Odocoileus hemionusmule deer

Ge­o­graphic Range

Odocoileus hemionus oc­curs over most of North Amer­ica west of the 100th merid­ian from 23 de­grees to 60 de­grees N. The east­ern edge of the usual range ex­tends from south­west­ern Saskatchewan through cen­tral North and South Dakota, Ne­braska, Kansas, and west­ern Texas. Iso­lated oc­cur­rences have been re­ported from Min­nesota, Iowa, and Mis­souri. Major gaps in ge­o­graphic dis­tri­b­u­tion are in south­ern Nevada, south­east­ern Cal­i­for­nia, south­west­ern Ari­zona, and the Great Salt Lake desert re­gion. Apart from these gaps, O. hemionus oc­curs in all of the bio­mes of west­ern North Amer­ica north of cen­tral Mex­ico, ex­cept the Arc­tic tun­dra (An­der­son 1984).

Habi­tat

Odocoileus hemionus is re­mark­ably adapt­able. Of at least sixty types of nat­ural veg­e­ta­tion west of the 100th merid­ian in the United States, all but two or three are or once were oc­cu­pied by O. hemionus. Sev­eral ad­di­tional veg­e­ta­tion types are in­hab­ited in Canada and Mex­ico as well. The veg­e­ta­tion types in Mex­ico are sim­i­lar to the types oc­cur­ring in the United States. How­ever, the trop­i­cal de­cid­u­ous veg­e­ta­tion at the tip of Baja Cal­i­for­nia is unique. In Canada, O. hemionus oc­cu­pies five bo­real for­est types that do not occur in the United States. O. hemionus oc­cu­pies a wide range of habi­tat provinces (re­gions of land con­tain­ing par­tic­u­lar veg­e­ta­tion types) in west­ern North Amer­ica. These habi­tat provinces in­clude the Cal­i­for­nia wood­land chap­ar­ral, the Mo­jave Sono­ran desert, the In­te­rior semi­desert shrub wood­land, the Great Plains, the Col­orado Plateau shrub­land and for­est, the Great Basin, the Sage­brush steepe, the North­ern moun­tain, and the Cana­dian bo­real for­est (Wallmo 1981).

Phys­i­cal De­scrip­tion

The pelage of Odocoileus hemionus ranges from dark brown gray, dark and light ash-gray to brown and even red­dish. The rump patch may be white or yel­low, while the throat patch is white (Geist 1981). The white tails of most mule deer ter­mi­nate in a tuft of black hairs, or less com­monly in a thin tuft of white hairs. On some mule deer, a dark dor­sal line runs from the back, down the top of the tail, to the black tail tip. All mark­ings vary con­sid­er­ably among O. hemionus, but re­main con­stant through­out the life of an in­di­vid­ual. O. hemionus pos­sess a dark V-shaped mark, ex­tend­ing from a point be­tween the eyes up­ward and lat­er­ally. This mark is more con­spic­u­ous in males. Growth in O. hemionus dur­ing the first year is roughly par­al­lel in males and fe­males. There­after, males, in gen­eral, ex­ceed fe­males in car­cass weight, chest girth, neck cir­cum­fer­ence, body length, head length, cra­nial breadth, shoul­der height, hind­foot length, and hoof length (An­der­son 1984). Car­cass weight ranges from 45 to 150 kg in males, and 43 to 75 kg in fe­males. Chest girth ranges from 80 to 117 cm in males, and 78 to 97 cm in fe­males. Neck cir­cum­fer­ence ranges from 30 to 65 cm in males, and 26 to 38 cm in fe­males. Body length ranges from 126 to 168 cm in males, and 125 to 156 cm in fe­males. Head length ranges from 28 to 35 cm in males, and 27 to 33 cm in fe­males. Cra­nial breadth ranges from 11 to 16 cm in males, and 10 to 14 cm in fe­males. Shoul­der height ranges from 84 to 106 cm in males, and 80 to 100 cm in fe­males (Wallmo 1981).

