Effects of partial cutting in winter on white-tailed deer
Transcription
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. 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