This page is the first in a series of future squirrel natural history pages .
   This paper was researched and written in the summer of 2002 © by Barbara Bellens- Picon and was
                              presented in November 2002 at New York State Wildlife Conference.All rights reserved.

                                Sciurus carolinensis

 

                                                   

 

The Sciuridae

Fossil Records

Diet

Distribution

Subspecies

General Characteristics

Nests

Predators

 

 



                                   
Eastern Gray Squirrel

                                    Sciurus carolinenis Gmelin, 1788

 

 Graceful, acrobatic, charismatic ,one of the more familiar yet fascinating tree squirrels in
North America is the eastern gray squirrel (Sciurus carolinensis)
Eastern gray squirrels belong
to the order Rodentia of the class Mammalia. Rodentia is the largest order of mammals in
the world and accounts for more than 1600 species, greater than one third of the
species of mammals living today. The word Rodent is derived from Latin, Rodere meaning;
to gnaw.
Some rodents 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. Most rodents feed on vegetation,
but a few species, notably the grasshopper mouse, feed extensively upon animal matter

Specialized anatomical features emphasize the diversity of the group and include the beaver
tail, porcupine quills, and the patagium of the flying squirrel. The order Rodentia is further
divided into three suborders, one of which, the sciuronathi, contains the squirrel-like rodents.
The 367 or so species in this sub-order are further divided into 7 families.
One of these, the Sciuridae, includes the tree squirrels, the ground squirrels and the flying
squirrels. Currently 26 species of tree squirrels in the genus sciurus are recognized with
 three species found in Eurasia and across North and South America (Wilson and Reeder 1993).

      The Sciuridae

 

The Sciuridae is made up of 267 species in 49 genera. A large family represented in both old and
 new worlds. The Sciuridae includes arboreal tree squirrels, terrestrial chipmunks and ground squirrels, and
semi fossorial-marmots and prairie dogs. In habits, members of this order are diverse. There are
 three basic body forms in squirrels: tree squirrels, ground squirrels and flying squirrels. Tree
squirrels have long, bushy tails, sharp claws and large ears. Some have well-developed ear tufts.
Flying squirrels have a furred membrane (patagium) extending between the wrist and ankle that
allows them to glide between trees.
Ground squirrels often have short, sturdy forelimbs that are used for digging. Their tails, while fully
furred, generally are not as bushy as those of tree squirrels.
 Like many other rodents, all squirrels have five functional toes on the hind feet and four on the
forefeet, with a well-developed claw on each digit.
Squirrels occur throughout the world except for Australia, Madagascar, southern South America,
parts of North Africa ,polar and some desert regions.( Gurnell ) They inhabit a wide variety of habitats
 from the tops of trees to underground tunnels.

There are seven native species of tree squirrels in North America which belong to the two genera,Tamiasciurus
and Sciurus; Pine squirrel,(Tamiasciurus hudsonicus),Douglas squirrel ( T.douglasii)
 Fox squirrel ( Sciurus niger) Eastern Gray squirrel( Sciurus carolinensis)Western gray
squirrel( S griseus)Arizona gray squirrel ( S arizonensis)and the Abert squirrel( S aberti )

Appearing on the USFW endangered species list are the Delmarva Fox squirrel
 (Sciurus niger cinereus) and mount graham red squirrel, (Tamiasciurus hudsonicus grahamensis). Two Appalachian Mountain subspecies of the Northern Flying Squirrel also appear on the US Endangered species list; the Carolina Northern Flying Squirrel (glaucomys sabrinas coloratus), classified as endangered in North Carolina and Tennessee, and the Virginia Northern Flying Squirrel (glaucomys sabrinus fuscus), classified as endangered in Virginia and West Virginia. Loss of habitat is the primary reason for the species decline
.

