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ORDER RODENTIA - Rodents  

         CHIPMUNK       WOODCHUCK    TREE SQUIRRELS  FLYING SQUIRRELS
           BEAVER                MICE                RATS            MUSKRAT
            VOLES          LEMMINGS      JUMPING MICE         PORCUPINE

 

Rodents are the largest order of mammals in the Appalachian region (as well as the world), both in species and actual numbers. In fact, 40 % of the world’s mammalian species are rodents.   The diversification of species has been a relatively recent and rapid evolutionary event, with few extinctions, causing much consternation among taxonomic botanists.

Rodents are the "gnawing" mammals. Rodents are known for their pair of large upper and lower chisel-like incisors.   Each incisor is a segment of a true circle, continuously being pushed out of the end of the jaw.  Some species have incisor growth rates of one inch per month. Unlike lagomorphs that have enamel covering the entire incisors, rodents only have hardened enamel on the anterior (front) side of the incisors.  This keeps a sharp edge on the teeth, since the relatively soft posterior (back) side of the teeth wears more quickly than the front. Also unlike lagomorphs, rodents have only one pair of upper incisors, instead of two permanent pair.   These incisors grow throughout the life of the rodent, kept in proper length by grinding of the teeth, not necessarily by gnawing plants.  Rodents lack canines, with the gap between the incisors and molars known as the diastema.  This large gap allows rodents to curl their lips backward into it while gnawing and chewing.  This behavior also permits the rodent to exclude unwanted soil and debris from its diet.  This also enables muskrats and beaver to eat underwater without inhaling water. The grinding molars, or "cheek" teeth,  are a combination of vertical layers of dentine and harder enamel, which produce peculiar ridged patterns on the crown of the teeth, useful in classification by zoologists.

Rodents have diversified to adapt to a variety of lifestyles including terrestrial (mice and voles), arboreal (squirrels), fossorial (woodchucks), semiaquatic (beavers and muskrats), and volant (flying squirrels). They range in size from a third of an ounce of the harvest mouse, to the eighty pound beaver. All are herbivores.  

With the cooling and drying out of much of North America during the Tertiary and the Eocene (40-55 million years ago), forests transformed into grasslands, with the concurrent evolution and diversification of mammals.  Grasses became the most important of all the flowering plants and became the  nutrient source base of wildlife herbivores, domestic grazing livestock, and, in fact, mankind.  As the plants provided an abundant energy source, herbivores expanded in number and evolved into the ecological niches.  The ungulates evolved in the Eocene, the grass-eating marsupials arose 25-38 million years ago, and the voles of the microtus genus were the last to emerge, about 10 million years ago.  They have rapidly evolved and diversified for the past 6 million years filling all grassland niches.  

In order for rodents to fill out the grassland niches, two adaptations were necessary. First, as a result of the abrasive silicas in the cells of grasses, teeth, easily worn down by the silicas, became ever-growing throughout the life of the rodent.  The second adaptation was the ability of the herbivores to digest the complex carbohydrates contained in the fibrous portion of grasses through fermentation in the animal's gut by microorganisms.

The success of the rodents became a problem in itself as carnivores evolved to take advantage of this abundant new food source.  In response, special reproductive adaptations evolved to enable massive birth rates within the rodent order. This includes "postpartum estrus" in mice, with females being mated within hours of birthing.

Rodents have four toes (sometimes five) in front and five toes in the rear.   

Rodents are known from fossils in North America dating back to late in the Paleocene Epoch (60 million years ago).

Worldwide, the rodent order is represented by 29 family, 468 genera and 2,052 species (Nowak's Walker's Mammals of the World).  In North American, there are 9 families,  43 genera, and 217 species (Jones' Checklist of North American Mammals).  In the eastern United States, there are about 38 species and five introduced species.

Rodents include twenty nine species in the Appalachian region within five families. These include the squirrel - which includes the groundhog - (Sciuridae), beaver (Castoridae), porcupine (Erethizontidae), mice, rat and vole (Muridae), and jumping mice (Zapodidae) families.

 

Family Sciuridae - Squirrels and Woodchucks

This family can be divided into three groups; tree squirrels, flying squirrels, and ground-dwelling squirrels (woodchucks and chipmunks). Tree squirrels are diurnal, leading solitary territorial lives, living on nuts and fruit.  The woodchuck and chipmunk are also diurnal.  Flying squirrels are nocturnal and tend to be more omnivorous.  The family name means "shade tail", alluding to the large bushy tail of the arboreal tree sciurids. Most of these family members stand on their haunches to view their surroundings. Most members of the sciurids have high birth rates to compensate for their high mortality, especially to the weasel family (R-selected strategy populations, like the lagomorphs).  Squirrels come in different sizes, habits, and habitats to maximize utilization of available resources and to minimize competition among species.  They have the largest brains relative to their body size of all small mammals and are the most intelligent of the rodents, as is reflected in their complex vocalizations and social systems.  Some are arboreal (red, gray, and fox squirrels), some are semifossorial - burrowers (prairie dogs, ground squirrels, woodchucks), and some do both (chipmunks).  Some hibernate (woodchucks), some enter deep torpor and feed occasionally on cached food supplies (chipmunks), and many are active year-round.   

Squirrels manufacture enzymes that enable them to eat the toxic amanita mushrooms.  However, like the pit viper that produces its venom and the skunk that produces its musk, the squirrels produce these enzymes at a physiological cost, thus “balancing” the benefit.  

The front foot (hand) has four toes and the rear foot  has five toes.

Worldwide, the 51 genera and 272 species of the squirrel family includes marmots, woodchucks, prairie dogs, ground squirrels, chipmunks and tree squirrels (Nowak's Walker's Mammals of the World).  They range in size from 1/3 ounce of the pygmy squirrel, to 16 1/2 pounds of the marmots.

Of 8 genera and 68 species found in North America, seven species are represented in the Appalachian region (Jones' Checklist of North American Mammals). 


EASTERN CHIPMUNK (Tamias striatus) (treasurer, or storer; striped)

