(DRAFT) - Taxonomy
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
TAXONOMY
NAME - SQUIRREL, FLYING, NORTHERN, VIRGINIA
OTHER COMMON NAMES - SQUIRREL, FLYING, NORTHERN, VIRGINIA; SQUIRREL, FLYING and NORTHERN
ELEMENT CODE -
CATEGORY - Mammals
PHYLUM AND SUBPHYLUM - CHORDATA,
CLASS AND SUBCLASS - MAMMALIA,
ORDER AND SUBORDER - RODENTIA,
FAMILY AND SUBFAMILY - SCIURIDAE,
GENUS AND SUBGENUS - GLAUCOMYS,
SPECIES AND SSP - SABRINUS, FUSCUS
SCIENTIFIC NAME - GLAUCOMYS SABRINUS FUSCUS
AUTHORITY -
TAXONOMY REFERENCES -
COMMENTS ON TAXONOMY -
Virginia Northern Flying Squirrel
Glaucomys sabrinus fuscus Miller, 1936
KINGDOM: Animal GROUP: Mammal
PHYLUM: Chordata CLASS: Mammalia
ORDER: Rodentia FAMILY: Sciuridae
The two endangered subspecies of northern flying squirrel,
Glaucomys sabrinus fuscus Miller and Glaucomys sabrinus coloratus
Handley are small, nocturnal, gliding mammals some 260-305 mm in
total length and 95-125 grams in weight (22,30,32,45). They possess
a long, broad, flattened tail (80% of head and body length), prominent
eyes, and a dense, silky fur (49). The distinctive patagia, folds of
skin between the wrist and ankle, are fully haried and supported by
slender cartilages extending from the wrist bones; these plus the
broad tail create a large gliding surface and are the basis for the
squirrels' characteristic aerial locomotion (41). Adults are
dorsally gray with a brownish, tan, or reddish wash and grayish white
or buffy white ventrally. Juveniles have uniform dark, slate-gray
backs and off-white undersides. The more southern subspecies G. s.
coloatus, is larger (avg. 286 vs 266 mm total length) than G. s.
fuscus, with a longer tail (avg. 134 vs. 115 mm) and brighter
Taxonomy - 1 (DRAFT) - Taxonomy
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
coloration (24). The northern flying squirrel can be distinguished
from the southern, G. volans, by its larger size (e.g. hindfoot 33-41
mm. vs. less than 33 mm. for G. volans; adult weight (85-125 grams vs.
50-80 grams)), the gray base of its ventral hairs as opposed to the
white base in the southern species, the relatively longer upper tooth
row, and the short, stout baculum (penis bone) of the males (49).
Wells-Gosling (48) provides numerous photographs of both Glaucomys
species.
There is no cuurrent disagreement on classification or
nomenclature, and synonyms since original descriptions of subspecies.
Glaucomys sabrinus fuscus was described by Miller in 1936 (32). A
full account of the species taxonomic history can be found in Howell
(25); Hall (21); and Wells-Gosling and Heaney (50). The type
specimen is in the U.S. National Museum (#360420).
Taxonomy - 2 (DRAFT) - Status
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
STATUS
Coded Status
Virginia; Federal Endangered
Virginia; State Recognized
Virginia; Unofficially Listed
West Virginia; Federal Endangered
West Virginia; State Recognized
West Virginia; Unofficially Listed
E: Federal Endangered
COMMENTS ON STATUS -
U.S. STATUSES AND LAWS:
The Virginia northern flying squirrel (Glaucomys sabrinus fuscus)
has been designated Endangered pursuant to the Endangered Species Act
of 1973 (50 CFR 17.11; P.L. 93-205, 87 Stat. 884; 16 U.S.C.
1531-1540), as amended. The subspecies has this status wherever found
including the States of Virginia and West Virginia.
This subspecies is protected by the Lacey Act (P.L. 97-79, as
amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to import,
export, transport, sell, receive, acquire, or purchase any wild animal
(alive or dead including parts, products, eggs, or offspring):
(1) in interstate or foreign commerce if taken, possessed,
transported or sold in violation of any State law or
regulation, or foreign law; or
(2) if taken or possessed in violation of any U.S. law,
treaty, or regulation or in violation of Indian tribal law.
It is also unlawful to possess any wild animal (alive or dead
including parts, products, eggs, and offspring) within the U.S.
territorial or special maritime jurisdiction (as defined in
18 U.S.C. 7) that is taken, possessed, transported, or sold in
violation of any State law or regulation, foreign law, or Indian
tribal law.
