Larter&Nagy.indd
131Larter & Nagy 2004: Polar Research 23(2), 131–140
Caribou (Rangifer tarandus) and muskoxen
(Ovibos moschatus) are the only ungulate spe-
cies successfully occupying Arctic tundra envi-
ronments. Caribou and muskoxen have differ-
ent morphological adaptations which presumably
enabled them to utilize forage resources with little
overlap (Klein 1992). Caribou, with their small
body size, narrow muzzle, small gut capacity,
and rapid passage rate (Tyler & Blix 1990; Klein
1992), are representative of a mixed feeder type,
intermediate between roughage feeders and con-
centrate selectors according to Hofmann’s (1989,
2000) classifi cation. Muskoxen, with their large
body size, broad muzzle, and large gut capac-
ity capable of processing large amounts of low
quality forage (Klein 1992) are representative
of roughage feeders (grazers) according to Hof-
mann’s (1989, 2000) classifi cation.
A number of studies have reported the diets
of muskoxen and caribou in Arctic Canada (e.g.
Wilkinson et al. 1976; Parker 1978; Shank et al.
1978; Thomas & Edmonds 1983; Oakes et al.
1992; Larter & Nagy 1997), Alaska (e.g. Svend-
sen 1992; Ihl 1999; Ihl & Klein 2001), and Green-
land (Thing 1984; Thing et al. 1987; Klein & Bay
1990). Many diet studies have shown willow (Salix
spp.) to be a noticeable component of the muskox-
en diet regardless of season (e.g. Wilkinson et
al. 1976; Thing 1984; Svendsen 1992; Larter &
Nagy 1997; Thomas et al. 1999). In early summer,
willow use was prominent among muskoxen on
Banks Island (Larter & Nagy 1997). Willow is
also a prominent component of their late winter
diet especially in years of deeper and harder snow
(Larter & Nagy 1997; Thomas et al. 1999).
Lichen is an important winter food for barren-
ground caribou on the mainland, but in the High
Arctic islands, which support low lichen biomass,
caribou must use other forages, usually willow
and graminoids (Reimers et al. 1980; Klein 1992).
Where food choices are limited or muskox densi-
ty is high, the two species may compete for food
Seasonal changes in the composition of the diets
of Peary caribou and muskoxen on Banks Island
Nicholas C. Larter & John A. Nagy
Caribou and muskoxen are the only ungulate species occupying Arctic
tundra environments. We analysed plant fragments found in fresh (< 4 hr
old) samples of faecal material to determine the diets of Peary caribou
(Rangifer tarandus pearyi) and muskoxen (Ovibos moschatus) on Banks
Island, Canada. Willow was a major component of the diets of both ani-
mals, dominating the caribou diet during summer and representing sub-
stantial proportions of the muskoxen diet during at least seven months of
the year. The diet of caribou was more diverse than that of muskoxen and
was dominated by sedge, willow (Salix arctica), legume (Astragalus spp.,
Oxytropis spp.) and Dryas integrifolia. The diet of muskoxen was domi-
nated by sedge and willow. There was substantial overlap (up to 70 %) in
the diets of these herbivores with the similarity more pronounced in areas
of high muskox density (ca. 1.65 animals/km2). We discuss herbivore
diets in relation to foraging behaviour and forage availability.
N. C. Larter, Government of the Northwest Territories, Department of Resources, Wildlife & Economic
Development, Box 240, Ft. Simpson, NT, Canada X0E 0N0, nic_larter@gov.nt.ca; J. A. Nagy, Government
of the Northwest Territories, Department of Resources, Wildlife & Economic Development, Bag Service #1,
Inuvik, NT, Canada X0E 0T0.
132 Seasonal changes in composition of diets of Peary caribou and muskoxen
(McKendrick 1981). On Banks Island in the Cana-
dian High Arctic, lichen biomass is low and the
muskox population represents a substantial pro-
portion of worldwide muskoxen numbers, with
some of the highest reported densities of muskox,
1.5 - 2.0 animals/km2 (Nagy et al. 1996; Larter
& Nagy 2001a). During the 1990s Peary caribou
numbers on Banks Island were at historic lows,
500 - 1000 animals, excluding calves (Nagy et al.