Appalachian Region Distribution: Throughout.
Continental Range: Eastern US and southern Canada.  Nine subspecies are recognized in eastern US, with three in the Appalachian region.
Abundance: Common
Population Density: 12 - 24/ acre; with great annual variation.  
Size and Molt: Head and body five to six inches; 2 ½ - 4 ½ oz. Two molts.
Mammae: Four pair.
Habitat: Open deciduous forests; especially rocky areas and forest/field bushy edges.
Active Period: Diurnal, with two peaks of daily activity, with a midday slump. Inactive in extremes of winter and summer. Whether chipmunks hibernate or not depends on your source and definition. They do not develop a layer of brown fat as does a true hibernator. Instead, they go through periods of torpor of variable length (rarely more than several days at a time) and frequency, normally starting in October or early November.   Not all animals in a population will become torpid.  During these periods of torpor, studies reveal chipmunk respiration rates drop from 60 to less than 20 breaths per minute, and temperature drops from 100 to 42 - 45 degrees. Chipmunks must wake about every two weeks to eat from their caches and defecate.  Will come out in mild weather any time.  (Mammal of Virginia states chipmunks have been seen in Blacksburg, VA at 15° F and three inches of snow on the ground.)  Individuals vary on length of torpidity, and can change from one year to the next.   Additionally, those of more northern habitat tend to spend more time “hibernating” while those of the south enter a torpid state only during severe winter weather.
Diet: Nuts primarily, with a preference for hickory. Also eats fruit, mushrooms, some invertebrates, annelids and arthropods (snails, worms, cicadas, and other insects). Chipmunks spend a lot of time caching food in their burrow, especially during fall. The food is used during waking time of winter's hibernation. Like the gray squirrel, the chipmunk disperses its food caches to deal with theft.  It has a primary storage site in its burrow and numerous smaller scatter hoards well hid throughout its territory.  As other chipmunks raid the primary storage site, the owner will refill it from the scatter sites.  If no thievery occurs, the scatter hoards often go unused, being left to rot or sprout. Carries food in its cheek pouches. One chipmunk was found to be carrying 70 sunflower seeds in its cheek pouches.  Another; 32 beechnuts.
Home Range: ½ - 2 acres, overlapping with others, with a smaller defended territory. Ranges may shift annually for food, winter protection, or for finding mates. Males have larger ranges than females. Within the home range is a protected territory, about 45 feet radius from the burrow opening.   Juveniles typically establish their own burrow systems near their birthplaces.    A PA study found a home range of .27 acre, with a 0.05 acre range when seeds were ripe.  An Adirondacks report was 0.5 - 1 acre.
Social Structure: Solitary and territorial mammals, although known to overwinter in small family groups. While ranges overlap, adults will strongly defend its smaller territory in order to protect it’s food caches and during breeding time.   There are no lasting pair bonds.
Life Cycle: Under favorable conditions, two seasonal breeding periods occur; with birthing in April and July or August, with usually four to five per litter. The second birthing period often consists of last years' young that failed to breed in the spring, although a number of chipmunks breed in both spring and summer, unusual in hibernators. This second birthing period does not appear in northern populations.  Females are in estrus for 3 to 10 days.  Gestation period of 31 days, weaned at 5 to 7 weeks, at which time they first leave the nest. At about 8 weeks, the female refuses to let the young re-enter the burrow and the young are on their own. This dispersal may be abrupt or take a week or two. At this time, adults will make continuous "chirps" near their burrows, perhaps advertising their own territories. Chipmunks can be sexually mature at approximately three months.  However, most individuals do not breed for the first time until they are one year old.  Life span of two to three years, although records of up to 13 years in captivity have been recorded .
Dens/Nest: Burrows fall into two categories.  They can be simple with one or two tunnels used for a hideaway or for food storage, or quite extensive with over 100 feet of tunnels (average is about 12 feet in length) used for nesting and storage.  Extensive tunnel systems include a ten inch diameter nest chamber and separate chambers for food caches approximately 3 feet underground. Surface opening is often straight down and 1 1/2 - 2 inches diameter. Openings are often plugged with soil each night and during the winter. Two side tunnels and openings may exist, but are often plugged and not used. May occupy the same burrow system for life. Burrows are not shared except briefly by the mother and young. Tree cavities are occasionally used.
Tracks: Four front and five hind toes approximately ½ to 5/8" wide and a long, depending on depth of imprint. Not common in winter since often will be hibernating. Straddle is 2 - 3". Holes of 2 inch diameter leading straight down with no trace of excavated soil is the diagnostic home of the chipmunk.
Scat: Small piles of four to six rice-sized droppings.

Remarks: Runs with bushy tail straight up. Perhaps best identified by it’s sharp chuck-chuck call in the woods as hikers pass. Well-developed cheek pouches stores have been known to hold up to 32 beechnuts or 70 sunflower seeds. Food caches also are kept throughout their range. Has been reported to overwinter with rattlesnakes, purportedly sharing the snake’s warmth. Understandably, the chipmunk enters the den after the snakes have become torpid, and exit in the early spring, before the snakes lose their lethargy.  However, chipmunks are a common prey of rattlesnakes.  The name chipmunk is of native Indian origin and probably relates to its chipping call.

Chipmunks from western Maryland are paler in color, and those from the Carolinas brighter than those from other areas.  The darkest individuals are found in the southern mountains, SW VA, Kentucky and Tennessee.   Both albinos and melanistic individuals are known.

There are twenty four species of chipmunks in North America (Nowak's Walker's Mammals of the World - the Checklist of North American Mammals lists 22 species).  There is one eastern species and 23 (or 21) western species.  The 1994 edition of Nowak's states the smaller western species and eastern species were formerly placed in one genus; Tamias, but recent evidence had caused the western species to be split into a separate genus, Eutamias.  Thus, at that time, the eastern chipmunk was the only species in the genus Tamias."  However, the 1999 edition of Nowak has the above two genera combined again in the same genus; Tamias.  It goes further to say, "There is general agreement that the issue is not settled and further investigation is needed on chipmunk systematics."  

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WOODCHUCK (Marmota monax) (marmot; solitary)

Appalachian Region Distribution: Throughout.
Continental Range: Most of Canada and eastern US.  Four recognized subspecies, with one in the study area.
Abundance: Common; in fact, more common than in pre-Colombian times.
Population Density: W/H says 1 per 11 acres, occasionally up to 1 per 2 acres. Nowak reports a Wisconsin average of 1/125 acres.  Other sources say 2-6/acres.    Populations tend to vary greatly year to year.  
Size and Molt: Head and body 12-20 inches; 5 - 12 pounds. Hibernation results in the loss of about 30% of fall body weight.  However, weight loss immediately after emergence from hibernation was much greater, with energy derived from remaining stored fat reserves.  Males tend to be slightly larger than females. One summer molt, taking 3 and ½ weeks, occurring from late May into September.
Mammae: Four pair. 
Habitat: Prefers open fields, but found in mature woods, pastures and hayfields.
Active Period: Primarily diurnal to crepuscular (morning and evening, like deer). Only spends one to three hours above ground each day during the summer. True hibernators, entering their dens in the first week of November  and emerging in early February (sometimes requiring help for Feb 2).  Populations further to the north may hibernate for up to four months, while more southern populations may only hibernate for six to eight weeks or not at all.  Winter activity has been documented, but is atypical.  Older and fatter woodchucks enter their hibernacula first and emerge first.
Diet: Herbivorous grazers.  Seasonal herbs and fruits, but known to taste grasshoppers, beetles and snails. Does not cache food.
Home Range: 2 - 5 acres.  Woodchucks exhibit little territoriality, except in the direct vicinity of their burrows.  Home range may shift from winter to summer den sites with corresponding shifts in home  range.  Same sexes do not generally overlap, but males may overlap 1-3 females.  Female ranges are generally smaller than males in the spring, but may become larger than males after young are born before contracting again in the fall.  
Social Structure: Solitary, but may live in small family groups with a male dominance hierarchy. Woodchucks are not territorial, relying on a dominance/submission hierarchy to establish avoidance of dominants by submissives where home ranges overlap.  Burrows are occupied by a single male, a mother and her young, or occasionally, a male and a female.  Contrary to other marmots, the woodchuck is fairly aggressive.  This is a function of the length of the growing season.  The shorter growing season of the western marmots (60 days) requires more time together to assure ample breeding (the Olympic marmot shares its burrow with an adult male, two adult females and offspring from the previous two litters).  The eastern woodchuck can afford to be more picky (150 day growing season in PA).
Life Cycle: Mating shortly after spring emergence (late February  to early April), one litter of four to five is produced in April to mid-May.  Gestation period of 32 days. Weaned in six weeks and on their own (chased off by mommy dearest) at two months of age (July).  By fall, the young have established their own burrows.  Occasional young females will overwinter with the mother, like other ground squirrels.  10 to 25 percent may reproduce in the subsequent spring at one year of age.  Most will mate in their second or, more commonly, third year. Life span of four to six years. (Nowak reports 13-15 years).
Den/Nest: Will normally have both a summer and winter den. The winter den (with normally only one opening) is often situated in brushy or gently sloping wooded areas, whereas the summer den (with several openings) is in open, flat fields. Occasionally, the winter den is used year round and by succeeding generations. The burrow system includes nest chambers, and/or a hibernaculum chamber (both about 15" wide and 10" high, lined with leaves and grass), and a latrine chamber.  The summer system has a number of surface openings, easily identified by the excavated soil and a number of nests. In addition to the main opening, there may be up to five more or less obvious openings for emergency entrance or exit needs.  At least one of these hidden entrances will be a “plunge hole”; having a two foot drop from the surface.  Burrows may be four to five feet deep (reportedly up to 16 feet deep) and 25 to 50 feet long.  Other mammals will use woodchuck burrows (skunks, foxes, opossums, raccoons, rabbits, mice, and others).  A woodchuck burrow can be distinguished from others by the fresh pile of soil at the main entrance, since woodchucks clean out their burrows several times a week.  Unlike foxes, woodchucks will not defecate outside the entrance.  Rather, they use separate chambers in the burrow.
Tracks: Four toes on front feet, 1 ¾" by 2" long, and five hind toes, the front a little longer than the rear.  Running gait shows four feet tracks within 12" and 12" between sets of prints.
Scat: Quite variable, depending on diet.  Often 2 1/2" long by 1/2" wide.  Can be longer strings (if not separated) or more loose piles.  Fecal pellets are often deposited in a specific fecal chamber underground or buried in the loose soil excavated outside of the burrow.  This burying behavior is unknown in other squirrels.

