Even-toed, Cloven-Hooved Mammals  

       WILD BOARS              BISON                  ELK                DEER

The order Artiodactyla contains all the even-toed ungulates. These are the mammals that basically led to man’s transition from hunter to farmer, including goats, camels, pigs, oxen, cattle, buffalo and sheep.  Other mammals in this order include hippopotamuses, llamas, giraffes, deer, pronghorn, and antelopes.  These mammals first appeared about 54 million years ago.  The canines are usually reduced or lost (as in the white-tailed deer), and the upper incisors are greatly reduced or absent (as in the case of deer and elk).   

This order and the order perissodactyla (horses, zebras, rhinos, and tapirs) make up a group called ungulates. Unlike the even-toed artiodactylas, the perissodactyla are odd-toed.  Although this group has no taxonomic status, it refers to a broad group of herbivorous mammals that are more or less specialized for cursorial (running) locomotion.  This refinement of running abilities is a function of their evolution on the expanding grasslands of the Miocene Period.  Speed became the primary means of avoiding predation, and seasonal movements to seek water or appropriate food probably became an important part of the ungulate mode of life. Many ungulates have become large, in part to minimize predator pressure, but also due to the large digestive systems required to process the voluminous quantities of low-nutritious vegetative material necessary for its energy needs.  

The Old World seems to have been the center of evolution of the Artiodactyla order, whereas the perissodactyls developed mainly in North America.

Three suborders exist in the artiodactyls, based on the tooth and horn structure.  Suiformes (pigs and hippopotamuses) have low-crowned teeth with low, rounded cusps and relatively large canines, adapted for an omnivorous diet.  They do not have horns or antlers.  They form a unique group that has evolved with short legs and a squat body.  Some species within this suborder have two chambered stomachs (pigs and peccaries), while others have three chambered stomachs (hippopotamuses).  None of these stomachs are ruminating.

The next two suborders have evolved long legs, made for running, a long set of broad molars and a specialized stomach for handling a purely vegetarian diet. These are the ruminants.  Their stomach utilizes microorganisms that ferment hard-to-digest molecules, converting them into useable forms.  It is this partially decomposed food that is regurgitated as “cud” to be further ground by the ruminant’s molars before further processing.  They swallow the food rapidly, with little chewing, and then may retire to some secluded spot to digest it more thoroughly.  Tylopoda (llamas and camels) have a three chambered stomach, with teeth cusps elongated longitudinally into crescents.  Ruminantia (deer, elk, caribou, moose, giraffe, pronghorn antelope, bison, sheep, goat, and others) have a four chambered stomach, with teeth similar to the Tylopoda, but generally sport horns or antlers.  Other adaptations of these vegetarians are flexible necks and lips.  Finally, as a concession for running, all ruminants run on their third and fourth toes, sheathed in thick, hard keratin, with the lateral toes reduced. 

Feeding in sunny, often open exposed areas, they have developed excellent senses of sight and smell to detect predators.  They are often gregarious (safety in numbers) and give birth to precocial young.  The exception is the white-tailed deer (and, to a lesser extent, the moose) of the woodlands, who give birth to altricial young, who are hidden in the brush until able to travel with the mother.  

For additional information on ungulates, go to MAMMOLOGY.

Worldwide, the Artiodactyla order is represented by 221 species in 86 genera in ten families (Nowak's Walker's Mammals of the World).  

North America supports 26 species in 19 genera, representing five families (Jones et al. 1997 Checklist of North American Mammals).  This includes fourteen exotic species that are known to support reproducing populations in the wild.

In the Suidae family, the European wild hog (Sus scrofa) is known to exist sporadically throughout the Southern Appalachians.   This is the source of the Arkansas razorback and the domestic hog, or pig.  It is not a native; there are no native pigs in North America.  Instead, North America is home to three species of peccaries, restricted to the New World; found in southern Texas and to the south.  These feral pigs now occur from Texas to Florida and the Carolina, in West Virginia and New Hampshire, throughout California, on eight of the major Hawaiian islands, and on Puerto Rico and the Virgin Islands.  It is estimated that these free-ranging pigs number 500,000 - 2,000,000. 

Within this Appalachian region, the wild hog is most notably present in the Great Smoky Mountains National Park.  Any hiker in the Smokies has seen the extensive rooting that decimates herbaceous and other groundcover vegetation.  These feral hogs are descended from an original population of thirteen young boars released in April of 1912 in a game preserve on Hooper Bald.  About 1920, an estimated 100 boars escaped and began it’s spread, breeding freely and hybridizing with feral domestic pigs along their way.  Kellogg (1939) stated: “So far as known to Arthur Stupka, park naturalist, no wild boars have come into the Great Smoky Mountains National Park.  He believes that the Little Tennessee River, which separates the park from the Cherokee National Forest, may constitute a real barrier against the northward spread of this introduced species.”  Unfortunately, this was not the case.  It is believed that they entered the southwest quadrant of the park in the late 1940’s, in the vicinity of Calderwood.  Since then, the boars have moved eastward at a rate of about 1.65 miles per year.   Their presence on Gregory Bald was first noted in 1958, with an estimated population of 500 concentrated in the region between Cades Cove and Fontana Lake in 1959.  Trapping began that same year.  By 1980, the estimated population in the park was 1500, with a density of 265 boar per square mile in northern hardwood forests in the western half of the park.  At that time, a moratorium was placed on shooting boars in the North Carolina side of the Park.  Instead, local volunteers were organized to trap and remove as many hogs as possible.  Unexplainably, those captured boars were transported and released in national forest in North Carolina.  From 1959 through 1977 (18 years), 1,143 boars were “removed” from the park.  From 1977 though 1993 (16 years), a total of 6,316 animals were removed.  However, recent annual harvest in the Park has substantially declined.  With the same amount of labor spent, the number of boars caught has fallen from 1200 in 1986, to only 203 in 2000.  These catch per manhour expended ratios indicate that the population in the Park in 2001 is somewhere between 400 and 600.