1996). Peary caribou rely on willow during the
short Arctic summers in order to regain fat prior
to winter and there was concern that the availabil-
ity of willow during summer may be impacted by
the increased annual use of willows by herbivores
(Larter & Nagy 1997; Larter 1999).
In order to describe, in detail, the seasonal diet
composition of Peary caribou and muskoxen on
Banks Island we embarked on a program to col-
lect fresh faecal samples deposited on known
dates during every month of the year. We hypoth-
esized that muskoxen would feed on willow to a
greater extent during the growing season when
willow was of higher nutritive quality (Larter
& Nagy 2001b) and/or during late-winter when
the availability of sedge (Carex spp., Eriopho-
rum spp.) would be lowest, due to the combina-
tion of grazing and snow conditions (Larter &
Nagy 2000, 2001c). We also hypothesized that
the diet composition of muskox would be dif-
ferent in areas of different muskox density. This
paper documents the monthly diets Peary cari-
bou, the monthly diets of muskoxen residing in
areas of both high and low muskox density, the
percent similarity in diet between the two ani-
mals, and relates the monthly changes in diet to
foraging behaviour.
Study area
Banks Island is the westernmost island in the
Canadian Arctic archipelago and covers approxi-
mately 70 000 km2 (Fig. 1). The climate is Arctic
maritime along coastal areas where weather sta-
tions are located, tending toward Arctic desert
inland (Zoltai et al. 1980). Winters are long and
cold; mean monthly temperatures are below 0 °C
from September through May; mean minimum
daily temperatures range from –30 °C to –40 °C
from December to March. Summers are short and
cool; mean maximum daily temperatures range
from 5 °C to 10 °C from June through August.
There is little precipitation; the annual mean is
90 mm (Zoltai et al. 1980). Sachs Harbour (pop-
ulation 125) is the only permanent settlement on
the Island.
Habitat descriptions were adapted from Kevan
(1974), Wilkinson et al. (1976), and Ferguson
(1991). There are four major terrestrial habitats:
1) wet sedge meadow, 2) upland barren, 3) hum-
mock tundra, and 4) stony barren. Muskoxen and
caribou forage in all habitats. Wet sedge mead-
ows (WSM) are generally level hydric and hygric
lowlands characterized by Carex aquatilis, Eri-
ophorum scheuchzeri, and Dupontia fi sheri.
Upland barrens (UB) are well drained sites found
on the upper and middle parts of slopes. Vegeta-
tion is dominated by Dryas integrifolia and Salix
arctica. Hummock tundra (HT) is found on mod-
erately steep slopes and is characterized by indi-
vidual hummocks which are vegetated primarily
by dwarf shrubs (D. integrifolia, S. arctica, with
some Cassiope tetragona). Stony barrens (SB)
have a coarse gravelly substrate and are found
in windblown areas, on ridges and on gravel and
sand bars; vegetation is sparse.
The areas of high and low muskox density
where samples were collected for this study cover
2700 and 18 400 km2, respectively, and have had
distinctly different densities of muskox since 1985
(Nagy et al. 1996). Similar vegetation types and
vegetation of similar quality are found through-
out both areas (Larter & Nagy 2001b). Howev-
er, in the low density area, vegetated habitats are
more patchily distributed with SB being more
prominent (Larter & Nagy unpubl. data) and late-
winter snow conditions tend to be harsher (Larter
& Nagy 2001c).
A more detailed description of the fl ora of
Banks Island can be found in Wilkinson et al.
(1976), Porsild & Cody (1980), and Zoltai et al.
(1980). Plant taxonomy follows Kartesz (1999).
Methods
Two fi eld camps were established in June 1993
on south central Banks Island (Fig. 1): one camp
(Coyote) is located in an area of high muskox
density (1.6 - 1.9 muskox/km2) about 90 km ESE
of Sachs Harbour, and the other (Bernard) in an
area of low muskox density (0.3 - 0.4 muskox/
km2) about 130 km ENE of Sachs Harbour (Fig.
1). From June 1993 to May 1998 we made six trips
to these camps annually in June, July, August,
November - December, February, and April - May.
133Larter & Nagy 2004: Polar Research 23(2), 131–140
Fig. 1. The study area, Banks
Island, Northwest Territories,
Canada. The hatched areas
indicate where samples were
collected from the high muskox
density (western) and low
muskox density (eastern) areas.
The location of the Coyote and
Bernard camps is indicated.
