Rabbits, Hares, and Pikas    




Lagomorphs are primitive placental mammals dating to the Paleocene times, about 62 million years ago in Asia. This order includes two families in the world.  The Ochotonidae family includes the pikas in the western US.  Hares and rabbits are in the Leporidae family.   The greatest radiation of species has occurred in North America.  Ears, used for defensive purposes, are largest in hares, who live in open habitats, and smallest in pikas, living in the protected environment of talus slopes.

Although similar in resemblance to rodents (with substantial evidence indicating they should both be put together in a superordinal group known as Glires), this is more a function of their similar herbaceous life style (convergent evolution) than direct evolutionary relations.  In fact, some studies now suggest that lagomorphs are more closely related to artiodactyls (even-toed hoofed mammals) than rodents. 

Both rodents and lagomorphs have a pair of large upper incisors that grow throughout their life, but the lagomorphs have a smaller pair right behind the first.   (In fact, even a third set of incisors is present in the lagomorphs at birth, but is soon lost.)  One slight difference exists in that the large incisors in lagomorphs are covered both front and back by enamel, while only the front of the incisors is covered by enamel in rodents.  A space exists between the incisors and the remaining cheek teeth. This space (diastema) is the space occupied by the canine tooth in many mammals. Canine teeth are absent in rabbits, hares, and all rodents. More significantly (at least to the rodents), rabbits and hares lack a penis bone (baculum), that is found in rodents.

Lagomorphs are herbivores that practice coprophagy, the re-ingestion of fecal droppings.  On first "pass", these soft, green pellets pass through the digestive system only partially digested. These pellets, rich in protein and B vitamins, are re-ingested directly from the anus.  After reingestion, the pellets are a drier brownish in nature.  This practice allows the animals to spend relatively little time exposed to predators while in the field actually feeding.  They consume green vegetation rapidly and then make optimum use of it in the safety of their cover.  This process is also called "pseudo-rumination", since it is functionally the same as cows chewing their cud.  Coprophagy is also practiced by beaver and voles and, apparently, by some shrews. 

Worldwide, there are only two families, with 13 genera and 81 species in the rather small lagomorph order (Nowak's Walker's Mammals of the World).  None were native to Australia.  

Five genera and 20 species are found in North America (including the introduced European rabbit and European hare), with three species in the Appalachian region; all in the Leporidae family (Jones).


Family Leporidae - Rabbits and Hares  


Rabbits and hares (leporids) differ in that rabbits give birth to altricial young (naked, closed eyes, requiring some time in the nest before ready to roam), while hares give birth to precocial young (born with hair, open eyes and ready to run) and don't 'make nests. Correspondingly, less postnatal care is necessary for hares than rabbits (however, hares have longer gestation periods).   Hares are creatures of open habitats and are specialized for running (up to 40 mph).  Rabbits inhabit more brushy habitats, have shorter legs, and thus, do not run as well.  Rabbits make their fur-lined nests in dense vegetation with smaller home ranges while hares live in simple depressions in open habitat and have large home ranges (Wilson says hares prefer rock crevices and caves).   While European rabbits make permanent extensive burrows, our North American rabbits and hares do not dig burrows (nor do they dig up carrots).  Instead, they make well-concealed shallow "forms" on the surface.  Rabbits and hares will use burrows of woodchucks and skunks during periods of bad weather, never venturing far inside.  Rabbits tend to run for cover when threatened, while hares, adapted to a more open habitat, will opt to out leap and outrun predators. 

Rabbits can be gregarious, while hares are often solitary.  In areas with high densities of rabbits, females form dominance hierarchies, with the dominant females suppressing the reproduction of subdominants by denying them access to nest sites and by physical intimidation.