 “Sciurus" the Latin word for squirrel comes from the ancient Greek  "skia," meaning shadow,
and "oura," meaning tail referring to the characteristic posture of a thick bushy tail held over its
head. The eastern gray squirrel (Sciurus carolinensis) was first observed in the Carolinas.
The earliest published record of this squirrel was in 1788 by J.F Gmelin. James J Audubon was
one of several naturalists who referred to the gray squirrel as the “migratory squirrel”
 (Sciurus migratorius) in reference to its tendency for mass migrations.

 

Fossil Records

 

Some biologists refer to tree squirrels as "living fossils" because their form and structure have
 been relatively unchanged for the last 5 million years. Discovered in fossil records from the
Oligocene periods in North America, Protosciurus  is the oldest known member of the squirrel
 family and its bones are very similar to those of the tree squirrel today. Fossils of some early
squirrels are on display in the Smithsonian Museum.

 

Diet

Classified as granivore/herbivores they primarily rely on the products of the trees in the
habitats in which they live. They are known to eat bark of dead trees for the fungal mycelia
within. Squirrels have a large cecum, optimized to digest plant protein, not animal protein .
However they will resort to occasional omnivory when necessary. Adults obtain minerals from
 bones, antlers and turtle shells. They feed on nuts flowers and buds of more than 24 oak
species, 10 species of hickory, and also pecan, walnut and beech. (Fagus grandifolia)
(Korschgen, 1981;Nixon et al, 1968) Other important foods include the fruits, seeds, buds or
flowers of maples (acer) mulberry (morus) hackberry (celtis) elms (ulnus) buckeye and horse
chestnuts (aesculus) wild cherries (prunus) dogwoods (cornus) hawthorn (crategus), black
gum (Nyssa sylvatica) hazelnut (Corylus americana) hop hornbeam (ostrya virginiana) and
ginko (Ginko biloba).
Seeds and catkins of gymnosperms are also eaten including cedar (juniperus) hemlock
(tsuga) pines (pinus) and spruce (picea) (Barkalow and Shorten 1973);
(Thompson and Thompson 1980) A variety of fungi are also eaten particularly in the summer
along with insects. Other lesser-consumed items are bones (Flyger and Gates 1982) bird
eggs and nestlings (bailey 1923) and frogs (Goodrum 1961) tree bark and cambium-
inner lining of trees.

 

Distribution/Habitat

Eastern gray squirrels are found throughout the eastern United States; their natural range
 extends from Florida, north to Canada, and west to where deciduous forests meet the
Great Plains grasslands. Due to introductions, they have become established in several
areas of western Us and Canada (Flyger and Gates 1982) Mature deciduous forests provide
an ideal habitat.
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 ( Sciurus niger). The more large trees an area has, the more gray squirrels will
 inhabit the area (larger cavities and more food). Adapted to a variety of wooded conditions,
included urban area parks and suburbia that include oak, maple and hickory trees.
Habitat loss worldwide greatly affects tree squirrels and is a central factor in their conservation.
For example since 1820, nearly 80 percent of forests in Illinois have been converted to agricultural
cropland (Iverson 1988) Habitat loss is the primary reason cited for the legal protection of several
 subspecies of fox squirrels in the eastern United States.

 

Home Range

The area where most of its foraging for food, nest building, and breeding/rearing occurs.
 The home ranges of males are larger than those of females. One to four acres, up to ten
acres (varies with food availability), Chapman and Feldhammer report 2 acres for males
and 1.25 acres for females in a Maryland study (Flyger).  Home ranges can overlap and
there is no evidence that of true territoriality (Don 1983) although core areas are
sometimes defended  (Kenward 1985)

Subspecies

There are 5 subspecies of eastern grey squirrels, S. carolinensis carolinensis is the
subspecies found in most of the south from northern Florida, to North Carolina, west to
Missouri, and eastern Texas. Others include S.cextimus,S.c.fuliginosus, S.c. hypophaeus
 
and S.c. pennslylvanicus
. Sciurus carolinensis also were introduced into various parts
of the pacific states and also into parts of Canada, England, Italy, Scotland, South Africa
and Australia. The overseas introductions occurred at the end of the 19th and beginning
of the 20th century. The 1880's introduction from England to Australia failed by 1973.
(Purdue, 1980)

