The relationship between habitat choice and lifetime reproductive success in female red deer

In non-territorial species, individuals can move freely and should be distributed in an ideal free manner between habitats and areas with respect to resources that influence lifetime reproductive success (LRS). Consequently, no relationship between diet quality and LRS should be expected. However, there have been no attempts to test this prediction. The present paper investigates the relationship between forage habitat use and LRS in red deer (Cervus elaphus) hinds within three neighbouring areas on the Isle of Rum which differed in their amounts of high-quality-forage habitat. Within areas, hinds move widely and have access to the same resources. We found no correlation between LRS of individual hinds and their use of high-quality-forage habitat (i.e. short Agrostis/Festuca grassland). Our analysis suggests that high hind densities on short Agrostis/Festuca grassland offset any advantages of increased access to preferred forage. These results support the hypothesis that red deer hinds are distributed in an ideal free manner with respect to the use of high-quality-forage habitat. However, hinds rarely leave areas where they are born, and the analysis suggests that constraints in changing areas hindered an ideal free distribution on a larger spatial scale. Consequently, mean LRS was not the same within the three investigated areas: one area, with a low amount of short Agrostis/Festuca grassland and a low hind density, contributed more male offspring (and more total offspring) per hind to the population than the other two areas. Received: 11 September 1998 / Accepted: 15 March 1999     

Long-term density-dependent changes in habitat selection in red deer (Cervus elaphus)

Understanding how habitat selection changes with population density is a key concept in population regulation, community composition and managing impacts on biodiversity and ecosystem services. At low density, it is expected that individuals select habitats in terms of their preference, but as population density increases, the availability of resources per individual declines on preferred habitats, leading to competition which forces some individuals to exploit less preferred habitats. Using spatial information of Scottish red deer (Cervus elaphus) winter counts, carried out in 110 areas across Scotland between 1961 and 2004 (a total of 1,206,495 deer observations), we showed how winter habitat niche breadth in red deer has widened with increasing population density. Heather moorland and montane habitats were most and least preferred for deer, respectively. Increasing density favoured the selection of grassland, to the detriment of the selection of heather moorland. The selection of heather and grassland decreased when temperature increased, while the selection of montane and peatland habitats increased. These findings are important for understanding how habitat use, density and population are likely to be affected by weather, and allow us to predict habitat impacts by large mammal herbivory and climate.     

Skewed distributions of lifetime reproductive success: beyond mean and variance

Lifetime reproductive performance is quantified here by the LRS (lifetime reproductive success), the random number of offspring an individual produces over its lifetime. Many field studies find that distributions of LRS among individuals are non‐normal, zero‐inflated and highly skewed. These results beg the question, what is the distribution of LRS predicted by demographic models when the only source of randomness is demographic stochasticity? Here we present the first exact analysis of the probability distribution of LRS for species described by age + stage models; our analysis starts with estimated vital rates. We illustrate with three examples: the Hadza, human hunter‐foragers (age‐only), the evergreen tree Tsuga canadensis (stage‐only) and Roe deer, Capreolus capreolus (age + stage). For each we obtain the exact distribution of LRS, but also calculate and discuss the first three moments. Our results point to important questions about how such LRS distributions affect natural selection, and life history evolution.     

Mating system, sexual dimorphism, and the opportunity for sexual selection in a territorial ungulate

In mammals, species with high sexual size dimorphism tend tohave highly polygynous mating systems associated with high variancein male lifetime reproductive success (LRS), leading to a highopportunity for sexual selection. However, little informationis available for species with weak sexual size dimorphism. Ina long-term study population, we used parentage analysis basedon 21 microsatellite markers to describe, for the first time,variance in male lifetime breeding success (LBS) of roe deer,a territorial ungulate where males weigh less than 10% morethan females. LBS ranged from 0 to 14 (mean = 4.54, variance= 15.5), and its distribution was highly skewed, with only afew males obtaining high LBS and many males failing to breedor siring only one fawn. As predicted for polygynous specieswith low sexual size dimorphism, the standardized variance inmale LBS was low (I_m = 0.75) and was only slightly higher thanthe standardized variance in female LRS (I_f = 0.53), suggestinga low opportunity for sexual selection. The I_m value reportedhere for roe deer is much lower than values reported for highlydimorphic ungulates such as red deer (I_m > 3). We suggestthat, along a continuum of opportunity for sexual selection,roe deer occupy a position closer to monogamous and monomorphicterritorial ungulates than to highly polygynous, sexually dimorphicungulates with dominance rank–based mating systems suchas harems or roving mating systems.     

