p139-146_4205.pdf ALCES VOL. 41, 2005 DUNGAN AND WRIGHT - MOOSE SUMMER DIET COMPOSITION 139 SUMMER DIET COMPOSITION OF MOOSE IN ROCKY MOUNTAIN NATIONAL PARK, COLORADO Jason D. Dungan1 and R. Gerald Wright2 1Department of Fish and Wildlife Resources, University of Idaho, P.O. Box 441136, Moscow, ID 83844-1136, USA; 2USGS Idaho Cooperative Fish and Wildlife Research Unit, Department of Fish and Wildlife Resources, University of Idaho, P.O. Box 441136, Moscow, ID 83844-1136, USA ABSTRACT: Summer diet composition of habituated adult moose (Alces alces) in Rocky Mountain National Park, Colorado, was determined using direct observations and fecal analysis. Direct ob- servations determined moose ate 20 different plant species, including 6 willow (Salix spp.) species, which comprised 91.3% of the overall diet from June through mid-September. Geyer willow (Salix geyeriana) accounted for 45.1% of summer diets. Other species included mountain alder (Alnus in- cana, 2.5%), quaking aspen (Populus tremuloides, 1.1%), and bog birch (Betula glandulosa, 1.0%). Aquatic plants accounted for 1.9%, forbs 1.1%, and grasses 0.9%. Moose ate 11 different species of woody browse, which comprised 96.9% of the diet. Species diversity in the diet peaked in July with 18 different species, including 7 species of non-woody browse. Fecal analysis showed moose consumed Carex spp. major genera (> 1%) contributing to moose summer diets that were indicated by direct observations, except quaking aspen (1.1%). ALCES VOL. 41: 139-146 (2005) Key words: Alces alces, diet, food habits, moose, willow, woody browse Historically, moose (Alces alces shirasi) were rare in Colorado and early sightings recorded in Colorado were believed to be moose that wandered into the state from northwestern Wyoming (Bailey 1944). The Colorado Division of Wildlife (CDOW) introduced two groups of 12 moose near the town of Rand, Colorado, in 1978 and 1979; 13 km west of Rocky Mountain National Park (RMNP). The objective was to establish a viable resident moose population in the area. - ter in the park was recorded in 1985-1986 (Stevens 1988). Presently between 61-66 moose are estimated to summer in the park (J. Dungan, University of Idaho, Moscow, Idaho, unpublished data). Large mammalian herbivores can cause major changes in plant community composi- tion and structure (Augustine and McNaugh- ton 1998). Herbivory is a major concern of RMNP where large numbers of Rocky Moun- tain elk (Cervus elaphus) occur. Managers are particularly concerned about effects on riparian willow (Salix spp.) and upland shrub communities, based on visual appearance of short-hedged willow on elk winter range. In a recent study, elk herbivory was found to suppress heights, leader lengths, and an- nual production of willow, and herbaceous productivity of willow sites within the park (Zeigenfuss et al. 1999). High densities of moose have also been of ecosystems. Bark stripping by moose in Denali National Park, Alaska, may increase the rate of succession in aspen-spruce com- MOOSE SUMMER DIET COMPOSITION - DUNGAN AND WRIGHT ALCES VOL. 41, 2005 140 munities by killing trees (Miquelle and Van Ballenberghe 1989). Similar results were found in Isle Royale National Park, Michigan, where high rates of moose browsing depressed nitrogen mineralization and net primary pro- duction of boreal forest ecosystems (Pastor et al. 1993). Furthermore, McInnes et al. (1992) showed that moose herbivory reduced tree biomass and production, and increased shrub and herb biomass at Isle Royale. Winter food supply is generally con- sidered the limiting factor in some moose populations (Crete 1989, Kufeld and Steinert 1990, MacCraken et al. 1997, Zheleznov-Chu- kotsky and Votiashova 1998). Woody browse is usually the only food supply available for moose during the winter, and therefore moose diet and habitat studies have focused on use, availability, and