134 Seasonal changes in composition of diets of Peary caribou and muskoxen
We collected fresh (< 4 hr old) Peary caribou
faecal samples opportunistically during all these
fi eld trips. Additional faecal samples were col-
lected during fi eld trips in March and May 1993
and January 2001, from animals killed by hunt-
ers (since 1990), and from animals taken during a
collection in winter 1993–94. In total 298 samples
were analysed from all months, range of 10 to 49
samples/month. We present monthly diet compo-
sition pooled across sex and age classes and years
with the individual caribou as the sample unit.
We assume inter-annual variation was minimal.
Previous analyses (Larter & Nagy 1997) indi-
cated that between-group variation was greater
than within-group variation in the diet of muskox.
Therefore, we assumed that a composite sample
of fresh (< 4 hr old) muskox faeces from a number
of individuals in a group was representative of
that group of muskox and that by sampling groups
instead of individuals we would get a better meas-
ure of the diet over a larger portion of the muskox
population. Thus, we used the composite sample
from a muskox group as the sample unit. Initial-
ly (March 1993) the sample unit consisted of fi ve
pellet groups; this was reduced to three pellet
groups/muskox group. Small pellets (i.e. from
calves) were not included in composite samples.
We collected faecal samples from mixed sex and
age groups during fi eld trips conducted between
March 1993 and May 1998 and during September,
October, and December 2000 and January 2001.
This provided data for all months of the year from
the high density area, and for all months except
January, September and December in the low
density area. We recorded the number of indi-
viduals present in each group we collected faecal
samples from. The number of groups sampled per
month during the sampling period ranged from 2
to 29, representing between 63 and 634 animals
in the high density muskox area, and from 1 to
23, representing between 30 and 371 animals, in
the low density muskox area. We present mean
monthly diet composition of muskoxen pooled
across years with the group as the sample unit,
weighted by the number of individuals in a group.
January and September data from the high den-
sity area and May and October data from the low
density area are from one year only. We assume
inter-annual variation was minimal.
Faecal samples were thawed, air dried for 24 hr,
oven dried at 60 °C for 48 hr, and ground through
a 1 mm screen with a centrifugal mill. Subsamples
(ca. 1 g dry weight) were forwarded to the Com-
position Analysis Laboratory, Ft. Collins, Colora-
do, for analysis. Diet composition was determined
by analysing plant fragments (Sparkes & Malech-
ek 1968) according to Hansen et al. (1976). The
microhistological technique has inherent limits,
such as an inability to separate some species, and
a limited percent of identifi able fragments in the
slides (Johnson et al. 1983; Barker 1986). How-
ever, we deemed this method suitable for this
study, since we were interested in the proportion-
al dietary contribution of different forage classes,
not of individual species. Moreover, this method
had been used in previous work on Banks Island
(Oakes et al. 1992; Larter & Nagy 1997). The diet
was subdivided into the following forage class-
es: sedge (Cyperaceae), willow (predominant-
ly Salix arctica), grass (Poaceae), rose/saxifrage
(Rosaceae and Saxifragaceae but predominant-
ly Dryas integrifolia), legume (Oxytropis spp.,
Astragalus spp.), fruticose lichen (Cetraria spp.,
Cladonia spp., Cladina spp., Peltigera spp., and
Thamnolia subuliformis), and other (other forbs,
moss, and Equisetum spp.). No samples had com-
pletely unidentifi able material. There were traces
(≤ 1.93 %) of unidentifi able grass, and forb mate-
rial. These were included in their respective cat-
egories. Trace (≤ 0.1 %) quantities of rush were
included as grass.
We used Simpson’s Index (SI) (Krebs 1989) as
a measure of monthly diet diversity. Annual diet
diversity was calculated as the mean of monthly
SI over twelve months for caribou and muskox-
en in high density areas, and over eight months
for muskoxen in low density areas. We used the
Renkonen index (Renkonen 1938; Krebs 1989)
to compare monthly percent diet similarity (PS)
between Peary caribou and muskoxen in high
and low muskox density areas. This index deter-
mines PS by summing the lower percentage of
each individual forage class in the diets being
compared. We compared monthly diet similarity
between Peary caribou and muskox both within
and between months. We used the same forage
classes as above except that the sedge class was
subdivided into Carex spp. and Eriophorum spp.
and that the moss component was reported sepa-
rately from the other category. We considered PS
> 50 % as substantial diet similarity. We used the
sign test (Conover 1980) to compare the amounts
of willow and sedge in the diet of muskoxen in
low and high density areas.