General Characteristics 

Sciurus carolinensis are medium sized arboreal dwelling quadrupeds weighing @ 300-700
grams with no sexual dimorphism in size or coloration.
 Body length measures@ 16-21 inches with the distinctive trademark
 tail measuring 8- 10 inches. They have strong back legs and their back
 feet contain 5 elongated claws with the 4th being the longest. Their back
 feet can turn 180 degrees, an adaptation that allows them to flexibly
climb head first, down a tall tree. Their front feet have 4 claws and the
appearance of a vestigial thumb. Diurnal in winter to crepuscular in
summer,  primarily arboreal mammals that are most active in
September and early October, burying acorns (this is also the peak time
for their arboreal activity). They do not hibernate like the woodchuck
 (Marmota monax), but may rest to conserve energy in response to
extreme temperatures. Active year-round,  they may remain in their nests
during inclement weather until hunger drives them to forage.  Activity
levels generally peak  @ 2 hours post dawn and 2-4 hours before sundown.

 

Appearing predominantly gray up close, the fur color palette actually is mixed with dark and
light gray hues. Underside fur color is whitish and tail guard hair tips are white tipped. White or
 buff colored fur ring the eyes .A buff tinted color appears on top of feet, shoulders and head
area. Pads of feet are pinkish. Sometimes in winter, whitish fur grows around the ears giving a
 "tufted" appearance and thicker fur grows on the inner bottom legs giving a “pantaloon”
appearance. In many areas of the country/state color variations exist. Gray squirrels can have
brown, dark gray, orange, erythristic (reddish) and tan color tone variations. Mixed breeding,
genetics and environmental factors could possibly attribute to the reason behind these
variations of color. Melanism (black) and albinism (white) occur throughout the species range,
although melanism  populations occur more commonly in the north.

Seasonal breeders, they are genetically timed to breed and reproduce coinciding with the
buds and flowering of indigenous trees .Two peaks in breeding occur resulting in spring and
 fall litters. Litter size ranges from 1-8,with an average of 2-3. Weather and food supply are two
 strong factors that affect their breeding behavior. Unfavorable weather and poor food supplies
 can delay or even offset breeding seasons. It can begin early as December and end around
July or August although litters of young are sometimes born well into fall and lactation frequently
 continues in September and October. Females can go into estrus and become sexually
mature at 9-15 months of age. Most females do not give birth during their first year. They have
 a duplex uterus ,os clitoris measuring 3.6 mm ( Burt 1960 ) and eight mammae (4 pectoral,
two abdominal, 2 iningual) that become black after the first pregnancy. (Flyger and Gates, 1982)
Gestation is @ 44 days with neonatal squirrels being born weighing 12-18 grams, and about 2
inches long, with claws and vibrissae but without fur, sight and hearing .The umbilicus remains
attached for @ 4 days. The dorsum darkens at 7-10 days preceding the development of velvet
like fuzz toward the end of the second week. It takes @3 to 4 months to achieve independence.

Most male eastern gray squirrels become sexually mature at @ 9 - 12 months of age. Inactive
testes weigh 1g, whereas active testes weight 6-7g. (Webley and Johnson 1983) The brain
weighs @ 7-10 grams (Meir, P.T. 1983) The cycle of testicular recrudescence and regression
 occurs twice a year. The baculum or os penis is thought to aid in copulation and is 9.2-12.3 mm
in length. ( Burt 1960 )Their heart beats 150-450 BPM. (E Smith and Johnson 1984) and their
body temperature ranges between 98 and 102 degrees F. (Bolls and Perfect 1972; Hoff Lassing
et al 1976)