quality of winter browse (LeResche and Davis 1973). Few studies have examined use of summer habitat, although summer diets are generally 1.5 – 3 times more nutritious than winter diets (Schwartz 1992), and summer is a key period in which moose build up fat reserves that take them through the winter. Knowledge of moose summer decisions on managing moose habitat. Peek (1974) reviewed 41 food habit stud- ies in North America, of which 18 were from the intermountain west, but none as far south as Colorado. Five of the 18 studies included summer food habits, and revealed even greater variation between areas than those on the winter range (Peek 1974). Summer diet stud- ies have increased (Joyal and Sherrer 1978, Butler 1986, Van Ballenberghe et al. 1989), but none have investigated the southern extent of the Rocky Mountains. In North Park, willow was the most commonly selected habitat by moose in all seasons between 1991 and 1995 (Kufeld and Bowden 1996). Similarly, willow comprised about 80-85% of the diet of wild adult moose in Denali National Park during the summer (Van Ballenberghe et al. 1989). The long- term effects of moose browsing on riparian communities from Alaska to RMNP, is not yet understood. The purpose of this study was to document summer diet composition of moose in RMNP. STUDY AREA This study was conducted in RMNP dur- ing the summers of 2003 and 2004. RMNP is located in north-central Colorado just west of Estes Park. RMNP covers an area of 1,075 km2 and ranges from 2,389 m to 4,345 m in elevation. The park lies astride the continental divide and the west and east side differ in climate. Annual precipitation ranges from 37.6 to 51.7 cm. Temperatures range from highs in July and August of 24°C to lows in December-February of –17°C (Monello and Johnson 2003). We conducted this study predominantly on the west side of the park within the Colorado River Drainage and within higher elevation meadows east of the Continental Divide. The study was conducted at 2 dis- tinct elevational strata, below 3,000 m (low elevation sites) and above 3,000 m (high elevation sites), because of differences in plant species composition, distribution, and plant morphology. Lower elevation sites tend to be comprised of a greater variety of plant species. Plants tend to be more sparsely distributed, and to be taller with longer leader lengths and greater biomass, than plants at high elevation sites. Low elevation riparian meadows are characterized by large stands of geyer wil- low (Salix geyeriana), mountain willow (Salix monticola), drummond willow (Salix drummondiana), plane-leaf willow (Salix planifolia), and smaller stands of whiplash willow (Salix lasiandra), and wolf willow ( ). Other common species are beaked sedge (Carex utriculata), bog birch (Betula glandulosa), mountain alder (Alnus incana), marsh reed grass (Calamagrostis canadensis), white clover (Trifolium repens), ALCES VOL. 41, 2005 DUNGAN AND WRIGHT - MOOSE SUMMER DIET COMPOSITION 141 and strawberry (Fragaria ovalis). These areas are surrounded by stands of ponderosa pine (Pinus ponderosa Pseudotsuga menziesii), quaking aspen (Populus tremuloi- des), and narrowleaf cottonwood (Populus angustifolia). High elevation meadows are characterized by large stands of plane-leaf wil- low, wolf willow, and bog birch. Surrounding trees include quaking aspen, lodgepole pine (Pinus contorta Abies lasiocarpa) (Beidleman et al. 2000). METHODS Over 3 million people visit RMNP an- nually. Viewing wildlife is a major visitor animals such as moose are visible from park roads. As a result, moose within RMNP are accustomed to people and their habituation enabled us to directly observe feeding be- havior and estimate diet composition. From 1 June through 15 September we observed the feeding behavior of moose at distances between 5-20 m. We recorded feeding data using hand-held voice