135Larter & Nagy 2004: Polar Research 23(2), 131–140
Results
The annual diet of caribou, Simpson’s Index (SI)
of 0.704, was more diverse than that of muskox-
en in both high (SI 0.433) and low (SI 0.314) den-
sity areas, and was dominated by four forage
classes: sedge, willow, legume, and rose/saxi-
frage (predominantly Dryas integrifolia) which
varied in their monthly contribution to the diet
(Fig. 2a). Willow was the dominant dietary com-
ponent during summer (June-August). Legumes
and Dryas integrifolia represented ≥ 50 % of the
diet from October to May. The sedge component
of the diet was generally between 15 and 25 %
for all months except January, April, May and
July. Lichen and grass use was negligible (≤ 5 %)
except during January. Diet diversity was great-
est in January and September (SI 0.811 and 0.809,
respectively), when the four dominant forage
classes made up < 75 % of the diet: 74 % in Janu-
ary and 66 % in September (Fig. 2a).
The annual diet of muskoxen in both high and
Fig. 2. Monthly diet composi-
tion, mean percent relative
density of faecal plant frag-
ments, of a) Peary caribou, b)
muskoxen in high density areas,
and c) muskoxen in low density
areas. Values over each histobar
are the number of samples.
For caribou, samples represent
individuals. For muskoxen,
samples represent groups. An
asterisk indicates data from one
year only.
136 Seasonal changes in composition of diets of Peary caribou and muskoxen
low density areas was dominated by two forage
classes, sedge and willow (Fig. 2b, c). Sedge
and willow made up ca. 95 %, or more, of the
diet in all months except July, when legume use
was noticeable. Muskoxen in high density areas
tended to have somewhat more willow and less
sedge (P < 0.01; 1-tailed) in their monthly diets
when compared to muskox in low density muskox
areas (Fig. 2b, c). In high density areas, substan-
tial willow use (48 - 75 %) occurred during fi ve
months of the year: January, May, June, Septem-
ber and October. In low density areas, willow use
was highest during October (46 %), May (25 %),
and June (20 %); there were no data for January
and September.
Only in June was there substantial overlap
(71 %) in diet between Peary caribou and muskox
in high density areas. However, there was sub-
stantial overlap between the summer (June - Au-
gust) diet of caribou and the fall diet (September
and October) of muskox and between the July diet
of muskox and the February, March and June diet
of caribou (Table 1). The April diet of Peary car-
ibou, with its low sedge and willow content, was
the least similar to the diet of muskoxen. Similar
patterns of diets were found for Peary caribou and
muskox in low density areas (Table 2). For any
month, the diets of muskoxen in high and in low
density areas had substantial similarity (Table 3).
In the three months with lowest similarity (May
57 %, June 51 % and October 67 %) the greater
use of willow by muskoxen in high density areas
relative to those in low density areas caused the
reduction in diet similarity (Table 3; Fig. 2b, c).
Discussion
At the start of the growing season the levels of
anti-herbivore defence compounds in deciduous
species are low (Chapin et al. 1980) and willow
leaves are of their highest quality (Murray 1991;
Larter & Nagy 2001b). Muskoxen grazed almost
exclusively on willow leaves during the start of
the growing season at Sverdrup Pass, Ellesmere
Island (Murray 1991; Raillard 1992). Similarly,
we found high use of willow by muskoxen during
May and June, consistent with the hypothesis that
foraging on willow occurs during the early growth
stages when leaves and young shoots are emerg-
ing and plants are of high nutritive quality. We
did not expect muskoxen would feed on willow
to such an extent during the fall (September and
October) and in January (Fig. 2b, c). The fi bre and
lignin content of willow leaves and stems increas-
es over the course of the growing season with a
corresponding decrease in digestibility (Hartley
& Jones 1997; Larter & Nagy 2001b). By fall and
winter, willow leaves and stems have lignifi ed and
are certainly not of the high nutritive quality they
were at the start of the growing season. Sample
sizes for these months were smaller than for other
months and may not have provided an accurate
refl ection of the diet; however, the percent willow
in any individual sample was ≥ 37 % which implies
willow was an important dietary component for
muskoxen during September, October and Janu-
ary.
