Journal of Mammalogy, 87(4):784–789, 2006

COMPARISON OF DIETS BETWEEN TWO SYMPATRIC

FLYING SQUIRREL SPECIES IN NORTHERN PAKISTAN
CHAUDHRY M. SHAFIQUE, SOHAIL BARKATI, TATSUO OSHIDA,* AND MOTOKAZU ANDO
Department of Zoology, University of Karachi, Karachi 75270, Pakistan (CMS, SB)
Laboratory of Wildlife Ecology, Obihiro University of Agriculture and Veterinary Medicine,

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Obihiro 080-8555, Japan (TO)
Laboratory of Wild Animals, Faculty of Agriculture, Tokyo University of Agriculture,
Atsugi 243-0034, Japan (MA)

To test the association between body size and degree of folivory and to examine the different food habits in
sympatric arboreal herbivorous mammals, we investigated diets of 2 flying squirrel species (Petaurista petaurista
and Eoglaucomys fimbriatus) sympatrically distributed in the Himalayan moist temperate forest of northern
Pakistan. P. petaurista and E. fimbriatus used 27 and 28 tree species, respectively. Both flying squirrel species
used the same 27 tree species. However, frequency of tree species used by P. petaurista was significantly
different from that used by E. fimbriatus. P. petaurista was significantly more folivorous than was E. fimbriatus.
Preference of P. petaurista for leaves may be an important factor preventing competition for resources between
these species. We conclude that larger body size in flying squirrels is associated with the tendency toward
a foliage diet.

Key words: diet, Eoglaucomys fimbriatus, folivory, Kashmir flying squirrel, northern Pakistan, Petaurista petaurista
albiventer, red giant flying squirrel, sympatric distribution

