Effects of partial cutting in winter on white-tailed deer

Color profile: Disabled
Composite Default screen
655
Effects of partial cutting in winter on white-tailed
deer
Antoine St-Louis, Jean-Pierre Ouellet, Michel Crête, Jean Maltais, and
Jean Huot
Abstract: We documented how commercial logging influenced the spatial behavior and nutritional ecology of northern
white-tailed deer (Odocoileus virginianus). Using periodic browse surveys, we estimated the additional biomass of
twigs available from felled trees to deer in the Pohénégamook wintering area (25 km2) at 55 kg/ha in a 43-ha cut conducted in 1995–1996 that aimed to favor conifers by removing overtopping deciduous trees. Over the entire winter,
deer used 54% of the browse made available by the felling residues. The use of the cutover, estimated by pellet group
census, was five times greater than the average recorded over the entire wintering area. Felled trees provided approximately 35% of the food intake of the animals that have used the cutover. Of 30 deer fitted with radio collars, the
cutover attracted only those whose range neighbored the logging area (<2 km). In preference tests carried out in the
winter of 1996–1997, deer showed no preference for twigs from newly cut trees over those from trees cut earlier in the
winter, nor for twigs from treetops (browse made accessible during the logging operation) over twigs from saplings
(browse usually accessible in winter). If commercial logging is conducted in winter as a means of providing emergency
food during snowy winters to enhance deer survival, our results suggest that partial cutting may be ineffective because
felling residues were used only by deer found near the cutover and because of the difficulties of logging in deep
snow.
Résumé : Nous avons évalué l’impact de la coupe forestière commerciale sur l’utilisation de l’espace et de la nourriture d’une population de cerfs de Virginie (Odocoileus virginianus). Des inventaires périodiques de brout ont permis
d’évaluer à 55 kg/ha la biomasse additionnelle des ramilles mise à la disposition des cerfs du ravage de Pohénégamook
(25 km2) dans une coupe de dégagement des conifères de 43 ha réalisée durant l’hiver 1995–1996. Pour l’ensemble de
l’hiver, les cerfs ont utilisé 54% du brout mis à leur disposition dans les résidus de coupe. La fréquentation du parterre
de coupe, estimée par un inventaire de fèces, était cinq fois plus importante qu’en moyenne dans le ravage. Nous
avons estimé que la biomasse additionnelle des ramilles provenant de la coupe représente 35% de la quantité de la
nourriture ingérée par les cerfs ayant fréquenté la coupe au cours de l’hiver. Le suivi télémétrique de 30 cerfs munis
de colliers émetteurs a montré que le rayon d’influence de la coupe forestière était limité aux cerfs dont le domaine vital avoisinait la coupe (< 2 km). Lors de tests de préférences réalisés à l’hiver 1996–1997, les cerfs n’ont montré aucune préférence entre des ramilles provenant d’arbres nouvellement abattus et d’autres abattus plus tôt en hiver, de
même qu’entre des ramilles de cimes d’arbres (brout mis à leur disposition lors des opérations de dégagement) et
d’arbustes (brout habituellement disponible en hiver). En raison de leur rayon d’influence limité et des difficultés
d’opération dans la neige épaisse, les coupes forestières commerciales pratiquées en hiver dans les ravages ne semblent
pas constituer une mesure efficace de production de nourriture d’urgence lors d’un hiver rigoureux afin d’accroître la
survie des cerfs. St-Louis et al. 661
Introduction
In northeastern North America, a survival strategy
adopted by deer consists of congregating in wintering areas
characterized by irregular mixed stands that provide food
and shelter (Huot 1974; Dumont et al. 1998). Deer benefit
from this aggregation at high density by maintaining a net-
work of trails that facilitate movements in the snow and improve protection against predators (Nelson and Mech 1981,
1991; Messier and Barrette 1985). Despite these adaptations,
food intake, which is dominated by terminal parts of deciduous twigs (Dumont et al. 1998), is not sufficient to meet
Received September 20, 1999. Accepted December 8, 1999.
A. St-Louis and J.-P. Ouellet.1 Département de biologie et des sciences de la santé, Université du Québec à Rimouski, Rimouski,
QC G5L 3A1, Canada.
M. Crête. Département de biologie et des sciences de la santé, Université du Québec à Rimouski, Rimouski, QC G5L 3A1,
Canada, Service de la faune terrestre, Faune et Parcs, 675 East René-Lévesque Boulevard, (P.O. Box 92) Québec, QC G1R 4Y1,
Canada, and Département de biologie, Pavillon Vachon, Université Laval, Sainte-Foy, QC G1K 7P4, Canada.