Eastern gray squirrels have both a summer and winter coat, and therefore molt twice each year,
coinciding with the changing of the seasons.  Molting lasts for @ 4-6 weeks but individual
differences vary. Spring molt begins at the face and head, progressing toward the rump. The tail
and tips of ears only molt once. Tail molt begins at base and ends at tip…this process can take
several months to complete. Fall molt occurs in the opposite direction of spring molt. Variations
of color coincide with the molt and most likely function in camouflage: to help them survive.
Squirrel fur is multi-layered providing insulation with short hairs, which lay close to the skin, and
long outer fur, called guard hairs, which grow longer during the winter months.

Like many other rodents lacking canine teeth, their incisors are hypsodont  (grow continuously
 throughout life). They need to gnaw in order to keep these teeth trimmed. Each incisor is a 
segment of a true circle, continuously being pushed out of the end of the jaw. 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. They have 22
 teeth in total. The total dental formula is 1 incisor, 0 canine, 1 premolar, and 3 molars on each 
side of the bottom mandible. A complimentary set of the same number of teeth occurs on each
 side of the top with a second premolar. Dental formula is expressed I 1/1,C 0/0, P 2/1 M 3/3 .

The front incisors are a yellowish, orange color. Adjacent to the incisors is a gap referred to as
 the diastema into which the lips can be drawn, sealing off the mouth from inedible food
 fragments and soil. Following the diastema are the grinding teeth, or "cheek" teeth, molars and
 pre-molars which 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.  
 Gray squirrels have 2 upper pre-molars. Other species like the fox squirrel do not have the first 
upper premolar. Permanent molars will replace the first premolars when the squirrel reaches 
adulthood.The molars are rooted.
Distinguished feature of sciuromorhs are a relatively primitive unspecialized 
arrangement of jaw muscles and parts of the skull. The lateral masseter muscle 
extends in front of the eye onto the nose, moving the lower jaw forward during 
gnawing. The deep masseter muscle is short and used only for closing the jaw. 

 

Reprinted with permission William Higgs-Dept of Archaeology, University of York

                                       http://www-users.york.ac.uk/~wjh101/hedbone/Squirrels/squirrels.htm

 

Squirrels have 4 sets of whiskers or vibrissae on the head. The whiskers are always black in
color. They are located above and below their eyes, in front of the throat and along side the
nose. There are also whiskers on the wrist (ulna) on the underside of the mammae, at the
base of the tail, and around the feet. (Laidler 1980, Shorten 1954). Some of these whiskers
 are even present at birth. The sensitive whiskers function as touch receptors that help relay
information about the squirrel’s immediate surroundings.  Eye- closed neonates use them to
locate the mammae in order to nurse.

Squirrels have very good eyesight with a wide field of vision and are able to see in dim light.
Evidence suggests that they possess dichromatic color vision which would akin them with a
red/green colorblind person. They possess a well-developed sense of smell, which aid in their
 instinct to survive. It allows them to help locate buried (cached) food and plays an important
 role in their social communication. Scent glands can be found on the underside of their paws
 and they also deposit urine on tree branches as a way to define their territory and home range.
 These marking sights can sometimes be found on the underside of large branches. Females
emit odor and deposit scent at the onset of estrus to announce their availability to the male
population. Males reciprocate by scooting across branches to deposit their own individual scent
on the branches and trees.

A squirrel’s glorious tail is its characteristic trademark providing multifunctional uses, which
 include protection, shade, balance, warmth, and communication. Stripes located on the
underside provide a rough estimation of a young squirrels age. Adults do not possess this
characteristic. The squirrel’s tail can also break at mid-point and allow the squirrel to escape
while being held by a predator. Squirrels with partial tails can function and survive successfully.
Depending on position and action tail movement reflects emotion; it can indicate fear, curiosity,
aggression, or anger. A limp non-responsive tail could mean injury.