The boars in the Smokies breed year-round with peaks in late fall/early winter and late spring/early summer, with litters averaging between 3 and 4 per litter (one litter per year).  Seasonal movements between higher and lower elevations occur (March/April and August/September). Studies have found 58% of their spring/summer diet consisting of spring-beauty corms (approximately 70% of their spring diet is subterranean in origin), an average of 1.75 salamanders per stomach (mainly red-cheeked), and hard mast comprising 60 to 85% of late summer diets.  

The North Carolina Wildlife Resources Commission has an excellent webpage on The Wild Boar in North Carolina(Go to the "species" link at the top row.  Then, "publications", and then you'll see it listed under the NC Wildlife & Wildlife Management Publications.)

The only member of this order found in the Appalachian study area, which has survived in the wild, is the white-tailed deer.  The deer were nearly extirpated in the 1920’s but have rebounded since that time, while being augmented by re-introductions in various locations.  

The last moose (Alces alces) were extirpated in Pennsylvania probably by the late 18th century.  However, the lack of historical records of moose in Pennsylvania suggests that the species was always poorly represented in this state, preferring the boreal spruce forests and aspen and willow thickets of New England and Canada.

Another ruminant, the elk, or wapiti, extirpated from the eastern US, has been re-established in the Appalachian study area in Pennsylvania and in the Great Smoky Mountains National Park (2001) in North Carolina and has been reintroduction in Tennessee, Kentucky, and Arkansas. (Actually, the native eastern elk subspecies has become extinct.  The introduced elk is a western subspecies.)  

Due to the controversy and extent of recent introduction efforts in many eastern states, the elk is included below as a separate listing.

In the Bovidae family, the American bison (Bos bison) has long been extirpated from the Appalachian study area.  Also known as the plains buffalo, this member of the Bovidae family (bison, muskoxen, antelope, goats, sheep, and cattle - differing from cervids by the presence of permanent unbranched horns on males and most females) was last seen east of the Mississippi in the early 1830’s.   Buffalo apparently were more abundant in Virginia than any other Atlantic state.  In fact, they were still abundant in the Charlottesville area at the time of Thomas Jefferson’s birth (1743).  Buffalo were particularly common in the Mount Rogers area of Virginia.  Even today, in the Elk Garden area of Mount Rogers, the remains of a once huge buffalo wallow can be found being used by cattle that now graze the land.

Buffalo, who can interbreed with domestic cattle, were themselves domesticated and bred in captivity in Virginia, with both bison-calves and mixed breeds commonly found among western settlements of Virginia in the late 1700’s.  (A little more about the domestication of buffalo can be read in the discussion of John James Audubon’s Quadrupeds of North America.)  Rockfish Gap (at the southern end of the Shenandoah National Park) was a major buffalo pass, leading across the Shenandoah Valley and through Great North Mountain.

In the Smokies and surrounding area, bison ranged throughout much of the mountains of North Carolina and Tennessee.  They traveled from North Carolina to the Great Valley of East Tennessee (the southern extension of the Shenandoah Valley) through the Cumberland and Great Smoky mountains along the Holston and French Broad Rivers.  Large southern populations were found in the Cumberland Valley bluegrass region of the Nashville, TN area. 

The last bison in North Carolina was recorded about 1760.  The last bison killed in Pennsylvania was in Union County, in 1801.  The last bison in Virginia was said to be killed by Nathan Boone, a son of Daniel Boone, in 1797 along the New River. The last bison killed in West Virginia was in 1815, 12 miles below Charleston, along the Great Kanawha River.  Stephenson says the last bison in West Virginia was killed near Valley Head, Randolph County in 1825.  The last bison in Kentucky are dated to 1800, and in Tennessee, between 1800 and 1810.  

In 1819, a male bison was sent to Paris to be a part of the world's first national zoo, under the Supervision of Frédéric Cuvier, younger brother of the museum's famous professor of comparative anatomy, George Cuvier.   After a disastrous first birth, where both the calf and cow died, Frédéric ultimately decided that bison were unsuitable for farming because "every time one turned one's back on the museum's bison, it charged."  

The bison of this region is the same plains bison of the western US (Bos bison bison).  It is believed to be a separate subspecies from the woods buffalo, Bos bison athabascae, generally limited to western Canada, north of approximately 55° N. latitude.  (The subspecies designation comes from the Athabasca River, which flows through northern Alberta; home of the 17,000 square-mile Wood Buffalo National Park.)  Recent mitochondrial DNA comparisons suggest that subspecific distinction may not be justified (i.e., they're both the same species with no subspecies existing).  The woods bison is generally larger, less shaggy, and has a more pronounced hump than the plains bison.