J. Maltais. Centre d’études nordiques, Université Laval, Sainte-Foy, QC G1K 7P4, Canada.
J. Huot. Département de biologie, and Centre d’études nordiques, Pavillon Vachon, Université Laval, Sainte-Foy, QC G1K 7P4,
Canada.
1
Corresponding author. e-mail: [email protected]
Can. J. For. Res. 30: 655–661 (2000)
I:\cjfr\cjfr30\cjfr-04\X99-248.vp
Tuesday, May 02, 2000 5:11:53 PM
© 2000 NRC Canada
Color profile: Disabled
Composite Default screen
656
Can. J. For. Res. Vol. 30, 2000
Fig. 1. Location of the Pohénégamook wintering area and boundaries of the 43-ha partial cut of overtopping deciduous trees to benefit
mature and immature conifers carried out during the winter of 1995–1996.
69°15'
QUEBEC
USA
289
47°
30'
47°
30'
Saint-Éleuthère
LOGGING AREA
Estcourt
DEER MONITORED
DEER WINTERING AREA
Pohénégamook
ABANDONED OR
CULTIVATED FIELDS
0
RESIDENTIAL AREA
5 km
69°15'
energy requirements, and deer are forced to draw on their
body reserves to survive until spring (Mautz et al. 1976).
This negative energy balance leads to death by starvation
during severe winters where snow impede movements
(Potvin et al. 1981; Verme and Ullrey 1984). Because of the
high energy demand associated to movement in the snow
(Mattfeld 1974; Parker et al. 1984), mortality rates associated with starvation often exceed 40% (Potvin et al. 1981).
White-tailed deer are not reluctant to eat food that human
activity can provide them in winter (Ozoga 1972, 1987;
Berteaux et al. 1998; Maltais and Crête 1998). One may distinguish two circumstances when deer can have access to
new sources of food in winter: (i) supplementary feeding,
where food is provided throughout the winter, to attract animals or to compensate for deficient browse supply and
(ii) emergency feeding, where the food is provided only
when the winter conditions are severe and the risk of starvation is high, generally towards the end of the season (Lenarz
1991; Doenier et al. 1997; Lewis and Rongstad 1998). During commercial logging in winter, branches from treetops
left on site, i.e., felling residue, represent a source of twigs
that might be used by deer (Ozoga 1972; Verme 1973;
Tierson et al. 1985; Hughes and Fahey 1991; Doenier et al.
1997; Van Deelen et al. 1998), but little is know about how
deer respond to the increased availability in browse in cutovers.
Our objective was to document how commercial logging
influenced the spatial behavior and nutritional ecology of
northern white-tailed deer.
Materials and methods
Study area
Field work was carried out during the winters of 1995–1996 and
1996–1997 in the Pohénégamook wintering area (PWA) (47°30′ N,
69°15′ W), approximately 250 km east of Québec City (Fig. 1). In
1996, PWA covered 25 km2 and supported a deer population estimated at 520 ± 62 animals (α = 10%) (Dumont et al. 1998). Deer
hunting had been prohibited since 1993.
Deer populations of the Bas-Saint-Laurent region live at the
northern edge of the species’ range and face severe winter weather
conditions (Potvin and Breton 1986). Sinking depths exceed 50 cm
on average 70 days per winter (Verreault and Côté 1994); sinking
depth beyond 50 cm considerably restricts movements (Potvin et
al. 1981). The winter of 1995–1996, however, was particularly
mild with a cumulative sinking depth of 2242 cm-days, whereas
this variable reached 8011 cm-days in 1996–1997.
PWA occupies a transition zone between the northern hardwood
forest and the boreal forest (Rowe 1972). It belongs to the balsam
fir (Abies balsamea (L.) Mill.) – yellow birch (Betula
alleghaniensis Britton) climatic domain, 5a region (moyennesAppalaches) (Ordre des ingénieurs forestiers du Québec 1996).
Mixed, coniferous, and deciduous stands make up 61, 8, and 31%
of PWA, respectively (Verreault and Côté 1994). Balsam fir – yellow birch stands cover over 40% of the area. Dominant coniferous
species are balsam fir, white spruce (Picea glauca (Moench) Voss),
and eastern white cedar (Thuja occidentalis L.) (Dumont et al.
1998). A major spruce budworm (Choristoneura fumiferana (Clem.))
outbreak killed most balsam fir between 1975 and 1983, which
considerably reduced conifer canopy closure (Verreault and Côté
1994). Yellow birch, paper birch (Betula papyrifera Marsh.), balsam
© 2000 NRC Canada
I:\cjfr\cjfr30\cjfr-04\X99-248.vp
Tuesday, May 02, 2000 5:12:04 PM
Color profile: Disabled
Composite Default screen
St-Louis et al.
poplar (Populus balsamifera L.), quaking aspen (Populus tremuloides Michx.), red maple (Acer rubrum L.), and American beech
(Fagus grandifolia J.F. Ehrh.) dominate among deciduous species.