Consequences of migratory strategy on habitat selection by mule deer

Ungulate behavior is often characterized as balancing selection for forage and avoidance of predation risk. Within partially migratory ungulate populations, this balancing occurs across multiple spatial scales, potentially resulting in different exposure to costs and benefits between migrants and residents. We assessed how availability and selection of forage and risk from predators varied between summer ranges of migrant and resident mule deer (Odocoileus hemionus; a species in which individual migratory strategies are generally fixed for life) in 3 study areas in western Montana, USA, during summers 2017–2019. We hypothesized that mule deer would face a tradeoff between selecting forage and avoiding predation risk, and that migration and residency would pose contrasting availability of forage and risk at a broad (summer range) spatial scale. We hypothesized deer exposed to lower forage at a given spatial scale would compensate for reduced availability by increasing selection of forage at the cost of reduced avoidance of predators, a mechanism whereby migrants and residents could potentially achieve similar exposure to forage despite disparate availability. We compared the availability of forage (kcal/m²) and predation risk from wolves (Canis lupus) and mountain lions (Puma concolor) between summer ranges of each migratory strategy, then assessed how selection for those factors at the home range (second order) and within-home range (third order) scales varied using resource selection functions (RSFs). As forage availability increased among mule deer summer ranges and individual home ranges, selection for forage decreased at the second-order (P = 0.052) and third-order (P = 0.081) scales, respectively, but avoidance of predators varied weakly. In 1 study area, summer range of residents contained lower forage and higher risk than summer range of migrants, but residents compensated for this disadvantage through stronger selection of forage and avoidance of risk at finer spatial scales. In the other 2 study areas, summer range of migrants contained lower forage and higher risk than residents, but migrants did not compensate through stronger selection for beneficial resources. The majority of mule deer in our study system were migratory, though the benefits of migration were unclear, suggesting partial migration may persist in populations even when exposure to forage and predation risk appears unequal between strategies.     

Early determinants of lifetime reproductive success differ between the sexes in red deer.

In polygynous, sexually dimorphic species, sexual selection should be stronger in males than in females. Although this prediction extends to the effects of early development on fitness, few studies have documented early determinants of lifetime reproductive success in a natural mammal population. In this paper, we describe factors affecting the reproductive success of male and female red deer (Cervus elaphus) on the island of Rum, Scotland. Birthweight was a significant determinant of total lifetime reproductive success in males, with heavier-born males being more successful than lighter ones. In contrast, birthweight did not affect female reproductive success. High population density and cold spring temperatures in the year of birth decreased several components of fitness in females, but did not affect the breeding success of males. The results confirm the prediction that selection on a sexually dimorphic trait should be greater in males than in females, and explain the differential maternal expenditure between sons and daughters observed in red deer. Differences between the sexes in the effects of environmental and phenotypic variation on fitness may generate differences in the amount of heritable genetic variation underlying traits such as birthweight.     