RESPONSIBLE FEDERAL AGENCIES:
USFWS -Responsible for the management/recovery, listing, and
law enforcement/protection of this species.
USFS -Responsible for the law enforcement/protection of this
species with applicable State and Federal laws on
public lands under their control. Also responsible
for management/recovery on Forest Service lands. The
Forest Service is responsible for integrating
management, protection, and conservation of Federally
listed species into the Forest Planning process
(36 CFR 219.19 and 219.20).
All Federal agencies have responsibility to ensure that any
action authorized, funded, or carried out by that agency is not likely
to jeopardize the continued existence of the species or result in the
destruction or adverse modification of Critical Habitat (50 CFR 402),
Status - 1 (DRAFT) - Status
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
and to utilize their authorities to carry out programs for the
conservation of the species.
STATE STATUSES AND LAWS:
STATE: Virginia
DESIGNATED STATUS: Recognized Endangered
ADMINISTRATIVE AGENCY: VA Comm. of Game and Inland Fisheries
STATE STATUTE: VA Code, Subsect. 29-230 to 29-237 and 29-2.1,
29-143.
UNOFFICIAL LIST:
STATE: West Virginia
DESIGNATED STATUS: Recognized Endangered
ADMINISTRATIVE AGENCY: Dept. of Natural Resources, Division of
Wildlife Resources
STATE STATUTE: WV Code, Subsect. 20-2-3, 20-2-4
Sec. 20-1-2, 20-2-50, and 20-2-12.
Co-operative agreement Sec. 6, in Appendix A.
INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS:
None.
ECONOMIC STATUSES:
The subspecies has no negative impact, and positive values are
unknown (species may be important in dispersal of hypogeous fungi
and/or in maintaining important mycorrhizal associations).
82/12/30:48 FR 58454/58460 - Notice of Review, Candidate species
84/11/21:49 FR 45880/45884 - Proposed listing, Endangered
85/07/01:50 FR 26999/27002 - Final Rule, Listed as Endangered
Status - 2 HABITAT ASSOCIATIONS
HABITAT - TERRESTRIAL
TERRESTRIAL
INLAND AQUATIC
SOCIETY OF AMERICAN FORESTRY TYPES
SAF TYPE STAGE CLOSURE
White spruce balsam fir young tree
White spruce balsam fir mature tree
White spruce balsam fir Old Growth
young tree
mature tree
Old Growth
LAND USE -
Evergreen Forest Land
Mixed Forest Land
Streams and Canals
NATIONAL WETLAND INVENTORY CODES
NWI NWICLS NWIMOD NWISPEC
Riverine, upper perennial
COMMENTS ON HABITAT ASSOCIATIONS -
Throughout its range the northern flying squirrel is usually
associated with conifer (especially spruce-fir) and northern hardwood
forests (02,08,21,22,26,31,44,47,49,50). The subspecies fuscus and
coloratus are commonly captured in mature hardwood stands of beech
(Fagus grandifolia), yellow birch (Betulia alleganiensis), sugar maple
(Acer saccharum) hemlock (Tsuga canadensis), black cherry (Prunus
serotina) and northern red oak (Quercus rubra) associated with red
spruce (Picea rubens) and fir (Abies fraseri and A. balsamea). These
squirrels have also been taken in deciduous areas some distance from
coniferous vegetation (14,20,48). Studies with captives indicate that
G. sabrinus will readily use both deciduous and coniferous habitats,
while G. volans has a marked preference for hardwood forest (47).
Although some northern flying squirrels have been captured in stands
of varying age and understory density, most records show a marked
preference for old growth forest with widely spaced, tall trees (50).
Such habitats seem well suited to the species' gliding form of
locomotion, while the abundance of natural cavities in old hardwoods
and their relative resistance to wind-throw (compared to many
conifers) may account for the northern flying squirrel's occupation of
the deciduous and mixed woodland just below the spruce-fir zone.
Since the northern hardwood - spruce-fir ecotone occurs at
progressively higher elevations from north to south, it is perhaps not
surprising that captures of G. sabrinus show a similar latitudinal
trend. While individuals have frequently been found at elevations
less than 800 meters in New England, New York and Pennsylvania, most
of West Virginia specimens have been taken at 1000-1350 meters) (41).