Body size of mammals is associated with their food habits
(e.g., Brown and Nicoletto 1991; Jxdrzejewski et al. 1989).
For instance, in ungulates, there are 3 feeding types: grazing,
browsing, and both grazing and browsing (Hoffman 1985, 1988,
1989; Hoffman and Stewart 1972). These feeding types are well
correlated with body size or weight (Gordon and Illius 1988;
Hobbs et al. 1982; Hoffman 1989), although a few exceptions
are recognized. The grazers (e.g., Bubalus bubalis, Syncerus
caffer, and Bison bonasus) are larger, mainly eating many
grasses of low nutritional quality that are processed in their
ruminant stomachs. Browsers, such as Capreolus capreolus,
Odocoileus hemionus, and Tragulus javanicus, have a smaller
body size and prefer browsing on shrubs of high nutritional
quality. Intermediate in size to grazers and browsers are
Aepyceros melampus, Gazella granti, and Ovis canadensis
(Hoffman 1989). To maintain their larger bodies, grazers effec-
tively use large quantities of grass, the most abundant resources
in their habitats. Smaller browsers cannot process so much grass,
but instead use less abundant but more nutritious resources.
* Correspondent: oshidata@obihiro.ac.jp
Ó 2006 American Society of Mammalogists
www.mammalogy.org
784
For arboreal herbivorous mammals, leaves may correspond
to grass in ungulates. Leaves are a very abundant, but less-
nutritious resource. Muul and Lim (1978), in their study of diets
of several genera of flying squirrels (Petinomys, Hylopetes,
Pteromyscus, Aeromys, and Petaurista) from Malaysia, sug-
gested that larger flying squirrel species tend to include more
foliage in their diet than do smaller flying squirrel species.
The Kashmir flying squirrel (Eoglaucomys fimbriatus),
endemic to Kashmir and Punjab, and the red giant flying
squirrel (Petaurista petaurista albiventer), an endemic sub-
species of Pakistan (and probably Afghanistan), are sympatri-
cally distributed in Himalayan moist temperate forests ranging
from 1,350 to 3,050 m in elevation (Corbet and Hill 1992;
Hoffmann et al. 1993; Roberts 1997). Although P. petaurista
albiventer is currently regarded as a distinct species (P.
albiventer) on the basis of molecular evidence (Oshida et al.
2004; Yu et al. 2006), to avoid any taxonomic confusion, we
here treat this flying squirrel as a subspecies of P. petaurista,
according to Wilson and Reeder (1993). Both flying squirrel
species are arboreal and nocturnal and have similar ecological
traits (Roberts 1997) resulting from their specialized gliding
behavior. These 2 species differ greatly in body size, with
P. petaurista (1,100–1,700 g) being 2–3 times larger than
E. fimbriatus (500–666 g—Roberts 1997). Therefore, these
2 species may exploit different foods in the Himalayan moist
temperate forest. We tested the hypothesis (Muul and Lim
August 2006 SHAFIQUE ET AL.—DIETS OF GIANT FLYING SQUIRRELS
FIG. 1.—Preferences of plant species eaten by Petaurista petaurista albiventer (gray) and Eoglaucomys fimbriatus (white) in the Himalayan
moist temperate forests of northern Pakistan. Asterisks indicate plant species not eaten.
1978) that the large species (P. petaurista) should consume sub-
stantially more foliage than the smaller species (E. fimbriatus),
which is predicted to eat more fruits and seeds.
MATERIALS AND METHODS
Feeding behavior of flying squirrels (E. fimbriatus and P. petaurista
albiventer) was observed within 5 km2 of Ayubia National Park
(348549N, 738229E), northern Pakistan, from March 1997 to February
2000 (10–15 nights per month) and for 1 week in March 2002. Total
number of nights of observation was 536. The study area was covered
with Himalayan moist temperate forests composed mainly of Acer
caesium, Pinus wallichiana, Populus cilliata, Quercus dilatata, and
Taxus baccata. The elevation of the study area was 2,439–2,591 m in
elevation. From January 1998 to December 1999, annual precipitation
ranged from 1,806 to 1,920 mm, and minimum and maximum
temperatures were about
58C to 128C and 128C to 298C. To find
flying squirrels, we walked at random along forest paths at night from
sunset to midnight or 0100 h. To watch the feeding behavior of flying
squirrels, we used binoculars (7  35, Pentax, Tokyo, Japan) with 6-V
searchlights. We recorded tree species and food items. Because it was
difficult to exactly evaluate other indices such as time eating each item
or the volume of items in our observation, we examined the percentage
of food items. A feeding record consisted of eating a single food item
until feeding was finished (Ando et al. 1985; Kawamichi 1997),
regardless of the time spent feeding or the quantity of food eaten. We
categorized foods into 9 items: cone, flower and floral bud, fruit, leaf,
lichen and bark, moss, seed, winter bud and shoot, and trunk gall.
Spearman rank correlation was calculated to assess correlation
between proportions of tree species used by each flying squirrel
species. In comparison between 2 flying squirrel species, chi-square
tests were used to determine significance of proportions of tree species
785
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used by each squirrel species and significance of proportions of tree
items used by each squirrel species. Statistical analyses were
computed using JMP IN version 5 (SAS Institute Inc. 2004) with
a significance level of 5%.
RESULTS
Feeding behavior of P. petaurista was observed 2,931 times
and that of E. fimbriatus 2,411 times. P. petaurista and
E. fimbriatus used 27 and 28 tree species, respectively (Fig. 1).
Both species used the same 27 tree species. Ranked use of the
tree species by the 2 species was highly correlated (r ¼ 0.82,
P , 0.001). However, the proportion of tree species used by
P. petaurista was significantly different from that used by
E. fimbriatus (v2 ¼ 523.30, d.f. ¼ 27, P , 0.001). Both flying
squirrel species frequently (75.0% of total diet) used 4 tree
species (Abies pindrow, P. wallichiana, P. cilliata, and Ulmus
wallichiana; Fig. 1). P. petaurista frequently used Quercus
floribunda (12.3% of total diet), but it was only 3.8% of total
diet of E. fimbriatus. Although Aesculus indica was well used
by E. fimbriatus (10.7%), it occupied only 4.5% of total diet of
P. petaurista. Pyrus pashia was only used by E. fimbriatus
(Fig. 1).
Both flying squirrel species used variable and seasonal tree
items (Tables 1 and 2; Fig. 2). Floral buds and flowers were
mainly eaten in the spring. Fruits were used in summer and
autumn. Winter buds and shoots were eaten in winter.
However, total item preference between 2 species was
significantly different (v2 ¼ 547.32, d.f. ¼ 8, P , 0.001).
P. petaurista was more folivorous, mainly using leaves of
786 JOURNAL OF MAMMALOGY Vol. 87, No. 4

TABLE 1.—Distribution of food items among various plants parts and food species in the diet of Petaurista petaurista albiventer in the
Himalayan moist temperate forests of northern Pakistan. Values indicate number of items of each type.

Flowers and Lichens Winter shoots

Species Cones floral buds Fruit Galls Leaves and bark Moss Seeds and buds Total
Conifers
Abies pindrow 31 74 13 14 87 34 253
Cedrus deodara 26 19 33 5 83
Picea smithiana 7 12 14 33
Pinus wallichiana 56 87 315 54 13 210 63 798
Taxus wallichiana 18 18

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Broad-leaved trees
Acer caesium 28 14 6 48
Acer pictum 4 4 5 13
Aesculus indica 22 88 23 133
Cormus macrophylla 17 119 7 143
Corylus colurna 3 3
Cotoneaster bacilliaris 5 13 18
Diospyros lotus 2 7 6 15
Ficus palmata 1 1
Juglans regia 6 37 43
Morus serrata 15 4 19
Populus cilliata 57 16 8 61 69 211
Prunus armeniaca 17 17
Prunus cornuta 19 14 70 103
Prunus persica 15 3 18
Pyrus malus 14 14
Quercus floribunda 34 86 22 161 56 359
Quercus glauca 17 42 11 76 13 159
Quercus incana 6 83 89
Rhododendron arboreum 8 12 20
Salix tetrasperma 7 24 11 42
Ulmus wallichiana 24 142 5 11 36 218
Climbers
Hedera nepalensis 14 31 6 51
Rosa moschata 1 8 9

