Paraguay where the world's largest floodplain exists.

habitat

Inhabit floodplains and marshlands, sometimes interchanged with islands of wet savanna covered with high grass and small forest. Connected to riparian habitats, migrating near grasslands in flood time.

behavior

Mostly nocturnal; start grazing at dusk at marshy clearings. They stay easily in water, though prefer shallow waters. During floods, they move to higher elevations. Live either solitarily or in small family groups (to six animals), including male, some females, and their offspring.

feeding ecology and diet

Feed on herbs, reeds, and aquatic plants.

reproductive biology

Polygynous. No definite rut season or definite time of antler shed. Fawns can be found year-round. Gestation period is about 260 days; usually one young per birth. Fawn follows the mother until one year old, though weaning is usual in about five months. Both sexes reach sexual maturity at one year. Females mate again just after parturition.

conservation status

Today considered an Endangered species, though in 1996 was regarded as Vulnerable. It is presumed that population of marsh deer is rapidly declining due to destruction of preferable habitats. Recent census shows that, in poorly accessible marshes in upper reaches of Rio Paraguay in south Brazil, still high density of marsh deer population. Status of marsh deer population is considered critical, as they vanished from Uruguay and became rare in Peru, Bolivia, Paraguay, and Argentina.

significance to humans

Livestock owners hunt for marsh deer to get rid of rival for livestock.

Chilean huemul

Hippocamelus bisulcus

taxonomy

Hippocamelus bisulcus (Molina, 1782), Chilean Andes.

other common names

English: Huemul, Chilean guemal; French: Cerf de andes méridionales, huémul des andes méridionales; Spanish: Ciervo andino meridional, huemul.

physical characteristics

Small. Shoulder height: 31–35 in (77–90 cm); body length: 55–65 in (140–165 cm); tail length: 4.5–5.5 in (11–13 cm); weight: 100–145 lb (45–65 kg). Coat color of both sexes year-long is monotonous, yellowish grizzly brown with whitish belly, a black Y-shaped stripe on muzzle, and a brown spot on rump. Bucks grow small two-pointed antlers. Both bucks and does have elongated canines, covered by a lip.

distribution

The Andes of central and southern Chile and Argentina.

habitat

Inhabit thick woods and bushes at altitudes 4,300–5,600 ft (1,300–1,700 m), on steep slopes, in rugged relief.

behavior

Linked to constant home ranges to 90–200 ac (36–82 ha). Buck and adult does live together, sometimes in a small group of eight, comprising a buck and does. In rut, buck marks range by butting bushes, thus dispersing secretion of head scent glands. A buck defends does from rivals.

feeding ecology and diet

Grass eater, feeding on cereals and sedges.

reproductive biology

Polygynous. Rut in winter (July–August). Gestation is eight months. Fawns appear at the end of rainy season February–April. Fawns stay hidden.

conservation status

Now considered Endangered. Recently, deer inhabited high altitude plateaus in Andes. Hunting pressure, competition with cattle, and pursuit by wild dogs decreased population numbers to 1,300 animals living in some localities in southern Chile and Argentina.

significance to humans

Game species in the past, now preserved as rare species.

Southern pudu

Pudu pudu

taxonomy

Pudu pudu (Molina, 1782), Chile.

other common names

English: Chilean pudu; French: Pudou du sud; German: Pudu; Spanish: Ciervo enano, venadito chileno.

physical characteristics

Smallest among Cervidae, with short legs and rounded body. Shoulder height: 14–18 in (35–45 cm); body length: 2.8 ft (85 cm); tail length: 3 in (8 cm); weight: 20–33 lb (9–15 kg). Thick bright coat is reddish brown, while lips and insides of ears are orangey. Fawns develop white spots. Males wear short spiked antlers, 2.8–4 in (7–10 cm) long.

distribution

Southern Chile.

habitat

Rainforests, bamboo groves, in mountains to the snow limit. Choose thickets to defend against human pursuit, though also prey of wild cats and foxes.

behavior

Live solitarily or in pairs, rarely in small groups (to three). Home grounds to 40–65 ac (16–26 ha) are well arranged with trails connecting feeding grounds and resting points. Dung piles often mark trails. Scent marks made by secretion of preorbital and frontal glands as well as urine spots on trails and tree branches play important role in pudu communications. Crepuscular and nocturnal, they are very cautious, regularly stop feeding to listen and sniff around. When in danger, they bark and flee in zigzag pattern through inaccessible thickets and steep rocks.

feeding ecology and diet

Feed on leaves, twigs, bark, buds, fruits, seeds, and rarely on herbs, To reach food, can stand on hind legs or scramble along fallen tree trunks.

reproductive biology

Polygynous. Rut starts in autumn; fawning takes place the next spring (or in November–January in Southern Hemisphere). Gestation period is 210 days, usually it is one young each birth. Weaning occurs after two months. Females are sexually matured and can participate in reproduction during first year, males on their second autumn. Life expectancy is 8–10 years, to 15 years in zoos.

conservation status

Endangered.

significance to humans

Game species.