  • Range mass
    43 to 150 kg
    94.71 to 330.40 lb

Re­pro­duc­tion

Odocoileus hemionus is a polyg­y­nous species, hav­ing a tend­ing-bond type breed­ing sys­tem. Courtship and mat­ing occur within the group (Geist 1981). A dom­i­nant male tends an es­trus fe­male until mat­ing or dis­place­ment by an­other male oc­curs. Dom­i­nance is largely a func­tion of size, with the largest males, which pos­sess the largest antlers, per­form­ing most of the cop­u­la­tions (Kucera 1978). Most O. hemionus fe­males con­ceive dur­ing their sec­ond year and only rarely dur­ing their first year. The breed­ing peak in O. hemionus oc­curs mainly from late No­vem­ber through mid-De­cem­ber. The av­er­age ges­ta­tion length is 204 days. The peak birth pe­riod in O. hemionus is es­ti­mated to be from June 16th to July 6th, with most births oc­cur­ring in June. The time of birth varies ac­cord­ing to the en­vi­ron­ment. Robi­nette (1977) cal­cu­lated that a 305-m rise in el­e­va­tion is as­so­ci­ated with a 7-day delay in the birth pe­riod. The mass at birth of O. hemionus ranges from 2 to 5 kg. Mass at birth is af­fected by lit­ter size and sex, with males being heav­ier. The com­mon liter size is two, with moth­ers in their first or sec­ond breed­ing year most fre­quently pro­duc­ing sin­gle­tons. Wean­ing be­gins at about 5 weeks of age and usu­ally is com­pleted at age 16 weeks. Full de­vel­op­ment of most skele­tal at­trib­utes oc­curs at about 49 months of age in males and 37 months of age in fe­males. How­ever, gains in car­cass mass are con­tin­u­ous until an age of 120 months in males and 96 months in fe­males. In O. hemionus, male neonates pre­dom­i­nate when poor nu­tri­tion pre­vails about 6 weeks be­fore, and dur­ing, the breed­ing pe­riod. Ovu­la­tion in fe­male O. hemionus oc­curs about 12 to 14 hours after es­trus ter­mi­nates. Ap­prox­i­mately 27 to 29 days elapse be­tween con­cep­tion and im­plan­ta­tion in fe­male O. hemionus. Among male O. hemionus, tes­tic­u­lar mass and vol­ume are max­i­mal dur­ing No­vem­ber and min­i­mal dur­ing April and May (An­der­son 1984).

  • Key Reproductive Features
  • gonochoric/gonochoristic/dioecious (sexes separate)
  • sexual
  • Average number of offspring
    1.5
    AnAge
  • Average gestation period
    207 days
    AnAge
  • Average age at sexual or reproductive maturity (female)
    Sex: female
    478 days
    AnAge
  • Average age at sexual or reproductive maturity (male)
    Sex: male
    503 days
    AnAge