From an original pool of 60-70 million bison, 30 million bison existed in 1870, with only 1,000 existing by 1889.  There are probably about 250,000 bison in North America today.  Most are in captivity, but at least three herds are free roaming. One free ranging herd is called the Wild Bunch, after Butch Cassidy’s gang, located in the Henry Mountains of southern Utah.  A second herd is in Alaska, and the third herd is actually the wood buffalo in the above mentioned Wood Buffalo National Park.  (According to a 1994 National Geographic article, 4,000 existed in Yellowstone National Park.  At the Flying D Ranch near Bozeman, MT, Ted Turner had about 5,700, with another 2,100 at a ranch in New Mexico.) 

Additional information on the bison can be found at the US FWS website.


Family Cervidae -Cervids

Members of the Cervid family (deer, caribou, moose, and elk) are best characterized by the presence of antlers, found generally only on males (caribou females also have antlers).  Cervids are ruminants; possessing four stomach chambers, thus enabling them to chew their cud.  Upper incisors are lacking.  The canine teeth are absent in the white-tailed deer, and are poorly developed in the elk.  They tend to be gregarious and all are herbivores; some specializing as grazers (grass eaters) and others as browsers (eating leaves and twigs).  The woodland species of moose and deer are much less gregarious than the caribou and elk, adapted to the more open habitat.  Only one genus  (Roe deer) are known to have delayed implantation.

Antlers are appendages of the skull, composed of a solid bony core and supported on permanent skin-covered pedicels.  In our temperate zones the antlers begin to grow early in the summer, during which time they are well-supplied with blood. They are soft and tender and are covered with a thin skin, which bears short fine hairs and has the appearance of velvet.  By late summer the antlers have attained their maximum size.  The blood then gradually recedes, and the thin skin with the velvety hair dries, loosens, and is rubbed off. By the time the velvet is shed all circulation of blood has ceased; thus, when shedding takes place there is no bleeding and probably not even discomfort for the animal.  After the velvet is rubbed off, the antlers serve as sexual ornaments and weapons.  The antlers are shed each year from January to April, following the mating season, taking about two to three weeks.  In a single deer both antlers are usually shed within several hours or days of each other.  First year antlers are unbranched straight spikes.  Throughout the deer's life, the size of the antlers is a function of the animal's diet.  

It is generally accepted that North American cervids arrived on this continent from Asia at various times from the middle Miocene to the late Pleistocene, between 1 million and 18 million years ago. 

The moose, extirpated in Pennsylvania in the late 18th century (and only known from Highland County, VA glacial age deposits), is the largest cervid, weighing up to 1300 pounds. 

There are 41 species of deer worldwide, representing 17 genera (Nowak's Walker's Mammals of the World).   There are nine species representing six genera found in the wild in North America (Jones et al. 1997 Checklist of North American Mammals).  However, only 5 species in 4 genera are native to North America (elk, black-tailed and white-tailed deer, moose and caribou).  

Two cervids are found in the Appalachian study area; the re-introduced elk, and the white-tailed deer.   


WAPITI or ELK (Cervus elaphus)