Red brocket

Mazama americana

taxonomy

Mazama americana (Erxleben, 1777), Cayenne, French Guiana.

other common names

French: Daguet rouge; German: Grossen Roten Spiesshirsch; Spanish: Corzuela roja.

physical characteristics

Small. Shoulder height: 14–30 in (35–75 cm); head and body length: 28–53 in (72–135 cm); tail length 3–6 in (8–15 cm); weight: 44 lb (20 kg). The hair on muzzle radiates in all directions from two whorls. Coat color is monotonous, light to dark brown; lighter in under parts and belly, tail end is white. The body is stout, limbs are slender, and the back is arched. Antlers are simple, spiked.

distribution

Mexico, Brazil, Argentina, Bolivia, and Paraguay.

habitat

Live in forests from sea level to altitudes of upper forest limit.

behavior

Live solitary, sedentary life; active day and night. Wary, hide when disturbed, often remain unnoticed due to camouflage colors. Easy prey for predators.

feeding ecology and diet

Grasses, vines, and tender green shoots. Important part of diet are fruits, cereals, and fungi.

reproductive biology

Polygynous. Bucks and does together only during rut, which is year long, though most of mating from July–September in main rainy season. Gestation period is 225 days, calving occurs in December–January, in short rainy period; usually one young per birth. Does participate in breeding from age one year. Life expectancy in captivity to 14 years.

conservation status

Data Deficient.

significance to humans

Experience severe hunting pressure and pursuit from farmers defending bean fields and corn crops.

Moose

Alces alces

taxonomy

Alces alces (Linnaeus, 1758), Sweden.

other common names

English: American moose; French: Elan; German: Elch; Spanish: Alce.

physical characteristics

Largest of contemporary Cervidae. European moose: bull body length 87–110 in (220–280 cm), cow body length 87–106 in (220–270 cm); weight: bull 620–930 lb (280–420 kg), cows 600–770 lb (270–350 kg). North American moose: bull body length 140 in (350 cm), cow 125 in (315 cm); weight: bull 880–1,400 lb (400–630 kg), cows 1,200 lb (550 kg). Short body with hump-like withers, sloping rump, and very short tail, all mounted on long legs to 31 in (80 cm). Head is huge, long, and narrow, with a square upper lip hangs over the lower one. Muzzle is hairy, with a small bald spot between nostrils. The ears are large, oval, and vividly express all moods from fear to aggression. Short and thick neck furnished with mane, a skinny pendant (the bell) hanging from the throat; in North American moose, bell reaches 14 in (35 cm). Hooves are long and narrow. Dewclaws are also long, functional fingers surrounded by a strong stretchable membrane that reaches ground surface while walking. There is also a stretchable web between hoof fingers. Usually moose antlers are wide, palmate, though there are individuals with deer-like antlers in the same habitats. Coat color in adults is dark-brown with lighter legs. Moose have no rump patch. Calves are reddish brown, with no spots. Teeth are adapted to soft vegetable forage. Incisors are straight, chisel-like, good at nipping off wood bark. Moose cannot feed on harsh steppe grasses.

distribution

In North America, moose area extends from Alaskan tundra to Minnesota. Range of moose distribution in Eurasia spans from Scandinavia and eastern Poland to Pacific Ocean.

habitat

Inhabits nearly total forest zone of the northern hemisphere, and penetrates along forested or bushy ravines or river valleys far to the north to tundra, or to the south to steppes. Altitude of habitats varies from seashore plains to mountain forest limits. Moose adapted to climate with drastically changing temperatures. Snow cover deeper than 28–31 in (70–80 cm) impedes traveling, though they survives in areas with snow cover deeper than 7 ft (200 cm). Inhabit vast marshlands of western Siberia where they cross the swampiest patches crawling on their belly with forelegs stretched out in front.

In winter, they prefer mature coniferous and mixed deciduous-coniferous forests, young coniferous forests, forested banks of rivulets and lakes, burnouts, and cut clearings. In northern areas of America and Eurasia, moose come to shrub tundra and reach shores of the Arctic Ocean migrating along river valleys.

behavior

Live solitarily or in small groups: mother, calf (calves), sometimes yearling(s). Two females with calves happen to make a congregation; sometimes a bull joins them. In areas rich in forages, there are sometimes large moose congregations, to 270 animals in Altai Mountains. No social bonds exist in those gatherings and, when disturbed, moose flee independently, without following any leader. Sedentary though summer and winter; home grounds might be separated. Migrate 18–24 mi (30–40 km) searching for more suitable habitat as seasons change. In winter, the deeper snow, the less desirable a home ground.