recorders. Feeding data were grouped into feeding periods (bouts) which the moose fed continuously. Miquelle and Jordan (1979) reported more than 95% of all bites recorded occurred during such bouts. We counted individual bites taken by plant species when possible. Individual moose were followed as long as possible, and each continuous span was considered a single observation set. Bite counts were not conducted at night for observer’s safety and compliance with park regulations. During feeding periods the sizes of all bites were recorded to estimate intake leaves), medium (5-10 leaves), or large (> 10 leaves). After each observation set, simulated moose bites were collected by clipping 10-20 samples/species in each of the three sizes as closely to the observed bite size as possible. These samples were bagged, oven-dried at 60ºC for 48 hours, and weighed in accor- dance with methods described by Renecker and Hudson (1985). Average dry weight per bite (g/bite) for each size and species was calculated. Diet composition was based on percentage dry weight of species consumed by moose. This was derived by multiplying the number of bites taken of each species in each size class by the average dry weight per bite (Van Ballenberghe et al. 1989). Fresh fecal samples were collected from observed moose during foraging bouts. Sample size ranged from 6 to 17 per month, and included pellets collected in October and November, while walking low elevation transects, to look at early winter diets. These samples were frozen and sent to the Wildlife Habitat Lab at Washington State University for microhistological analysis. Sub-samples of each fecal pellet group were combined with others from the same month and year. Fecal samples were blended with water, stained with a lactophenol blue stain. Rela- tive cover (Korfhage 1974, Davitt 1979) of plant cuticle and epidermal fragments were - scope views on each of four slides (total 100 views) per month. A 10 square x 10 square grid was used to measure area covered by area covered were recorded by plant genus and species. Percent diet composition was calculated by dividing cover of each plant by total cover observed for all species, then multiplying by 100. We compared differences in percentages of major plant species between years, eleva- tions, months, and sexes for bite count data using a 1-way ANOVA. Salix species were pooled, and all percentages were Arcsine transformed in accordance with Krebs (1999). Experimental units were observations (n = 54), and pair-wise differences were located MOOSE SUMMER DIET COMPOSITION - DUNGAN AND WRIGHT ALCES VOL. 41, 2005 142 using the Tukey HSD procedure. Differences and all statistics were performed using SAS 8.3 (SAS Institute Inc., Cary, North Carolina) statistical software. RESULTS We recorded feeding data on 11 female and 43 male moose for a total of 54 observa- tion sets (Table 1). Over 75,000 bites were counted during 177 feeding bouts (Table 1). Data from direct observations showed no difference between sexes and elevations (P (F = 3.51; df = 3,48; P = 0.02) among months for - ly more mountain alder in late summer (Sep- tember) than early summer (June). Likewise, years for western dock (Rumex aquaticus) (F = 4.15; df = 1,48; P = 0.04) and trian- gular leaf Senecio (Senecio triangularis) (F = 5.71; df = 1,48; P = 0.02). Moose ate 2004. Other than the forementioned species, moose ate similar diets among months (P > 0.05) and years (P > 0.05). All bite count data were therefore pooled, for all animals, sexes, months, and years to estimate summer diet composition (Table 2). Moose consumed 11 different species of woody browse, which comprised 96.9% of the diet. Six willow species comprised 91.3 % of moose summer diets. Geyer willow accounted for 45.1% of summer diets followed by plane-leaf willow (22.7%), mountain willow (11.7%), and drum- Month Site (Elevation) Observations