Remarks: The common name comes from the Cree Indian word, wuchak.  The largest member of the squirrel family, with a flattened, bushy tail. Also, the most widely distributed and best known of all the marmots.  The only representative of a genus of circumpolar distribution. An accomplished climber and swimmer. True hibernators, with the oldest (and fattest) entering the burrow first (late October). In Autumn, after gaining 30% of it’s summer weight (mostly in the form of a half-inch layer of brown fat stored over much of the body, especially the back and shoulders), the ground hog will line it’s hibernaculum with grass and leaves and then plug the entrances (to maintain constant temperatures and to keep out curious visitors) before curling into a ball. During hibernation, heart rate drops from 100 to four beats per minute (W/H says 100 to 15), respiration rate drops to one breath per three or four minutes, and the body temperature drops from 98 to about 40 degrees (W/H says 96 to 47). Males usually exit a few days earlier than females in early March (similar to the hibernating jumping mice) (W/H says one month earlier). This allows males to compete for good home range and dominance over other males.  The period of hibernation is about three to four months, with each 10 miles northward adding one day of hibernation; given similar elevation.  Due to the shortened northern growing season, the emergence is much more precise than in the southern range, thus enabling the well-known "groundhogs day" of Punxsutawney PA on Feb 2.  In an eight-year period, the PA emergence date fell between January 29 and February 8, regardless of the weather.  They will awaken occasionally throughout the winter (about every two weeks) to defecate. This may occur in special chambers below ground or, less frequently, on the surface. They will emerge having lost approximately one half their fall weight (W/H says 20 to 37%).  Ground hogs have been observed every month of the year, even occasionally with snow on the ground and temperatures in the 20’s.

Melanistic (black) or erythristic (reddish-cinnamon) groundhogs are not uncommon. Albinos are uncommon. Is known to climb trees for fruit and is a good swimmer. It’s loud, shrill whistle gives it the common name "whistle pig" (call is made with vocal chords; thus, not truly a whistle).  At the beginning of European man's introduction, woodchucks were scarce.  As a result of human disturbances, the range and population of ground hogs have increased over the last 200 years.  

In Kentucky, between 1964 and 1971, some 267,500 ground hogs were taken each year!

There are six other North American species in the genus Marmota, all located in the west, called marmots. 

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EASTERN GRAY SQUIRREL (Sciurus carolinensis) (shadow tail; belonging to Carolina)

ARTICLES

Appalachian Region Distribution: Throughout.
Continental Range: East of the Great Plains and south of the Great Lakes. Basically, their range correlates with the eastern hardwood forests.   Five subspecies recognized in eastern US, with only one in the study area. 
Abundance: Common at lower elevations; rare at higher elevations. Populations reflect "mast" years (acorn supply).   Much higher numbers were found when the native American chestnut tree was common.                     
Population Density: Highly variable, with highest populations in late August and September.  Chapman and Feldhamer report four WVA state forest densities over a six year period ranging from 0.44-1.0, 0.24-0.84, 0.2-0.44, and 0.32-1.4 per acre.  Our friend, Dr. Vagn Flyger of Silver Spring, MD, found a range in his suburban yard backing up to the Northwest Branch in Quaint Acres from 0.8-4.0 per acre over a three year period.   2-20/acre in some studies, others report 0.1-6/acre; max up to 20 / acre (public park densities can exceed 50 per acre).  
Size and Molt: Head and body is 9 to11 inches; ¾ - 1 ¾ poundsSexes are alike in size and color.  Two molts; paler winter pelage, although the tail molts only once in mid summer.  Northern individuals are larger than southern populations.  
Mammae: Four pair.
Habitat: Mature oak and hickory hardwood forests.  Land use may be highest in forested river bottomlands, river valleys and swamp hardwoods.  Extremely adaptable, they are found anywhere, except spruce/fir forests, where red squirrels dominate.  Gray squirrels tend to prefer more dense woods than fox squirrels.  The more large trees an area has, the more gray squirrels will inhabit the area (larger cavities and more food).
Active Period: Diurnal in winter to crepuscular in summer, non-hibernating, primarily arboreal mammals.  Active year-round, but will will remain in their nests in very cold or stormy weather until food forces them out.  W/H states that females are more active in spring and summer, with males being more active in autumn and winter.  Most active in September and early October, burying acorns (this is also the peak time for their arboreal activity).  Additionally, spring born (and some adult) fox and gray squirrels frequently move to new locations at this time.  This dispersal period is also referred to as the “fall reshuffle”, with movement of 6 or more miles.  As noted in the remarks below, this fall reshuffle, in conjunction with a failed hard mast season, can result in massive fall migrations, more appropriately called movements.
Diet: Nuts are the mainstay, with buds, flowers, and roots in spring; maple samaras, fruits, berries, mushrooms, and insects in the summer; nuts and fungi in fall and winter (specifically oak and hickory nuts and maple samaras). They usually feed on just one food at a time, changing the item as additional sources come along.  They are known to eat bark of dead trees for the fungal mycelia within.  Shedd even notes gray squirrels are known to eat bird eggs and frogs.  In February and March, they chew twigs of maples, oaks and pine. Winter food storages, or caches, consists of each acorn buried separately, decreasing the risk of losing an entire stash.  Adults obtain minerals from bones, antlers and turtle shells.  See Articles above for more information.
Home Range: Most studies indicate one to four acres, up to ten acres (varies with food availability), larger for males than females.  Chapman and Feldhammer report 2 acres for males and 1.25 acres for females in a Maryland study (Flyger).  Nowak reports 1.3 acres for males and 1 acre for females in a Virginia study.  Male home range will overlap that of several females.  Overlaps other squirrel home range.  Range of gray squirrels are smaller than the fox squirrel.  Basically, their home range focuses on (and around) one nest tree, shifting as  food sources become available.
Social Structure: Usually solitary, shy and non-aggressive.    During the mating season, males will be territorial and exhibit antagonism toward one another.  Males are polygamous.  The same applies to females on a nest.  The male provides no parental care.  Mammals of Virginia say gray squirrels are gregarious and often congregate in considerable numbers, while W/H says they  are not social, only gathering in winter denning situations..   Chapman and Feldhamer says both fox and gray are relatively nonaggressive toward either their own species or the other species, but may share dens with a number of individuals of their own species (Shedd says only males and juveniles will den together in winter, with females – probably pregnant - being cantankerous and denning by themselves).  Wilson says at night and during bad weather, as many as seven or eight squirrels may occupy a communal den, where they will groom each other and conserve heat during cold winter nights.  Among squirrels with overlapping home range, especially at high densities, there is an established dominance hierarchy, with the oldest (male or female) normally assuming dominance (W/H says males over females, adults over juveniles, and residents over immigrants).  Dominance (i.e., aggression) appears at mating time, food concentrations (including bird feeders), and at den trees. 
Life Cycle: Normally two litters of about three (1-9) per litter per year (parturition about March and August).  Forsyth says only 20 to 40% have two litters.  Gestation period of 45 days, weaned in two months; about the same time they first leave the nest.  Breeding can begin as early as December.  Males play no role in child-rearing.  The spring brood stays with the mother until late summer, and the second brood will often stay with the mother over winter. Young make leaf nests at about 18 weeks of age.  Young breed the next year (may only have one litter).  Life span has been up to 15 years, although 6 years is more the maximum. Wilson says more like one year for the average.  One captured animal lived for 23 years.
Dens/Nest: Winter and summer nests are made.  Will make winter nests in tree cavities and summer leaf nests in branches (occasionally a winter leaf nest is maintained).  A favored den tree may be used for many years, but an animal always has several other nests where it can escape enemies.  The tree cavity must be 12 inches deep and have an opening at least 3 inches in diameter.  Often, several leaf nests (called dreys) are built around different food sources. Gray squirrels average 1 1/2 to 2 leaf nests per squirrel. The leafy nest is composed of leaves and twigs forming a water-resistant form lined on the inside with moss, grass, and shredded bark. The main entrance is positioned to face the main tree trunk.  The winter nest is high off the ground in a tree hollow, lined with vegetation.  Normally, the first brood is had in the tree cavity with the second brood in a leaf nest.
Tracks: Has five toes on hind feet (2 5/8" l x 1 1/4" w) and four on the front feet ( 2" l x 3/8" w).  Straddle is 4 to 5". Running stride of 16 - 36".  Front and rear tracks are often lined like a square, so it can give the appearance of two question marks (!!), with the front tracks behind the rear tracks (or somewhat behind and between the rear tracks) .
Scat: Pea-sized droppings (1/4 – 3/8” x 3/16”).