A Comparison of Two Modeling Approaches for Evaluating Wildlife-Habitat Relationships

ABSTRACT Studies of resource selection form the basis for much of our understanding of wildlife habitat requirements, and resource selection functions (RSFs), which predict relative probability of use, have been proposed as a unifying concept for analysis and interpretation of wildlife habitat data. Logistic regression that contrasts used and available or unused resource units is one of the most common analyses for developing RSFs. Recently, resource utilization functions (RUFs) have been developed, which also predict probability of use. Unlike RSFs, however, RUFs are based on a continuous metric of space use summarized by a utilization distribution. Although both RSFs and RUFs predict space use, a direct comparison of these 2 modeling approaches is lacking. We compared performance of RSFs and RUFs by applying both approaches to location data for 75 Rocky Mountain elk (Cervus elaphus) and 39 mule deer (Odocoileus hemionus) collected at the Starkey Experimental Forest and Range in northeastern Oregon, USA. We evaluated differences in maps of predicted probability of use, relative ranking of habitat variables, and predictive power between the 2 models. For elk, 3 habitat variables were statistically significant (P < 0.05) in the RSF, whereas 7 variables were significant in the RUF. Maps of predicted probability of use differed substantially between the 2 models for elk, as did the relative ranking of habitat variables. For mule deer, 4 variables were significant in the RSF, whereas 6 were significant in the RUF, and maps of predicted probability of use were similar between models. In addition, distance to water was the top-ranked variable in both models for mule deer. Although space use by both species was predicted most accurately by the RSF based on cross-validation, differences in predictive power between models were more substantial for elk than mule deer. To maximize accuracy and utility of predictive wildlife-habitat models, managers must be aware of the relative strengths and weaknesses of different modeling techniques. We conclude that although RUFs represent a substantial advance in resource selection theory, techniques available for generating RUFs remain underdeveloped and, as a result, RUFs sometimes predict less accurately than models derived using more conventional techniques.     

A machine‐learning approach for extending classical wildlife resource selection analyses

Resource selection functions (RSFs) are tremendously valuable for ecologists and resource managers because they quantify spatial patterns in resource utilization by wildlife, thereby facilitating identification of critical habitat areas and characterizing specific habitat features that are selected or avoided. RSFs discriminate between known‐use resource units (e.g., telemetry locations) and available (or randomly selected) resource units based on an array of environmental features, and in their standard form are performed using logistic regression. As generalized linear models, standard RSFs have some notable limitations, such as difficulties in accommodating nonlinear (e.g., humped or threshold) relationships and complex interactions. Increasingly, ecologists are using flexible machine‐learning methods (e.g., random forests, neural networks) to overcome these limitations. Herein, we investigate the seasonal resource selection patterns of mule deer (Odocoileus hemionus) by comparing a logistic regression framework with random forest (RF), a popular machine‐learning algorithm. Random forest (RF) models detected nonlinear relationships (e.g., optimal ranges for slope and elevation) and complex interactions which would have been very challenging to discover and characterize using standard model‐based approaches. Compared with standard RSF models, RF models exhibited improved predictive skill, provided novel insights about resource selection patterns of mule deer, and, when projected across a relevant geographic space, manifested notable differences in predicted habitat suitability. We recommend that wildlife researchers harness the strengths of machine‐learning tools like RF in addition to “classical” tools (e.g., mixed‐effects logistic regression) for evaluating resource selection, especially in cases where extensive telemetry data sets are available.     

Mating frequency and inclusive fitness in Drosophila melanogaster

In many species, increased mating frequency reduces maternal survival and reproduction. In order to understand the evolution of mating frequency, we need to determine the consequences of increased mating frequency for offspring. We conducted an experiment in Drosophila melanogaster in which we manipulated the mating frequency of mothers and examined the survival and fecundity of the mothers and their daughters. We found that mothers with the highest mating frequency had accelerated mortality and more rapid reproductive senescence. On average, they had 50% shorter lives and 30% lower lifetime reproductive success (LRS) than did mothers with the lowest mating frequency. However, mothers with the highest mating frequency produced daughters with 28% greater LRS. This finding implies that frequent mating stimulates cross-generational fitness trade-offs such that maternal fitness is reduced while offspring fitness is enhanced. We evaluate these results using a demographic metric of inclusive fitness. We show that the costs and benefits of mating frequency depend on the growth rate of the population. In an inclusive fitness context, there was no evidence that increased mating frequency results in fitness costs for mothers. These results indicate that cross-generational fitness trade-offs have an important role in sexual selection and life-history evolution.     