Active mostly in mornings and evenings, and switch to nocturnal life in summer, as insect harassment increases. Moose usually walk or trot (to 9.3 mph [15 km/h]) and can rush to gallop (18.6 mph [30 km/h]) for a short distance; they are good swimmers and divers.

feeding ecology and diet

Winter diet is wood and shrub bark and branches, while in summer they feed on leaves of trees and shrubs, on aquatic plants, forest under story grasses, and various herbs. It strongly prefers willow, poplar, aspen, mountain ash, blueberry, bird cherry tree, and buckthorn. An adult in winter consumes daily 22–30 lb (10–13 kg) of forage, and more than 66 lb (30 kg) during summer and spring. Aquatic plants make an important part of diet to supply animals with necessary nutritive components. Moose easily digest many toxic plants, the latter comprising up to 30% of diet. The highly unusual structure of the moose nose is apparently a specialization for feeding on aquatic vegetation. It evolved late in the Pleistocene in Eurasia.

reproductive biology

Polygynous. Beginning of rut in August is signed by broken bushes, striped bark, as bulls are getting rid of antler velvet. Bulls produce typical sounds similar with groans. After one to two days together, they part and a bull starts searching for another female. The hairy skin flap under the jaw, the bell, has been identified as a scent distribution organ. It is splashed with urine when bulls dig rutting pits and serves to attract females, which are greatly attracted to bull moose scent. From end of August to mid October, during rut, females come to heat every 18–21 days. Gestation period varies 215–243 days, calving lasts from mid April to mid July. There are average 1.2-1.6 young per birth. Females from 3–7 years old often give birth to twins. Moose reach sexual maturity at one and a half years, bulls mate from an age of two and a half years.

conservation status

Not threatened.

significance to humans

One of the most important game animal of the northern hemisphere. Many people use moose harvest as main source of food and skin. In Russian army of the eighteenth century, all horsemen wore trousers made of moose skin. Currently, moose became subject of sport game for sustainable use. Annual harvest in North America is more than 50,000; in Sweden 164,000; in Russia 80,000.

Reindeer

Rangifer tarandus

taxonomy

Rangifer tarandus (Linnaeus, 1758), Swedish Lapland.

other common names

English: Caribou; French: Renne; German: Rentier, Wildren; Spanish: Reno.

physical characteristics

Medium-sized. Shoulder height: 33–59 in (85–150 cm); body length: males 70–84 in (180–214 cm), females 64–81 in (162–205 cm); tail length: 6–8 in (14–20 cm); weight: males 200–460 lb (92–210 kg), females 174–256 lb (79–116 kg). A dwarf subspecies (Rangifer tarandus platyrhynchus) of 31–37 in (80–95 cm) shoulder height inhabits Spitzbergen Islands. Reindeer have elongated body and short legs. Muzzle is covered by hair, and a thick mane along the lower part of neck adorns both sexes. Hooves are very large, wide, and flat, accompanied by wide and long dewclaws, with long coarse hairs between fingers.

Unlike all other deer, both males and females wear antlers. Antlers are forked, reaching 39 in (100 cm) width and 53 in (135 cm) length of beam in males. Coat color is nearly white to a shady light in winter and grayish brown in summer. Tail and rump patch are white. In winter, hairs are air-filled to supply thermal insulation. Construction of teeth is particularly adapted for grazing and to eat soft lichens; teeth are not fitted for browsing on woody vegetation.

distribution

Wide areas in Eurasia and North America from the Arctic Ocean and arctic islands southward to 50°N in Scandinavia, Finland, European part of Russia, and to 58°N in western Siberia.

habitat

Inhabit arctic deserts such as on Arctic Ocean islands, in lichen or lichen-mossy tundra, covered by dwarf bushes of birch, willow, alder; in high mountains, tundra with dwarf vegetation, in forest-tundra where clearings with dwarf vegetation inter-change with larch clumps. Thin coniferous forests (pine, larch) with abundant woody lichens are also common habitat as well as vast marshlands (in western Siberia) and swamps amidst forests. Reindeer are perfectly adapted to life on seaside plains as well as in coniferous forests on hillsides and in open, woodless mountain plateaus to 8,850 ft (2,700 m).

behavior

Reindeer live in families (female with calf), herds, and gatherings. Most typical are herds of 2,500–3,000 individuals coordinated in movements and following a single leader when disturbed. Gatherings during migrations reach 80,000–100,000 animals and more, consisting of several herds; each herd coordinated in movements. Forest herd is three to eight (maximum 55) animals, as visual contact between more animals is hard in a forest.

Following the leaders is a typical behavioral reaction of disturbed deer. Actually, experienced females and bulls are the first to leave a herd's protection to escape on their own.