In Virginia, G. sabrinus, commonly occupies forests 1170-1630 meters
in altitude, but one population is known from a riparian hemlock-
hardwood-rhododendron habitat at 615 meters (19). In North Carolina
all captures have occurred above 1540 meters with the exception of a
single specimen taken in the Great Smokey Mountains at 1230 meters
(21,44). Thus, in the southern part of its range, G. sabrinus shows a
Habitat Associations - 1 relict distribution and trends to occupy rather small potentially
vulnerable islands of high elevation habitat.
While the above vegetational and altitudinal information provide
an indication of the habitat preferences of this species it is not
adequate or complete in itself. The species' absence from many
apparently suitable areas, its apparent population oscillations in
particular areas over extended periods and its occupation of only
cool, moist areas suggest that variety of habitat requirements remain
unknown. The inflence of hypogeous fungal crops and other animal
species on its habitat needs further study.
During the cooler months, squirrels commonly occupy tree cavities
and woodpecker holes (02,09,22). Their tendency to form small
aggregations in such hollows appears to reduce winter thermoregulatory
costs. In the summer, animals may continue to use cavities, may
construct leaf nests, or may use those of other squirrel species (34,
44). Data from captive squirrels (44) and a study in progress in
West Virginia (43) indicate that G. sabrinus sometimes enters burrows
in the ground, although the extent of their use is not yet known.
Telemetry studies reveal that animals use secondary refuges
intermittently during the night, possibly for feeding or concealment
(48).
Habitat Associations - 2 (DRAFT) - Food Habits
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
FOOD HABITS
TROPHIC LEVEL -
HERBIVORE
LIFESTAGE FOOD FOOD PART
General Myxomycophyta
General Deciduous Shrubs-Flowers/Fruit/Seed
General Evergreen Shrubs-Flowers/Fruit/Seed
General Deciduous Trees-Leaves/Twigs
General Deciduous Trees-Flowers/Fruit/Seed
General Coniferae
General Evergreen Trees-Flowers/Fruit/Seed
General Arthropods
General Aves
General Mammalia
General Carrion
Food Habits - 1 (DRAFT) - Environment Associations
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
ENVIRONMENTAL ASSOCIATIONS
G = General A = Adult
LIM = Limiting RA = Resting Adult
J = Juvenile FA = Feeding Adult
RJ = Resting Juvenile BA = Breeding Adult
FJ = Feeding Juvenile P = Pupae
L = Larvae E = Egg
RL = Resting Larvae
FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS
G Terrestrial Features: Burrows
G Terrestrial Features: Downed logs
G Snags: Unknown
G Tree Cavities: Unknown
Environment Associations - 1 (DRAFT) - Life History
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
LIFE HISTORY
FOOD HABITS:
Because of its rarity, nocturnal and secretive habits and the
remoteness of its habitat, little is known of the ecology of the
northern flying squirrel in the southern Appalachians. Unlike
Glaucomys volans, it is less dependent on seeds and nuts for food
(44,47), and in fact, may not be able to use conifer seeds effectively
in some boreal habitats (05,06). Over much of it's range it can
apparently subsist on lichens and fungi (30,32,35,44), but also eats
certain seeds, buds, fruit, staminate cones, insects, and other
animal material (21,22,24,26,44,50). The year-round abundance of
lichens and many species of hypogeous (underground) fungi may provide
a steady and almost exclusive food supply and reduce food competition
with other squirrel species. Periodic dependence on fungi and
lichens may also be a factor in restricting the species to high
elevation, mesic habitats.
HOME RANGE/TERRITORY:
Northern flying squirrels are relatively gregarious and are known
to share nests; however, the spectacular winter nesting aggregations
reported for G. volans (up to 50 in a nest) are unknown for this
species. Northern flying squirrels apparently live in family groups
of adults and juveniles; in those areas where the species has
been located, it has often been possible to trap two to six
individuals within a 300 x 400 meter area (46). Such data may also
indicate an approximate density of one squirrel per two hectares.
Home range determinations based on summer telemetry data suggest
individual home ranges of 2-3 ha in North Carolina (37,48) and 5-6 ha
in West Virginia (43). Given the preliminary nature of the above
density and home range data, reasonable estimates of the amount of
habitat necessary to support viable populations of northern flying
squirrels are undetermined.