The diet of Cape Bathurst caribou (Rangif-
er tarandus groenlandicus), found on the main-
Table 1. The Renkonen index of percent similarity between the monthly diets of muskox and
caribou in high muskox density areas. Substantial overlap (> 50 %) indicated by bold print.
Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sep. Oct. Nov. Dec.
January 20.8 27.4 27.9 28.7 27.5 26.9 43.4 26.7 27.4 23.7 21.9 26.5
February 25.6 26.7 27.2 26.8 21.7 51.3 26.0 28.1 24.0 22.6 25.2
March 38.1 38.9 37.6 33.0 62.2 36.8 33.2 29.5 27.6 34.9
April 13.6 12.9 12.4 37.4 12.1 16.9 13.0 10.5 11.6
May 24.8 24.2 49.3 24.0 28.6 24.9 22.3 23.9
June 70.8 53.1 37.7 60.6 57.2 27.7 52.4
July 24.9 18.0 58.9 84.2 21.9 41.0
August 29.5 67.5 63.7 32.1 50.7
September 36.0 32.2 30.4 37.5
October 27.1 25.3 28.7
November 25.8 32.1
December 31.1
137Larter & Nagy 2004: Polar Research 23(2), 131–140
land of the Northwest Territories to the south of
Banks Island, was > 85 % willow during late June
and July (N. Larter & J. Nagy unpubl. data), and
we expected Peary caribou, being a mixed feeder,
also to forage on high quality patchily distrib-
uted forage such as willow during the growing
season. During July the Peary caribou diet was
least diverse (SI 0.276) and willow made up 85 %
of the diet. In June and August, willow constitut-
ed ca. 45 % of the diet of Peary caribou on Banks
Island (Fig. 2a). Willow use peaked in July when
the available biomass of willow leaf crude pro-
tein was at its greatest (Larter & Nagy 2001b; N.
Larter & J. Nagy unpubl. data). This pattern of
foraging is consistent with the multiplier effect in
ruminant physiology (White 1983), where selec-
tive foraging on high quality foods has a positive
infl uence on animal production, and it occurred
during a period of milk production (for females)
and fat deposition. Body size and fat reserves have
a powerful effect on calf production and age of fi rst
reproduction of reindeer and caribou (Dauphine
1976; Leader-Williams 1980; Thomas 1982).
Some studies indicated that legumes were
an important component of the summer diet of
muskoxen on Banks Island and that their forag-
ing behaviour may shape regrowth and repro-
duction of Oxytropis borealis (Oakes et al.
1992; Mulder & Harmsen 1995). We found a fair
amount of legume in the July diet of muskoxen,
but negligible use during the other months of the
year. Legumes were at their highest digestibili-
ty in July (Larter & Nagy 2001b). Selective feed-
ing on highly digestible legumes in July may rep-
resent a multiplier effect (White 1983) for Banks
Island muskoxen. The proportion of legume in the
July diet was higher for muskoxen in high densi-
ty areas (Fig. 2b, c). Legumes made up substantial
portions of the monthly diet of caribou from Octo-
ber through May; July was the only month caribou
did not feed on legumes (Fig. 2a). Legumes are an
ideal forage for mixed feeders like caribou during
winter. They are patchily distributed throughout
upland habitats (Larter & Nagy 2001d) and have
a substantially higher crude protein content (ca.
12 %) than other available forages (Larter & Nagy
2001b).
Larter & Nagy (1997) indicated that the dif-
ferences in the diet of muskoxen between areas
of high and low density may have been related
to forage availability. Greater use of legumes by
muskoxen in high density areas may be a result of
Table 2. The Renkonen index of percent similarity between the monthly diets of muskox and
caribou in low muskox density areas. Substantial overlap (> 50 %) indicated by bold print.
Feb Apr May Jun Jul Aug Oct Nov
February 21.6 27.4 25.4 28.2 36.9 27.0 22.7 23.7
April 13.8 11.9 14.3 23.1 13.2 11.5 9.9
May 23.8 26.2 35.0 20.9 23.2 19.2
June 49.6 54.2 33.5 56.2 22.1
July 24.2 12.7 53.7 13.2
August 25.3 62.6 23.5
October 25.4 27.9
November 24.2
Table 3. The Renkonen index of percent similarity between the monthly diets of muskox in
high and low muskox density areas. Substantial overlap (> 50 %) indicated by bold print.