Female Male Feeding Bouts Bites June Low 15 4 11 27 9,807 July Low 17 4 13 60 20,324 High 3 0 3 15 11,013 August Low 7 3 4 32 15,277 High 4 0 4 17 10,544 September Low 6 0 6 21 7,692 High 2 0 2 5 2,322 Totals 54 11 43 177 76,979 Table 1. Summary of moose feeding data collected in Rocky Mountain National Park, Colorado from June 1st through September 15th, 2003 and 2004. Species Diet % Salix geyeriana 45.10 S. planifolia 22.70 S. monticola 11.70 S. drummondiana 9.90 Alnus incana 2.50 1.50 Aquatic spp. 1.30 Populus tremuloides 1.10 Betula glandulosa 0.97 Grasses 0.91 Cirsium spp. 0.71 Rumex aquaticus 0.47 Senecio triangularis 0.32 S. lasiandra 0.29 Nuphar lutea ssp. Polysepala 0.21 0.16 Psychrophila leptosepala 0.03 Mentha spicata 0.02 Pincus contorota Trace1 Shepherdia canadensis Trace1 Table 2. Percentage of plant species consumed by moose from June 1st through September 15th 2003 and 2004 in Rocky Mountain National Park, Colorado. Percentages were based on - sociated weights, measured in grams/bite. 1Trace species represented less than 0.01 % of the diet. ALCES VOL. 41, 2005 DUNGAN AND WRIGHT - MOOSE SUMMER DIET COMPOSITION 143 mond willow (9.9%). Geyer willow ranked it ranked third behind mountain willow and drummond willow. Other woody species included mountain alder (2.5%), quaking aspen (1.1%), and bog birch (1.0%). Aquatic species accounted for 1.9%, forbs 1.1%, and grasses at 0.9%. Species diversity peaked in July with moose eating 18 different species, including 7 species of non-woody browse. Monthly fecal analysis for 2003 showed moose relied more heavily on species other than willow in June, most notably Carex spp. (46.4%, Table 3), then increased willow con- sumption in July with a peak in August at 90%. In September, willow consumption began to decrease and early winter diets included spe- cies other than willow, most notably conifer needles (34.1%), shrubs (18.1%), and Carex spp. (11.6%, Table 3). Fecal analysis data were pooled for all months and years, with the exception Octo- ber/November 2003, to show summer moose diet composition (Table 4). Fecal analysis showed moose consume 79.3% willow, which is 11.9% less than direct observations Carex spp. as a major contributor to moose by direct observation. Fecal analysis was not able to identify forbs, willow, or shrubs - tions with the exception of quaking aspen (1.1%, Table 4). With the exception of Carex spp., both techniques showed similar results (Table 4). DISCUSSION Moose summer diets in RMNP consisted of 11 different species of woody browse, which accounted for roughly 97 % of the over- all diet. These results are similar to those found by Van Ballenberghe et al. (1989) in Denali National Park, Alaska, where woody species made up 96% of moose summer diets, and Joyal and Scherrer (1978) in Mont-Tremblant Park, Quebec, where moose summer diets consisted of 100% woody browse. Moose in RMNP use riparian willow communities dur- ing the summer, which contain little aquatic vegetation or forbs. Moose were observed eating 9 different non- woody species. Stud- ies in less mountainous habitats have found moose consume larger proportions of forbs (25%, LeResche and Davis 1973; 70.6%, Knowlton 1960) and aquatics (9.3%, McMil- lan 1953) than those in RMNP. Forbs and aquatics may contain higher concentrations of important minerals (Belovsky 1978), and have higher digestibility levels than woody browse (LeResche and Davis 1973). Forbs and aquatics only accounted for 2.4% of the overall summer diet of moose in RMNP. Moose ate 6 willow species comprising 91.3% of their summer diets, with geyer willow accounting for 45.1%. Results were similar to those found by McMillan (1953) for Plants Jun-03 Jul-03 Aug-03 Sep-03 Oct/Nov 03 2004 Grasses 2.10% 0.00% 0.60% 0.80% 8.90% 7.70% Carex spp. 46.40% 5.00% 0.10% 3.80% 11.60% 3.60% Salix spp. 48.00% 88.20% 90.00% 82.90% 25.30% 