Silviculture treatment
From mid-January to the end of February 1996, a partial cut was
carried out near the northern tip of Lake Pohénégamook (Fig. 1).
The cutover covered 43 ha of hillsides at an altitude varying from
300 to 400 m and affected three tree stands: a stand of shadeintolerant hardwood trees (8 ha), a mixed stand dominated by paper birch (7 ha), and a mixed stand dominated equally by paper
birch and balsam fir (28 ha) (Verreault and Côté 1994). The partial
release cut aimed to free mature and immature conifers from overtopping deciduous trees that reached up to 20 m in height in the
case of quaking aspen. The harvest consisted mainly of paper birch
and quaking aspen, although yellow birch, balsam poplar, red maple, and sugar maple (Acer saccharum Marsh.) were occasionally
cut. The operation removed about 40% (280 stems/ha, 80 m3/ha) of
the deciduous stems. Conifers and most understory deciduous trees
were retained.
Influence of the cutover on feeding behavior and space
use of deer
Browse availability and use from felled trees
Twig availability and use from felled trees were measured using
one hundred and fifty-one 1 m × 4 m plots, distributed systematically according to a constant 30-m sampling interval. Between
February 5 and March 27, 1996, five surveys were carried out at
2-week intervals, and a sixth one was carried out on April 28,
1996, following snowmelt and the departure of deer from the wintering area. Because the logging operation took place from midJanuary to the end of February 1996, we gradually added plots to
follow the expansion of the cutting area. Accordingly, three series
of plots were considered. On February 5 and 6, 39 plots were sampled. From February 12 to 15, those 39 plots were resampled, and
65 new plots were sampled for the first time. From February 29 to
March 2, all pre-established plots were sampled and 47 new plots
were added, for a total of 151. All plots were sampled three more
times (March 13, March 26 and 27, and April 28). During each
survey, available and browsed twigs from felled trees were counted
in each plot. A twig was considered available if it was located at a
height of less than 2.5 m and was at least 10 cm long. We estimated browse use at each plot using the proportion of the number
of browsed twigs over the total of twigs (Potvin 1978).
To convert twig availability and browse use into biomass, we
used the methodology developed by Potvin (1978). The mean dry
mass of 100 browsed paper birch and 100 browsed quaking aspen
(predominant species) twigs was estimated for twigs collected over
the cutting area. Then, for each of the 100 selected browsed twigs,
a corresponding unbrowsed twig was cut at the same diameter at
point of browsing (DPB). Twigs (i.e., bites) were dried at 70°C for
24 h and weighed (±0.1 g). We computed a common twig biomass
for the two species, because there was no difference in biomass between the two (t test, P > 0.05); we assumed that the biomass of
the other species was similar.
Deer movements and use of the cutting area
To determine deer movements, 30 deer were captured throughout the wintering area, fitted with radio collars and monitored from
mid-January to early April 1996. These animals had been marked
in January between 1994 and 1996, using Stephenson traps
(Rongstad and McCabe 1984); during the study, 13 adult (>1-yearold) females, 10 adult males, four young (<1-year-old) females and
three young males were monitored. The position of each deer was
estimated by triangulation using a nine-element yagi antenna
mounted through the roof of a vehicle (Amlaner and Macdonald
657
1979; Lovallo et al. 1994). Bearings were measured with an Azimuth 1000 digital compass (KVH Industries, Inc.; Lovallo et al.
1994). For each location, at least three readings were made from
permanent stations. Positions were estimated using LOCATE II
software (Nams 1990). Following blind tests carried out with 21
radio collars, the precision of each location was evaluated at 212 ±
42 m (mean ± SE). Every 10 days, from mid-January until early
April, each deer was located once during the day and once at night.
Radio-tracking alternated from one survey to the next between
morning and afternoon (06:00–12:00 and 12:00–18:00) and between evening and night (18:00–24:00 and 00:00 and 06:00).
Individual deer locations monitored by telemetry were transferred onto a digitized map using the ArcView GIS software (Environmental Systems Research Institute Inc. 1996) (see Fig. 1). The
home range size of each deer was calculated with the program
CALHOME (Kie et al. 1996), using the minimum convex polygon
method (Mohr 1947) including 70 and 100% of the locations. The
deer were then separated into two groups, according to the proximity of their home range to the cutting area. Deer likely to have frequented the cutover were identified as those that were visually
identified in the cutover (n = 2), those that were directly located in
the cutover by telemetry (n = 2), and those located outside the cutting area but whose center of activity was situated at a distance
from the cutting area that was less than the diameter of their home
range (n = 4). For the analyses, these eight deer were identified as
living near the cutover while the other 22 radio-collared animals in
the wintering area were assumed to be unlikely to use the logging
area. Daily movements were expressed as the distance covered between the daytime and nighttime positions recorded on the same
day divided by the time interval (hours) multiplied by 24.