PERIODICITY:
Northern flying squirrels are nocturnal, non-hibernators. The
activity patterns of the genus Glaucomys have been the subject of
intensive laboratory study (11,12,13,16,38,44) and it appears that
photoperiod and levels of illumination are major controlling factors
in their behavior (12,38). Temperature-telemetry studies in the
southern Appalachians during the summer have revealed a marked
biphasic nocturnal activity pattern with peaks between sundown and
midnight and 1-3 hours before sunrise (37,48). During these times
animals are extremely active in trees and on the ground and enter a
number of different nests or refuges. While heavy fog (cloud),
rain, and high winds delay the onset and decrease the intensity of
the activity, they do not suppress it altogether. At such times
flying squirrels appear to spend more time moving along branches
than gliding. In the ongoing study in West Virginia, one third of
the captures have been in ground sets, indicating that the squirrels
may spend a significant amount of time foraging on the ground (27,43).
Such an observation is consitent with the species' dependence on
hypogeous fungi.
Life History - 1 (DRAFT) - Life History
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
Little is known about seasonal variation in activity in this
species. Although squirrels take refuge in nests during extreme
weather conditions, they are known to be active throughout the winter
(26,49).
MIGRATION PATTERNS:
No evidence of migratory behavior.
COVER/SHELTER REQUIREMENTS:
Because of the squirrel's small size, the climatic severity of
it's habitat, and it's vulnerability to predators, nests and refuges
represent critical resources (44). During the cooler months,
squirrels commonly occupy tree cavities and woodpecker holes (02,09,
22). Their tendency to form small aggregations in such hollows
appears to reduce winter thermoregulatory costs. In the summer,
animals may continue to use cavities, may construct leaf nests, or
may use those of other squirrel species (34,44). The interior of
both winter and summer nests is lined with lichens, moss, sedges, and
finely chewed bark. Data from captive squirrels (44) and a study in
progress in West Virginia (43) indicate that G. sabrinus sometimes
enters burrows in the ground, although the extent of their use is
not yet known. Telemetry studies reveal that animals use secondary
refuges intermittently during the night, possibly for feeding or
concealment (48).
REPRODUCTIVE SITE REQUIREMENTS:
Almost nothing is known about reproductive requirements in the
wild for this species (10,26). Recent studies (49,50) indicate that
young are born and reared in tree cavities (and nest boxes) lined with
the same materials described above in "COVER/SHELTER REQUIREMENTS"
above.
REPRODUCTIVE CHARACTERISTICS:
While most data on reproduction in Glaucomys sabrinus has been
obtained from captive animals, a limited amount of data has come to
light from field studies (19,39,41). In the southern Appalachians
females have been observed to be in estrus in late March, April, and
early May, but not later in the year, suggesting the production
of only one litter per year (50). After a gestation period 37-42 days
(36), 2-5 young are born (19,50). Litter sizes for the two
subspecies, fuscus and coloratus, have ranged from 2-4 for both
captive females and those observed in the wild (24,41,46). Weaning
apparently occurs in approximately 8 weeks (50). The small size of
the grayish juveniles in middle and late summer suggests that females
are not capable of breeding until the beginning of their second year.
PARENTAL CARE:
Although Muul (36) and Ferron (18) have described aspects of
early development and growth of offspring, little is known of parental
care (10). It appears that young are raised almost exclusively by
females but after weaning associate with both parents in small family
groups (24,46).
Life History - 2 (DRAFT) - Life History
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
POPULATION BIOLOGY:
Not known, see Wells-Gosling (50) for brief summary of available
information. Until recently, no field study has been designed to
obtain demographic data.
SPECIES INTERRELATIONSHIPS:
Species interrelationships can be divided into the categories:
1) basic requirements, 2) potential competitors, 3) pathogens, and
4) predators.
1) Basic requirements - The tree species that provide or support
critical foods such as seeds, insects, mycorrhizal fungi, lichens,
etc. and nesting sites represent critical resources. To the extent
that these trees also benefit from the dispersal of hypogeous
fungal spores and seeds, the relationship with G. sabrinus might
be considered mutalistic.
2) Potential competitors - While the red squirrel (Tamiasciurus
hudsonicus) is often cited as a potential competitor, its diurnal
habits, dependence on seeds, and long evolutionary history with
G. sabrinus argue against a severe competitive relationship.
Glaucomys volans, on the other hand, has many similar requirements
and behaviors, has demonstrated a degree of competitive superiority
in the laboratory, and has apparently increased its range in high
elevation forests in the southern Appalachians. The two species
have been captured in the same areas but not usually in successive
years and thus may not have established any kind of stable
relationship.