Gray squirrels are scatter hoarders, placing single food items at different locations. Food is
buried (cached) one half to 1 inch beneath the soil. Scatter hoarding is an important survival
strategy that allows gray squirrels to remain active through the winter and spring. They primarily
 use their keen sense of smell to locate the buried food but often, the squirrels do not locate
caches and the stored seeds germinate to produce trees and other forest plants. Because fungi
 is part of their diet, the Eastern grey squirrel may play as important role in the dispersal of
hypogeous mycorrhizal spores. All conifers and many hardwoods strongly depend on
mycorrhizae (fungus roots) for growth and survival.

Generally socially tolerant, a dominance hierarchy in both females and males is maintained by
 bluffing or chasing rather than by actual fighting. Usually considered solitary animals, studies
suggest that they operate within a complex social system.
(Thompson 1978a, 1978b:Koprowski 1991a, 1993a)
Males dominate over females, adults dominate over immature sub adults. Pregnant and lactating
 females are aggressive toward all
. Individual squirrels are often seen feeding close to each other
 without any aggressive activity

 

Nests     

 

Three general types: Summer, winter dreys and winter den. The leaf nests are called dreys and the
dens are formed in tree cavities and crevices. Winter dreys are usually more sturdy and elaborate
 then summer ones. Dreys that are @12- 15 inches wide and one foot high are designed to be
waterproof, constructed with an outer layer of coarse natural items such as larger twigs and
 branches. The inner layer consists of leaves, bark and other insulating materials .In urban
metropolitan areas unnatural items have been used as construction material such as; ceiling
insulation, cereal boxes, cigarette cartons and plastic bags. It may take squirrel one or two days
 to complete building a nest. They can be seen adding to it over time with the advent of inclement
weather conditions. A nest will always have one entrance and sometimes a second, which probably
 functions as an escape route. Groups of individuals can share a nest. Unlike flying squirrels, grays
generally do not use their nest as a latrine or to store food. The nest is usually clean. When the
problem of fleas arise they vacate and set up housekeeping elsewhere. Gray squirrels can have
more than one nest. Perhaps this helps to control external parasites, elude predators or establish
 territory. Gray squirrels also use tree dens primarily in winter. These dens exist in the main trunk of
 large mature or dead trees. They offer substantial protection against cold and inclement weather.

 

 

Predators

 

Natural predators include Largemouth bass (Micropterus salmoides) Timber snake
(crotalus horridus) Diamond back rattlesnake (C.adamanteus), Black rat snakes
(Elaphe obsoleta): moore 1941) Bullsnakes (pituophis melanoleucus) Red tail hawk,
 (Buteo jamaicensis) Red shouldered hawk (B Lineatus), Broad winged hawk (Buteo platypterus)
 Marsh Harrier (Circus aeruginosus) Cooper’s hawk, (A. Cooperi) Goshawk (Accipiter gentilis),
Great horned owl (Bubo virginiacus), Long-eared Owl  (Asio otus), Barred owl, (Strix varia)
American crow, (Corvus brachyrhynchos)Common raven,(Corvus corax) Raccoon ( procyon lotor),
Opossum,( Didelphis virginiana) American martin(Martes americana),
Weasel( Mastela frenata;hamilton 1934), Red fox,(vulpes,vulpes),
Gray fox( Urocon cinereoargenteus) Bobcat,( lynxs rufus) Coyote,( C Latrans) mink(M.Vison),
 Red Squirrel (tamiascirus hudsonicus)
Unnatural/introduced predators include;  House cat,(Felis catus) Dog,(Canis familiaris )
and Human beings.(Homo sapiens)

 

 

 

References and Literature cited

Bailey, B 1923             Meat eating propensities of some rodents in Minnesota.
                                    Journal of Mammalogy, 4: 129.

Barkalow F, M Shorten 1973  The World of the Gray Squirrel. Lippincott Co.