To assess the use of the cutover area during winter relative to
the rest of the PWA, a census of pellet groups was made on April
27 and 28, 1996 (Potvin 1978). Within the cutover, eighty-four 2 m
× 40 m plots were systematically distributed 30 m apart. Defecation rate in winter was assumed to be 22.3 pellet groups/day (Rogers 1987). Pellet-group density for the entire PWA was calculated
as follows: 520 deer wintering 135 days in a 25 km2 area and defecation at a rate of 22.3 pellet groups/day.
Palatability of twigs from felled trees
Preference tests were carried out in the PWA in the winter of
1996–1997 to evaluate the palatability of twigs from felling residues. The first experiment dealt with the effect on selection of paper birch on the time elapsed since the tree was felled. At the time
of the experiments, both old and fresh twigs were presented to deer
simultaneously on branches 1.5 m in length, driven into the snow
at three clumps per site so that several animals could have access
to twigs simultaneously (Grenier 1997). The experiment was carried out at seven sites frequented by deer, spread out over the entire PWA. At each site, 300 twigs per treatment were offered to the
deer. Browsed and unbrowsed twigs were counted at the end of the
first day and on the three following days, both in the morning and
afternoon, to monitor the progress of percentage twig consumption.
The DPB of 30 browsed twigs per treatment, chosen randomly,
was measured (±1 mm) for each site at the time of the surveys.
Comparisons were made between twigs originating from trees cut
in early January 1997 with twigs collected in early February,
March, and April 1997. Thus, on January 7 and 8, ten paper birch
stems had been cut to provide experimental twigs for each of the
three tests. Fresh twigs were collected over the previous 2 days before each experiment from at least three different mature trees.
Two other experiments, dealing with the relative preference of
treetop paper birch twigs (i.e., twigs made accessible during logging operations) over paper birch sapling twigs (i.e., twigs normally accessible to deer) and preference for paper birch over
quaking aspen twigs took place from March 31 to April 5 1997,
at the same sites and following the same procedure as for the
© 2000 NRC Canada
I:\cjfr\cjfr30\cjfr-04\X99-248.vp
Tuesday, May 02, 2000 5:12:05 PM
Color profile: Disabled
Composite Default screen
658
Can. J. For. Res. Vol. 30, 2000
Table 1. Use of twigs (mean ± SE) by white-tailed deer according to date on which trees were cut and time elapsed since felling (2, 4, and 6 weeks) in the 43-ha selective cut, PWA, winter
1995–1996.
Table 2. Home range size (mean ± SE) obtained by the convex
polygon method (CPM) at 70 and 100%, for white-tailed deer
monitored in the PWA in the winter of 1995–1996 according to
age and proximity to the logging area.
Use rate (%)
Home range size (ha)
Starting date
2 weeks
4 weeks
6 weeks
Age
January 15
January 30
February 15
19±4.7a
30±3.7a
53±5.3b
47±5.7a
38±4.5a
66±6.4b
50±5.8a
48±4.6a
73±6.5a
70% CPM
Fawns
29.1±10.19 (n=4)a
Adults
16.9±2.41 (n=16)c
100% CPM
Fawns
184.2±81.06 (n=4)ns
Adults
63.8±9.62 (n=17)ns
Note: Values with different letters within a column are significantly
different (P < 0.05) in browse use.
previous experiment. Twigs originated from at least three stems
per treatment and were cut down the day before the experiment.
Statistical analyses
Effects of proximity of the cutover (near or far) and deer age
(fawn or adult) on home range size were tested using ANOVA
(procedure GLM; SAS Institute Inc. 1992). Effects of cutover
proximity and deer age on daily movements were assessed using
ANOVA for repeated measures (procedure MIXED; SAS Institute
Inc. 1992), because this variable was recorded several times for
each individual. Influences of date (mid-January, late January, and
mid-February) when trees were cut and of time elapsed since felling (2, 4, or 6 weeks) on deer use of the browse (all species combined) was also evaluated using ANOVA for repeated measures.
Pairwise comparisons were conducted using the LSMEANS statement (SAS Institute Inc. 1992).
Deer food preferences between the various types of twigs in the
winter of 1996–1997 were analyzed using paired t tests (SYSTAT,
version 7.0, SPSS 1997). The same test served for comparing DPB
(i.e., bite size) during preference tests.
Twig biomass, daily movements, and home range size were log
transformed prior to statistical analyses. For each of the statistical
tests carried out, we ensured the normality of residuals with the
Shapiro–Wilks test and the absence of a pattern by visual examination of the plots. Statistical means are given with their standard
error.