81.40% Shrubs 0.40% 6.00% 9.30% 11.50% 18.10% 6.90% Conifer needle 0.70% 0.00% 0.00% 0.00% 34.10% 0.20% Forbs 2.40% 0.60% 0.00% 1.00% 2.00% 0.00% Sphagnum Moss 0.00% 0.00% 0.00% 0.00% 0.00% 0.20% Insect 0.00% 0.20% 0.00% 0.00% 0.00% 0.00% Table 3. Percent of major plant species consumed by moose in Rocky Mountain National Park, Colo- rado per month for 2003, and by year for 2004, as indicated by fecal analysis. MOOSE SUMMER DIET COMPOSITION - DUNGAN AND WRIGHT ALCES VOL. 41, 2005 144 moose in Yellowstone National Park (willow 88.5%) and Van Ballenberghe (1989) in Denali National Park (81.5%). Similarities of diet composition of moose in RMNP and moose in Denali National Park suggest use of very similar habitats during the summer. Willow habitats not only provide a high quality woody browse for consumption (LeResche and Davis 1973) but are also used extensively for cover (Kufeld and Bowden 1996). Fecal analysis showed moose rely more heavily on grasses, sedges, and forbs (50.9%) in early summer than woody browse (49.1%). Peek (1974) found that grasses and sedges were seldom consumed other than in spring, when digestibility and nutrient content are high. Consumption of willow species in- creased in July, peaked in August, and started to decline in September. Both techniques showed the same trend, with exception of June bite count data, suggesting moose eat willow when available biomass and nutrients are high (Stumph 2005) and rely less heavily on willow in spring and fall when available biomass and nutrients are lower. Early winter fecal analysis showed willow comprised only 25.3 % of the diet, whereas conifer needles made up 34.1%, and grasses, forbs, and sedges made up 22.5 %. Forbs and grasses comprised relatively larger percentages of fall diets than winter diets (Peek 1974). During both years we observed moose moving out of the lower riparian willow communities, with the onset of the rut, and moving into higher more heavily forested areas of the park, possibly giving a reason for the large percent of conifer needles in early winter diets. In a study conducted from December 1991 to November 1995, moose from North Park, Colorado, tended to winter at lower elevations and move to higher elevations during spring, summer, and fall (Kufeld and Bowden 1996). Winter transect data showed little use of low elevation riparian willow communities from October through December for both years. Willows were the primary forage plants in 5 out of 6 winter diet studies reviewed by Peek (1974), and Risenhoover (1987) reported that willow comprised 94.3% of winter diets in Denali National Park. Stevens (1970) reported timber types received 82% of total use during the winter in the Gallatin region of Montana. October/November fecal analyses coupled with winter transect data suggest that willow may not constitute a large proportion of moose winter diets in RMNP, although a comprehensive study of winter diets has not been performed. Similar results were achieved using bite count data and fecal analysis. Direct observa- tions failed to detect the large amount of sedges consumed by moose in early summer, which was largely affected by adult movements during June. Older, more habituated moose had not yet moved to summer range from Table 4. Percent diet composition of major plant species consumed by moose in Rocky Mountain National Park, Colorado from June 1st through September 15th, 2003 and 2004, using direct observations and fecal analysis. All months and years were pooled for both techniques. 1Shrubs consisted of Alnus incana, Betula glan- dulosa, , and Shep- herdia Canadensis. 2Aquatics consisted of Aquatic spp., Nuphar lutea spp. Polysepala, and Rumex aquaticus. 3Forbs consisted of Cirsium spp., Mentha spi- cata, Psychrophila leptosepala, and Senecio triangularis. 