3) Pathogens - Studies in North Carolina suggest that G. sabrinus is
adversely affected by the nematode parasite Strongyloides robustus.
While this parasite is only rarely found in wild G. sabrinus, it
has frequently been identified in G. volans and other low elevation
squirrels (15). Thus, it is possible that the relatively recent
invasion of second growth or disturbed montane forests by G. volans
has affected G. sabrinus survival or competitive abilities and
thus reduced its present range (45).
4) Predators - Northern flying squirrels are eaten by a wide range of
predators and are particularly vulnerable while on the ground. In
the Southern Appalachians, these predators include a variety of
avian raptors, especially owls, and such mammals as bobcats, foxes,
and weasels (44,50).
OTHER LIFE HISTORY DESCRIPTORS:
None.
Life History - 3 (DRAFT) - Management Practices
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
MANAGEMENT PRACTICES
RESULT MANAGEMENT PRACTICE
Beneficial Controlling/Restricting Mining
Beneficial Controlling/Restricting Road Maintainance Actions
Beneficial Land Acquisition
Beneficial Controlling pollution [thermal, chemical, physical]
Beneficial Controlling/Restricting Pesticide Use
Beneficial Controlling/Restricting Herbicide Use
Beneficial Creating Artificial Habitat/Nesting Structure
Beneficial Restricting Timber Harvest
Beneficial Maintaining Later Stages of Succession
Adverse Climate Alteration
Existing Climate Alteration
Adverse Low Gene Pool
Existing Low Gene Pool
Adverse Parasites
Existing Parasites
Adverse Surface Mines
Existing Surface Mines
Adverse Recreational development
Existing Recreational development
Adverse Soil compaction by heavy equipment in mine areas
Existing Soil compaction by heavy equipment in mine areas
Adverse Environmental Contamination/Pollution
Existing Environmental Contamination/Pollution
Adverse Competition
Existing Competition
Adverse Fire
Existing Fire
Adverse Forest Alteration
Existing Forest Alteration
Adverse Harvesting
Existing Harvesting
COMMENTS ON MANAGEMENT PRACTICES -
The two subspecies of Glaucomys sabrinus found in the southern
Appalachians occur as relict populations in mesic, boreal, mixed
forest habitats. During the late Pleistocene such forests covered
much of the Appalachian highlands and it is clear that the range of
this species was correspondingly greater in the past (28,29,40,44).
Just as post-Wisconsin warming has tended to confine boreal vegetation
to high altitudes, north slopes, or cool upland valleys, the range of
G. sabrinus has also contracted so that the species now occupies
disjunct islands of remaining suitable habitat. Such limited range
and small local population size might make a species vulnerable to
stochastic genetic factors (such as drift, progressive homozygosity,
etc.), to continued climatically-driven vegetation change, and to
human disturbance in the form of direct impacts (habitat destruction)
(i.e. forest clearing, forest alteration, forest fire, etc.)
or indirect influences (pollution, and facilitation of competitors and
Management Practices - 1 (DRAFT) - Management Practices
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
pathogens) (04).
In the southern Appalachians habitat destruction and modification
are the major threats to the northern flying squirrel. While the
effects of timber harvest, mining, and recreational development pose
obvious threats to habitats, forest decline associated with
destructive insects such as the balsam wooly aphid (Adelgis piceae)
(03) and gypsy moth (Lymantria dispar) and with the postulated
"acid rain" phenomenon (01,07) must be considered as factors
increasing the vulnerability of this species (51). Finally, it is
possible that human modification of forest habitats has facilitated
the spread of potential competitors such as the southern flying
squirrel, Glaucomys volans, and its parasites (eg. Strongyloides) and
pathogens (15,44,45).
The above paragraphs include the known past, present, and
potential threats to these subspecies. The only additional threat --
at present unsubstantiated - would stem from the bio-magnification
of heavy metals and other contaminants derived from "acid
percipitation". Since Glaucomys sabrinus is a lichen and mycorrhizal
fungus consumer, it could obtain large quantities of certain
contaminants concentrated by its food organisms (04).
UNAPPROVED PLAN:
A Technical Draft is scheduled for completion by 01/01/1988.
A major thrust of the recovery recommendations currently being
developed by the northern flying squirrel recovery team is to
increase our knoweledge of the distribution, life history and ecology
of these little-known subspecies.
Distributional information will be obtained by delineating and
surveying potential habitat. Surveys will be accomplished by
live-trapping and placing nest boxes in potential habitat. Obtaining
data on life history and ecology will require in-depth, long term
studies of known populations, possibly including radio telemetry.