Results
Availability and use of browse provided through the
silviculture treatment
The overall browse available and use of felled trees was
estimated using the last survey carried out at the end of
April, which averaged 161 376 ± 15 653 twigs/ha for all
species combined. Given that mean bite weighed 0.34 ±
0.03 g (n = 200), the cutover provided 54.8 ± 5.32 kg/ha of
food. Paper birch accounted for 34 ± 5 kg/ha and quaking
aspen for 14 ± 2 kg/ha; all other species were marginal as a
food source. Deer consumed 27 ± 4.1% (n = 68 plots) of
quaking aspen twigs and 56 ± 3.5% (n = 90) of paper birch
twigs. Although red and sugar maple were rare, deer use was
high for these species (98 ± 1.2%, n = 13).
Twig use by deer differed according to the date on which
trees were cut during winter (F4,226 = 2.57, p = 0.039). Two
weeks post-logging, consumption rates were significantly
greater for the browse produced mid-February (series 3)
than the browse produced earlier in the winter (series 1 and
2) (series 1 vs. 3, P < 0.001; series 2 vs. 3, P < 0.001)
(Table 1). This trend remained after a time period of
Away from logging area
Near logging area
6.0±1.74 (n=4)b
11.7±3.81 (n=3)c
76.3±34.59 (n=4)ns
66.1±28.14 (n=4)ns
Note: Values with different letters are significantly different (P < 0.05)
in home range size for a given estimator (70 or 100% CPM).
4 weeks (series 1 vs. 3, P = 0.026) but disappeared 6 weeks
post-logging (series 1 vs. 2 vs. 3, P > 0.05). Based on the
pellet group census carried out in the spring, total use of the
cutover area for the winter of 1995–1996 reached 15 170
deer-days/km2.
Influence of the cutover on space use
The plotting of deer locations indicated that no radiocollared animal shifted its home range to use the cutover
area. Deer that used the cutover were those that had their
center of activity within 2 km of the cutover. The cutover
appeared to influence the pattern of space use by reducing
home range size when computed by the 70% minimum convex polygon method (F1,20 = 9.490, P = 0.006). Fawns located near the cutover area had a smaller home range than
those located away from the cutover (F1,20 = 5.743, P =
0.027), unlike adults where no significant difference was detected (P > 0.05) (Table 2). The cutover proximity, however,
did not influence home range size of fawns or adults when
computed with the 100% minimum convex polygon method
(P > 0.05).
Mean daily movements averaged 1086 ± 144 m. Proximity to the cutover and age had no significant impact on daily
movements of the deer monitored (P > 0.05).
Selection of twigs
No preference was detected between fresh twigs and those
coming from trees felled earlier in the winter (January–
February: t = 1.90, df = 6, P = 0.102; January–March: t =
0.96, df = 6, P = 0.384; January–April: t = 0.16, df = 6, P =
0.883) (Table 3). Similarly, bite size did not differ between
twigs type (January–March: t = 0.08, df = 6, P = 0.932;
January–April: t = 0.15, df = 6, P = 0.881). Deer showed no
preference for twigs originating from mature trees (89 ±
5.2%) over saplings (89 ± 5.8%) (t = –0.232, df = 6, P =
0.820). However, deer preferred twigs from paper birch
(80 ± 11.7%) over those from quaking aspen (47 ± 13.9%)
(t = –3.39, df = 6, P = 0.015).
Discussion
Overall, deer intensively used the study cutover as a feeding site throughout the winter of 1995–1996. Based on deer
pellet-group density, the use of the cutover was five times
© 2000 NRC Canada
I:\cjfr\cjfr30\cjfr-04\X99-248.vp
Tuesday, May 02, 2000 5:12:06 PM
Color profile: Disabled
Composite Default screen
St-Louis et al.
659
Table 3. Percent use (mean ± SE) by white-tailed
deer of fresh twigs and twigs cut earlier in the winter during preference tests carried out in the PWA,
winter 1996–1997.
Use rate (%)
Time elapsed since
felling (months)
Twigs cut
earlier
Fresh
twigs
1
2
3
40±11.2
70±10.7
71±12.8
30±6.4
69±10.7
71±12.2
Note: Use rates between fresh twigs and twigs cut earlier
were not significantly different.
higher than that observed on average for the entire PWA.
The cut was only used by deer living close to the logging
site, as no major movement towards the cutover was observed among the 30 deer fitted with radio collars. Only
eight of the 30 monitored deer were likely to have frequented the cutting area, and those eight deer were located
within 2 km of it. Lewis and Rongstad (1998) obtained similar results with feeders, which only attracted deer living
within 2.5 km. The social and spatial organization of whitetailed deer, characterized by matrilineal family groups,
partly explains the high site fidelity shown by individual
deer (Van Deelen et al. 1998). Similarly, no short-term influence of logging operations on home range location was
observed for wapiti (Cervus canadensis) or roe deer
(Capreolus capreolus) (Edge et al. 1985; Linnell and
Andersen 1995).