Barkalow F.S. Jr. and R.F.Soots Jr. 1975 life span and reproductive longevity of
               the gray squirrel sciurus carolinensis.Journal of Mammology 56:522-524

Bolls,N.J and J.R.Perfect 1972  Summer resting metabolic rate of the gray squirrel.
                   Physiological Zoology 45: 54-59

Burt W.H. 1960            Bacula of North American Mammals.
                  Misc publications of the museum of Zoology.U of Michigan 113; 1-75

Chapman J, Feldhammer G 1982. Wild mammals of North America JHU Press
Daniel H.J and L.K Roberson 1987 The ossicles of the eastern gray squirrel,
                Sciurus Carolinensis. Journal of the Elisha  Mitchell  Scientific
                                                                                  Society 103:21-27

Dunaway,P.B 1969           " Perfect" polydactylism in hind feet of a gray  squirrel.
                                            The American Midland Naturalist,81:244247

Don,B.A.C. 1983. Home range characteristics and correlates in tree squirrels.
                                      Mammal Review 13:123-132
Flyger and Gates 1982        Fox and gray squirrels
                                             pgs 209-229    WILD MAMMALS of N America

Goodrum, P.D. 1961 The gray squirrel in Texas.Texas parks and

                     wildlife Dept bulletin 42:1-43

Gurnell,J ( 1987 ) The Natural history of Squirrels  Facts on File,N.Y

Lavenex,P.,W Shiflett,R.K.Lee,and L.F.Jacobs.1998    Spatial versus

    Nonspacial relational learning in free-ranging fox (sciurus niger) Journal of Comparative Psychology 112:127-136

Kalman B, Langille J   What is a Rodent ?  2002. Crabtree Publishing Company

Koprowski,John L,1994   Sciurus Carolinensis  Mammalian Species.
                                           American Society of Mammologists.No 480 pp 1-9

Korschgen, L.J. 1981  Food of Fox and Gray squirrels in Missouri.
                                        Journal of wildlife management 45:260-266

MacClintock D. 1970 Squirrels of North America. Van Nostrand Rheinholdt Co

MacDonald, D. (Ed.). 1987. The Encyclopedia of Mammals.
                                              Facts on File Pubs. New York.

MacDonald ,I.M.V. 1992  Gray squirrels discriminate red from green in a
                                          foraging situation Animal Behavior 43:694-695

Meier,P.T. 1983        Relative brain size within the North American Sciuridae.
                                  Journal of Mammology 64;642-647

Muchlinski,A.E. and K.A.Shump,Jr 1979   The Sciurid tail:
        a possible thermoregulatory mechanism. Journal of Mammalogy 60:652-654

Short H.L.and W.B.Duke 1971 Seasonal food consumption and body weights
                 of captive tree squirrels.The Journal of Wildlife Management 35;435-439

Steele M, Koprowski J,  North American Tree Squirrels 2001.
                                      Smithsonian Institution Press

Steele M, Merritt J,Zegers D,1998 Ecology and  Evolutionary Biology of  Tree Squirrels

Stone W.B. 1995-96  Annual Wildlife Pathology Report ,New York

Stupka, A.  1935 - 63. Nature Journal, Great Smoky Mountains National Park.

Thorington ,R.W. Jr 1966 The Biology of Rodent Tails; a study of form
               and function Fort Wainwright, Alaska; Artic Aeromedical Laboratory,137p

Webley G.E. and E.Johnson 1983 Reproductive physiology of the gray squirrel
                            ( sciurus carolinensis) Journal of Endocrinology 89:107-116

Wilson, D. E., and D. M. Reeder. 1993. Mammal Species of the World,
                                     A Taxonomic and Geographic Reference. 2nd edition.
                                    Smithsonian Institution Press, Washington.  xviii+1206 pp. 

Woods S.E., The Squirrels of Canada 1980 .The National Museums of Canada

 

 

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