Lifes­pan/Longevity

  • Average lifespan
    Status: captivity
    22 years
    AnAge

Be­hav­ior

In­di­vid­u­als of Odocoileus hemionus tend to con­fine their daily move­ments to dis­crete home ranges. Most mule deer with es­tab­lished home ranges use the same win­ter and sum­mer home ranges in con­sec­u­tive years. Dis­per­sal in O. hemionus in­volves move­ments be­yond the home range to dis­tances of up to 8 km. This move­ment re­sults in the es­tab­lish­ment of a new home range. Sea­sonal move­ments in­volv­ing mi­gra­tions from higher el­e­va­tions (sum­mer ranges) to lower win­ter ranges are as­so­ci­ated, in part, with de­creas­ing tem­per­a­tures, se­vere snow­storms, and snow depths that re­duce mo­bil­ity and food sup­ply. Deep snows ul­ti­mately limit use­able range to a frac­tion of the total. Mule deer in the arid south­west may mi­grate in re­sponse to rain­fall pat­terns. Com­mon preda­tors of O. hemionus in­clude pumas, coy­otes, bob­cats, golden ea­gles, feral dogs, and black bears. The so­cial sys­tem of O. hemionus con­sists of clans of fe­males re­lated by ma­ter­nal de­scent. These clans are the fac­ul­ta­tive re­source de­fend­ers. Males dis­perse as in­di­vid­u­als or ag­gre­gate in groups of un­re­lated in­di­vid­u­als. Dur­ing win­ter and spring, the sta­bil­ity of fe­male clans and male groups is main­tained with dom­i­nance hi­er­ar­chies. In­creases in strife and alarm be­hav­ior, and de­creases in play among fawns, occur as pop­u­la­tion den­sity in­creases. The fre­quency of ag­gres­sive be­hav­ior be­tween the sexes re­mains low year round in O. hemionus. Com­mu­ni­ca­tion among O. hemionus is fa­cil­i­tated by the se­ba­ceous and su­dorif­er­ous se­cre­tory cells of five in­tegu­men­tary glands. The cells of each gland pro­duce spe­cific scents (pheromones) that elicit spe­cific re­ac­tions in con­specifics. The metatarsal gland pro­duces an alarm pheromone, the tarsal gland aids in mu­tual recog­ni­tion, the in­ter­dig­i­tal gland leaves a scent trail, and the func­tion of the tail gland is un­known. Urine has a pheromone func­tion at all ages and for both sexes. It is de­posited on tufts of hair sur­round­ing the tarsal glands. In fawns, it func­tions as a dis­tress sig­nal, while in adults, it func­tions as a threat sig­nal (An­der­son 1984). O. hemionus has sev­eral dis­tinct strate­gies for avoid­ing preda­tors. O. hemionus spe­cial­izes in de­tect­ing dan­ger at a very long range by means of large ears and ex­cel­lent vi­sion. Males can quickly de­tect and vi­su­ally track an­other an­i­mal as far as 600 me­ters. Once dan­ger is de­tected, O. hemionus may choose to hide, or move into cover and cau­tiously out­ma­neu­ver the preda­tor. An­other strat­egy is to de­part while the preda­tor is still a long way off and move sev­eral miles to an­other area. O. hemionus, in­stead, may bound rapidly up­hill, im­pos­ing on pur­su­ing preda­tors an un­ac­cept­ably high cost per unit time of lo­co­mo­tion. In yet an­other strat­egy, O. hemionus may bound off and then trot away, stop­ping fre­quently to gain in­for­ma­tion on the dis­tur­bance. This ini­tial bound­ing, com­bined with re­lease of metatarsal scent that in­hibits feed­ing, is highly ad­van­ta­geous in that, by alarm­ing oth­ers, it causes other mule deer to bound off as well, re­duc­ing the con­spic­u­ous­ness of the deer who bounded off first. This strat­egy would also trig­ger group for­ma­tion. Fi­nally, when a preda­tor closes in, O. hemionus ini­ti­ates eva­sive ma­neu­vers based on sud­den un­pre­dictable changes in di­rec­tion and on plac­ing ob­sta­cles be­tween it­self and the preda­tor. This strat­egy, how­ever, does not work against group-hunt­ing preda­tors. O. hemionus is an ex­cel­lent swim­mer, but water is rarely used as a means of es­cap­ing preda­tors (Geist).

Com­mu­ni­ca­tion and Per­cep­tion

Food Habits

Odocoileus hemionus is a small ru­mi­nant with lim­ited abil­ity to di­gest highly fi­brous roughage (Short 1981). Op­ti­mum growth and pro­duc­tiv­ity of in­di­vid­u­als and pop­u­la­tions are de­pen­dent upon ad­e­quate sup­plies of highly di­gestible, suc­cu­lent for­age. Diets con­sist­ing pri­mar­ily of woody twigs can­not meet the main­te­nance re­quire­ments of O. hemionus. Based on its stom­ach struc­ture and its diet of woody and herba­ceous for­age in ap­prox­i­mate equal pro­por­tions, O. hemionus is clas­si­fied as an in­ter­me­di­ate feeder. Be­cause nu­tri­tious for­age is in poor sup­ply for much of the year, O hemionus has an an­nual cycle of meta­bolic rates. A higher en­ergy flux and food in­take in the sum­mer en­ables O. hemionus to cap­i­tal­ize on abun­dant high-qual­ity for­age for growth and fat stor­age. A lower en­ergy flux in the win­ter per­mits O. hemionus to sur­vive on a lower in­take of poor-qual­ity for­age while min­i­miz­ing the ca­tab­o­lism of stored fat for body func­tions. The es­ti­mated rate of food in­take is about 22 g/kg body weight/day. In adult males, food in­take drops abruptly with the onset of rut (An­der­son 1984). O. hemionus fre­quently browses leaves and twigs of trees and shrubs. Green leaves are very suc­cu­lent and, ex­cept for epi­der­mal tis­sue and struc­tural ribs, con­sist largely of eas­ily di­gestible cell con­tents. Dead and weath­ered leaves have lit­tle pro­tein and high cell-wall val­ues. As a re­sult, they are of very low di­gestibil­ity. O. hemionus also eats acorns, legume seeds, and fleshy fruits, in­clud­ing berries and dru­pes, that have mod­er­ate cell-wall lev­els and are eas­ily di­gested (Short 1981).