The quantity of browse per surface area provided by
felled trees was 2.7 times that of twigs normally accessible
to deer during winter (PWA in 1994–1995: 60 485 twigs/ha
(Dumont et al. 1998); cutover in 1995–1996: 161 376
twigs/ha (this study)). Moreover, the rate of twig consumption produced by felling residues (54%) was greater than the
browsing rates observed in the entire PWA during the two
previous years (1993–1994: 42.3%; 1994–1995: 33.8%),
which were among the highest observed in Québec (Dumont
et al. 1998). The concentration of twigs at one site may have
favored such a high utilization rate, because we did not find
that deer preferred twigs from felled trees over twigs normally accessible in winter (i.e., from saplings).
Deer consumed a greater fraction of twigs made available
in mid-February than at the beginning of logging in midJanuary. This could be partly explained by the fact that deer
took some time to become accustomed to logging operations
and to use the browse made available. Other white-tailed
deer provided with artificial feed reacted in a similar manner, often taking many weeks before accepting the new food
source (Maltais et al. 1998; Berteaux et al. 1998). The delay
in forage consumption did not depend on browse quality, because our preference tests showed that deer did not prefer
twigs from saplings to felled residues. There are also reasons
to believe that the addition of easily available natural food
(felling residues) would result in a greater response by deer
in the middle or at the end of the winter because of the annual decline of accessible browse as winter progresses combined with the reduction of deer body reserves (Mautz 1978;
Verme and Ullrey 1984; DelGiudice et al. 1990).
The daily food intake of PWA deer averaged 629 g/day
and 522 g/day during the winters of 1993–1994 and 1994–
1995, respectively (A. Dumont, unpublished data). Based on
this estimate, the felled trees provided approximately 35% of
the food intake of the animals that have used the cutover, assuming that the food was shared evenly among animals.
This estimate may appear relatively low, since other studies
on food addition revealed a greater use of feeders by deer
during adverse winters (Doenier et al. 1997; Lewis and
Rongstad 1998). However, the 1995–1996 winter was mild,
and one might expect a greater consumption of felling residues under difficult winter conditions, as major snow accumulations make woody browse less accessible (Dumont et
al. 1998). For the entire wintering area (i.e., 25 km2), felling
residues provided only slightly less than 5% of the overall
forage availability (wintering area: 60 485 twigs/ha over
2500 ha (Dumont et al. 1998); cutover: 161,376 twigs/ha
over 43 ha (this study)).
Fawns located near the cutting area had smaller home
ranges than fawns located away from the cutover. Fawns
face high risk of winter mortality because of their small size
and limited body reserves (Messier 1979) and, thus, may
have benefited by reducing the extent of their home range to
lower foraging costs (Mattfeld 1974; Parker et al. 1984).
Management implications
During winters with deep snow, cutting deciduous trees
permits the use of felling residues by deer, thereby making
cutting a potential management tool for populations that suffer high mortality due to deep snow in some winters. The
fact that deer are sedentary, however, meant that the additional food was not available to many deer. In addition, logging operations carried out in deep snow are subject to
increased frequency of mechanical breakdowns and to a
50% reduction in timber harvest in comparison with the
same work carried out in summer (A. Lapierre, personal
communication). As a result, it would likely be difficult to
carry out commercial logging as part of winter feeding for
deer. On the other hand, because browse on trees felled in
early winter remains palatable to deer throughout the winter,
commercial logging in wintering areas could be carried out
in late fall or early winter. Deer would be able to consume
felling residues during the following months, provided that
twigs remain accessible. We observed that many twigs remain above the snow. Deer would have the opportunity to
eat this browse early in winter or during snowmelt.
Acknowledgments
This study was supported by le ministère de
l’Environnement et de la Faune, le ministère des Ressources
naturelles, la Fondation de la faune du Québec, les Fonds
pour la formation de chercheurs et l’aide à la recherche, la
Fondation de l’Université du Québec à Rimouski, and
la Forêt Modèle du Bas-Saint-Laurent Inc. We sincerely
thank Jean-François Gaudreault, Céline Meunier, Nicolas
Bergeron, and Sylvain Gagnon for their invaluable assistance in the field. Our thanks also go to André Dumont,
Alain Caron, Alain Lapierre, and Éric Aubert for their precious collaboration on the project. We are grateful to
Dominique Arseneault, Marc Bélisle, André Dumont, and
© 2000 NRC Canada
I:\cjfr\cjfr30\cjfr-04\X99-248.vp
Tuesday, May 02, 2000 5:12:06 PM
Color profile: Disabled
Composite Default screen
660
François Potvin for their comments on a previous version of
this manuscript. Gaétan Daigle and Hélène Crépeau of Service de consultation statistique de l’Université Laval kindly
advised on the statistical analysis.