P. wallichiana, U. wallichiana, and Q. floribunda (Table 1). In
contrast, E. fimbriatus preferred seeds, fruits, flowers, and floral
buds, especially seeds of A. indica and P. cilliata and floral
buds of P. wallichiana (Table 2). Only E. fimbriatus used trunk
galls of P. wallichiana, and it did so only infrequently.
DISCUSSION
Dietary differences of 2 flying squirrel species.— The
2 flying squirrel species exploited almost the same tree species
in the Himalayan moist temperate forests of northern Pakistan
(Fig. 1). Both squirrel species also used seasonal tree items such
as buds, cones, seeds, and flowers (Tables 1 and 2). Because of
the similarities in diet, these 2 species may compete for food.
However, their diets differed significantly in the proportion
(in percentage) of tree parts used (Fig. 1). Leaf items occupied
20.6% of the diet of P. petaurista but only 2.3% of that of E.
fimbriatus (Fig. 2). The proportion of seeds, flower and flower
buds, and fruits was greater in E. fimbriatus than in P. petaurista
(Fig. 2). In contrast to the folivory of P. petaurista, E. fimbriatus
was more likely to use these seasonal items.
There are few reports of diet preferences of sympatric flying
squirrel species. In a study of food preference of 2 giant flying
squirrel species that are sympatric in Taiwan, Lee et al. (1986)
reported that Petaurista philippensis grandis fed on more tree
species than Petaurista alborufus lena. In North America,
2 marginally sympatric Glaucomys species essentially used
different resources (Weigl 1978; Wells-Gosling 1985). Glauc-
omys volans is a granivore (e.g., Muul 1968) and G. sabrinus is
a fungivore (e.g., McKeever 1960) that also uses a small
amount of other diet items such as seeds and nuts (Carey 2000;
Thysell et al. 1998). Thus, the use of different foods is an
effective strategy to divide restricted forest resources among
sympatric flying squirrel species. Sympatric mammalian spe-
cies with similar ecological characteristics often are considered
to compete for resources (e.g., Vaughan et al. 2000). To avoid
competition (exploitation), sympatric mammalian species
should partially or completely differ in their resource
preferences, as in the cases of primates on Sumatra Island
(Ungar 1995), rodents living in the Arizona desert (Brown and
Harney 1993), and carnivores in India (Karanth and Sunquist
1995) and the United States (Neale and Sacks 2001). In the
Himalayan moist temperate forest of northern Pakistan, the
August 2006 SHAFIQUE ET AL.—DIETS OF GIANT FLYING SQUIRRELS 787

TABLE 2.—Distribution of food items among various plant parts and food species in the diet of Eoglaucomys fimbriatus in the Himalayan moist
temperate forests of northern Pakistan. Values indicate number of items of each type.

Flowers and Lichens Winter shoots

Species Cones floral buds Fruit Galls Leaves and bark Moss Seeds and buds Total
Conifers
Abies pindrow 38 77 36 27 178
Cedrus deodara 11 18 22 13 64
Picea smithiana 14 12 11 9 46
Pinus wallichiana 14 164 5 13 34 13 144 58 445
Taxus wallichiana 103 103

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Broad-leaved trees
Acer caesium 31 21 8 60
Acer pictum 9 6 15
Aesculus indica 25 222 11 258
Cormus macrophylla 18 112 8 138
Corylus colurna 6 10 16
Cotoneaster bacilliaris 7 21 28
Diospyros lotus 13 8 21
Ficus palmata 4 4
Juglans regia 12 82 94
Morus serrata 6 5 11
Populus cilliata 45 38 11 103 55 252
Prunus armeniaca 17 17
Prunus cornuta 14 10 70 11 105
Prunus persica 8 8
Pyrus malus 16 16
Pyrus pashia 22 22
Quercus floribunda 14 5 56 16 91
Quercus glauca 5 7 25 16 53
Quercus incana 26 26
Rhododendron arboreum 11 8 19
Salix tetrasperma 8 44 19 71
Ulmus wallichiana 37 31 15 53 46 182
Climbers
Hedera nepalensis 12 21 27 60
Rosa moschata 8 8