Both rabbits and hares produce relatively large litters and are prodigious reproducers, with large populations maintained; supplying a significant food source for many predators (high fecundity is balanced by high mortality).  This is known as an r-selected strategy.  (The other primary reproductive strategy is k-selected species which produce low numbers of young but a much greater proportion of the young enters the breeding populations.) Reproduction is increased by a phenomenon known as induced ovulation.  Basically, ovulation is induced by copulation (like cats).  In both cases, the scentless young are left alone in the nest for the first few weeks, with the mother nearby feeding, in order to protect them from predators. The mother only approaches for a short period, often at night, for nursing and then leaves the young again.  The females (does) are known to grunt or purr while nursing.  The males (bucks) offer no parental care.   

Northern forms frequently have larger litter sizes and shorter gestation length than do southern ones.  This adaptation takes advantage of the shorter growing season while maximizing the food resources.

The phrase, ďMad as a March hareĒ comes from the springtime mating antics of female hares, sizing up the courting males.  Females stand up on their hind legs and cuff males in the face and ears.   Female hares and rabbits are normally larger than males, indicative of the evolution of selection for combative females.

Regarding the courtship behavior of lagomorphs, the following describes the snowshoe hare, ďThe male snowshoe approached the female, sniffed her and jumped into the air.  After landing, the male urinated on the female and left.  The male re-approached the female, and the female jumped into the air twice, after which the male left.  The male returned, jumped into the air and urinated on the female.  Both snowshoes then went into the bushes, where more jumping occurred  Similar courtship behavior is practiced by other hares and rabbits. 

In contrast to the case in most mammalian families, female leporids are usually larger than males.

Scat resembles chocolate M& Mís; dark brown, round and slightly flattened. In addition to these fecal pellets, soft, greenish pellets are formed that are only partially digested and will be re-ingested directly from the anus through the process known as coprophagy, as noted above.  Found in groups of 5 to 10.

Rabbit tracks have four front and rear toes, often with claws showing.

There are 11 genera and 54 species of rabbits and hares in the world (Nowak), with 5 genera and 18 species in North America (Checklist of North American Mammals), of which two species are introduced. 

Two species of rabbits (Sylvilagus) and one hare (Lepus) exists in the Appalachian region. These are the eastern and Appalachian cottontails and the snowshoe hare. 

EASTERN COTTONTAIL (Sylvilagus floridanus) (forest, hare; from Florida)