The states of Virginia, West Virginia and North Carolina plan to
cooperate in conducting this study.
Once life history requirements are better understood, the
recovery plan will call for formulating management guidelines, for
use on both public and private lands. Other recovery actions include
developing public education materials, encouraging public acquistion
of occupied habitat and protecting individual squirrels and
populations through law enforcement. Recovery actions may include:
maintaining later stages of succession; restricting timber harvest;
creating artifical nesting structures; controlling/restricting -
herbicide use, pesticide use, pollutants, road maintenance, and
mining; and land acquisition.
On-going recovery activities include nest box surveys in Virginia
and West Virginia, and live-trapping/preliminary ecological studies in
West Virginia
Management Practices - 2 (DRAFT) - References
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
References
***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE *****
01 Adams, H.S., S.L. Stephenson, T.J. Blasing, and D.N. Duvick. 1985.
Growth trend declines of spruce and fir in mis-Appalachian
subalpine forests. Environ. Experim. Bot. 25(4):315-325.
02 Backer, H.R. 1983. Michigan mammals. Mich. St. Univ. Press.,
East Lansing. 642 pp.
03 Barry, P.J. and T.M. Oprean, III. 1979. Evaluation of balsam
wooly aphids on Roan Mountain, Toecane Ranger District, Pisgah
National Forest, North Carolina. U.S. For. Serv., Forest Insect
and Disease Mgmt. Rept. No. 79-1-12, Atlanta, GA. 7 pp.
04 Bogle, M.A. and R.R. Turner. 1984. Lead in vegetation, forest
floor material and soils of the spruce-fir zone, GSMNP. In: White,
P.S. 1984. The southern Appalachian spruce-fir ecosystem: its
biology and threats. Nat. Park Serv., Research/Resource Mgmt.
Rept. Ser-71. Pp. 211-224.
05 Brink, C.H. 1965. Spruce seed as a food of the squirrels
Tamiasciurus hudsonicus and Glaucomys sabrinus in interior Alaska.
Unpubl. MS thesis, Univ. of Alaska, College, AK. 73 pp.
06 Brink, C.H. and F.C. Dean. 1966. Spruce seed as a food of red
squirrels and flying squirrels in interior Alaska. J. Wildl.
Mgmt. 30:503-512.
07 Bruck, R.I. 1984. Decline of montane boreal ecosystems in central
Europe and the southern Appalachian Mountains. Res. and Dev.
Conf., TAPPI, Tech. Park, Atlanta, GA. Pp. 159-163.
08 Connor, P.F. 1960. The small mammals of Otsega and Schoharie
Counties, New York. NY St. Mus. Sci. Serv. Bull. 382:1-84.
09 Cowan, I.M. 1936. Nesting habits of the flying squirrel Glaucomys
sabrinus. J. Mammal. 17:58-60.
10 Davis, W. 1963. Reproductive ecology of the northern flying
squirrel. Unpubl. MA thesis, Univ. Saskatchewan, Canada. 87 pp.
11 DeCoursey, P.J. 1959. Daily activity rhythms in the flying
squirrel, Glaucomys volans. Ph.D. thesis, Univ. Wisconson.
12 DeCoursey, P.J. 1960. Daily light sensitivity rhythm in a rodent.
Science 131:33-35.
13 DeCoursey, P.J. 1961. Effect of light on the circadin activity
rhythm of the flying squirrel, Glaucomys volans. Z. Vergl.
Physiol. 44:331-354.
14 Doutt, J.K., C.A. Heppenstall, and J.E. Guilday. 1977. Mammals
of Pennsylvania. PA Game Comm., Harrisburg. 283 pp.
15 Eckerlin, R.P. 1974. Studies of the life cycle of Strongyloides
robustus Chandler, 1942, and a survey of the helminths of
Connecticut sciurids Unpubl. Ph.D. Diss., Univ. Conn. 116 pp.
16 Ferron, J. 1981. Comparitive activity patterns of two sympatric
sciurid species. Nat. Can. (Que.) 110(2):207-212.
17 Ferron, J. 1983. Scent marking by cheek rubbing in the northern
flying squirrel (Glaucomys sabrinus). Can. J. Zool.
61(11):2377-2380.
18 Ferron, J. and J. P. Ovellet. 1985. Postnatal physical
development in northern flying squirrel, Glaucomys sabrinus. Can.