Appalachian Region Distribution: Originally, throughout, although totally extirpated from eastern US by the late 1800’s.  Now re-established in several north-central counties in Pennsylvania and in the Great Smoky Mountains National Park (see Remarks below).
Continental Range: Throughout, with the exception of it’s southernmost reaches.
  This species includes what is known as the red deer of Europe and northeastern Asia.
Abundance:  Limited to areas of reintroductions in Pennsylvania and the Great Smoky Mountains National Park. (see Remarks below.) 
Population Density: In Michigan, densities vary up to 4 to 5 elk per square mile. 
Size and Molt: 5 feet at the shoulders; head and body length of 8 feet for the male, (7.5 feet for the female); Bulls weigh 580 – 1000 lbs; cows, 500 – 650 lbs. The wapiti is the second largest cervid in the world, second only to the moose.  They are heaviest in late summer.   Two annual molts. 
Mammae: Four mammae.       
Forested mountain edges and open meadows.   The wapiti shares its habitat with the white-tailed deer, although it is more of a wilderness animal.  
Active Period
Crepuscular, or nocturnal, in eating habits.   
Grazers (feeding on woody and herbaceous plants) in spring and summer, and browsers (feeding on woody stems) in winter.   Some mushrooms will be eaten.  Being a ruminant (thus, having a four-chambered stomach), wapiti regurgitate their food and chew it as cud before being reingested.         
Home Range: In one Scottish study, females had home ranges of about 150 acres in summer and 100 acres in winter, whereas males had corresponding ranges of 100 acres and 62 acres.  Eastern elk apparently do not migrate, while western populations often do, depending on habitat.  In such cases, migration is elevational, with much smaller winter ranges (thus the reason for smaller cougar winter ranges).  About a fourth of the Jackson Hole herd remains in the same area and even most of the northern Yellowstone mountainous herds remains in the same high elevation range.
Social Structure:
A highly gregarious species, known to maintain herds up to 200 in open western habitat, with smaller groups in wooded areas.  Within each herd, a rank hierarchy is maintained.  During winter, males and females are in separate herds in their own discrete areas, the females and immature calves in large herds, the males in smaller herds. As spring approaches, the large winter herds break up, sorting into smaller bands, which follow the snowline up to higher elevations.  Following parturition in early June (in PA), the mother and newborn live alone for several weeks.  By early July, the cows and their young, along with immature calves of both sexes, begin to congregate.  By mid-July, herds of as many as 400 have formed and are led by a dominant female, the matriarch.  Bulls are living separately, or in small groups, numbering up to about six, on the outskirts of the female herds.  These separate herds break up during rutting (female herds restrict their ranges in September, while males leave their ranges and join the females in October) when males and females come together and the bulls compete to form harems of 15 to 20 cows (up to 60).  Around the third week of August, evidence of the rut begins, with bulls shedding the velvet from their fully-formed antlers.  By Labor Day, bulls begin to bugle, and by mid-September, full rut is in motion in Pennsylvania.  After mating is completed, by about mid-October, the sexes separate.  Often,  numerous herds of both sexes will concentrate in large aggregations of up to 1,000 animals in regions of limited winter range.    
Life Cycle
After a mid September or early October breeding, one calf (twinning is rare) is produced in late May or early June (in PA) after an eight and a half month gestation period (249 – 262 days). Like white-tailed deer, the altricial calf is born with a spotted coat and left alone in hiding for the first two or three weeks, only approaching five or six times a day for nursing.  Weaning is complete by late summer (4-7 months).  Bulls are polygynous; mating with as many females as possible.   However, in a dense forested habitat, the bull will normally search out females and stay with each until she is in estrus, while in a more open habitat, the bull more often will advertise his presence by bugling and attract females to him.  Rutting begins in autumn, with males practicing the ritual characteristic of bugling.  The bulls can round up as many as 60 cows (a Michigan study showed harems of one to 21).  Regardless of the harem, seldom does a male actually father more than four per season.  Bulls will not normally be able to hold a harem until at least four years of age.  It is believed that it is really the cow that picks the bull, and not vice versa.  Cows are seasonally polyestrous, the cycle being about 18 days. It is believed that there may be as many as four estrous periods during the rut.  They are in heat for only about 18 hours.  A healthy bull can lose as much as 100 pounds during the rut, due to lack of sleep, food, and frenzied mating for several days.  Rutting involves roars (bugling), thrashing and spraying of urine, with frequent fighting and injuries.  In fact, about 5% of males in a population may be expected to die annually from fighting. Rutting ends quickly after the cows are mated, with the harems breaking up shortly later.  Young females usually adopt a home range overlapping that of their mother, while males will generally leave the area at 2-3 years of age to join a stag group.  Cows reach sexual maturity at three years of age; bulls usually acquire harems and mate at four to five years of age, although they are sexually mature at two years of age.  Life span of 14 to 20 years.
Similar, but larger (3 ½ to 4 ½ “ in length) and rounder than white-tailed deer.  Stride of 30 - 60"; up to 14' when running.
Scats are fairly distinctive, but variable, being larger than deer pellets (1.5” long).  Summer diets produce flat, elongated or circular chips similar to cattle, but smaller in diameter (5 to 6”), and occasionally longer.  Winter scat is the common ungulate elongated “sawdust “ pellet form.

Remarks: The wapiti was originally described as Cervus canadensis, but is currently believed to be the same species as the Eurasian red deer, C, elaphus.  In Europe, the common name “elk” applies to the what we call “moose”; thus, the reason for the preferred common name “wapiti”, meaning “white rump” in Algonquian language. 

Wapiti are the most vocal of eastern US ungulates.  The wapiti has a higher pitched voice than woodland cervids like moose.  This is because deep, low-frequency sounds travel better through vegetation than higher pitched sounds and, conversely, higher pitched sounds travel best across unobstructed open space.  Bugling is an accurate sign of male strength, and is a major attraction used by females in picking a mate.  (As mentioned above, it is believed that the cow selects the harem to which it will belong, and not the bull that selects the harem.  The cow has ample opportunity to leave a harem if she so desires.) 

Antlers are also a quality of males that females observe in their selection.  Antlers can weigh up to thirty pounds and have a spread of five feet.  Reproductive hormones are excreted in the bull’s urine, and it, too, is a reliable signal of reproductive condition.  Therefore, bulls will spray themselves with their own urine and wallow in it to impress the females.

All this bugling, growing large antlers and, well, ok, so wallowing in urine doesn’t use much energy, but the first two items do require the bull to utilize a lot of energy in the late summer period.  This means a lot of foraging in the summer must occur, and this implies a large foraging range in search of browse. 

After the rut, the stressed bulls are weakened and more vulnerable to predation.  At this point, the antlers are a liability, both in terms of weight and visibility, and will fall off quicker for populations living in forested habitats than those living in more open habitat, who may need them for defense and competition among other elk for more limited food sources. 

The last recorded in Virginia were killed in Clarke County in October, 1854, and January, 1855.  In Pennsylvania, the last recorded was in Elk County, in 1867.   The last elk in West Virginia were found near the headwaters of the Tygart and Greenbrier Rivers in 1875, but were gone by 1890.  Also, every county in Tennessee had elk.  The last one shot was in 1849. 

