Alces (A Journal Devoted to the Biology and Management of Moose)
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STRUCTURE OF MOOSE (ALCES ALCES) POPULATIONS IN RUSSIA WITH SPECIAL REFERENCE TO COMMUNICATION DISTANCES
The structure of moose populations was studied in the southern subzone of taiga and mixed forests. Moose were distributed irregularly in small groups. Some migrations were recorded. The distance between moose in compact groups (5–9.3 m) and the distance between groups were compared to orientation vectors a–k, representing one of the parameters of the inner activity rhythms. The distances and the vectors correspond to the definition of the critical levels of the development of natural systems. The spatial pattern of moose distribution was revealed and a canonic distribution model developed
GENETIC RELATIONSHIPS DEDUCED FROM CYTOCHROME-b SEQUENCES AMONG MOOSE
We studied variation in nucleotide sequences of the mitochondrial cytochrome-b gene to assess the phylogeny of moose (Alces alces) in general, and the position of North American moose within that phylogeny in particular. We combined North American, Asian, and European haplotypes generated for this study with 3 Eurasian haplotypes obtained from GenBank. No nucleotide variation occurred within moose from North America, whereas 3 haplotypes were present in European moose and 4 haplotypes in Asian moose. Clade structure was consistent over 6 most-parsimonious trees, with Asian haplotypes composing 1 clade, and North American and European haplotypes composing a second, albeit poorly supported clade. Low diversity of nucleotides in cytochrome-b indicated a recent ancestry among moose worldwide. Existence of 1 North American haplotype is strong evidence of a single, recent entry into the New World via the Bering land bridge, rather than multiple entries through >1 corridors. Furthermore, no phylogenetic support existed for the theory of distinct lineages of European versus Asian-North American moose
ECOSYSTEM MANAGEMENT AND MOOSE: CREATING A COHERENT CONCEPT WITH FUNCTIONAL MANAGEMENT STRATEGIES
Ecosystem management is a popular but poorly defined concept in conservation biology. Current vague, non-operational definitions provoke criticism of the concept and undermine credibility of its associated principles. We propose a definition of ecosystem management that emphasizes essential qualities of the concept rather than its accidental associations or properties, and that explains functional and operational attributes of ecosystem management rather than its descriptive characteristics. Based on these criteria, we offer a definition of ecosystem management as “a pattern of prescribed, goal-oriented environmental manipulations that: (1) treat a specified ecological system of identifiable boundaries as the fundamental unit to be managed; (2) has, as its desired outcome, the achievement of a state or collection of states in the ecosystem such that historical components, structure, function, products, and services of the ecosystem persist within biologically normal ranges and with normal rates of change; (3) uses naturally occurring, landscapescale processes as the primary means of management; and (4) determines management objectives through cooperative decision-making of individuals and groups who reside in, administer, and/or have vested interests in the state of the ecosystem”. Achieving workable ecosystem management is currently hindered by the lack of a unified vision and system of values for ecosystems, the absence of permanent inter-agency bodies with authority to manage ecosystems across multiple jurisdictions, and the lack of administrative mechanisms for the translation of ecosystem research findings into ecosystem management policies. We propose strategies to overcome these obstacles and examine moose (Alces alces) as an example of a species that is both important to ecosystem management and may benefit from it
MODELING OF MOOSE HUNTING: PROTECTION OF COWS WITH TWINS
I developed a simulation model to evaluate moose hunting strategies. The model incorporated age and sex-specific schedules for natural (nonhunting) mortality and fecundity. I evaluated 2 strategies for harvesting cow moose. In the first, cow moose were harvested irrespective of whether they were accompanied by calves. In the second, hunters were not allowed to kill cows accompanied by twin calves. Simulation results indicated that 500 calves/1000 cows could be saved under the second harvest strategy