J. Zool. 63(11):2548-2552.
References - 1 (DRAFT) - References
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
19 Fies, M. 1986-87. Pers. comm. Virginia Game Commission, Stanton,
Va.
20 Grimm, W.C. and R. Whitebread. 1952. Mammal survey of
northeastern Pennsylvania. PA Game Comm., Harrisburg. 82 pp.
21 Hall, E.R. 1981. American flying squirrels. In: The mammals of
North America (2nd ed.). John Wiley & Sons, New York. 1181 pp.
22 Hamilton, W.J., Jr. and J.O. Whitaker, Jr. 1979. Mammals of the
eastern United States (2nd ed.). Cornell Univ. Press, Ithaca,
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23 Handley, C.O., Jr. 1953. A new flying squirrel from the southern
Appalachian Mountains. Biol. Soc. wash., Proc. 66:191-194.
24 Handley, C.O., Jr. 1980. Northern flying squirrel. In: D.W.
Linzey, ed. Endangered and threatened plants and animals of
Virginia. Pp. 513-516. VPI and St. Univ., Balcksburg, Va.
25 Howell, A.H. 1918. Revision of American flying squirrels. N.
Amer. Fauna No. 44:1-64.
26 Jackson, H.H.T. 1961. Mammals of Wisconsin. Univ. of Wisc.
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27 Keith, L.B. and E.C. Meslow. 1966. Animals using runways in
common with snowshoe hares. J. Mammal. 47:541.
28 Kurten, B. and E. Anderson. 1980. Pleistocence mammals of North
America.Colombia Univ. Press. 442 pp.
29 Lundelius, E.L., Jr., R.W. Graham, E. Anderson, J. Guilday, J.A.
Holman, D. Steadman and S.D. Webb. 1983. Terrestrial vertebrate
fauna. In: S.C. Porter, Jr., ed. Late-Quaternary environments
of the United States. Vol I, The Late Pleistocence. Pp. 311-353.
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30 Maser, Z., C. Maser, and J.M. Trappe. 1985. Food habits of the
northern flying squirrel (Glaucomys sabrinus) in Oregan. Can. J.
Zool. 63:1084-1088.
31 McKeever, S. 1952. A survey of West Virginia mammals. W.VA Cons.
Comm. Pittman-Robertson Proj. 22-R. 126 pp.
32 McKeever, S. 1960. Food of the northern flying squirrel in
northeastern California. J. Mammal. 41(2):270-271.
33 Miller, G.S., Jr. 1936. A new flying squirrel from West Virginia.
Proc. Biol. Soc. Wash. 49:143-144.
34 Mowrey, R.A. and J.C. Zasda. 1982. Den tree use and movements of
northern flying squirrels in interior Alaska and implications for
forest management. In: Fish and wildlife relationships in
old-growth forests: Proceedings of a symposium. (April, 1982,
Juneau, AK) (W.R. Meehan, T.R. Merrell, Jr., and T.A. Hanley, eds).
BookMasters, Ashland, Ohio.
35 Mowrey, R.A., G.A. Laursen, and T.A. Moore. 1981. Hypogeous
fungi and small mammal mycophagy in Alaska taiga. Proc. Alaska
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36 Muul, I. 1969. Mating behavior, gestation periods, and
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37 Osgood, D.W. and P.D. Weigl. 1972. Monitoring activity of small
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38 Radvanyi, A. 1959. Inherent rhythms of activity of the northern
flying squirrel in relation to illumination and to lunar and solar
photoperiodism. Unpubl. Ph.D. diss., Univ. British Columbia,
Canada.
References - 2 (DRAFT) - References
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
39 Rapheal, M.G. 1984. late fall breeding of the northern flying
squirrel, Glaucomys sabrinus. J. Mammal. 65:138-139.
40 Semken, H.A., Jr. 1983. Holocene mammalian biogeography and
climatic change in the eastern and central United States. In: H.E.
Wright, Jr., ed. Late-Quaternary environments of the United
States. Vol. II, The Holocene. Pp. 183-207. Univ. Minnesota
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41 Stihler, C., J. Cum, and K. Knight. 1986-87. Pers. comm. West
Virginia Dept. of Natural Resources, Elkins.
42 Thorington, R.W., and L.R. Haeny. 1981. Body proportions and
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43 Urban, V. and E. Michael. 1987. Pers. comm. W. VA Univ.,
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44 Weigl, P.D. 1968. The distribution of the flying squirrels
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exclusion idea. Unpubl. Ph.D. diss., Duke Univ., Durham, NC.