Wapiti have been introduced to north-central Pennsylvania, dating back to 1913.  The stocking program, which totaled 177 elk, ended in 1936.  145 of these elk came from Yellowstone National Park.  A hunting season ensued from 1923 to 1932, when low stock ended the harvest.  By 1965, the population may have been 35 individuals (or less).   As of 1982, an estimated population of 125 existed within an 80 square mile area within Cameron, Elk, and McKean counties.  By 1997, the resident wapiti population exceeded 300 for the first time.  In the winter of 1998, 33 wapiti were transferred east to the Kettle Creek Valley area within the Sproul State Forest (Clinton County).   It is expected by the year 2000, the total state population will reach 500, inhabiting an area of 800 square miles.  (The best viewing is the Elk Viewing Area on the top of the mountain beyond the village of Benezette (presumably, this is Winslow Hill), or anywhere along the local roadways near that town.  To find the Elk Viewing Area, take the road across from the Benezette Store, on Route 555, north a short distance to where the pavement forks at the Benezette Hotel.  Keep to the right and follow the road to the top of the hill.  The Elk Viewing Area is on the left side of the road.  Recently, a 217 acre tract located in Elk State Forest, near Benezette, has been acquired for the protection of the wapiti, and is a popular area for viewing.)  The best time of the year is September.  (A 10/28/01 Washington Post article comments you can stay at the Towne House Inn in St. Marys (800 851-9180), or a Comfort Inn and Best Western, both on the south side of St Marys.  The Bavarian Inn is recommended for German food and rooms (877 351-3624).  Also, there is a mandatory stop at Straub's brewery (814 834-2875), where free beer is served 9-5 weekdays and 9-1 on Saturday!)

Wapiti were introduced into Virginia in 1917.  Between 140 and 150 were taken from Yellowstone National Park and released in Bland, Botetourt, Cumberland, Giles, Montgomery, Princess Anne, Pulaski, Roanoke, Russell, Warren and Washington counties, with the first limited hunting allowed in late December of 1920.  By 1922, an estimated population of 500 elk existed in Virginia.  An additional 56 wapiti were imported from Yellowstone in 1935, released in the Sugar Hollow section of Giles County and Botetourt County near Natural Bridge.  By 1940, the Giles-Bland herd was estimated to contain approximately 100 animals and the Botetourt-Bedford herd approximately 25 animals.  At this time, there were no white-tailed deer in these areas, those being extirpated by the late 1800’s.  A total of 85 white-tailed deer were released in the 1950-1956 period that flourished.  However, these deer were heavily infested with roundworms (brainworm), a nematode parasite, which proved lethal to the wapiti.  The last wapiti was seen in the Botetourt-Bedford range in 1970, and in the Giles-Bland range in 1974.  

After much study and public hearings, Virginia decided in 2000 not to proceed with re-introduction studies of elk for the state.  This was based on agricultural concerns expressed by many farmers as well as the acknowledgement of existing, albeit, unrequested, entry into SW Virginia of reintroduced elk from Kentucky.

An experimental release of elk in the Great Smoky Mountains National Park began with the introduction of 25 in the Cataloochee area in 2001.  25 more are planned for release in both 2002 and 2003, for a total of 75.  This will be a five year experimental release program.  Their impact on the vegetation of the  Park and the rare plants of the unique high-elevation balds will be assessed as part of this experiment.  This highly publicized effort, like the red wolf re-introduction effort, is being driven by the Park's stated mission of returning native species to the area.  (Apparently, the Smokies were near the southern limits of the elk’s former range, and never heavily populated this area.  None-the-less, elk were common inhabitants of the North Carolina northern Piedmont and all of the mountain counties.)   


However, the most ambitious elk reintroduction effort attempted in the east goes to Kentucky.  The Kentucky Department of Fish and Wildlife Resources (KDFWR) has begun the release of wapiti into the Robinson Forest (University of Kentucky property) in eastern Kentucky.  Since 1997, more than 750 elk have been released in this area.  An estimated 200 wapiti will be released each year until 2007 in this area and land owned by Cyprus-Amax Coal Company, managed by the KDFWR, for a grand total of 1,800 elk.  Elk from this release in southeastern Kentucky, have already migrated into Virginia, and, in part, has caused the Commonwealth of Virginia to shelf any plans to reintroduce elk to Virginia at this time.

The reintroduction of elk into eastern US states is becoming a major attraction to state hunting programs.  Funding for much of the state studies to consider reintroducing elk is supported, both financially and politically, by the Rocky Mountain Elk Foundation, a private organization.  Additionally, most of the elk are being supplied through the Rocky Mountain Elk Foundation.  While hunters, State game commissions, and the Rocky Mountain Elk Foundation are active supporters of such reintroduction efforts, some question the net ecological benefits of introducing these large herbivores to our eastern woodlands.  Acknowledging the current pressure on the plant community presented by the deer population, the addition of new, and larger grazers is viewed by some to only amplify this problem.  With limited natural predation, population control can only be had through human “harvesting”, which is not a viable option in the National Parks, and other areas where hunting is not permitted.  Additionally, residents in Pennsylvania are already seeing damage to their crops by the elk population and are receiving state compensation.  Not surprisingly, the Rocky Mountain Elk Foundation is providing funds to the states for this compensation program.

