Alces alces (Linnaeus, 1758) Gray, 1821

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 : 431

34.

Moose

Alces alces

French: Elan / German: Eich / Spanish: Alce

Other common names: Elk (in Europe); Caucasian Moose (caucasica)

Taxonomy. Cervus alces Linnaeus, 1758 ,

Sweden.

Early Alceini appeared in the Middle Miocene of central Siberia, about three million years ago. Modern Moose first occurred early in the Late Pleistocene, about 100,000 years ago, in Eurasia, and entered North America only about 10,000-14,000 years ago. The analysis of mtDNA revealed three haplogroups, one entirely Asian, one primarily European, and one North American. North-eastern Siberia can be identified as the origin of all the extant lineages. East Siberian and Alaskan Moose have similar body size and antler design but are not closely related. The race caucasica (Vereshchagin, 1955) from the Caucasusis extinct. Eight extant subspecies are recognized.

Subspecies and Distribution.

A.a.alcesLinnaeus,1758—fromScandinavia,Finland,BalticstatesandPolandEtotheYeniseyRiver.

A.a.americanaClinton,1822—ECanada(COntariotoNewfoundland).

A.a.andersoniPeterson,1950—fromBritishColumbiatoMinnesotaandOntario.

A.a.buturliniChernyavsky&Zhelesnov,1982—NESiberiaandKamchatka.

A.a.cameloidesMilne-Edwards,1867—NMongolia,Ussuriland,NManchuria.

A.a.gigasMiller,1899—AlaskaandYukon.

A.a.pfizenmayeriZukowsky,1910—fromCSiberiatoStanovoyMtsandCherskiyMts.

A. a. shirasi Nelson, 1914 — from S Alberta to Wyoming and Utah.

Descriptive notes. Head—body 250-300 cm for males (bulls) and 240-290 cm for females (cows), tail 12-16 cm, shoulder height 190-210 cm (bulls) and 185-200 cm (cows); weight of adult bulls 300-600 kg and adult cows 280-460 kg. The heaviest animals occur in eastern Siberia and Alaska (with records of more than 770 kg for bulls and 600 kg for cows), the lightest in the Amur region and Manchuria (with some bulls weighing just 260 kg). Bulls are on average 25-30% heavier than cows. The largest extant deer, with massive trunk, short neck, shoulder hump, long and pendulous nose, very small bare rhinarium, long dewlap (“bell”) from the throat; large head and elongated snout, long legs. The coat is prevailingly dark brown with grayish legs. The tail is short and of the same color as the rump. Molts in May and September. Newborn calves have a reddish-brown unspotted coat. Suborbital, tarsal, and interdigital glands are present. Permanent dentition of 32 teeth. Permanent teeth erupt at 6-19 months of age. Antlers of adults are typically palmated, but a simpler design without any flattening is the rule in Manchuria and common in Europe. Pedicles appear at 4-5 months after birth; their apices, generally simple, minute buttons, ossify at 6-7 months and cast at 10-11 months. The second antler set, simple spikes or forked beams (in Alaska sometimes already with traces of palmation) develop at about 13-14 months and are cast the following February—March. At 7-11 years antlers attain their full size. In good quality populations of Europe, antlers of adult bulls have a span of 105 cm on average and a brutto (skull plus antlers) weight of 10 kg (with records of 145 cm and 18.5-20 kg); in Alaska mean antler span is 145 cm and antler mass is about 20 kg, with records of 205 cm and 30-35. 8 kg. In adult bulls velvet shedding occurs in September, antler casting in December, and the beginning of the new antler growth in April. Hooves are 13-18 cm long and 11-14 cm broad in adult bulls, with the pads extending right outto the tip of the hoof.

Habitat. Boreal forests, mixed forests of the northern temperate zone, especially with damp, marshy areas. In Alaska also tundra. Up to 1700 m above sea level on the Altai Mountains. Snow depth more than 70 cm and mean temperature in summer higher than 14°C are limiting factors. Males tend to select habitats with greater food abundance. Females select areas with better concealment cover.

Food and Feeding. As a browser, it feeds on the vegetative parts of trees, shrubs, dwarf shrubs, herbs, and aquatic plants, preferring birch, alder, and willow. Adult daily consumption is about 10-30 kg of fresh vegetation.