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45 Weigl, P.D. 1975. Parasitism as a possible biological weapon
affecting the ranges and interactions of the flying squirrels,
Glaucomys volans and G. sabrinus. Abstract paper presented at
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46 Weigl, P.D. 1977. Northern flying squirrel. Pp. 398-300. In:
Endangered and threatened plants and animals of North Carolina.
Cooper, J.E. and J.B. Funderburg (eds.). Pp. 398-400. NC St. Mus.
Nat. Hist., Raleigh.
47 Weigl, P.D. 1978. Resource overlap, interspecific interactions
and the distribution of the flying squirrels, Glaucomys volans
and G. sabrinus. Amer. Midl. Nat. 100:83-96.
48 Weigl, P.D. and D.W. Osgood. 1974. Study of the northern flying
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49 Wells-Gosling, N. 1985. Flying squirrels, gliders in the dark.
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50 Wells-Gosling, N. and L.R. Heaney. 1984. Glaucomys sabrinus.
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51 White, P.S. 1984. The southern Appalachian spruce-fir ecosystem:
its biology and threats. Nat. Park Serv., Research/Resources Mgmt.
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***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY *****
01 Fies, M. 1986-87. Pers. comm. Virginia Game Commission, Stanton,
Va.
02 Guilday, J.E. 1971. The Pleistocene history of the Appalachian
mammal fauna. Pp. 223-262. In: The distributional history of the
biota of the southern Appalachians. Part III: Vertebrates (P.C.
Holt, ed.). Virginia Poly. Inst. and State Univ., Blacksburg, Va.
03 Hall, E.R. 1981. American flying squirrels. In: The mammals of
North America (2nd ed.). John Wiley & Sons, New York. 1181 pp.
04 Handley, C.O., Jr. 1953. A new flying squirrel from the southern
Appalachian Mountains. Biol. Soc. wash., Proc. 66:191-194.
References - 3 (DRAFT) - References
Species SQUIRREL, FLYING, NORTHERN, VIRGINIA
Species Id ESIS055002
Date 14 MAR 96
05 Handley, C.O., Jr. 1980. Northern flying squirrel. In: D.W.
Linzey, ed. Endangered and threatened plants and animals of
Virginia. Pp. 513-516. VPI and St. Univ., Balcksburg, Va.
06 Kurten, B. and E. Anderson. 1980. Pleistocene mammals of North
America. Colombia Univ. Press. 442 pp.
07 Linzey, D.W. 1983. Status and distribution of the northern water
shrew (Sorex palustris) and two subspecies of northern flying
squirrel (Glaucomys sabrinus coloratus and Glaucomys sabrinus
fuscus). Final Rept. Submitted to the U.S. Fish and Wildl. Serv.
Contract No. 14-16-0005-79-068.
08 Lundelius, E.L., Jr., R.W. Graham, E. Anderson, J. Guilday, J.A.
Holman, D. Steadman and S.D. Webb. 1983. Terrestrial vertebrate
fauna. In: S.C. Porter, Jr., ed. Late-Quaternary environments
of the United States. Vol I, The Late Pleistocence. Pp. 311-353.
Univ. Minnesota Press, Minneapolis.
09 McKeever, S. 1960. Food of the northern flying squirrel in
northeastern California. J. Mammal. 41(2):270-271.
10 Miller, G.S., Jr. 1936. A new flying squirrel from West Virginia.
Proc. Biol. Soc. Wash. 49:143-144.
11 Semken, H.A., Jr. 1983. Holocene mammalian biogeography and
climatic change in the eastern and central United States. In: H.E.
Wright, Jr., ed. Late-Quaternary environments of the United
States. Vol. II, The Holocene. Pp. 183-207. Univ. Minnesota
Press, Minneapolis.
12 Stihler, C., J. Cum, and K. Knight. 1986-87. Pers. comm. West
Virginia Dept. of Natural Resources, Elkins.
13 Weigl, P.D. 1977. Northern flying squirrel. Pp. 398-300. In:
Endangered and threatened plants and animals of North Carolina.
Cooper, J.E. and J.B. Funderburg (eds.). Pp. 398-400. NC St. Mus.
Nat. Hist., Raleigh.
14 Wells-Gosling, N. and L.R. Heaney. 1984. Glaucomys sabrinus.
Mammalian Species 247. Am. Soc. of Mammalogists.
References - 4