4Trace species represented less than 0.01 % of the diet. Species Fecal Analysis Bite Count Salix spp. 79.3 91.3 Carex spp. 8.7 - Shrubs 6.8 3.61 Grasses 4.3 0.91 Aquatics 0.11 1.982 Populus tremuloides - 1.12 Forbs 0.51 1.083 Conifer needles 0.19 Trace4 Insects 0.02 - ALCES VOL. 41, 2005 DUNGAN AND WRIGHT - MOOSE SUMMER DIET COMPOSITION 145 winter range, necessitating that our sample be focused on younger, less habituated moose. This caused us to miss a large proportion of feeding data per day, which could account for some of the variation. Grasses were also slightly higher (3.4%) in fecal analysis than direct observations. Consumption of grasses and sedges is hard to observe because of their low growth form, and bites were not recorded which could also lead to differences between the two techniques. Wallmo et al. (1973) reported bite count methods estimated more use of shrubs as a class and less use of grass and forbs than actually occurred for mule deer (Odocoileus hemionus). Fecal analysis could not differentiate willow, forb, or shrub species, and failed to detect Populus tremuloides. Other species not detected by fecal analysis included Cirsium spp., Rumex aquaticus, Senecio triangularis, and Nuphar lutea ssp. Polysepala, but these species were less than 1 % of the observed diet. Monthly fecal sample size was smaller than similar studies, which could have contributed to not detecting these species. ACKNOWLEDGEMENTS John Skinner, Basil Iannone, and Mandy Cluck who spent countless hours observing moose night and day. Park service staff were more than helpful all 3 years of the study, with special thanks going to Ryan Monello. Funding for this project was provided by Rocky Mountain National Park through the USGS Idaho Cooperative Fish and Wildlife Research Unit. This manuscript is dedicated to the memory of Dr. Francis Singer. REFERENCES AUGUSTINE, D. J., and S. J. MCNAUGHTON. 1998. Ungulate effects on the functional species composition of plant communi- ties: herbivore selectivity and plant tol- erance. Journal of Wildlife Management 62:1165-1183. BAILEY, A. 1944. Records of moose in Colo- rado. Journal of Mammalogy 25:192- 193. BEIDLEMAN, L. H., R. G. BEIDLEMAN, and B. E. WILLARD. 2000. Plants of Rocky Mountain National Park. Falcon, Helena, Montana, USA. BELOVSKY, G. 1978. Diet optimization in a generalist herbivore: the moose. Theoreti- cal Population Biology 14:105-134. BUTLER, C. E. 1986. Summer food utiliza- tion and observations of a tame moose, Alces alces. Canadian Field Naturalist 100:85-88. CRETE, M. 1989. Approximation of K car- rying capacity for moose in eastern Quebec. Canadian Journal of Zoology 67:373-380. DAVITT, B. B. 1979. Elk summer diet composi- tion and quality on the Colockum Multiple Use Research Unit, Central Washington. M.S. Thesis. Washington State University, Pullman, Washington, USA. JOYAL, R., and B. SCHERRER. 1978. Summer movement and feeding by moose in west- ern Quebec. Canadian Field Naturalist 92:252-258. KNOWLTON, F. F. 1960. Food habits, move- ment, and populations of moose in Gravely Mountains, Montana. Journal of Wildlife Management 24:162-170. KORFHAGE, R. C. 1974. Summer food habits of elk in the Blue Mountains of north- eastern Oregon based on fecal analysis. M.S. Thesis. Washington State University, Pullman, Washington, USA. KREBS, C. J. 1999. Ecological Methodology. Second edition. Benjamin/Cummings, New York, New York, USA. KUFELD, R. C., and D. C. BOWDEN. 1996. Movement and habitat selection of Shiras moose (Alces alces shirasi) in Colorado. Alces 32:85-99. _____, and S. F. STEINERT. 1990. An estimate of moose carrying capacity in willow MOOSE SUMMER DIET COMPOSITION - DUNGAN AND WRIGHT ALCES VOL. 41, 2005 146 habitat in North Park, Colorado. Colorado Division of Wildlife Unpublished Report. Fort Collins, Colorado, USA. LERESCHE, R. E., and J. L. DAVIS. 