Feb Apr May Jun Jul Aug Oct Nov
February 79.5 87.3 37.9 66.5 77.4 86.0 64.5 87.8
April 94.0 43.0 72.2 81.5 90.3 63.1 82.7
May 56.9 71.9 67.3 55.8 82.7 47.8
June 50.7 46.4 35.2 58.8 24.9
July 83.4 63.0 61.4 55.6
August 88.3 61.6 72.7
October 66.6 28.1
November 91.3
138 Seasonal changes in composition of diets of Peary caribou and muskoxen
availability. The frequency of occurrence (Larter
& Nagy 2001d) and aboveground standing crop
(N. Larter & J. Nagy unpubl. data) of legumes
was greater in habitats in the high muskox den-
sity area. This was not the case for willows or
sedges in wet sedge meadows, which had simi-
lar availability in areas of high and low muskox
density; yet the willow proportion of the diet in
summer is greater for muskoxen in high densi-
ty areas. Larter & Nagy (2001a) indicated that a
per capita decrease in summer sedge availability
at peak muskox densities was the likely cause of
the larger willow component of the summer diet.
Greater intraspecifi c competition for sedge would
explain why sedge remained a lower proportion
of the summer diet of muskoxen in high density
areas (Fig. 2b).
During winter, muskoxen in the High Arctic gen-
erally forage in moister habitats where there is an
abundance of monocots, generally sedge (Urquhart
1973; Schaefer & Messier 1995a, 1995b; Thomas
et al. 1999; Larter and Nagy 2001c). This would
be expected of a grazing animal. If muskox select
wet sedge meadow habitat throughout winter, why
do muskoxen in high density areas have so much
willow in the diet during some months? Sedges
do constitute the majority of forage biomass in
wet sedge meadows on Banks Island; however,
willow is often found in this habitat (Larter &
Nagy 2001d). We believe that the decrease in the
sedge component and the increase in the willow
component of the diet from November to January
is caused by changing forage availability in wet
meadows as a result of foraging by muskoxen, and
the onset of 24-hr darkness. Muskox move into
wet meadows once the ground is frozen in Novem-
ber and actively forage on sedges, thus decreas-
ing sedge biomass relative to willow biomass and
making the snow cover harder as a result of cra-
tering for forage. During the period of 24-hr dark-
ness and after the onset of extreme cold (December
through mid-January) there appears to be a reluc-
tancy to search for ungrazed meadows. In Febru-
ary, with the rapid return of daylight, we believe
that the animals search for ungrazed meadows to
meet foraging requirements and again remain in
these localized areas through March, as indicat-
ed by a decrease in the sedge and an increase in
the willow component of the diet. By mid-April
the extreme cold has ended, daylight lasts almost
24 hr and we believe muskoxen then actively seek
ungrazed meadows in which to forage. This would
coincide with the time (April and May) when
muskox show an increase in energy expenditure
(Lawler & White 1997). Whether the pattern is the
same in areas of low muskox density is unknown
because of a lack of data.
The similarity between the diets of Peary car-
ibou and muskoxen in high density areas during
any given month was < 40 % except during June,
when there was substantial diet similarity (71 %;
Table 1). However, diet similarity in any given
month does not completely address the potential
for food competition. Foraging during the non-
growing season clearly has a direct impact on the
availability of forage in the non-growing season,
but it may also impact availability in the following
growing season. The June diet of Peary caribou
shows substantial similarity to the June, July, Sep-
tember, October and December diets of muskox-
en—particularly in their willow components—
and the September and October diets of muskoxen
have substantial overlap with the June, July and
August diets of Peary caribou. Therefore for fi ve
months of the year the diet of muskoxen is similar
to the summer diet of caribou. Given this similar-
ity between their diets, even though the two spe-
cies may not forage in the same areas at the same
time, there is potential for competition should
food become limiting.