Remarks: Can be blonde, black (melanistic), erythristic (reddish), or albinos (Olney, IL is famous for its protected albinos - Trenton, NJ and Greenwood, SC also have albino populations). Black squirrels are more common in the northern portions of their range.  Gray squirrels from MD and the western parts of VA and NC are slightly darker and larger than those from eastern VA and most of the Carolinas. It is common for one of the color variants to be dominant in an urban setting, since the predatory natural selection (culling out) of these colored aberrants does not exist.

Melanistic gray squirrels of the Washington DC area owe their existence to two shipments of black squirrels from Ontario being released in the National Zoological Park in 1906.  It appears that the dark pelage gives them an advantage over the gray winter coats of the normal squirrels in winter through energy savings due to the additional absorbed solar heat (lower heat loss and lower basal metabolic rates).

Formerly known to make massive movements in search of food.  Ernest Thompson Seton estimated one mass movement at more than one billion individuals in 1920.   Such migrations are necessitated by local high populations and erratic annual acorn mast crops.  A more recent mass movements on a reduced scale was reported in October of 1968 in the southern Appalachians of TN, GE and NC.  This followed a mast crop the previous year, resulting in  an abundance of young squirrels.  Unfortunately, 1968 was a poor mast year, resulting in a major food shortage.   Such a phenomenon is incorrectly called a migration, when it actually is nothing more than a big fall reshuffle.

Caching of seeds and nuts ensure tree generation. Nuts are buried separately, about 1/4" to an inch deep and are recovered by any squirrel that may smell them (one study found 85% of buried nuts were found over the winter). Nuts buried by scientists were recovered at the same rate as nuts buried by the squirrels, indicating that memory is not involved in nut recovery.  The nuts can be smelled when buried under a foot of snow.  A thicker pelage and layer of fat help insulate the squirrel in winter. While fox squirrels and gray squirrels inhabit similar habitats and share common foods, they are rarely found in the same area (mutually exclusive, or allopatric) and do not interbreed. Fox squirrels prefer the more open woods, the gray squirrels in the more dense woods. When 10% of the land is wooded, both species seem to be equally abundant.  When 70% of the land is wooded, fox squirrels are absent.  Gray squirrels are a major prey species of timber rattlesnakes. In the late summer, swellings under the skin are signs of the developing larvae of the parasitic botfly, which emerges in the fall. They do not seriously endanger the squirrel. 

Due to the extraordinary amount of tannins in their systems afforded the gray and fox squirrels by their acorn digestion, they are unusual in rarely having either tapeworms or roundworms, which is a deadly poison to these parasites.  

Much more abundant a century ago.  Gypsy moth defoliation and subsequent loss of acorns in the 1980's practically wiped out squirrel populations in southern PA.  Important game species with 40 million "harvested" each year in the US.  In Kentucky, an average of 1,309,000 gray and fox squirrels were taken annually from 1964 to 1971.

There are seven species of the genus Sciurus in North America.

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EASTERN FOX SQUIRREL (Sciurus niger) (shadow tail; dark)

Appalachian Region Distribution: Throughout.  
Continental Range: East of the Great Plains and from PA south.  The range extends slightly farther west than that of the gray squirrel but not as far northward.  W/H says to the west of a line from Harrisburg PA, Harpers Ferry WV, Roanoke VA, and along the Blue Ridge to the Smokies, with a disjunct subspecies found on the eastern shore, although other sources show  the fox squirrel ranging throughout all SE US.  However, declining populations are found in the piedmont and coastal plains regions of both Virginia and North Carolina.  Eight subspecies recognized in the eastern US, with two subspecies found in the study area.. (The well-known federally-designated endangered species, the Delmarva fox squirrel; Sciurus niger cinereus, is one of these eight subspecies.  It is only found on the Delmarva peninsula of Virginia and Maryland.)
Abundance: Uncommon in central PA and the piedmont and mainland coastal plain of VA and NC, common to the west of these locations, but with declining numbers.  Due to the decrease in the open, mature forest habitat in the east preferred by fox squirrels, their numbers are declining and often confined to small and shrinking localities.
Population Density: Highly variable.  Reported densities of 0.4 - 2/acre, up to nearly 10 per acre.  Weigl, working with piedmont and coastal plain populations in North Carolina, found densities averaging 0.002/acre.  
Size and Molt: Head and body 10 to 15 inches; 1 ¼ - 3 pounds.  (Weigl's NC piedmont and coastal plain populations averaged 2.2 lbs.)  Sexes are alike in size and color.  Largest tree squirrel in the Western Hemisphere.   Chapman and Feldhamer says two molts, with one tail molt.  Nowak says there are two molts per year in some,  perhaps all, species (of the scuirus genus), but the tail fur is shed only once yearly.  W/H, PA Mammals, says one molt in late spring.  Twice the size of gray squirrels.
Mammae: Four pair.
Habitat: Fox squirrels prefer higher ground and larger trees than gray squirrels.  Open hardwood forests and pine woods where occasional fires promote grassy surface growth are ideal. Will not be found in the closed forests of the gray squirrel.
Active Period: Crepuscular in summer, diurnal in winter; non-hibernating, they start foraging later in the morning than gray squirrels. Tends to aestivate (summer dormancy) in summer if hot.  Most active in summer in light rain or just after rains (cooler temperatures, nuts and fungi are easier to smell, and the ground is easier to dig).  Most active in September and early October, burying acorns.  Additionally, fox and gray squirrels frequently move to new locations at this time.  This dispersal period is also referred to as the “fall reshuffle”, with movement of over 6 or more miles.  However, mass movements by fox squirrels have never been as massive as gray squirrels.
Diet: No apparent difference is observed between the gray and fox squirrels.  Preference is for hickory and oak acorns.  Pine seeds from cones; green or mature and the cambium tree layer are preferred in some habitats. Also buds and berries, fungi, insects, bird eggs, and corn.  Nuts and seeds are buried individually or two or three together like gray squirrels, versus the caching done by red squirrels.  Early summer may be the hardest time for fox squirrels to find food, especially in pine forests.  The buds, flowers and young fruits of tulip trees are heavily foraged when available.  Studies on recovery of cached acorns reveal tremendous range from 33% to 99%.  Succulent vegetation will normally satisfy their moisture requirements. Water is utilized if present, but the lack of it is not a limiting factor.
Home Range: Varies by densities, food supplies, and habitat quality.  7 - 24 acres and greater are commonly reported.  Weigl's NC piedmont and coastal plains populations had home ranges estimates of 66 and 42 acres, or 107 and 62 acres, respectively for males and females.  A study of a Florida population revealed a home range of 106 and 41 acres for males and females.  This is larger than the gray squirrel, not just due to size, but also the relative paucity of food in the preferred more open woods habitat.  Home range may overlap.
Social Structure: Solitary, except occasionally to feed in common areas and share winter dens among family members.
Life Cycle: Similar to gray squirrels (but breeds slightly earlier), with two litters (from two year olds) with an average of two or three (1-6) per litter per year. (Weigl's NC piedmont and coastal plain populations revealed no evidence of two litters per year.)  If  a female produces 2 litters per year, they may produce none the next year if food conditions are bad.  Birthing usually occurs in  March and again in July, although pregnant females have been found every month of the year.  Weaned, able to eat solid food in eight weeks, and soon thereafter are independent.  Gestation of 44 days.  Sexual maturity in one year (first year females usually having only one brood).  Will disperse a distance of 9 miles or more from their birthing site.  Life span up to 12 years.
Dens/Nest: Prefers hollows in trees for winter and natal nests, but will make large leaf nests in summer (and winter if tree cavities are scarce). Cavity openings are 3 inches wide with a hollow of six inches wide by sixteen inches deep. Outside leaf nests vary greatly, averaging 20 inches in diameter. Like gray squirrels, the opening faces the trunk. Winter nests have an outer layer of twigs with leaves attached, a series of inner layers of damp leaves pressed together, with a lining of shredded bark and leaf fragments.  A leaf nest may be used for several years.  Fox squirrels average about three to six active leaf nests per individual (one study reported in Nowak found an average of nine nests were used per year).  Platforms, made just for sitting, are sometimes made.
Tracks: Four-toed front feet are 1 and 1/2" and five-toed rear feet are 2" in length.  While running/hopping, the front foot prints are parallel and behind the rear feet.  Straddle is 4 1/2" with 2' stride.  Food debris is often found scattered about the base of a tree used as a feeding perch by fox squirrels.  
Scat: Pea-sized droppings.