1973. Im- portance of nonbrowse foods to moose on the Kenai Peninsula, Alaska. Journal of Wildlife Management 37:279-287. MACCRACKEN, J. G., V. VAN BALLENBERGHE, and J. M. PEEK. 1997. Habitat relation- ships of moose on the Copper River delta in coastal south-central Alaska. Wildlife Monographs 136. MCINNES, P. F., R. J. NAIMAN, J. PASTOR, and Y. COHEN. 1992. Effects of moose browsing on vegetation and litter of the boreal forest, Isle Royale, Michigan, USA. Ecology 73:2059-2075. MCMILLAN, J. F. 1953. Some feeding habits of moose in Yellowstone National Park. Ecology 34:102-110. MIQUELLE, D. G., and P. A. JORDAN. 1979. The importance of diversity in the diet of moose. Proceedings of the North Ameri- can Moose Conference and Workshop 15:54-79. _____, and V. VAN BALLENBERGHE. 1989. Impact of bark stripping by moose on aspen-spruce communities. Journal of Wildlife Management 53:577-586. MONELLO, R., and T. JOHNSON. 2003. The elk herd in Rocky Mountain National Park. Rocky Mountain National Park Unpublished Report. Estes Park, Colo- rado, USA. PASTOR, J., B. DEWEY, R. J. NAIMAN, P. F. MCINNES, and Y. COHEN. 1993. Moose browsing and soil fertility in the boreal forest of Isle Royale National Park. Ecol- ogy 74:467-480. PEEK, J. M. 1974. A review of moose food habit studies in North America. Naturali- ste Canadien 101:195-215. RENECKER, L. A., and R. J. HUDSON. 1985. Estimation of dry matter intake of free- ranging moose. Journal of Wildlife Management 49:785-792. RISENHOOVER, K. L. 1987. Winter foraging strategies of moose in subarctic and bo- real forest habitats. Ph.D. Dissertation. Michigan Technical University, Hough- ton, Michigan, USA. SCHWARTZ, C. C. 1992. Physiological and nutritional adaptations of moose to north- ern environments. Alces Supplement 1:139-155. STEVENS, D. R. 1970. Winter ecology of moose in the Gallitin Mountains, Mon- tana. Journal of Wildlife Management 34:37-46. _____. 1988. Moose in Rocky Mountain National Park. Rocky Mountain National Park Unpublished Report. Estes Park, Colorado, USA. STUMPH, B. P. 2005. The summer forage qual- - ence on moose forage ecology in Rocky Mountain National Park, Colorado. M.S. Thesis. University of Idaho, Moscow, Idaho, USA. VAN BALLENBERGHE, V., D. G. MIGUELLE, and J. G. MACCRACKEN. 1989. Heavy utili- zation of woody plants by moose during summer at Denali National Park, Alaska. Alces 25:31-35. WALLMO, O. C., R. B. GILL, L. H. CARPENTER, and D. W. REICHERT. 1973. Accuracy of of Wildlife Management 37:556-562. ZEIGENFUSS, L. C., F. J. SINGER, and D. BOWDEN. 1999. Vegetation responses to natural regulation of elk in Rocky Mountain National Park. Biological Science Report USGS/BRD - 1999 - 003. U.S. Govern- USA. ZHELEZNOV-CHUKOTSKY, N. K., and E. S. VO- TIASHOVA. 1998. Comparative analysis of moose nutrition of the Anadyrsky and Omolonsky populations (far north east) in different seasons. Alces 34:445-451. << /ASCII85EncodePages false /AllowTransparency false /AutoPositionEPSFiles true /AutoRotatePages /All /Binding /Left /CalGrayProfile (Dot Gain 20%) /CalRGBProfile (sRGB IEC61966-2.1) /CalCMYKProfile (U.S. Web Coated \050SWOP\051 v2) /sRGBProfile (sRGB IEC61966-2.1) /CannotEmbedFontPolicy /Warning /CompatibilityLevel 1.4 /CompressObjects /Tags /CompressPages true /ConvertImagesToIndexed true /PassThroughJPEGImages true /CreateJDFFile false /CreateJobTicket false /DefaultRenderingIntent /Default /DetectBlends true /DetectCurves 0.0000 /ColorConversionStrategy /LeaveColorUnchanged /DoThumbnails false /EmbedAllFonts true /EmbedOpenType false /ParseICCProfilesInComments true /EmbedJobOptions true /DSCReportingLevel 0 /EmitDSCWarnings false /EndPage -1 /ImageMemory 1048576 /LockDistillerParams false /MaxSubsetPct 100 /Optimize true /OPM 1 /ParseDSCComments true /ParseDSCCommentsForDocInfo true /PreserveCopyPage true /PreserveDICMYKValues true /PreserveEPSInfo true /PreserveFlatness true /PreserveHalftoneInfo false /PreserveOPIComments false /PreserveOverprintSettings true /StartPage 1 /SubsetFonts