Appalachian Region Distribution: Throughout.                                                                 
Continental Range: East of the Rockies and south of Canada throughout Mexico and Central America.  Seven subspecies have been recognized by W/H, however, Chapman and Feldhamer note that subspecies are somewhat meaningless, due to the wide transplanting of this species that has occurred with time.
Abundance: Common.  The widest distribution of all of the cottontails.  The range of this species overlaps that of seven other cottontails and six species of hares. 
Population Density: 2-5 /acre.  Novak reports 4/acre in Maryland.  W/H reports up to 9/acre, but normally much lower.  Periodic cycles may have occurred in historic populations, but recent land use changes has more profound impact on population densities than any natural functions.  Nesting density is habitat-specific.  A PA study found nests every 1.5 acres in unkempt orchards, 7 acres apart in hayfields, 13.5 acres in woodlands, and 14 acres required for nests in pasture lands. 
Size and Molt: Head and body 14 - 19 inches; 2 - 3 pounds. Females slightly larger than males. Two molts.
Mammae: Four pair.
Habitat: Everywhere but wet areas. Prefers heavy brush. (On the southern and southeastern peripheries of the region (lower elevations), the swamp and marsh rabbit will inhabit these areas.)
Active Period: Chiefly crepuscular to nocturnal, active from early evening to late morning. Spends days resting in a "form"; a small, scratched-out depression in a clump of grass. Can be observed "sunbathing" on warm days.  Active year-round.
Diet: Herbivorous, green vegetation in summer, twigs in winter (especially dewberries and blackberries). Easier to identify what is not eaten by rabbits. Practices coprophagy (see Order Lagomorpha above).
Home Range: 2 Ĺ to 5 acres is normal, with a range from 1 to10 acres (depending on populations, habitat quality, season and sex, with the male having a slightly larger range).  Ranges for males are the largest in the main breeding period of late spring-early summer, while females have the smallest range at this same time.  Dominant males have the largest ranges.  Ranges are generally smaller in spring (before breeding) and winter reflecting lush vegetation and severe limiting weather.  Not being territorial, ranges of individuals often overlap (up to 50% for males and 25% for females in spring), especially in winter, when they tend to concentrate in areas offering the best combination of food and cover.  Chapman and Feldhamer report females have little or no overlap of home ranges during breeding season.
Social Structure: Solitary, generally not territorial, except for females in the immediate vicinity of a nest.  Males have a dominance hierarchy in which the most dominant have more aggressive encounters with other males and do most of the mating. Dominance hierarchy of males allows the strongest males to fertilize more females than subordinates and also minimizes fighting. Most aggressive behavior is exhibited between the dominant male and the individual immediately below it in social status.  Females have a less rigid hierarchy.  Females exhibit dominance over males except during estrus.
Life Cycle: Breeds throughout the growing season (from February through September) with three to five litters per year (can be up to 7 litters in warmer seasons) with an average of  5 per litter (can be as many as 12). Throughout its North American range, litter size increases with latitude (from 3 to 5) and gestation period of about 28 days. The female comes into estrus immediately after giving birth (post-partum estrus and pregnancy).  Like all cottontails, they are synchronous breeders (all members of a population tend to breed at the same time).  After four to five weeks, the altricial young are weaned and leave the nest and the mother gives birth again. Thus, the mother is pregnant most of the season.  45% mortality of newborns within the first month, with 80% mortality by the end of the first year.  In our Appalachian region, most females and virtually all males will not breed until the following spring.  Up to half of the newborn female young will breed their first season in the southern states.  Usually, about 80 - 85% of the cottontail population are juveniles.  Life span of two years is reached by only 25 % of adults.
Nest: North American rabbits don't dig or live in "rabbit holes", as some slower European species do, although they will occasionally utilize groundhog holes.  Other places of shelter include brush piles.  Nesting is done in well-hid surface depressions, or ďformsĒ.  This elaborate nest is more of a slanting hole; six to seven inches long, five inches wide, four to five inches deep, lined with grass and then covered with the motherís hair.  Newborn are covered with nest material during the day while the mother feeds nearby.  The mother returns to the nest at night to feed the young.  She does not actually reside in the nest; she merely crouches above it, and the babies climb to the top of the nest to nurse.  They will use underground dens of woodchucks as a temporary home, especially during winter storms.  
Tracks: The normal hopping pace produces the well-known "Y" form, with the front two feet forming the stem of the "Y" and the larger rear feet landing in front forming the two top sides. Usually, the snowshoe hare does not overlap the range of the eastern Cottontail. Normally will make straddle trails  4-5 inches wide.  
Scat: Similar in shape to "M & Mís"; about 3/8" in diameter.  One rabbit may void 250 to 500 pellets in a day.

Remarks: The eastern cottontail can jump 10 to 15 feet.  They can often be differentiated from the Appalachian cottontail (next listing) by a white spot on its forehead, whereas, the Appalachian cottontail often has a black spot between the ears.  Urinating on the partner is a part of the mating ritual.   Rabbits are r-selected, exhibiting massive births with few surviving to adulthood versus k-selected, which emphasizes few young with high percentage making adulthood.

Valued as the most important game species in many parts of the eastern US, the eastern cottontail can be infected with the disease tularemia, a bacterial disease that can be transmitted to man handling the rabbit with open cuts.  

In flight, will stay in its known range, often circling in a large loop back to its original site and often jump sideways to break their scent trail.  

Many millions of rabbits are killed each year.  