Remarks: Fox squirrels get their common name from their fox-like tails.  Species name comes from southern subspecies, which is much darker than other regional populations.   

The populations of the Appalachians and to the west differ from the eastern populations in being smaller (~2 lbs) and more consistently reddish in color.  The eastern populations of the piedmont and coastal plains averaged more like 3 pounds in size and were quite variable in coloration, being silver, gray, black and gold, often with black masks and with distinct white marking on the nose, ears, and feet.  Additionally, western populations inhabited deciduous forests while the eastern populations occupied mature pine-oak woodlands.  Weigl and others suggest that the southeastern and western fox squirrel populations evolved in isolation in separate refugia during the Pleistocene, eventually colonizing different regions to the north and subsequently establishing variable zones of overlap and interbreeding.

Fox squirrels have the widest range of colors of any other mammal.  Fox squirrels can be found in several dorsal (back) color morphs (subspecies); brown, brownish orange, gray-black, blackish brown, steel gray, or all black (color variations tend to match local habitat - yellow midwestern squirrels blend with the yellows of the hickories in fall, black of southeastern squirrels match periodic burning of the pine forests, etc.- its just someones' thought). Along the coastal area of Virginia and Delaware (S. niger cinereus) the population is colored steel gray with no tawny. The ventral (belly) can also vary from creamy yellow to rusty orange to black.  Fox squirrels have a darker head (than gray squirrels), some white on the muzzle and white on the back of the ears, and a larger, more fluffy tail than the gray squirrel.

More terrestrial than gray squirrels (less agile in trees than gray squirrels).  Fox squirrels spend more time foraging and running about on the ground than do gray squirrels and will more often run on the ground between trees while grays will jump from one tree branch to another tree branch.  Wilson says fox squirrels are more “easy going”, getting up later in the morning and turning in earlier in the evening.  Fox squirrels have home ranges of up to ten times the size of grays, due to the more open (poorer food supply) habitat.

Fox squirrel bones are pink (due to the accumulation of porphyrin compounds); gray squirrel bones are white. Of the three tree squirrels, the fox is the quietest, the gray more vocal, the red squirrel, the most vocal. Mass movements, well-documented among gray squirrels, are not common, nor as large, among the fox squirrels.

Due to the extraordinary amount of tannins in their systems afforded the gray and fox squirrels by their acorn digestion, which is a deadly poison to tapeworms or roundworms, they are unusual in rarely having either of these parasites.

An account by Seton (1953) states; Arlington Cemetery had always been a haven of fox-squirrels.  About 20 years ago, they had increased to surprising numbers.  Then one day, they seemed to be possessed of a migration craze; they all set out eastward.  At once, they were met by the broad Potomac; but plunged in, swimming away toward Analostan Island (Roosevelt Island), the nearest wooded tract.  Here many of them stayed; but many moved on, and were lost sight of.  They still frequent Arlington Cemetery.

Approximately 18 Fox-squirrels were released in the National Zoological Park from 1899 to 1916 from various states across the country (see remarks for gray squirrels above).  

When Audubon and Bachman were conducting their research for the Quadrapeds of North America, they noted that in South Carolina the fox squirrel "takes possession of the deserted hole of the ivory-billed woodpecker."

In Kentucky, an average of 1,309,000 gray and fox squirrels were taken annually from 1964 to 1971.

There are five species of the genus Sciurus in North America.

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RED SQUIRREL (Tamiasciurus hudsonicus) (steward, shade tail; the Hudson Bay)