Breeding. In good quality habitats most females attain puberty at 16-17 months of age. Males reach physiological sexual maturity at the same age, but in naturally structured populations yearling and subadults are excluded from breeding by dominant bulls and begin to reproduce only at five years of age; in heavily hunted populations with a low proportion of adults, subadults have a good chance of mating. Rutting season is in September—October. Typically an adult male locates and defends an estrous female until he mates with her, and then he moves on in search of other females (“tending bond mating system”). In tundra the dominant bull tends to herd and defend a group of females (“harem mating system”). Bulls thrash antlers on vegetation, rub trees, dig rutting pits, and urinate and wallow,salivating copiously. Females are polyestrous, with cycles on average of 24 days. During estrus they are receptive for 15-26 hours. Estrous females rub trees and visit rutting pits dug by males and wallow in them. Bulls lose 9-18% of their weight during the rut. Gestation lasts on average 234 days, with the birth season in May-June. Cows give birth to one or two calves, weighing on average 10-12 kg in Europe and 14-18 kg in Alaska; their mass doubles in about two weeks. Mothers remain within 50 m of their offspring for 5-7 days. Calves are nursed 7-9 times during the first days and four times at about 50 days of age. At 2-3 weeks calves begin to ingest solid food and at one month, half of their diet consists mostly of plants. Lactation lasts about 4-5 months. Final weight is attained at about eight years by bulls and four years by cows. Senescence occurs from 12-13 years onward. Normally they live to a maximum of 16-19 years, with records of 21 years for bulls and 25 for cows. In captivity the maximum longevity recorded is 27 years of age. In populations subjected only to natural mortality, average life expectancy in protected areas can be about seven years. In populations subjected to hunting, average life expectancy can decline to about three years. Gray Wolves (Canis lupus) and Brown Bears (Ursus arctos) are the main predators.

Activity patterns. It is active though the day, with peaks at dawn and dusk and 5-8 feeding bouts for a total of 9-13 hours in summer and 6-8 hours in winter.

Movements, Home range and Social organization. Moose have a specialized trotting gait, by which they glide over obstacles with little energetic cost. They can reach a speed of 60 km /h. Sedentary Moose have home ranges of 2-90 km®. A segment of the population can be migratory, moving 10-30 km on average, normally to reach areas with shallower snow and better food availability in winter. Migrations can sometimes be much longer, up to 200 km in North America, up to 300-500 km in Siberia. Natal dispersal can be up to 180-250 km. The Moose is not particularly gregarious. The basic social unitis a female with her young offspring. Bulls are generally solitary and tend to spatially segregate from cows for most of the year. In open terrain in Alaska predation risk favors some grouping behavior.

Status and Conservation. Classified as Least Concern on The [UCN Red List. In the early Holocene the European Moose extended its range to Great Britain, the Pyrenees, and the Alps. In the early Middle Ages it disappeared from the last relict areas of western Central Europe,its range restricted to Fennoscandia and Eastern Europe. At the beginning of the 19" century Moose suffered a drastic decline due to overhunting and disappeared from the Caucasus. In the last 50 years European Moose populations have increased dramatically. In Europe there are at least 440,000 individuals, possibly more than half a million, of which about 214,000 are harvested every year.Vagrant individuals are currently recorded in Central and South-eastern Europe, from Germany to Croatia and Romania. A few very small populations are now established in South Czech Republic and North Austria. In North America the original population was estimated at about one million; overexploitation caused a decline to around 500,000 in 1948, but in 1960 the number had grown to more than 900,000 and is now again about one million. Annual harvest in late 1990s was about 85,000 individuals.

Bibliography. Apollonio et al. 2010, van Ballenberghe (1982), Baskin & Danell (2003), Bowyer, Stewart et al. (2001), Bowyer, van Ballengerghe & Kie (2003), Cederlund et al. (1988), Crete & Daigle (1999), Ericsson et al. (2001), Franzmann et al. (2008), Garel et al. (2006), Gasaway et al. (1987), Hundermark & Bowyer (2004), Nygrén (1986), Nygrén et al. (2007), Schmolcke & Zachos (2005).

Figures

34. Moose (Alces alces), 35. Caribou (Rangifer tarandus)

Subspecies and Distribution. A.a.alcesLinnaeus,1758—fromScandinavia,Finland,BalticstatesandPolandEtotheYeniseyRiver. A.a.americanaClinton,1822—ECanada(COntariotoNewfoundland). A.a.andersoniPeterson,1950—fromBritishColumbiatoMinnesotaandOntario. A.a.buturliniChernyavsky&Zhelesnov,1982—NESiberiaandKamchatka. A.a.cameloidesMilne-Edwards,1867—NMongolia,Ussuriland,NManchuria. A.a.gigasMiller,1899—AlaskaandYukon. A.a.pfizenmayeriZukowsky,1910—fromCSiberiatoStanovoyMtsandCherskiyMts. A. a. shirasi Nelson, 1914 — from S Alberta to Wyoming and Utah.