APPALACHIAN COTTONTAIL (Sylvilagus obscurus) (forest, hare; obscure)

Appalachian Region Distribution: Throughout higher elevations from PA to GA.
Continental Range: Same. The type locality is Dolly Sods, WV.  No subspecies are described. It is postulated that unbroken woods, rather than high elevation, delineates the distribution of the Appalachian cottontail.  See remarks for a detailed WVA range.
Abundance: Uncertain, never abundant, but declining due to deforestation.  A candidate for designation as federal endangered species.
Population Density: Unknown
Size and Molt:
Head and body 14 - 16"; 1 Ĺ - 2 Ĺ pounds. Females are slightly larger than males. One molt, unlike the Eastern Cottontail.
Mammae: Four pair.
Habitat: Dense forests and thickets at high elevations, especially birch/red maple forests, hemlock and rhododendron areas within oak-hickory forests, blueberries, mountain laurel and coniferous forests. Has a noted preference for six to seven year old clear cuts and old overgrown farmsteads and pockets of heath-conifer habitat.  More specialized than eastern cottontail.  
Active Period: Active from early evening to late morning (nocturnal).
Diet: Less varied than eastern cottontail, although some studies conclude that dietary preferences are nearly identical.  The only cottontail that feeds extensively on conifer needles. 
Home Range: Ĺ to 1 Ĺ acre.
Social Structure: Exhibits both solitary and social behaviors.  They groom and dust alone, but vocalize amongst themselves and may establish hierarchies.  
Life Cycle: Three to six litters with an average of four per litter per year, starting in March to September.  Gestation of 28 days. Altricial young.  Like all cottontails, they are synchronous breeders (all members of a population tend to breed at the same time).  WV studies show 18% of females bred in the first year, while males were not able to breed until the following season.  Life span of 3 - 4 years.
Nest: Similar to eastern cottontail.
Tracks: Similar to Eastern Cottontail.
Scat: Similar in shape to "M & Mís"; about 3/8" in diameter.

Remarks: W/H and NAS calls this the Allegheny cottontail.  Smaller than the eastern cottontail, commonly with a black patch between the ears, which are shorter and rounder than the eastern cottontail and a greater amount of black on the back.  Unlike the eastern cottontail, the Appalachian cottontail undergoes only one late summer molt.

A WVA DNR staff member says the Appalachian cottontail is abundant in clear cuts of higher elevations in WVA, including Dolly Sods and Spruce Knob.  They love dense heath protection. (Mentioned Mower tract, clear cut 15 years ago, now thick with 8' spruce and thick with Appalachian cottontails.)

Hybridization occurs with the eastern cottontail and New England cottontail at overlapping ranges. The New England cottontail (Sylvilagus transitionalis) has recently been split from the Appalachian cottontail (1992). Although they both have very similar mountain habitat, they are allopatric (don't live in same area), so interbreeding cannot be tested. When captured, the Appalachian cottontail is more aggressive than the eastern cottontail.  However, in the wild, it is the eastern cottontail that out-competes the smaller and less aggressive Appalachian cottontail.  The New England cottontail is restricted to boreal habitat in Maine, New Hampshire, Vermont, Massachusetts, Connecticut, Rhode Island and New York as far west as the Hudson River.  



SNOWSHOE HARE (Lepus americanus) (hare; from America)