Appalachian Region Distribution: Throughout.
Continental Range: A boreal species of the north across Canada, south along the Rockies to southern Arizona and New Mexico, and down the Appalachians to Smokies.  Five subspecies recognized in eastern US, with two found in the study area.  In fact, one of the widest distributions of any North American squirrel.
Abundance: Common, especially in conifer forests.
Population Density: 1-3 / acre. W/H says up to18 per acre with great seasonal variability.  Wilson notes up to 17 – 20 per acre is possible in good habitat.  Nowak reports from 2 - 11 per acre.  Population densities reach a high every three to four years in the Adirondacks of New York.
Size and Molt: Head and body 7 to 9 inches; 5 to 9 oz.  No differences in size or color by sex.  Two molts, although, the tail only molts once in the middle of summer.  
Mammae: Four pair.
Habitat: Highly arboreal.  Apparently, it originated in coniferous forests, but now is found also in mixed hardwood forests and swamps of higher elevations.  In southern limits of its range (southern Appalachians), the red squirrel is found in the same areas as gray and fox squirrels, with resultant competition for food and den sites.
Active Period: Diurnal in winter, crepuscular in summer, spending much time on the ground as well as arboreally; year-around.  Red squirrels do not hibernate, but may spend several days in their dens during inclement winter weather. 
Diet: Quite diverse diet (more omnivorous than the gray squirrel).  Due to high metabolism, a high energy content diet is required.  Some brown fat is present in winter.  Most of the energy is derived from the hemlock, pine and other conifer cones, which are stored in mass food caches ("middens") if not eaten on the spot.  Also acorns, hickory and beechnuts, tulip tree and sycamore seeds, when seasonally available, buds, twigs, occasional insects and mushrooms (including the very poisonous amanita; aka, destroying angel or fly agaric).  In the Smokies, fruits of the cucumber magnolia, mountain holly, silverbell, beech, buckeye, serviceberry, black walnut, American chestnut, and blackberry; seeds of mountain maples, hemlock, and pines; cones of fir and spruce; mushrooms; the buds of the rosebay rhododendron and the buckeye and roadside garbage cans are eaten.  Fungi are cut and placed in trees to dry for later winter eating, or later cached in middens.  Terminal evergreen buds are a main stay in winter diets.  Have been known to eat young birds and bird eggs.  Unlike the single nut burial philosophy of the gray and fox squirrels, the red squirrel goes for middens, used for winter food, which is vigorously defended. These middens (made up primarily of pine cones) may grow over the years to amass several bushels of nuts. Red squirrels can afford to make this middens where they only have other red squirrels to defend against.  However, in regions cohabited with gray squirrels, red squirrels must scatter their hordes, since gray squirrels are not intimidated by red squirrels’ threats.  Hole eaten into nut is irregular in shape.  They have favorite feeding sites, as opposed to gray squirrels, that pretty much eat their acorns where they find them.  The size of this debris pile is measured in bushels and varies with conifer species, seeds, proportion of diet, forest age, squirrel population and the feeding place, but can be up to 20 feet by 12 feet and 3 feet deep.  Red squirrels are also known to "tap" maple trees for sap. 
Home Range:  2 - 5 acres, with no overlap of range (unlike gray squirrels), but only 0.5 to 3 acres defended in the area of the nest and prime feeding areas.  In high quality habitat, red squirrels have been observed to be non-territorial.  
Social Structure: Solitary. Very territorial and aggressive in protecting their core home range (1/2 to 2 acres), especially in late summer when juveniles are dispersing.  Females allow males into their territories in late winter for mating.  Not known for communal denning in winter like gray squirrels. 
Life Cycle:  One or two litters per year (April - May) with an average of five or six (1-7) per litter. Females are receptive for only one day, at which time, she allows males to enter her territory.  Gestation period of 35 days.  A second litter is had in August in good seasons. Weaning at 9 - 11 weeks of age (W/H says 6 to 7 weeks, Novak says 7 - 8 weeks). (PA mammals says weaning occurs after the female makes a nest on the periphery of her range for her young. W/H says the female is known to occasionally leave the area to the young, who then subdivide it.)  Spring young disperse at around 18 weeks (September).   Second brood overwinters with the mother. Young are sexually mature the following season.  Life span two or three years common, with ten years known in captivity.  Only 5% of a cohort live beyond five years.
Dens/Nest: A winter and summer nest is common.  Tree cavities are preferred for winter nests, while leaf nests (~15 feet high) in summer are common (about the size of a basketball, often made of grapevine bark, while gray squirrel nests are larger and coarser), especially in conifer woods, where cavities are less numerous.  Will also build on abandoned bird nests or make use of underground dens.  Often, a second winter nest is made in the form of a weather-tight structure located in the densest foliage of a tree.  Can tunnel and make nests in the ground under rocks or stumps in the winter. Burrow holes are 2 ½ to 3 inches in diameter, often with cone remains near the entrance.
Tracks: Four front toes (1/2" l x 3/8" w) and five rear toes (7/8" l x 5/8"w).  Straddle is 3 to 4".  Running stride is 9 - 30".  In winter, makes extensive runways through the snow. 
Scat: Small, 1/4" - 1/2" elongated droppings.

Remarks: Known in WV as the "fairy diddle".  A northern species, the red squirrel is smaller than the gray squirrel, with somewhat different summer and winter pelages. In summer, a lateral black stripe separates the dorsal brown from the white ventral fur. In the winter, although the black stripe disappears, the back and tail is rusty red and tufts of hair appear on the ears. Red squirrels are distinctly darker on the head and sides in the spruce and fir forests of the higher southern Allegheny Mountains.  Both albinism and melanism is known among red squirrels.  Very vocal and active; usually heard before seen. Smaller than the gray and fox squirrels. Vocally protects it’s territory, many hunters have had their hiding place announced by the red squirrel.  Noted for its food caches, which may accumulate in size over several years, usually located in hollow trees. The seed caches (middens) are favorite sources of high quality seed used by foresters, which are cleaned and used in nursery propagation.  

Hickory nuts eaten by red squirrels will have a ragged hole opening.  Flying squirrels have smooth edges of the opening, while white-footed and deer mice will have several openings.  Gray and fox squirrels crush the nut.

The western Douglas squirrel is the only other species of this genus in North America.  Both species are known as pine squirrel, mountain boomer, chickaree, or fairydiddle.

Several million red squirrels are trapped each year for the fur trade.  

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NORTHERN FLYING SQUIRREL (Glaucomys sabrinus) (gray, mouse; Severn River)