THE IMPACT OF MOOSE ON ASH PRODUCTIVITY
Data characterizing the impact of moose (Alces alces) on ash (Fraxinus spp.) seedlings in the broad–leaved forests of the Tula region are given. Resistance of ash to the strong browsing pressure is shown and also the resulting peculiarities of crown structure, including the position of shoots and branches. When ash is isolated from moose, annual accretion of shoot phytomass is 5 times more than in locations where ash is exposed to browsing; in leaves, phytomass is 10–12 times more than browsed ash. The large number of shoots and a great quantity of large leaves are considered as an adaptation of the ash tree to survival under the browsing pressure of moose
EFFECTS OF ANIMAL ACTIVITY ON GPS TELEMETRY LOCATION ATTEMPTS
Interpretation of habitat use from GPS collar locations could be biased if the activity of animals wearing GPS collars affects the probability of obtaining a successful location. We tested for this bias with GPS location attempts made by collars deployed on free-ranging moose (Alces alces) in northern Minnesota, USA. We classified moose as being either inactive or active during each GPS location attempt based on activity counts recorded by the collar. Only 69% of GPS location attempts were successful while moose were active, compared to 88% of when moose were inactive. Moose activity reduced success of location attempts in both summer and winter. We also estimated the precision of GPS locations while collars were deployed on free-ranging moose. When moose were inactive 50% of 3-dimensional locations were within 5 m of the estimated location, and 95% were within 17 m of the estimated location. When moose were inactive, 50% of 2-dimensional locations were within 7 m of the estimated location, and 95% were within 26 m of the estimated location. Despite the bias induced by animal activity, GPS telemetry is the most precise method currently available to obtain locations of free-ranging large mammals such as moose. Sampling biases in GPS units resulting from animal activity should be accounted for when interpreting habitat use by free ranging animals
SEASONALITY OF SEXUAL SEGREGATION IN DIMORPHIC DEER: EXTENDING THE GASTROCENTRIC MODEL
We extend our gastrocentric model for sexual segregation in dimorphic deer (Cervidae) to include seasonal metabolism of energy and protein in reproductive female, nonreproductive female, and male reindeer and caribou (Rangifer tarandus). We define sexual segregation as the differential use of space by the sexes of animals outside the mating season. Minimum density of energy and protein in the diet is predicted to favor sexual segregation in late winter and summer, but not during autumn. Demands for energy during gestation and for energy and protein during lactation increase dietary minima for reproductive females above levels required by nonreproductive females and males. Projections of gross energy and total protein in diets based on estimated metabolizabilities for males and nonreproductive females predicted that males could subsist on the lowest-quality diets during summer. Projections of minimal dietary content of gross energy indicated that reproductive females probably would feed longer than nonreproductive females and males during summer, as suggested by field observations. Differences in feeding activity between the sexes may be a consequence of metabolic demands and, therefore, and outcome of sexual segregation rather than its cause. The gastrocentric model provides and explanation of behavioral ecology of deer with physiological data for 1 species. Further integration of these disciplines may predict the consequences of size and sex on niche partitioning in relation to environmental changes in forage quality and quantity. Nonetheless, simplistic models are only as valid as data upon which they are constructed. Our model indicated the dire need for studies of diet selection and nutritional physiology in the context of both reproductive status and seasonal demands for both sexes of cervids.