Lifetime reproductive success of female mountain gorillas

Studies of lifetime reproductive success (LRS) are important for understanding population dynamics and life history strategies, yet relatively little information is available for long-lived species. This study provides a preliminary assessment of LRS among female mountain gorillas in the Virunga volcanoes region. Adult females produced an average of 3.6 ± 2.1 surviving offspring during their lifetime, which indicates a growing population that contrasts with most other great apes. The standardized variance in LRS (variance/mean(2) = 0.34) was lower than many other mammals and birds. When we excluded the most apparent source of environmental variability (poaching), the average LRS increased to 4.3 ± 1.8 and the standardized variance dropped in half. Adult lifespan was a greater source of variance in LRS than fertility or offspring survival. Females with higher LRS had significantly longer adult lifespans and higher dominance ranks. Results for LRS were similar to another standard fitness measurement, the individually estimated finite rate of increase (λ(ind) ), but λ(ind) showed diminishing benefits for greater longevity.     

Population dynamics and resource use of red deer after release from harvesting in New?Zealand

Despite periods of extensive government-funded control, fluctuating commercial exploitation and ongoing recreational hunting, little is known about how red deer (Cervus elaphus scoticus L?nnberg) in New?Zealand respond to the cessation of harvesting in terms of population growth rate and resource use. We describe the population dynamics and resource use of red deer in a montane catchment over 5?years (1962?67) following cessation of intensive government-funded control in 1961. Locations and sex?age classes of deer were observed monthly along a fixed route in the Harper-Avoca catchment, inland Canterbury. A total of 2036 red deer groups were observed. The number of groups observed annually increased during the study but no trends in median (2 or 3) or modal (1 or 2) group sizes were found. Population growth rates (r) of deer were extraordinarily high in the first two years (e.g. 2.33 ? 0.22 for adult females and 1.61 ? 0.23 for adult males), but decreased in subsequent years and were not biologically possible without substantial immigration and/or changes in detectability of deer. Sexual segregation and selection of vegetation types (alpine grassland, montane grassland, and forest) and 10 topographic landforms showed stronger intra-annual than inter-annual patterns. Segregation was greatest in spring and summer, least in the rut, and variable in winter. In all seasons, sexual segregation was greatest at 25- and 50-ha scales, moderate at 100-ha, and absent at the 500- and 1000-ha scales. Selection of vegetation types also varied seasonally, with deer of both sexes preferring montane grasslands in spring and summer and alpine grasslands in the rut. Backslopes were preferred landforms in spring and summer, spurs during spring and the rut, and hollows during the rut. Our results highlight the need to consider spatial scale, immigration, and detectability in the design of red deer culling and harvesting programmes. Studies of home-range size and use, migration patterns, dispersal rates and distances are required to better understand the impacts of red deer on New?Zealand ecosystems and the effects of management on red deer populations.     

Activity patterns and resource use by sheep and red deer grazing across a grass/heather boundary

Diet selection and activities of sheep and red deer grazing alone and together across a grass (primarily Lolium perenne/Agrostis capillaris ) and heather ( Calluna vulgaris ) boundary, between July and October, are described.
Throughout the experiment, both animal species had a much higher proportion of grass than heather in their diet. Time spent grazing on each type of vegetation was similarly greater on grass, and the bite rates of the animals were also more rapid on the grass than on the heather.
Red deer spent more time grazing per 24 hours than did sheep, with sheep grazing less during darkness hours than red deer. Total grazing time of both species did not decrease with decreasing daylength from July to October.
There were no consistent differences in the diurnal patterns of activity of sheep and red deer, with both species showing 2-3 large grazing peaks per day.
There were no consistent effects of mixing sheep and deer on their grazing behaviour and diet composition.     