Appalachian Region Distribution: High elevations (> 2,680') of the Appalachian Mtns. as far south as the Smokies. Can be found continuously from Maine to central PA.  South of central PA, only two disjunct relic populations exists.  Both, by their isolation, are recognized as subspecies and designated as Federally listed Endangered Species.  In six eastern WV counties (Greenbrier, Pendleton, Pocahontas, Randolph, Tucker and Webster) and Highland County, VA, the subspecies Glaucomys sabrinus fuscus was first described in 1936.  The type locality is Mill Point, Cranberry River, Pocahontas County.  The second subspecies, Glaucomys sabrinus coloratus, first described in 1953, is found in SW Virginia near Mount Rogers, VA (Whitetop Mountain) and along the Appalachian Mountains of the North Carolina/Tennessee border through the Smokies as far south as Georgia.   Populations identified for study by Weigl included Roan Mountain, Bald Knob Ridge, Grandfather Mountain, the Black Mountains (including Craggy Mountains), and the Great Smoky Mountains, among others.  For the first time since 1958, three northern flying squirrels were found in the Smokies in 1987 on Clingman's Dome. The type locality is Bald Knob, 5,000 feet south of the summit, Mt. Mitchell, NC.
Continental Range: Throughout Canada, extending south along the Rockies and the Appalachians.  Four subspecies recognized in eastern US, with the two Endangered Species mentioned above and a third found in PA making up the population of the Appalachian region. 
Abundance: Becoming rare - listed federally as Endangered in VA, NC and WV.  In WV,  the characteristic spruce-fir and northern hardwood habitat of the G. s. fuscus subspecies is relatively abundant, potentially more continuous and at lower elevations, thus suggesting some potential for sustained populations.  The subspecies G. s. coloratus, resides only on high elevation ridges and peaks, and thus occupies habitat islands which are both highly disjunct and of limited size.   It is very limited from southern Virginia to the Smokies due to the more aggressive southern flying squirrel (and, to some degree, a parasite that is transmitted by the southern flying squirrel).  This limited range, along with the possible impact of warming trends, recent droughts and high elevation loss of conifers from introduced insects and acid-rain impacts, suggests a more vulnerable status of this southern subspecies.   These disjunct populations are relics of former broader populations that existed in historically cooler climatic conditions.  Populations have been shown to fluctuate, with peaks reached in the late 1980's in areas studied in WV, VA and NC.
Population Density: <1 to 4/acre.
Size and Molt: Head and body 5 ½ to 6 2/5 inches; 2 - 4 ½ oz. (Weigl's study populations in NC and TN had a mean head and body length of 6" and weight of  3.85 oz.  These are the subspecies G. s. coloratus.)  Females are generally larger than males.  The northern flying squirrel is slightly larger than the southern flying squirrel. (In addition, the northern subspecies, G. s fuscus, is smaller with a more muted pelage than the southern subspecies, G. s. coloratus.  One molt.  Shedd says two molts, unlike the southern flying squirrel.  Shedd also says the northern’s tail molts only once, like the gray, red and southern flying squirrel.  
Mammae: Four pair.
Habitat: In the southern Appalachians, a mixed ecotone of dense coniferous forests and northern hardwood forests (birch/beech/maple/cherry) are favored habitat.  The WV subspecies (G. s. fuscus) is generally limited to habitat over 3,000', while the more southern G. s. coloratus is most commonly found at elevations over 4,500'.  Habitat in this Appalachian region is limited by extent of  spruce forests and competition with the southern flying squirrel.  See remarks below.
Active Period: Nocturnal, primarily in the evening and just before sunrise. Active year-round.  It is known to withstand cold and wet conditions lethal to red squirrels and southern flying squirrels.  Has been found foraging on the forest floor even to -24
° F (Weigl reports -4° F).  However, Weigl found that when inactive and exposed to cold temperatures, the northern flying squirrel will drop it's temperature from ~98 to 86° F and assume a curled position.  Red squirrels under similar conditions are not able to reduce their body temperatures, and thus are not able to survive similar conditions without adequate store.  Southern flying squirrels show an intermediate response, with a slightly reduced metabolism and body temperature.  Weigl notes this reduced metabolism and body temperature is not torpor, since the northern flying squirrel is capable of immediate activity at these reduced levels.
Diet: Somewhat more omnivorous than the southern flying squirrel.  Lichens and fungi, including mycorrhizal fungi (which is a major source of minerals, such as sodium and phosphorus, as well as energy), can constitute a major part of their diet .  The staminate cones of conifers are a major part of the spring diet.  Nuts are eaten, however, less so than the southern flying squirrel due to the absence of oaks and hickories in its range. Also eats buds, fruit, various seeds and a number of insects (beetles and moths). They are known to raid the middens of nocturnal red squirrels during the night.  Will eat meat (young rodents, birds and bird eggs, but less than southern flying squirrel); are known for stealing bait. Creation of food caches in the summer for winter use is contradicted by various authors (as a minimum, the behavior is not as common as with the southern flying squirrel).  It has been found that many of the mycorrhizal forms of fungi that provide food for the forest trees, depend on the dispersal of fungal spores through the defecation of these squirrels.  In fact,  a northern California study correlated the fungal biomass positively with the numbers of northern flying squirrels.
Home Range: 2 ½ - 7 ½ acres. (W/H says 5 - 19 acres or more.) One North Carolina/ Pennsylvania study reported a home range of about 5 acres.  Weigl's North Carolina work found ten squirrels had a mean of 23 acres, with a winter range of 29 acres and a summer range of 15 acres, possibly a function of remote conifer food sources and mate-finding scenarios.
Social Structure: Very gregarious, although not as much so as the southern flying squirrel.  Weigl's study found 33 instances of nest sharing, averaging 2.8 per nest (2-6).  Most groups consisted of only two individuals (13 out of 33) and 15 groups were composed of adults of both sexes.  About equal numbers of all male or all females groups (7 and 4 respectively) were observed.  In groups of three or more involving juveniles, the adult was always a female.  Males do not help in child care.
Life Cycle: Usually one litter with two or four (1-6) per litter per year.  Parturition from April through September, after a gestation period of 37 - 40 days. Weaned in eight to nine weeks, often spending the first winter with the mother, normally breeding the following spring.  Life span 3 to 5 years, with 13-14 years in captivity.
Den/Nest: Prefers woodpecker holes or tree cavities, although even witch’s brooms and mats of moss have been used in northern regions.  Will also use leaf nests (dreys) in spruce trees.  (Being larger than the southern flying squirrel, it must make more leaf nests, since less suitable trees can be found.)  Also uses old crow or red squirrel nests.   Weigl found 12 summer nests included 7 cavities and 5 dreys, with 27 winter nests including 15 cavities and 12 dreys.  Urban (1988) found an almost exclusive reliance on drey nests in WV.  Nests are covered with firmly matted spruce twigs, lined with shredded yellow birch bark, lichens, grasses, sedges, and moss.  Outside nests are 8" - 10" in diameter, and 6" - 8" deep, usually high (14-28') in red spruce (exclusively spruce for Weigl's study).  A single 2" opening leads to the nest.  Weigl found an average of 3.25 nests per squirrel were used.  70% of Weigl's nests were found within 300' of the coniferous/hardwood borders.  A third category of nest is recognized as a retreat nest, of marginal nest quality and serving as sheltered breeding stations, food storage areas or defecatoria.
Tracks: The give away is the "sitzmark", or landing spot, with the pattern of feet leaving the site, much like a gray squirrel, only smaller (rear foot 1 5/8" in length, with a straddle of 3 3/4 inches).
Scat: Indistinct piles, each dropping less than 1/4" in length.

Remarks: The northern and southern flying squirrels are the only two species in the genus.  The northern flying squirrel is much larger and heavily built than the southern flying squirrel. Belly hairs basically white with gray at the base (but occasionally white throughout), while the southern flying squirrel belly hairs are a solid white.  Individuals from the southern Appalachians tend to be darker than northern individuals. Mean average glide is 66 feet, although flights up to 270 feet are recorded (the distance is more a function of height; the glide course is about three horizontal feet for every vertical foot drop).  Their tail allows them a good deal of maneuverability.  In fact, flying squirrels have been observed turning more than 180° during a glide.   Less vocal than the southern flying squirrel.  

The northern and southern flying squirrel are generally not sympatric (occurring in the same area; thus, they are allopatric); the smaller southern flying squirrel being more aggressive and tending to displace the larger northern flying squirrel.  Where they overlap, populations of both are limited and unstable.  Overlap areas generally support one species or the other.  However, where there are good populations of the northern flying squirrel, red squirrel populations are almost always abundant.   There is also evidence of a nematode parasite of the southern flying squirrel that is lethal or debilitating to the northern flying squirrel.  When northern flying squirrels come down from their mountain habitats and come into contact with the southern flying squirrel, it would very likely pick up the parasite and perhaps suffer either sickness or at least a competitive disadvantage.  On the other hand, when southern flying squirrel populations invade the spruce forest habitats of the northern flying squirrel, it cannot store adequate winter reserves of acorns to enable it to competitively survive the winters.  Still, another factor comes into play regarding the nematode.  The staminate spruce cones that make up a significant portion of the northern flying squirrel's spring diet are filled with highly aromatic oils that are known to have some potential as "vermifuges".  It is possible that these cones provide the northern flying squirrel with some protection in the high elevation spruce forests which is lost when it leaves the coniferous forests for the lower elevation oak forests.  Interestingly, it has also been found that the success of the nematode is severely limited at colder temperatures.

With southern flying squirrels found at the highest elevations in Virginia, it does not look good for the future of the northern flying squirrel in Virginia.  

The introduced balsam wooly adelgid will have a major impact on these squirrels by the loss of the nesting sites and food source.  High concentrations of heavy metals found at high elevations in NC (lead, copper, zinc, manganese) can also be harmful to these squirrels.

At night, the eyes of the flying squirrel will shine red by the light of a flashlight.