Appalachian Region Distribution: Confined to the higher elevations of the Appalachians as far south as the Allegheny mountain range into TN in disjunct populations (no documentation of these recorded in the Smokies, although some authors indicate that it was a former resident).   See remarks for details of distribution.  In 1989, 26 hares trapped in West Virginia were released with radio collars in the Laurel fork region of Highland County of Virginia. Twenty-four survived an average of 23.8 days post release. They were lost primarily to predation.  They are not known in Maryland or North Carolina.
Continental Range: Throughout Canada, extending south along the Rockies and Appalachians.  Four subspecies are recognized.  
Abundance: Somewhat common in good habitat, but decreasing in numbers due to deforestation and increase in white-tailed deer population (reducing available habitat and competition for food).  Captured Canadian hares have been released in VA and MD, although apparently not very successfully.  In MD,  no individuals have been taken in the last 30 years.  See remarks below for the status of the snowshoe population in WV and VA.
Population Density: 1-5/acre.  Densities in the Appalachians are probably highest in early succession forest stages after fires and clear cuttings.  The Appalachian region populations do not tend to exhibit the cyclic nature found in more boreal habitats.  Stephenson suggests that the cyclic nature of the northern populations is a result of the extensive uniform vegetation of aspen, alder and spruce found in the north, dictating a uniform foraging habit among the hares.  The broken forests, thus diverse vegetation of the Appalachians, results in varied food sources of our hare populations, thus eliminating any mass population effects.  (Canadian populations exhibit 6 to 13 year population cycles, averaging ten years.  An Alberta study has found population densities ranging from .1/acre in low years to 4/acre in high years during a 16-year period. Nowak reports a northwestern Canadian study that found a range from 6/square mile to 3,380/square mile, and even 10,400/square mile.  These cyclic patterns have been explained as an interaction between hares and their food supply, followed by a hare-predator interaction.  Specifically, when the shortage of vegetation ultimately causes a population crash, the predators reduce the populations below the level dictated by limiting vegetation alone, increasing the drop of hare numbers.)  
Size and Molt: Head and body 13.5 - 18.5 inches; 2.2 - 5.0 pounds, averaging 2.8 pounds. Females are slightly larger than males.  The only lagomorph that changes pelage color.  Two molts, with the winter molt taking 70 to 90 days beginning in October and being complete in December; the spring molt being complete in May.  Not only a defensive aid, the hollow white hairs, without the pigment melanin, have more air spaces within the hairs and thus has greater insulation.  Snowshoe hares' white winter pelage has 27% better insulative qualities than the summer brown coat.  The molts are triggered by day length.  Populations of a subspecies of L. americanus (oregonus and washingtonii) in the Pacific Northwest, where snow does not generally cover the ground, do not turn white in winter.  Some melanistic hares in the Adirondack Mountains remain black all year.  
Mammae: Three pair.
Habitat: This is a forest species, never far from dense woods, including swamps and thickets. Prefers coniferous forests.  Often found in dense second growth beech/birch/maple forests and young spruce stands of WV.  Rhododendron and mountain laurel thickets are its habitat in the southern mountains.
Active Period: Largely nocturnal and crepuscular. Active year-round. Sits in "form" during the day, located in bushes, or under logs or stumps.
Diet: Herbaceous plants in summer, twigs in winter. Maurice Brooks (1955) found snowshoe hares prefer southern highbush cranberry, but also fed on rosebay rhododendron, red spruce, and hemlock in West Virginia.  Unlike most lagomorphs, Brooks also found that the snowshoe hare is fond of meat and will eat carrion, especially in winter (he found two feeding on a deer carcass).  Apparently, occasionally a nuisance to trappers by stealing bait.  Diet closely corresponds to that of white-tailed deer, with direct competition likely in times of high pops/low food resources.  Hares practice coprophagy, ingesting soft, mucus-covered pellets directly from the anus, high in proteins and Vitamin B. 
Home Range: 15 - 25 acres, dependent on population density and vegetative cover.  A maleís territory typically overlaps that of several females and may or may not be larger than the females home range. A Montana report finds an overall home range of about 25 acres for males and 19 acres for females and a Colorado/Utah study found 20 acres for both sexes.  
Social Structure: Solitary, but will tolerate others in feeding areas.  Novak reports that this hare is social, with several adults often living in close proximity and sharing the same forms.  Novak further states as many as 25 hares have been seen in a clearing at night.  Known to exhibit elaborate courtship behavior during the breeding season, including much fighting between both bucks and between bucks and perspective doe mates.  Females have been known to be injured by the overly-excited kicking and biting of the testosterone-laden males.   Premating activities include both sexes jumping while urinating on its mate.  (Note information presented in discussion of the Leporidae family above)
Life Cycle: In this study area, on the southern limits of the range, two to three litters, with three per litter being common.  Breeding begins in March with broods occurring from April to August.   Higher numbers of litters (3) and young per litter, averaging 6.5 in Michigan, are found in the central portion of the hareís range.  Numbers of litters at the northern limits of the range are smaller (2), but litter size increases (4-5).  First litters of newly mature females are the smallest.  It has been found that largest litters result after winters of greatest snows, presumably due to the increased depth allowing the hares to reach higher for the younger green nutritious branches.  Females are seasonally polyestrous, with a gestation period of 36 days. Precocial young (born active and alert, fully haired with open eyes) are able to hop on their first day, and are weaned in 25 to 28 days.  Mothers are characterized by postpartum estrus and are often impregnated within a day after giving birth.   Until weaned, the "leverets" hide separately (on their own, or by the mother; depends on who you read) during the day, returning to the nest in the evening for one nursing, which only lasts for 5 - 10 minutes.  Young Lepus usually do not breed within their first calendar year of life.   Life span of 3 - 5 years, although probably not more than 15% survive to breed during more than one season.
Dens/Nest: Snowshoe hare do not dig or occupy burrows, relying on their speed to escape danger.  Nor do they make nests, even for birthing; rather, depressions (forms) are made. Sits in thickets during the day.
Tracks: Hindprints 4-5" long.  Straddle about 6".  Like a large rabbit, with the rear feet placed just in front of the front feet. Normally, the front feet will be oriented in a diagonal manner, while squirrels will have their front feet aligned straight; perpendicular to itís direction of travel. Has a well worn system of trails in it's range. Many other animals use these trails, especially in winter when the snow is hard packed from the hare's travels.
Scat: Similar in shape to "M & Mís"; about 1/2" in diameter.