References
Amlaner, C.J., Jr., and Macdonald, D.W. 1979. A handbook of
biotelemetry and radio-tracking. Pergamon Press. Oxford.
Berteaux, D., Crête, M., Huot, J., Maltais, J., and Ouellet, J.-P.
1998. Food choice by white-tailed deer in relation to protein and
energy content of the diet: a field experiment. Oecologia, 115:
84–92.
DelGiudice, G.D., Mech, L.D., and Seal, U.S. 1990. Effects of
winter undernutrition on body composition and physiological
profiles of white-tailed deer. J. Wildl. Manage. 54: 539–550.
Doenier, P.B., DelGiudice, G.D., and Riggs, M.R. 1997. Effects of
winter supplemental feeding on browse consumption by whitetailed deer. Wildl. Soc. Bull. 25: 235–243.
Dumont, A., Ouellet, J.-P., Crête, M., and Huot, J. 1998.
Caractéristiques des peuplements forestiers recherchés par le
cerf de Virginie en hiver à la limite nord de son aire de
répartition. Can. J. Zool. 76: 1024–1036.
Edge, W.D., Marcum, C.L., and Olson, S.L. 1985. Effects of logging activities on home range fidelity of elk. J. Wildl. Manage.
49: 741–744.
Environmental Systems Research Institute Inc. 1996. Using
ArcView GIS. Environmental Systems Research Institute Inc.,
Redlands, Calif.
Grenier, D. 1997. Accès à la nourriture chez des cerfs de Virginie
(Odocoileus virginianus) fréquentant des sites d’alimentation
hivernaux de la région du Bas-Saint-Laurent. M.Sc. thesis,
Université Laval, Sainte-Foy, Que.
Hughes, J.W., and Fahey, T.J. 1991. Availability, quality, and selection of browse by white-tailed deer after clearcutting. For. Sci.
37: 261–270.
Huot, J. 1974. Winter habitat of white-tailed deer at Thirty-one
Mile Lake, Québec. Can. Field-Nat. 88: 293–301.
Kie, J.G., Baldwin, J.A., and Evans, C.J. 1996. CALHOME : a
program for estimating animal home ranges. Wildl. Soc. Bull.
24: 342–344.
Lenarz, M.S. 1991. Simulation of the effects of emergency winter
feeding of white-tailed deer. Wildl. Soc. Bull. 19: 171–176.
Lewis, T.L., and Rongstad, O.J. 1998. Effects of supplemental
feeding on white-tailed deer, Odocoileus virginianus, migration
and survival in northern Wisconsin. Can. Field-Nat. 112: 75–81.
Linnell, J.D.C., and Andersen, R. 1995. Site tenacity in roe deer:
short-term effects of logging. Wildl. Soc. Bull. 23: 31–35.
Lovallo, M.J., Vercauteran, K.C., Hedge, N.C., Anderson, E.M.,
and Hygnstrom, S.E. 1994. An evaluation of electronic versus
hand-held compasses for telemetry studies. Wildl. Soc. Bull. 22:
662–667.
Maltais, J., and Crête, M. (Editors). 1998. Développement d’un
programme de nourrissage d’urgence du cerf de Virginie au Bas
Saint-Laurent en prévision d’hivers catastrophiques. Service de
la faune terrestre, ministère de l’Environnement et de la Faune,
Québec, Que., Rep. No. 3956-98-09.
Maltais, J., Huot, J., Crête, M., and Ouellet, J.-P. 1998. Mise au
point d’une méthode pour faire accepter de la moulée à des cerfs
naïfs comme nourriture d’urgence lors d’hivers catastrophiques.
In Développement d’un programme de nourrissage d’urgence du
cerf de Virginie au Bas Saint-Laurent en prévision d’hivers
catastrophiques. Edited by J. Maltais and M. Crête. Service de la
Can. J. For. Res. Vol. 30, 2000
faune terrestre, ministère de l’Environnement et de la Faune,
Québec, Que., Rep. No. 3956-98-09.
Mattfeld, G.F. 1974. The energetics of winter foraging by whitetailed deer: a perspective on winter concentration. Ph.D. thesis,
University of Syracuse, Syracuse, N.Y.
Mautz, W.W. 1978. Sledding on a bushy hillside: the fat cycle in
deer. Wildl. Soc. Bull. 6: 88–90.