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SOUTHERN FLYING SQUIRREL (Glaucomys volans) (gray, mouse; flying)

Appalachian Region Distribution: Throughout.
Continental Range: All of eastern US.  Three eastern subspecies recognized in eastern US, with only one in the Appalachian region.
Abundance: Common wherever there are oak/hickory forests.  In fact, often outnumbers the diurnal gray squirrel in mature woodlands. 
Population Density: 1-3/acre, up to 5/acre.     
Size and Molt: Head and body 5 ½ to 5 2/3 inches; 1 - 3 oz. One fall molt.  Females are generally larger than males.  Smaller than the northern flying squirrel.    
Mammae: Four pair.
Habitat: Prefers mature hardwood forests, as opposed to the preferred coniferous forest of the northern flying squirrel, since they rely on stored middens of acorns, unlike the northern flying squirrel.
Active Period: Nocturnal. Does not hibernate, but undergoes periods of torpor and inactivity, more often than the northern flying squirrel (who, apparently, is more used to the cold weather). NAS says body temperatures can drop to 22°F and will take up to 40 minutes to wake.  Tends to aggregate in winter nests (in one study of 841 winter nests, 676 - 81% - had two or more in the nest, and, of those, 79% housed only adults).  16 were found curled together in a wood duck box one February day at the Patuxent Research Refuge in Maryland (50 have been recorded in one nest).  Low body temperatures seem to coincide with larger denning numbers.
Diet: Most carnivorous member of the squirrel family; eating insects, birds, eggs, small mice and shrews, and even carrion. Animal sources dominate summer diet. Winter and spring diet includes buds, catkins, and shoots of trees.  Also mushrooms, berries, buds and maple sap. Forages mainly arboreally. Known to cache food (unlike the scurids, this squirrel hides its nuts above ground in tree hollows. The southern flying squirrel may store up to 15,000 nuts in a season.  Hole eaten in nut is circular with a smooth edge. (It is a single hole versus numerous smaller holes of the mouse.)
Home Range: 1 - 7 acres.  Females are territorial in summer.  Males are less territorial, with broadly overlapping ranges.  A Maryland study a home range of 6 acres for males, 5 acres for females and 1.5 acres for juveniles.   Ranges of males overlapped extensively, with females overlapping very little with other females or males.  
Social Structure: Somewhat gregarious, especially in winter nests.  Very shy.  Males overlap ranges extensively, but females ranges overlapped little among themselves or with male ranges during the breeding period.   Reports vary on male role in rearing young; one says they help raise young, SE Mammals says male kicked out at child birth; raised by the mother, then male reappears for second mating when young are weaned, others make no comment about male role. Gregarious in winter months, with usually three to eight sleeping in the same cavity (although 28 and 50 – in an Illinois tree hollow - have been recorded).   Shedd states the southern flying squirrel is considerably more aggressive than the northern, and generally is dominant where the two species are sympatric.    
Life Cycle: Usually two litters, with two to four per litter per year, born in April and July. Females are polyestrus, in estrus for only one day, with a gestation period of 41 days.  Weaned (and can glide) at eight weeks. Mother and young will stay together until next litter.  Will breed in one year (although many studies show summer births often bear young the following spring). Young females usually have one litter, while older females often have two litters.  Life span of three to five years, with records of up to 13 years in captivity.
Dens/Nest: Dependent on cavities in trees, often  made by woodpeckers (15 - 20 feet above the ground). Has a primary nest and several secondary nests used intermittently as feeding stations and, even some for defecating stations, with openings of 1 ½ to 2 inches. Are known to make summer leaf nests, but being smaller than the northern flying squirrel, it can more often find a suitable tree cavity.  Nests are lined with lichens or finely chewed bark.  Outside nests are 8" - 10" in diameter.  Known to reuse fox or gray squirrel nests, or even bird houses.
Tracks: The give away is the "sitzmark", or landing spot, with the pattern of feet leaving the site, much like a gray squirrel, only smaller (rear foot 1 5/8" in length, with a straddle of 3 3/4 inches).
Scat: Indistinct piles, each dropping less than 1/4" in length.

Remarks: A very small tree squirrel with loose skin between the limbs. Belly hairs totally white, which differentiate the southern from the northern flying squirrel (whose white belly hairs have a gray base). Extremely arboreal, can glide up to 260 feet (normally at a 30 degree slope), although 30 to 50 feet is more common, and capable of turning 90 degrees in mid-flight. Nut-caching fall habits are photoperiodically triggered (short days). Those from SC and western NC are darker than those from MD, VA, and eastern NC. Shy and docile, they make good pets, except they can be quite noisy at night.  

Audubon and Bachman reported seeing at least 200 in and under several large beech and oak trees near Philadelphia.

At night, the eyes of the flying squirrel will shine red by the light of a flashlight.

The northern and southern flying squirrels are the only two species in the genus. 

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Family Castoridae – Beaver

With only two species worldwide, this is truly a small family.  The other species, Castor fiber, is native to portions of Europe and Asia.  Numerous closely related family members existed in prehistoric times, including the Pleistocene and late Tertiary.  One was a 700 - 800 pound, muskrat-like animal that fed on marshy vegetation.  Another extinct group constructed deep, spiraling burrows. 

The beaver, the only species in North America, is the largest rodent in North America, weighing 88 pounds, or more.  Although it is frequently reported that a beaver will continue growing throughout its life, studies indicate that growth basically terminates at age four or five. 

AMERICAN BEAVER (Castor canadensis) (beaver; from Canada)  

ARTICLES

Appalachian Region Distribution: Throughout, due to restocking. Was basically extirpated from eastern US by the turn of the century.
Continental Range: All of North America.  Only one recognized eastern US subspecies.
Abundance: Becoming more common.
Population Density: Depending on habitat quality, studies show .2 to 2 colonies/square mile (up to 8/square mile in ideal habitat)  (one colony equals 4-8 individuals), or one colony per 0.4 - 1.5 miles of stream habitat, up to a maximum of 4.2 per stream mile.  
Size and Molt: Head and body 25 to 35 inches. The largest rodent in the Appalachian region, weighing 35 to 70 pounds, with a record as of 1981 of 115 pounds.
  Second largest rodent in the world, with only the capybara of South America being larger. While many references state that beaver grow continuously throughout their life, others say they reach their practical full size after 3 -5 years.  Beaver are somewhat smaller in the southern part of their range.   It can be assumed beaver found in Appalachian streams and rivers will be on the small size, based on food quality and quantity compared with coastal marsh/ broad piedmont-coastal plain rivers.  One molt.
Mammae: Two pair.
Habitat: Live along water courses having adequate tree supply.
Active Period: Nocturnal and crepuscular, year-round. Most active in the fall to repair the dam and create a food cache.
Diet: Vegetarian, feeding on the cambium layer of buds and twigs in winter, aquatic plants and grasses in summer. A colony of 6 to 8 beaver will consume about 1,000 trees a year.  Known to "mow down" an entire aspen forest before other trees are eaten.  Coprophagy is known to occur among beaver.  Beaver are unable to digest cellulose any better than any other non-ruminant mammal, thus, large quantities of woody fibrous material must be eaten.  Will cache branches in fall underwater (cache can be ten feet high and forty feet wide) for winter food, which, in the north, must last for up to six months.  Caches are not cut until after the sap has returned to the roots, minimizing chance of spoilage.  In the south, such caches are not made.  In addition, brown fat, stored in the tail, is utilized to provide needed winter energy.  One study found witch hazels were cached for food, while maple was eaten on the spot.  Beaver will discriminate among aspen trees, “testing” the bark on aspens, and selecting those individuals with a lower concentration of natural toxic plant compounds in the bark.
Home Range: Within 660 feet from the lodge.
Social Structure: Monogamous, gregarious and very territorial (as a colony).  The social structure of a beaver colony is based on the mating pair.  Living together in a lodge, beaver live in social units of about 6, consisting of a mating pair, the yearlings, and the kits (and occasionally one or more that are about 2.5 years old).  Larger colonies (up to 12 members) are found in higher quality habitat.  Most sources say beaver mate for life (monogamous).  PA mammals says the female is reported to mate for life, although males are polygynous.  There usually is only a single breeding female in a colony.  The female appears to be dominant over both males and young most of the time.  Bonds normally last only a few years, with the death of one of the mates.  At this time, a member of a younger generation from another colony replaces the adult.  This difference in age in the couples is common. It is normally the female that selects the home site.  If the paired male dies, the female remains at the existing site.  If the female dies, the territory may be abandoned.  The two-year olds are driven off (or instinctively disperse) a few miles to establish their own range (one to five miles upstream or downstream), at which time they have reached sexual maturity. Castor scent mounds are made to stake a claim, which may lead to territorial fights, or the finding of a unmated beaver of the opposite sex.