More information about the introduction efforts and the Rocky Mountain Elk Foundation can be viewed by going to ROCKY MOUNTAIN ELK.

Studies as of 1995, suggest there are some 400,00 elk that presently roam the western mountain and plains areas, and 782,500 in the US and Canada.  About 100,000 are killed each year (as of 1982).


WHITETAIL DEER (Odocoileus virginianus) (hollow tooth; of Virginia)  


Appalachian Region Distribution: Throughout.                                                                 
Continental Range: Most all of North America, with the exception of the SW US, reaching down to northern South America.)  Two subspecies are recognized in the study area.
Abundance: Often too common.                                                                                         
Population Density: 13-65 (or more)/ square mile.  Wilson reports an average between 51 and 77 per square mile, dependant on forest quality.  A Texas density of 130/square mile is referenced in Nowak.

Size and Molt: Height 3 to 3 ½ feet, 6 foot length, averaging 100-250 pounds. Males are slightly larger than females. The largest deer are in the northern region (a mature deer on good range can attain a weight of over 400 pounds), and the smallest are the Florida Key deer (weighing a mere 50 pounds).  Actually, equally small deer are found in Central America and coastal South America.  Two molts.  Fawns molt in late summer or early fall, at about three to four months of age.
Mammae: Two pair.
Habitat: Everywhere, especially forest-field margins and suburban areas.
Active Period: Crepuscular, year-round.  They tend to increase activity before storms and restrict activity during bad weather.  They usually bed down during the day, often in the woods, moving into open areas at dusk to begin feeding.  Feeding often continues throughout the night, with a short siesta not uncommon.  Bedding areas are cool, shady areas in summers, sunny areas in cool weather, and more protected areas in bad weather.  Bedding areas are usually not the same spot day after day, but, in winter, small herds will repeatedly use winter yards.
Diet: Strictly herbivorous.  They have a tremendous adaptability in the diversity of foods they eat.  Deer are mainly grazers in the spring and summer (eating herbaceous plants - including sensitive fern and bracken fern), but in winter become a browser, eating twigs, buds and fruit.  In spring, green succulent leaves and stems of both woody and herbaceous species were the dominant food items.  In summer, materials from succulent green plants continue to dominate, with mushrooms also added.  In the fall, acorns were the dominant food item along with  mushrooms, grapes, apples, sumac, and blueberry.  In winter, acorns, grasses, mushrooms, grapes,  and sumac were common food items.  Even rhododendron leaves have been found to be a dominant food item in the more southerly Appalachian Mountains in winter.  Deer have an uncanny ability to select the most nutritious foods available.  Agriculturists tell of observing deer select green bean seedlings that were fertilized over unfertilized ones.  Its four-chambered stomach allows for quick foraging in open fields and later regurgitation for better chewing.  This enables the deer (like all ruminants) to eat fast in the open; ready to bolt from predators, and later, "chew it's cud" in the safety of the forest.  By late winter, bark of the slippery elm is eaten as a last-resort survival food source.  (Deer utilize microorganisms in the stomachs to help digest its food.  Interestingly, while the deer’s diet changes from herbaceous plants in summer to buds and twigs in winter, so, the microorganisms in the gut change species abundance.  This explains why a deer in winter can die with a stomach full of grasses or hay.)  Mushrooms are major food source, after green leaves and buds of woody plants and herbaceous plants.  Woody plants and grasses hold much less nutrient value after the last frosts, thus reducing their value as a major food source for deer.  However, site fidelity overrules food supplies.  Many deer die in winter with their stomachs more or less full of food having little nutritive value, rather than move to areas of less shelter, but more sustaining foods.  Attempts to drive deer from their chosen winter sites with depleted food supply to other sites with abundant food supplies usually fail.  Shedd mentions that deer can put on fat in the fall to help the deer survive the winter season.  The last fat vestiges are found in the marrow of the bones.  Thus, to determine if a deer died from malnutrition, all a biologist has to do is break open a bone and check the marrow.  Marrow from healthy deer is white and fatty, almost like suet.  A moderately malnutrient deer’s  marrow is somewhat less solid with a pinkish tinge, while a deer that was in the last stages of malnutrition has marrow that is thin and red.
Home Range: Highly variable.  A 1985 study in the Shenandoah National Park found, for deer along the Skyline Drive, females averaged 1.7 square miles and males 7.2 square miles, and for deer in backcountry areas, females had an average home range of 3.4 square miles and males a range of 6.1 square miles.  Other statistics found a home range for males may be 1 to 1 ½ miles while females only may range in a ½ to 3/4 mile area.  Nowak says most ranges fall within the ranges of 0.1 - 0.5 square miles for females and 0.4  - 1.4 square miles for males.  W/H says male range varies from 40 acres to 330 acres (~1/2 square mile), dependent on habitat quality.  Winter range is much larger, approaching up to 1285 acres.  Scent glands on the legs near the hooves are used to mark home territories.   Deer occupy the same home range year after year, but are not territorial.  They will, however, defend their bedding sites.            