Remarks: One of the smallest of the hares, and the only eastern hare, also called the varying hare. Larger than the cottontails, with much larger hind feet, adapted for running on snow. Hares will run in circles when frightened, returning to the original site. Known to leap 12 feet at a bound and run at 30 mph. Known to take "dust baths". 

Populations in the Pacific Northwest, where snow does not generally cover the ground, do not turn white in winter.

Populations seem to be declining in the southern Appalachians.

Pennsylvania has snowshoe hare populations limited to the state's mountainous regions, with the exception of the SW PA Appalachian plateau, where it is rare.   It's habitat is ridges and slopes of mountain laurel, rhododendron and hemlock.

West Virginia has a disjunct population, restricted to spruce and northern hardwood forests above 3,000 feet.  Snowshoe hare range in the Allegheny Mountains of West Virginia generally extends north to the vicinity of Mt. Storm Lake, Grant County; to the east the range follows the main Allegheny Mountain to the south in the vicinity of U.S. Route 250; to the west, starting as far south as Cold Knob in Greenbrier County, following the west side of the higher elevations of the Elk River, Greenbrier River, Williams River, and Cheat River Watersheds in Pocahontas, Randolph, and Tucker Counties.   Seems well established in all high elevation areas of snow cover and brush.  Will be found in snow covered spots when snow cover is limited.   Summer range is the same.  Habitat varies from open to very closed.  

In Virginia, historic habitat closely coincided with the red spruce forests, now depleted.  Current habitat is second growth birch/red maple forests with a significant and dense shrub layer.  Such habitats are declining as secondary growth woods are maturing, limiting the shrub layer.  Audubon and Bachman were unsuccessful in their attempts to confirm the existence of snowshoe hare in VA in 1846.  Efforts in the 1960ís and 70ís to introduce nonnative hares from New Brunswick were, fortunately, unsuccessful.  Beginning in 1989, a restocking program using native hares from WVA introduced 26 hares to the Laurel Fork area in the GW National Forest.  Apparently, none survived, due to predation by bobcats and other predators. By 1991, no more than a few dozen hares existed in a 15 square mile area in northwestern Highland County.   At that time, it was acknowledged that habitat management is urgent and critical for snowshoe hare in Virginia.  It was also acknowledged (in 1991) that, even with a crash program to improve habitat, the snowshoe hare probably will disappear from Virginia in the next decade.   

The mating behavior of the snowshoe hare is unique in that hares will jump in front of prospective mates and commonly urinate on their partners during premating activities.   

Between 200,000 and 400,000 hares are killed annually by hunters in Michigan alone, with any one years' harvest reflecting the population level.