Inter-annual variation in diet could potential-
ly affect our results, particularly if there were
directional changes in diet over time or if great-
er or lesser sampling intensity occurred during a
year in which diet was different from that in most
years. Changes in the quality and/or availabili-
ty of forages would be important factors in inter-
annual variation in diet. Larter & Nagy (2001b)
found annual variability in the quality of forag-
es on Banks Island, but these changes were con-
sistent across all forages and across areas of both
high and low muskox density. There was annual
variation in the standing crop of forages on Banks
Island, but this was not consistent across habitat
types (N. Larter & J. Nagy unpubl. data). Given
the variability in diet of individual caribou and
groups of muskoxen, and the lack of evidence for
any directional change in forage quality or avail-
ability, any inter-annual variability in diet would
not drastically affect the dietary patterns we
describe.
Although data were not collected specifi cally
to investigate changes in diet that may be relat-
ed to winter forage availability, many faecal sam-
ples were collected during a period when winter
snow conditions (Larter & Nagy 2000) and winter
139Larter & Nagy 2004: Polar Research 23(2), 131–140
foraging behaviour (Larter & Nagy 2001d) were
investigated. Winters 1996–97 and 1997–98 were
less severe with shallower and softer snow cover
than previous winters; there was very little snow
cover in November 1997 (Larter & Nagy 2000;
2001d). Shallower snow conditions in early winter
may have increased lichen availability, as suggest-
ed by the observation that the amount of lichen
in the November diet of Peary caribou in 1997
was larger than November in other years, 10.3 %
(n = 5) in 1997, versus mean of 1.4 % (n = 44) in
1991, 1993–96. Less severe snow conditions may
also increase the availability of legumes relative
to Dryas integrifolia for caribou. The absolute
amount of legume and the ratio of D. integrifolia
to legume in the November diet of caribou during
1996 and 1997 (n = 10) was greater than for four
(n = 39) of the previous years (41 versus 27 % and
1.1 versus 0.9, respectively). Acquiring a higher
proportion of legumes and consequently a diet
of higher protein content during winter could be
advantageous for Banks Island Peary caribou.
The numbers of both muskoxen and Peary cari-
bou on Banks Island have remained high over the
past decade: 50 000 to 65 000 muskoxen over the
age of one year, and 500 to 1200 caribou over the
same age (Nagy et al. 1996; Larter & Nagy 1997;
J. Nagy & N. Larter unpubl. data). In addition to
Peary caribou and muskoxen herbivory, Arctic
willows make up a considerable amount of the
winter diets of Arctic hares (Lepus arcticus) and
ptarmigan (Lagopus mutus, L. lagopus) (Klein
& Bay 1990; Larter 1999; N. Larter & D. Hik
unpubl. data). Can willows sustain the substan-
tial amounts of herbivory to which they are cur-
rently being exposed? Willows that were released
from browsing on Banks Island did not alter the
ratio of leaf biomass to stem diameter compared
to browsed plants; however, plants that were not
browsed had larger diameter stems resulting in an
increase in total leaf biomass of 30 % (Larter &
Hik in press). Arctic willows appear to be rela-
tively tolerant of herbivory, but there is some indi-
cation that vegetative growth is becoming dom-
inated by younger and narrower stems. Should
herbivory on willows result in reduced willow bio-
mass, the Peary caribou population would incur
more of a negative impact than the muskoxen pop-
ulation because caribou feed selectively on high
quality willows during summer when the animals
are growing and putting on fat reserves. A reduc-
tion in willow biomass would reduce the potential
for caribou numbers to increase.
Acknowledgements.—We thank the following for fi eld and
lab assistance during the course of the study: S. Baryluk, A.
Esau, E. Esau, L. Gordon, S. Gray, J. Lennie, J. Lucas Sr., T.
Lucas, F. Raddi, L. Raddi, P. Raddi, W. Raddi, and D. Semple.
We thank all the hunters in Sachs Harbour who provided cari-
bou faecal samples. T. Foppe and colleagues at the Composi-
tion Analysis Lab are gratefully acknowledged for comments
and insight into the microhistological technique. S. Carrière
provided comments on early drafts of this manuscript. Major
funding for this project was provided through the Inuvialuit
Final Agreement.
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/NOR <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>
/SVE <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>
/ENU <FEFF002000500044004600200069006e006e007300740069006c006c0069006e00670065007200200066006f00720020004f00660066007300650074007400720079006b006b0069006e006700200068006f00730020004c0075006e00640062006c006100640020004d006500640069006100200041005300200034002e00310032002e00300033>
>>
>> setdistillerparams
<<
/HWResolution [2400 2400]
/PageSize [595.276 822.047]
>> setpagedevice