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Odocoileus hemionus is of tremen­dous in­ter­est to hunters. Pop­u­la­tions of O. hemionus that are large enough to sup­port hunt­ing dur­ing two or three weeks in au­tumn offer count­less recre­ational op­por­tu­ni­ties for the pub­lic. This de­sire to hunt gen­er­ates rev­enue for the econ­omy (Wallmo 1981).

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

Dou­glas fir and Pon­derosa pine are of major eco­nomic im­por­tance for com­mer­cial tim­ber. How­ever, these trees are browsed heav­ily by O. hemionus. Brows­ing of other trees is sel­dom con­sid­ered an eco­nomic prob­lem. In the Dou­glas fir re­gion, O. hemionus browses on trees dur­ing both the dor­mant and grow­ing sea­sons. Prac­tices that en­cour­age the growth of O. hemionus pop­u­la­tions can there­fore also en­cour­age dam­age. Dou­glas fir is har­vested mainly by clearcut­ting and is re­gen­er­ated by plant­ing with nurs­ery-grown stock. O. hemionus is at­tracted to clear-cuts, and Dou­glas fir is an ac­cept­able and some­times pre­ferred for­age species. This sit­u­a­tion in­vites brows­ing of suf­fi­cient in­ten­sity to in­flu­ence for­est re­gen­er­a­tion in many areas (Wallmo 1981).

Con­ser­va­tion Sta­tus

All fed­eral, state, and provin­cial land and wildlife man­age­ment agen­cies rec­og­nize the fun­da­men­tal need to main­tain O. hemionus ranges and keep them hab­it­able. To counter the trend of agri­cul­tural de­vel­op­ment, range­land con­ver­sion, min­ing, road and high­way con­struc­tion, and the de­vel­op­ment of hous­ing tracts, many states and provinces have pur­chased crit­i­cal areas, es­pe­cially win­ter ranges, to main­tain the var­i­ous habi­tats of O. hemionus. But, due to po­lit­i­cal op­po­si­tion to gov­ern­ment ac­qui­si­tion of pri­vately owned lands, plus a scarcity of funds for this pur­pose, only a small frac­tion of O. hemionus ranges has been ac­quired by the gov­ern­ment. The ef­fects of re­duced O. hemionus ranges can be mit­i­gated by bet­ter man­age­ment of the re­main­ing lands to max­i­mize their pro­duc­tiviy for O. hemionus. Var­i­ous habi­tat man­age­ment pro­grams in­clude the ma­nip­u­la­tion of live­stock graz­ing, the ma­nip­u­la­tion of cul­ti­va­tive com­mu­ni­ties, and the ma­nip­u­la­tion of veg­e­ta­tive com­mu­ni­ties. For O. hemionus, the op­ti­mal suc­ces­sional stages are sub­cli­max plant com­mu­ni­ties that can be per­pet­u­ated only through the in­flu­ence of hu­mans. Since O. hemionus pro­duc­tion is not the pri­mary man­age­ment goal on most pri­vate or pub­lic lands in west­ern North Amer­ica, O. hemionus habi­tat im­prove­ment pro­grams typ­i­cally in­volve a com­plex process of co­or­di­na­tion among bu­reau­cra­cies with mis­sions that are usu­ally not com­pat­i­ble (Wallmo 1981).