Herd life increases competition for feeding in snow. Adult bulls shed antlers soon after rut, in the beginning of winter. Calves follow mothers during winter to feed in their crater.

Barren females and young bulls shed antlers in March, so only pregnant females wear antlers in springtime.

Reindeer make seasonal migrations. Each population uses particular calving grounds where females return year after year. Summer pastures are also constant for each population, though position of those pastures might change, depending on natural conditions of the year. Reindeer use constant migration routes, especially points of river crossings. They move to points of most favorable combinations of snow cover and abundance of food.

Reindeer run with high speed. Good swimmers, they can cross sea straits that are 75 mi (120 km) wide. In areas under high hunting pressure, deer run 1,640–1,980 ft (500–600 m), but in places of low human pressure, they will allow a human to approach within a distance of 32–39 ft (80–100 m).

feeding ecology and diet

In Eurasia tundra, winter food consists of 20–90% lichens. Daily diet of an animal is about 11 lb (5 kg) of lichens. Also, winter diet is comprised of dry plants, green shoots of cotton grass, sedges, horse-tail, and green mosses. Important source of protein in autumn are mushrooms. Feeding in winter is mostly on lichens, which consist of 50–80% carbohydrates. At the same time, animals experience crucial lack of vitamins, proteins, and fat. They consume young leaves of birch and willow, twigs, buds, and flowers, cotton grass, and sedges.

reproductive biology

Polygynous. Rut time coincides with autumn migrations in September–October. Fights between bulls are frequent. Winners, the strongest animals control 7–8 females. Rutting bulls eat rarely, and soon are exhausted, changing the hierarchy. Bulls younger than one and a half years do not take part in rut due to rivalry of elder ones. Most large females take part in rut; among those, more than 80% get pregnant. Gestation period is 192–246 days, mother gives birth to one calf. A newborn stands in one hour and first suckles in five hours. After 5–7 hours, mother can leave the birthing ground, followed by calf. Strong link between mother and calf lasts three months. Lifespan of bulls is 4–5 years, maximum to 14 years; of females, 6–7 years up to 19 years.

More than 40% of calves die in the first year and about 30% of the rest die at the second year. Wolves are the main predators, then brown bear, raven, golden eagle and sea eagle. Also, calves die during spring migration while crossing big rivers or from cold.

conservation status

Not threatened.

significance to humans

Reindeer is the principal source of survival for indigenous people of the north. There are more than three million wild reindeer in North America and Eurasia and, also, about 2.5 million of tame reindeer. Reindeer meat is of specific quality. Deer pelts provide clothes necessary to survive in harsh northern climate. Velvet antlers are used in Asian medicines.

Resources

Books

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

Bubenik, G. A., and Anthony B. Bubenik, eds. Horns, Pronghorns, and Antlers. Evolution, Morphology, Physiology, and Social Significance. Heidelberg: Springer-Verlag, 1990.

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

Geist, Valerius. Deer of the World. Mechanicsburg, PA: Stackpole Books, 1998.

Geist, Valerius, and Fritz Walther, eds. The Behaviour of Ungulates and Its Relation to Management. Gland: International Union Conservation Nature Publications, 1974.

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

Redform, K. H., and J. F. Eisenberg. Mammals of the Neotropics: The Southern Cone. Chicago and London: The University of Chicago Press, 1992.

Serret, A. El Huemul. Fantasma de la Patagonia. Buenos Aires: Zagier and Urruty Publishers, 2000.

Vislobokova, Inessa A. Fossilized Deer of Eurasia. Moscow: Nauka, 1990.

Whitehead, G. K. The Whitehead Encyclopedia of Deer. Stillwater, MN: Voyager Press, 1993.

Periodicals

Cowan, I. McT., and V. Geist. "Aggressive Behavior in Deer of the Genus Odocoileus." Journal of Mammalogy 42, no. 4(1961): 522.

Dusek, G. L. "Ecology of White-Tailed Deer in Upland Ponderosa Pine Habitat on Southern Montana." Prairie Naturalist 19, no. 1 (1987): 1.

Hershkovitz, P. "Neotropical Deer (Cervidae). Part 1. Pudus, Genus Pudu Gray." Fieldiana Zoology New Series 11 (1982): 1.

Pac, H. I., W. F. Kasworm, L. R. Irby, and R. J. Mackie. "Ecology of the Mule Deer, Odocoileus hemionus, along the Cast Front of the Rocky Mountains, Montana." Canadian Field-Naturalist 102, no. 2 (1988): 227.

Povilitis, A. "Characteristics and Conservation of a Fragmented Population of Huemul Hippocamelus bisulcus in Central Chile." Biological Conservation 86 (1998): 97.

Sher, Andrey V. "History and Evolution of Moose in USSR." Swedish Wildlife Research "Viltrevy" 1, no. 1 (1987): 71.

Leonid Baskin, PhD