Social Structure: Social, however, not to the degree (or the large herds) shared by the elk and caribou.  There are two basic social groupings among deer.  Family, or matriarchal groups consist of a doe and her offspring, which remain together for most of the year.  Most of the year, does, her young, and last year’s young form groups. In spring, the doe will drive away the young to bear her young. In fall, the last year’s young doe rejoin the parent doe, but the young bucks join other bucks or remain alone.  The other group is the buck, or fraternal group, consisting of several adult and juvenile bucks.  Males can also remain alone.  Older bucks may form groups of their own.  Normally, the buck groups will form in the winter after mating, and will disband after the velvet is lost; 4 to 6 weeks prior to the fall rutting.  At this time, sparring will establish the new dominance hierarchies prior to the actual breeding.  Occasionally, mixed feeding groups of both males and females will occur.  In the autumn rut, bucks can either remain together, except when pursuing a doe in heat, or separate for the duration of the rutting season.  Bucks are polygynous; forming a harem of up to ten does, although, with group members constantly moving in and out of herds, the concept of a harem is really a misnomer.  (In fact, Nowak states that a male does not attempt to gather a group of females or to defend a territory, but rather, follows a single doe until mating (or driven away by contenders).  A dominance hierarchy exists, both in the male and female groups.  Dominance hierarchies are maintained through complex, stereotyped behavior and threat displays.  Such displays include stares, head bobbing, kicking, chasing, and various vocal sounds.  Winter herds may number up to 150 individuals, centering around areas of food abundance called “yards”.  These yards, better called “deer wintering areas”, are not cleared areas, but rather areas with numerous overlapping paths, sloping south or west, with considerable vegetation, especially coniferous vegetation.  In such circumstances, leadership is matriarchal (different from dominance, which is always the largest male).  Even though it appears to be one large group of deer, they are actually a concentration of groups of either bucks or does and their offspring.  (The Shenandoah National Park study found winter group size ranging from one to 28, with a mean of 3.4 and late summer group sizes ranging from 1 to 10, with a mean of 2.0.)  Herding allows pathways to remain open and to enable more individual protection from predators.  As the fawning season approaches, the wintering groups disperse, with the pregnant does seeking secluded places for birthing.
Life Cycle: One litter per year with two per litter for mature adults (10-15% bear triplets; four are rare, but recorded). Single fawns will be produced by first litters, old does, and in poor seasons. Does are in heat for 24 hours, and, if not impregnated, will not be receptive for another lunar month (28 days) (seasonally polyestrous from October to January). The main breeding is in November (some reach estrus in October, with non-mated females achieving estrus again in November and a possible final third time in December).  In our Appalachian study area, the peak period of heat is the second and third week of each of the three months. Gestation period is 200 days. Most young are born in late May and early June in this Appalachian region.  Being altricial, the young hide in the tall grass or bushes motionless for the first three to four weeks while the mother feeds. The mother will have little contact with the fawns for the first four weeks, except for nursing, which occurs 3 or 4 times a day.  Twins will hide in separate locations.  During this time, the fawns have little or no scent. After a month, they follow the mother and start eating solid foods. By about September, the fawns are weaned (three to four months of age - Wilson says 8 to 10 weeks, Nowak says completely weaned at four months) and lose their spots (four to five months of age). They will stay with the does (who form winter herds) over the first winter before males disperse in the spring, with some female fawns staying with the mother for two years.  Dispersal is on the order of 6 to 120 miles.  Some young does can breed in the first fall, often late in the season, and will usually bear only one fawn.  As many as 60 – 70% of females of this age class do breed according to some studies.  Such successful reproduction is directly related to nutritional conditions.  A mean of 1.3 young per female was found for 130 first fall birthings.   Males, and most of the spring females, don’t breed until the second fall. Most deer are killed at two or three years of age. Females (and those lucky males not shot) can live to about 10 to 15 years in the wild.
Nest: Deer make no permanent dens or nests, moving from site to site on a daily basis. Bedding sites are simply concealed sites, forming three-foot diameter depressions in the vegetation.
Tracks: Two hooves; about 2 ¼ to 2 ¾” long.  In mud or deep snow, dew claws can also be seen.  Straddle of 4-6".  Stride of 1' when walking, up to 6' when running.  Can leap up to 30' (so NAS says).   Favored “buck rubs”, are made with antlers in fall on trees about an inch in diameter and six feet high.  They can be made rubbing off velvet, marking territory, or by venting hormone-induced frustration, waiting for a doe to go into estrus.  Bucks also make “buck scrapes” to establish their territory.  Scrapes are made by pawing the ground and thrashing low bushes. 
Scat: Scat is commonly 1" long; 3/8" diameter; cylindrically shaped, with a flat end and a tapered end, in groups of 20 to 30.

Remarks: As noted above, the species name means "hollow tooth".  Here is another case of zoological names being misleading.  Linaeus was given a tooth that was, in fact, hollow.  However, this was from an old deer with decaying teeth.  Normally, the teeth are not hollow.

White-tailed deer evolved in North America approximately 4 million years ago.  Their ancestoral Asian ungulate stock evolved more like 60 million years ago, shortly after those dastardly dinosaurs were deemed dispensable.   