Other Com­ments

The an­nual cycle of antler growth in O. hemionus is ini­ti­ated and con­trolled by changes in day length act­ing on sev­eral cell types of the an­te­rior pi­tu­itary. These cell types se­crete growth-stim­u­lat­ing hor­mones that act mainly on the antlers and in­ci­den­tally on the testes. Antler hard­en­ing, shed­ding, and the breed­ing pe­riod are me­di­ated by de­creas­ing day length through the ac­tion of go­nadotropins on Ley­dig cells, thus pro­duc­ing an­dro­gens. An­dro­gens in­duce sec­ondary os­si­fi­ca­tion, ac­cel­er­ate mat­u­ra­tion, in­duce be­hav­ioral changes that re­sult in shed­ding antler vel­vet, and aid in the main­te­nance of os­teoblasts and os­teo­cytes to main­tain antlers in hard bone con­di­tion. With­drawal of an­dro­gens at the end of the breed­ing sea­son per­mits re­sorp­tion of bone at the pedi­cel-antler junc­tion and antler shed­ding. O. hemionus has ex­cel­lent binoc­u­lar vi­sion. While un­able to de­tect mo­tion­less ob­jects, O. hemionus is ex­tra­or­di­nar­ily sen­si­tive to mov­ing ob­jects. The sense of hear­ing is also ex­tremely acute. O. hemionus is a tar­get for var­i­ous viral, bac­te­r­ial, and par­a­sitic dis­eases. For ex­am­ple, heavy amounts of gas­troin­testi­nal ne­ma­todes may cause death in O. hemionus. This par­a­sitic dis­ease is usu­ally in­dica­tive of such pre­dis­pos­ing fac­tors as high mule deer den­sity and mal­nu­tri­tion. In­fec­tion by the par­a­sitic meningeal worm can cause fatal neu­ro­logic dis­ease in O. hemionus. Live­stock may trans­mit viral dis­eases to O. hemionus as seen in foot-and-mouth dis­ease. This in­fec­tion is char­ac­ter­ized by blis­ters in the mouth, above the hooves, and be­tween the dig­its (An­der­son 1984).

Con­trib­u­tors

Michael Mis­uraca (au­thor), Uni­ver­sity of Michi­gan-Ann Arbor.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chaparral

Found in coastal areas between 30 and 40 degrees latitude, in areas with a Mediterranean climate. Vegetation is dominated by stands of dense, spiny shrubs with tough (hard or waxy) evergreen leaves. May be maintained by periodic fire. In South America it includes the scrub ecotone between forest and paramo.

chemical

uses smells or other chemicals to communicate

desert or dunes

in deserts low (less than 30 cm per year) and unpredictable rainfall results in landscapes dominated by plants and animals adapted to aridity. Vegetation is typically sparse, though spectacular blooms may occur following rain. Deserts can be cold or warm and daily temperates typically fluctuate. In dune areas vegetation is also sparse and conditions are dry. This is because sand does not hold water well so little is available to plants. In dunes near seas and oceans this is compounded by the influence of salt in the air and soil. Salt limits the ability of plants to take up water through their roots.

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

motile

having the capacity to move from one place to another.

mountains

This terrestrial biome includes summits of high mountains, either without vegetation or covered by low, tundra-like vegetation.

native range

the area in which the animal is naturally found, the region in which it is endemic.

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

tactile

uses touch to communicate

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

Ref­er­ences

Geist, V. 1981. Be­hav­ior: adap­tive strate­gies in mule deer. Pp. 157-224, in Mule and Black-tailed deer of North Amer­ica (O. C. Wallmo, ed.). Univ. Ne­braska Press, Lin­coln, xvii + 605 pp.

Kucera, T. E. 1978. So­cial be­hav­ior and breed­ing sys­tem of the Desert mule deer. J. Mamm., 59:463-476.

Short, H.L. 1981. Nu­tri­tion and me­tab­o­lism. Pp. 99-127, in Mule and Black-tailed deer of North Amer­ica (O. C. Wallmo, ed.). Univ. Ne­braska Press, Lin­coln, xvii + 605 pp.

Wallmo, O. C. 1981. Mule and Black-tailed deer dis­tri­b­u­tion and habi­tats. Pp. 1-25, in Mule and Black-tailed deer of North Amer­ica (O. C. Wallmo, ed.). Univ. Ne­braska Press, Lin­coln, xvii + 605 pp.

An­der­son, A., O. Wallmo. 1984. Odocoileus hemionus. Mam­malian Species, 219: 1-9. Ac­cessed Oc­to­ber 30, 2006 at http://​www.​science.​smith.​edu/​departments/​Biology/​VHAYSSEN/​msi/​default.​html.