true /TransferFunctionInfo /Apply /UCRandBGInfo /Preserve /UsePrologue false /ColorSettingsFile () /AlwaysEmbed [ true ] /NeverEmbed [ true ] /AntiAliasColorImages false /CropColorImages true /ColorImageMinResolution 300 /ColorImageMinResolutionPolicy /OK /DownsampleColorImages true /ColorImageDownsampleType /Bicubic /ColorImageResolution 300 /ColorImageDepth -1 /ColorImageMinDownsampleDepth 1 /ColorImageDownsampleThreshold 1.50000 /EncodeColorImages true /ColorImageFilter /DCTEncode /AutoFilterColorImages true /ColorImageAutoFilterStrategy /JPEG /ColorACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /ColorImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000ColorACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000ColorImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasGrayImages false /CropGrayImages true /GrayImageMinResolution 300 /GrayImageMinResolutionPolicy /OK /DownsampleGrayImages true /GrayImageDownsampleType /Bicubic /GrayImageResolution 300 /GrayImageDepth -1 /GrayImageMinDownsampleDepth 2 /GrayImageDownsampleThreshold 1.50000 /EncodeGrayImages true /GrayImageFilter /DCTEncode /AutoFilterGrayImages true /GrayImageAutoFilterStrategy /JPEG /GrayACSImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /GrayImageDict << /QFactor 0.15 /HSamples [1 1 1 1] /VSamples [1 1 1 1] >> /JPEG2000GrayACSImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /JPEG2000GrayImageDict << /TileWidth 256 /TileHeight 256 /Quality 30 >> /AntiAliasMonoImages false /CropMonoImages true /MonoImageMinResolution 1200 /MonoImageMinResolutionPolicy /OK /DownsampleMonoImages true /MonoImageDownsampleType /Bicubic /MonoImageResolution 1200 /MonoImageDepth -1 /MonoImageDownsampleThreshold 1.50000 /EncodeMonoImages true /MonoImageFilter /CCITTFaxEncode /MonoImageDict << /K -1 >> /AllowPSXObjects false /CheckCompliance [ /None ] /PDFX1aCheck false /PDFX3Check false /PDFXCompliantPDFOnly false /PDFXNoTrimBoxError true /PDFXTrimBoxToMediaBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXSetBleedBoxToMediaBox true /PDFXBleedBoxToTrimBoxOffset [ 0.00000 0.00000 0.00000 0.00000 ] /PDFXOutputIntentProfile () /PDFXOutputConditionIdentifier () /PDFXOutputCondition () /PDFXRegistryName () /PDFXTrapped /False /Description << /CHS /CHT /DAN /DEU /ESP /FRA /ITA /JPN /KOR /NLD (Gebruik deze instellingen om Adobe PDF-documenten te maken voor kwaliteitsafdrukken op desktopprinters en proofers. De gemaakte PDF-documenten kunnen worden geopend met Acrobat en Adobe Reader 5.0 en hoger.) /NOR /PTB /SUO /SVE /ENU (Use these settings to create Adobe PDF documents for quality printing on desktop printers and proofers. Created PDF documents can be opened with Acrobat and Adobe Reader 5.0 and later.) >> /Namespace [ (Adobe) (Common) (1.0) ] /OtherNamespaces [ << /AsReaderSpreads false /CropImagesToFrames true /ErrorControl /WarnAndContinue /FlattenerIgnoreSpreadOverrides false /IncludeGuidesGrids false /IncludeNonPrinting false /IncludeSlug false /Namespace [ (Adobe) (InDesign) (4.0) ] /OmitPlacedBitmaps false /OmitPlacedEPS false /OmitPlacedPDF false /SimulateOverprint /Legacy >> << /AddBleedMarks false /AddColorBars false /AddCropMarks false /AddPageInfo false /AddRegMarks false /ConvertColors /NoConversion /DestinationProfileName () /DestinationProfileSelector /NA /Downsample16BitImages true /FlattenerPreset << /PresetSelector /MediumResolution >> /FormElements false /GenerateStructure true /IncludeBookmarks false /IncludeHyperlinks false /IncludeInteractive false /IncludeLayers false /IncludeProfiles true /MultimediaHandling /UseObjectSettings /Namespace [ (Adobe) (CreativeSuite) (2.0) ] /PDFXOutputIntentProfileSelector /NA /PreserveEditing true /UntaggedCMYKHandling /LeaveUntagged /UntaggedRGBHandling /LeaveUntagged /UseDocumentBleed false >> ] >> setdistillerparams << /HWResolution [2400 2400] /PageSize [612.000 792.000] >> setpagedevice