Spatio-temporal variation of natural and sexual selection in breeding populations of the coreid bug,Colpula lativentris (Heteroptera: Coreidae)

Spatio-temporal variations of lifetime reproductive succes (LRS) of both male and female individuals of a coreid bugColpula lativentris were measured and analyzed using the multiple regression method of Arnold and Wade (1984a, b). The standardized variance of LRS was larger in males than that in females as males often to secure mates for a long period whereas females could easily find mates and oviposit simply dependent on ovarial maturation. LRS was partitioned into 4 consecutive fitness components: (1) reproductive lifespan, (2) copulating efficiency, (3) guarding efficiency (for males) or oviposition efficiency (for females), and (4) number of eggs per clutch. In males copulating efficiency was the largest determining factor of LRS, whereas in females reproductive lifespan was the most important factor. Such tendencies were stable on both a yearly and local basis. Patterns of relative contribution of natural selection (reproductive lifespan and number of eggs per clutch) and sexual selection (copulating efficiency and guarding or oviposition efficiency) to LRS were clearly different between males and females. This sexual difference is, at least to some extent, thought to be brought about by sexual selection among males for mating opportunity, though no physical fight was observed among males. Directional selection on body length was found only in relation to the clutch size of females because large females tended to lay larger clutches. No significant directional selection was found in other fitness components.     

Natural selection on age-specific fertilities in human females: comparison of individual-level fitness measures.

Lifetime reproductive success and timing of reproduction are key components of life-history evolution. To understand the evolution of reproductive schedules, it is important to use a measure of fitness that is sensitive both to reproductive quantity and reproductive timing. There is a contradiction between the theory, which mainly focuses on the rate measures of fitness (r and lambda), and empirical studies, which mainly use lifetime reproductive success (LRS), or some of its correlates, as a fitness measure. We measured phenotypic selection on age-specific fertilities in three pre-modern human populations using individually estimated finite rate of increase, er (lambda). We found that lambda and lifetime reproductive success ranked individuals differently according to their fitness: for example, a female giving birth to four children at a young age may actually have a higher fitness than a female giving birth to six children at a greater age. Increase in fertility at the young age classes (15-19 years) was favoured by selection, but the intensity of selection on fertility was higher in the older age classes (20-30 years), where the variance in fertility was highest. Hence, variation in fertility in the older age classes (20-30) was actually responsible for most of the observed variation in fitness among the individuals. Additionally, more than 90% of variation in fitness (lambda) was attributable to individual differences in LRS, whereas only about 5% of all variation in fitness was due to differences in the reproductive schedule. The rate-sensitive fitness measure did not significantly challenge the importance of total fertility as a component of fitness in humans. However, the rate-sensitive measure clearly allowed more accurate estimation of individual fitness, which may be important for answering some more specific questions.     

BREEDING RESOURCE DISTRIBUTION AFFECTS SELECTION GRADIENTS ON MALE PHENOTYPIC TRAITS: EXPERIMENTAL STUDY ON LIFETIME REPRODUCTIVE SUCCESS IN THE BITTERLING FISH (RHODEUS AMARUS)

The spatial distribution of breeding resources can have pronounced demographic and evolutionary consequences. We used 20 experimental groups of the bitterling ( Rhodeus amarus ), an annual fish with a promiscuous, resource-based mating system, and extended breeding season to investigate how the spatial distribution (clumped or regular) of bitterling oviposition sites (live freshwater mussels) affected offspring production, variation in reproductive success, and directional selection on phenotypic traits over their entire reproductive lifetime. We did not detect any effect of resource distribution on offspring production or variation in reproductive success among individual fish, although variation between replicates was higher with a clumped distribution. This finding is discussed with regard to the incidence of alternative mating behaviors (sneaking) within the limitations imposed by our experimental design. Breeding resource distribution had a significant effect on selection on male phenotypic traits. Stronger directional selection on traits associated with intrasexual competition for fertilizations, gonad mass (an indicator of sperm competition), and the extent of red, carotenoid-based pigment in the iris (an index of dominance status), was detected with a clumped resource distribution. With a regular resource distribution, a stronger positive selection on male body size was detected. We discuss the implications of our results for natural populations.     