preference of P. petaurista for leaves may prevent competition
for resources between it and E. fimbriatus.
Association between body size and degree of folivory.—
Leaves are generally a low-quality food resource (e.g.,
Wischusen 2000); nevertheless, Petaurista species have
a strong preference for foliage. Leaves account for 40% of
the diet of the Japanese giant flying squirrel (Petaurista
leucogenys—Ando and Imaizumi 1982) and leaves and buds
make up 36% of the diet of P. leucogenys (Kawamichi 1997).
Leaves are frequently used by P. leucogenys (Baba et al. 1982)
and leaves were eaten in 61% of feeding observations of
P. philippensis grandis and 71% of observations of
P. alborufus lena (Lee et al. 1986).
On the other hand, with respect to small-sized flying
squirrels, the diet of the Siberian flying squirrel (Pteromys
volans), widely distributed throughout northern Eurasia,
includes leaves, seeds, buds, and catkins (Airapetyants and
Fokin 2003; Hanski et al. 2000), although proportions are not
known. G. volans in North America is a typical granivore
(Harlow and Doyle 1990; Weigl 1978), and Iomys, Hylopetes,
Petaurillus, and Petinomys, distributed in Southeast Asia, are
recognized as mainly being granivores (Muul and Lim 1978).
Miyao (1972) suggested, based on its longer colon and
cecum, that P. leucogenys could be a highly developed
folivore. Muul and Lim (1978), from their study of diets of
several flying squirrel species (Petinomys, Hylopetes, Ptero-
myscus, Aeromys, and Petaurista) from Malaysia, suggested
that larger flying squirrel species tend to include more foliage
in their diet than smaller flying squirrel species. They also
observed that cecum length is relatively greater in the larger
flying squirrel species. The body size of Petaurista species,
700–1,500 g (e.g., Corbet and Hill 1992; Lekagul and McNeely
1988), is larger than that of other flying squirrel species.
Therefore, our observations of folivorous tendency in P.
petaurista support the hypothesis of Muul and Lim (1978). The
woolly flying squirrel (Eupetaurus cinereus), which weighs
1,400–2,500 g, is the most massive of the gliding mammals
(Zahler 2001; Zahler and Woods 1997). Although its exact diet
is still unclear, Zahler and Khan (2003) indicated that, based on
fecal sample analysis, this species feeds mostly or entirely on
pine needles. That report also would support a folivorous
tendency in larger flying squirrel species.
There are few gliding mammalian folivores other than
species of Petaurista. The greater glider (Petauroides volans),
788 JOURNAL OF MAMMALOGY
FIG. 2.—Proportion of food items used by Petaurista petaurista albiventer and Eoglaucomys fimbriatus in the Himalayan moist temperate
forests of northern Pakistan. Trunk galls are not shown, because there were only 5 observations of use by Eoglaucomys fimbriatus and none by
Petaurista petaurista.
which feeds almost exclusively on the foliage of Eucalyptus,
weighs approximately 1,400 g (Goldingay 2000; Kavanagh and
Lambert 1990). The Philippine flying lemur (Cynocephalus
volans), a highly selective folivore, weighs 1,000–1,800 g
(Wischusen 2000; Wischusen and Richmond 1998). All these
gliding mammals are both large and folivorous. Because
P. petaurista albiventer is much larger than E. fimbriatus, we
were not surprised to find it included more foliage in its diet.
Interactions between large body size and the tendency
toward a foliage diet in gliding mammals such as flying
squirrels deserve greater attention. There is strong evidence that
the division of food resources by gliding mammals parallels
that of ungulates, which have 3 feeding types: grazing, brows-
ing, and both grazing and browsing (Hoffman 1985, 1988,
1989; Hoffman and Stewart 1972). For both gliding mammals
(Goldingay 2000; Kavanagh and Lambert 1990; Muul and Lim
1978; Wischusen 2000; Wischusen and Richmond 1998; this
study) and ungulates (Gordon and Illius 1988; Hobbs et al.
1982; Hoffman 1989), there is a correlation between body size
and food habits. Therefore, studies of locations with 3 or more
species of gliding mammals may reveal a stronger correlation
with patterns observed in ungulates.
It is worth examining how much of the food preferences of
P. petaurista and E. fimbriatus truly depend on competition
between these 2 species. Food preferences of these species also
may vary in locations where 1 species is absent. In contrast,
P. petaurista may well be an obligate folivore, supplementing
its leaf diet with the occasional fruit or seed. In that case,
evidence for this also will appear in the structure and function
of its digestive organs.
ACKNOWLEDGMENTS
We thank M. M. Malik (Department of Wildlife and National Park,
Northwest Frontier Province, Pakistan) for his courtesy for our study.
We are grateful to I. Shah, S. Ahmad, S. Hussain, Riaz, and G. Ahmad
for their assistance to the research project in Pakistan. We thank
C. L. Bridgman for her critical reading of the manuscript.
Vol. 87, No. 4
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Submitted 29 June 2005. Accepted 20 January 2006.
Associate Editor was Gerardo Ceballos.