UTILITY OF STABLE ISOTOPE ANALYSIS IN STUDYING FORAGING ECOLOGY OF HERBIVORES: EXAMPLES FROM MOOSE AND CARIBOU
Recently, researchers emphasized that patterns of stable isotope ratios observed at the individual level are a result of an interaction between ecological, physiological, and biochemical processes. Isotopic models for herbivores provide additional complications because those mammals consume foods that have high variability in nitrogen concentrations. In addition, distribution of amino acids in plants may differ greatly from that required by a herbivore. Thus, substantial amounts of amino acids would be synthesized or recycled by herbivores. At northern latitudes, were the growing season of vegetation is short, isotope ratios in herbivore tissues are expected to differ between seasons. Summer ratios likely reflect diet composition, whereas winter ratios would reflect diet and nutrient recycling by the animals. We tested this hypothesis using data collected from blood samples of caribou (Rangifer tarandus) and moose (Alces alces) in Denali National Park and Preserve, Alaska, USA. Stable isotope ratios of moose and caribou were significantly different from each other in late summer-autumn and winter. Also, late summer-autumn and winter ratios differed significantly between seasons in both species. Nonetheless, we were unable to evaluate whether differences in seasonal isotopic ratios were a result of diet selection or a response to nutrient recycling. We believe that additional studies on plant isotopic ratios as related to ecological factors in conjunction with investigations of diet selection by the herbivores will enhance our understanding of those interactions. Also, controlled studies investigating the relation between diet and physiological responses in herbivores will increase the utility of isotope analysis in studying foraging ecology of herbivores
EFFECTS OF MARCESCENT LEAVES ON WINTER BROWSING BY LARGE HERBIVORES IN NORTHERN TEMPERATE DECIDUOUS FORESTS
Presence of marcescent leaves during winter is a common phenomenon in northern-temperate deciduous forests across the Holarctic, but the ecological significance of marcescence on woody vegetation has received little attention. Especially, implications for browsing by cervids during winter have not been determined. Therefore, I conducted a feeding trial using free-roaming red deer (Cernus elaphus), fallow deer (Dama dama), and sika deer (Cervus nippon) to evaluate effects of marcescent leaves on herbivory of European beech (Fagus sylvatica), hornbeam (Carpinus betulus), and common oak (Quercus robur) during winter. The feeding trial was conducted in Jægersborg Dyrehave, Denmark, during February 1992. Forty small-diameter (< 6 mm) branches of each tree species with < 2 years of growth and marcescent leaves present were divided into 2 groups. One group had their leaves removed (treatment) and the other group had the leaves retained (control). Branches were labeled, weighed, and randomly placed in a grid (0.5 x 0.5 m apart) in the field. After 7 days, branches were counted and weighed to determine amount of browse removed. In addition, 12 branches of each species were collected for biomass and chemical analyses. Stems and leaves were analyzed separately for crude protein, neutral-detergent fiver, acid-detergent fiber, lignin, and cellulose. Both beech and hornbeam were browsed significantly more by weight and by number of branches when the leaves were removed. Oak branches were browsed the same regardless of leaf presence. Biomass or marcescent leaves of beech and hornbeam were a significantly greater proportion of the overall branch biomass compared with oak. Chemical analyses showed that stems of hornbeam and beech, when compared with their marcescent leaves has a higher lignin content. Leaves compared with stems of common oak had a higher protein and lower lignin content compared with other species. These results indicate that marcescent leaves greatly reduced the nutritive value of winter browse, which was reflected in the lower browse preference for their leaves. Therefore, marcescent leaves may be viewed as an anti-herbivore mechanism
REGIONAL VARIATION IN MINERAL CONTENTS OF PLANTS AND ITS SIGNIFICANCE FOR MIGRATION BY ARCTIC REINDEER AND CARIBOU
Ten minerals, 6 macro (Na, K, Ca, P, Mg, Cl) and 4 trace elements (Co, Cu, Mo, Mn), were analysed in 13 forage plants of reindeer and caribou (Rangifer tarandus) to compare differences between coastal and interior areas. Samples were collected in northern Norway (coastal and interior regions), southern Norway (interior), and Alaska, USA (interior). We tested the hypothesis that the domestic herding practice of moving reindeer to spring-summer pasture on the coast is to allow reindeer to make up mineral balances that are negative during winter. This hypothesis was supported by data for Na and Cl, which were higher in 12 of 13 forage plants from the coastal region compared with inland areas. Analyses of other minerals, however, indicated a higher variability among plants than between regions. Aquatic plants from the coast and inland were higher in Na and Cl that terrestrial species. A high concentration of Co in willows was independent of region. Graminoids were low in Na and Cl, independent of region. Lichens were low in all macro minerals but were high in trace minerals. This study supported hypotheses based on salt hunger; namely, that the primary reason to move in coastal regions was to compensate for Na deficiency in winter. We suggest this movement also would maximize milk synthesis, which would otherwise be limited because of high Na content of reindeer milk. Selective foraging within coastal vegetation allows animals to meet requirements of macrominerals. Selective use of willows not only supports the high protein requirements of lactation and growth, but an adequate Co intake is required for synthesis of vitamin B12, critical for animal growth and rapid development of rumen function in young reindeer and caribou