There are two species in the genus Odocoileus, the white-tailed, and the mule, or black-tailed deer.  However, keep in mind the tail of the white-tailed is dark and the black-tailed is white.  Actually, the tail of the white-tailed is brown above and white laterally and below, while the black-tailed is white or black above and tipped with black, and is a bit smaller in size.  Also, the antlers of the white-tailed has one main beam with minor branches, while the antlers of the black-tailed branches into two nearly equal parts.

Deer are the most popular big-game mammal in eastern US.  Bucks become quite aggressive when in rut. While young deer make good pets, a newly mature buck has often attacked its owners due to the increase in testosterone, responsible for antler growth and maturation. Antler growth is directly related to genetics and nutrition, with nutrition playing the greatest role in the size of antlers.   Antler growth is triggered by a change in photoperiod, or day length. As day length increases, the  pituitary gland in a buck's brain produces a hormone which stimulates the pedicels to begin growing antlers.  Concurrently, the pituitary triggers the testicles to begin producing testosterone in small amounts.  Antlers grow from March and continues through late August, supported by a vascular, soft skin, supplying nutrients necessary for growth, called "velvet".   By late September, the hormones change; the velvet shuts down it’s growing process, and the male sloughs off the dead skin, producing the common scrapings found on bushes and young trees in the bucks’ range.  By late December to February, a pituitary hormone called androgen, which has maintained the point of connection between the antler and the pedicel, now decreases in levels,  causing a portion of the base of the antler to be reabsorbed, causing an abscission layer (like leaf bases in the fall), resulting in the antler falling off, usually one at a time.  In another six to eight weeks, the new buds start the cycle over again.

While antlers are growing ('in velvet'), they are susceptible to injury and subsequent malformation.  This is why a buck can be found with one normal antler and one deformed.  Interestingly, injuries to the large leg bones may cause abnormal development in the opposite side antler.

Antlers, replaced annually (as opposed to permanent horns), are first produced at 1 1/2 years of age, and are the largest at the peak of the breeding age (usually 5 1/2 - 7 1/2), not the oldest age. In the eastern US, all points on both antlers are counted, while in the western US, only a single antler is counted, so that a 10 point buck in VA is a 5 point buck in CO.   Females with excess testosterone may produce small antlers (one study states one out of 4,000 does; Chapman and Feldhamer reports  1 in 1,000).  Antler size increases with age, but cannot be used to age a deer, due to habitat quality and calcium intake variations.   It is not unusual to encounter yearling and two year old bucks without antlers due to poor nutrition.  Similarly, yearlings have been found with small eight-point racks in areas of high quality habitat.  Normally,  1 1/2 year old bucks have forked antlers, or, less often, spikes.  Spikes would be more likely in yearlings bred in December, rather than the normal November rut.  For eastern US deer populations, a rack with eight points is most common.  Racks with 11 or more points is found in one buck per 20 to 70.  One buck per 300/400 will have 13 points or more, and one in about 1,000 to 1,300 will have 15 or more points.  Only one in about 4,500 to 5,000 will have 17 or more points.

Deer deal with winter by putting on body fat in the fall, reducing their metabolic rate,  producing a heavy winter coat, and moving to “deer yards”.   Their winter coat hairs are tubular, stiff and brittle.  For this reason, the pelts float and have been used for life preservers. 

Contrary to popular opinion, fawns are not scentless.  Research biologists have used dogs to locate fawns by scent.  Their scent is much less pronounced than an adult, but does exist. 

It is not clear why deer raise their white flag tail as they run away.  A warning to other deer, a distraction to the predator to protect the young, a “follow me” sign to the young, or a startle effect to confuse the predator are all proposed explanations.  The explanation given the most weight is a “I know you’re there, so you’ve lost your advantage, and you might as well give up”.  (Perhaps a liberal use of anthropomorphism, but you get the idea.)

Partially white (piebald) deer are not uncommon.  Albinos are more uncommon, with several being reported at Fort Pickett, VA.  Melanism is extremely rare, but has been observed. 

The Key deer of Florida is a subspecies of the white-tailed deer (O. virginianus clavium), averaging 50 pounds.  As of 1945, only 26 individuals were known to exist.  Full legal protection has allowed this number to swell to an estimated 350-400 in the early 1970's.  However, continued loss of habitat, killing by automobiles, and other mortality has caused an drop to about 250 animals in 1999, with the trend suggesting the likelihood of extinction within the next century.  

Hybridization between mule deer and white-tailed deer occurs in the wild as well as in captivity, where most of the offspring are sterile.  

While white-tailed deer had to be reintroduced in the Shenandoah National Park in the 30’s (the last deer were chased down by packs of wild dogs), the Smokies maintained a scarce population that has repopulated the park.   

West Virginia never totally lost their deer population, but only scattered herds could be located in the remote high mountain areas by the early 1900's.  Reintroductions starting in 1933 and continued until 1957.

Legal hunting accounts for 2 million deer harvested in the US, with another 500,000 deer killed each year in collisions with cars. 

The following presents the total number of white-tailed deer in Canada and the US :

Precolonial     23 to 40,000,000

1908                           300 - 500,000

 1948                   6,000,000

1978                                15,000,000

 2000               25 to 27,000,000

In certain areas, like the Great Lakes region, where logging has created favorable habitat, there are more deer now then in pre-Colombian days.