Mautz, W.W., Silver, H., Holter, J.B., Hayes, H.H., and Urban,
W.E. 1976. Digestibility and related nutritional data for seven
northern deer browse species. J. Wildl. Manage. 40: 630–638.
Messier, F. 1979. Étude de la prédation du cerf de Virginie par le
coyote dans le ravage d’Armstrong, Beauce sud. M.Sc. thesis,
Université Laval, Sainte-Foy, Que.
Messier, F., and Barrette, C. 1985. The efficiency of yarding behavior by white-tailed deer as an antipredator strategy. Can. J.
Zool. 63: 785–789.
Mohr, C.O. 1947. Table of equivalent populations of North American small mammals. J. Wildl. Nat. 37: 223–249.
Nams, V.O. 1990. LOCATE II user’s guide. Pacer, Truro, N.S.
Nelson, M.E., and Mech, L.D. 1981. Deer social organization and
wolf predation in northeastern Minnesota. Wildl. Monogr.
No. 77.
Nelson, M.E., and Mech, L.D. 1991. Wolf predation risk associated
with white-tailed deer movements. Can. J. Zool. 69: 2696–2699.
Ordre des ingénieurs forestiers du Québec. 1996. Manuel de
foresterie. Les Presses de l’Université Laval, Sainte-Foy, Que.
Ozoga, J.J. 1972. Aggressive behavior of white-tailed deer at winter cuttings. J. Wildl. Manage. 36: 861–868.
Ozoga, J.J. 1987. Maximum fecundity in supplementally-fed northern Michigan white-tailed deer. J. Mammal. 68: 878–879.
Parker, K.L., Robbins, C.T., and Hanley, T.A. 1984. Energy expenditures for locomotion by mule deer and elk. J. Wildl. Manage.
48: 474–488.
Potvin, F. 1978. L’inventaire de brout : revue des méthodes et description de deux techniques. Direction générale de la faune,
ministère du Tourisme, de la Chasse et de la Pêche, Québec,
Que., Spec. Rep. No. 9.
Potvin, F., and Breton, L. 1986. Sommaire des conditions
d’enneigement pour le cerf au Québec de 1973 à 1985. Direction de la faune terrestre, ministère du Loisir, de la Chasse et de
la Pêche, Québec, Que., Spec. Rep. No. 1208.
Potvin, F., Huot, J., and Duchesneau, F. 1981. Deer mortality in the
Pohénégamook wintering area, Québec. Can. Field-Nat. 95: 80–
84.
Rogers, L.L. 1987. Seasonal changes in defecation rates of freeranging white-tailed deer. J. Wildl. Manage. 51 : 300–333.
Rongstad, O.J., and McCabe, R.A. 1984. Capture technique. In
White-tailed deer ecology and management. Edited by L.K.
Halls. Stackpole Books, Harrisburg, Pa. pp. 655–676.
Rowe, J.S. 1972. Les régions forestières du Canada. Service canadien des Forêts, ministère de l’Environnement, Ottawa, Ont.,
Publ. No. 1300F.
SAS Institute Inc. 1992. SAS/STAT software: changes and enhancements, release 6.07 edition. SAS Institute Inc., Cary, N.C.,
Tech. Rep. No. P-229.
SPSS, Inc. 1997. SYSTAT: statistics, version 7.0 edition. SPSS,
Inc., Chicago, Ill.
Tierson, W.C., Mattfeld, F., Sage, R.W., Jr., and Behrend, D.F.
1985. Seasonal movements and home range of white-tailed deer
in the Adirondacks. J. Wildl. Manage. 49: 760–769.
Van Deelen, T.R., Campa, H., III, Hamady, M., and Haufler, J.B.
1998. Migration and seasonal range dynamics of deer using adjacent deeryards in northern Michigan. J. Wildl. Manage. 62:
205–213.
© 2000 NRC Canada
I:\cjfr\cjfr30\cjfr-04\X99-248.vp
Tuesday, May 02, 2000 5:12:07 PM
Color profile: Disabled
Composite Default screen
St-Louis et al.
Verme, L.J. 1973. Movements of white-tailed deer in upper Michigan. J. Wildl. Manage. 37: 545–552.
Verme, L.J., and Ullrey, D.E. 1984. Physiology and nutrition. In
White-tailed deer ecology and management. Edited by L.K.
Halls. Stackpole Books, Harrisburg, Pa. pp. 91–118.
661
Verreault, G., and Côté, B. 1994. Plan d’aménagement du ravage
du lac Pohénégamook : plan d’intervention 1994–1999. Ministère
de l’Environnement et de la Faune et ministère des Ressources
naturelles, Rivière-du-Loup, Que.
© 2000 NRC Canada
I:\cjfr\cjfr30\cjfr-04\X99-248.vp
Tuesday, May 02, 2000 5:12:07 PM