Helper effects on pup lifetime fitness in the cooperatively breeding red wolf (Canis rufus)

The evolutionary maintenance of cooperative breeding systems is thought to be a function of relative costs and benefits to breeders, helpers and juveniles. Beneficial effects of helpers on early-life survivorship and performance have been established in several species, but lifetime fitness benefits and/or costs of being helped remain unclear, particularly for long-lived species. We tested for effects of helpers on early- and late-life traits in a population of reintroduced red wolves (Canis rufus), while controlling for ecological variables such as home-range size and population density. We found that the presence of helpers in family groups was positively correlated with pup mass and survival at low population density, but negatively correlated with mass/size at high density, with no relation to survival. Interestingly, mass/size differences persisted into adulthood for both sexes. While the presence of helpers did not advance age at first reproduction for pups of either sex, females appeared to garner long-term fitness benefits from helpers through later age at last reproduction, longer reproductive lifespan and a greater number of lifetime reproductive events, which translated to higher lifetime reproductive success. In contrast, males with helpers exhibited diminished lifetime reproductive performance. Our findings suggest that while helper presence may have beneficial short-term effects in some ecological contexts, it may also incur long-term sex-dependent costs with critical ramifications for lifetime fitness.     

Natural selection on female life-history traits in relation to socio-economic class in pre-industrial human populations

Life-history theory predicts that resource scarcity constrains individual optimal reproductive strategies and shapes the evolution of life-history traits. In species where the inherited structure of social class may lead to consistent resource differences among family lines, between-class variation in resource availability should select for divergence in optimal reproductive strategies. Evaluating this prediction requires information on the phenotypic selection and quantitative genetics of life-history trait variation in relation to individual lifetime access to resources. Here, we show using path analysis how resource availability, measured as the wealth class of the family, affected the opportunity and intensity of phenotypic selection on the key life-history traits of women living in pre-industrial Finland during the 1800s and 1900s. We found the highest opportunity for total selection and the strongest selection on earlier age at first reproduction in women of the poorest wealth class, whereas selection favoured older age at reproductive cessation in mothers of the wealthier classes. We also found clear differences in female life-history traits across wealth classes: the poorest women had the lowest age-specific survival throughout their lives, they started reproduction later, delivered fewer offspring during their lifetime, ceased reproduction younger, had poorer offspring survival to adulthood and, hence, had lower fitness compared to the wealthier women. Our results show that the amount of wealth affected the selection pressure on female life-history in a pre-industrial human population.     

Lifetime reproductive success in common buzzard, Buteo buteo: from individual variation to population demography

We studied annual and lifetime reproductive success (LRS) of both sexes of common buzzard Buteo buteo in eastern Westphalia, Germany. We followed a bottom‐up approach starting from individual breeding attempts, over lifetime reproductive success to derive population demography. Annual breeding performance and survival followed a quadratic relationship with breeding experience; individuals starting their breeding career were less likely to survive and breed successfully than birds of intermediate breeding experience. According to an analysis of selection gradients, both the opportunity and intensity of selection peaked in the early stages of the breeding career. The distribution of both LRS and another fitness measure, λ, was highly skewed, with ca 17% of adult birds producing 50% of fledglings in both sexes. Besides breeding life span and number of breeding attempts, habitat quality and plumage morph were significant predictors of LRS. There were strong differences in LRS and λ between the plumage morphs in both sexes: intermediate pigmented buzzards were much more successful than either dark or light ones. There was no significant difference between buzzard cohorts either in LRS or λ, nor did these fitness measures differ between individuals starting their breeding career at different conditions of food availability. Based on individual life histories, we formed a transition matrix and analysed its properties to study the population as a whole. This analysis showed that the population growth rate was close to unity (0.906, bootstrapped 95% confidence limits: 0.834 and 0.962). Analysis of reproductive values and elasticities further emphasised colour morph differences: the contribution of intermediate individuals to population growth greatly exceeded that of dark or light individuals. Thus most phenomena on all levels from individual breeding attempts over lifetime reproductive success to population demography can be explained by the fitness differences between the colour morphs with the intermediate morph maintaining the current population renewal potential.     

Lifetime reproductive success,selection on lifespan,and multiple sexual ornaments in male European barn swallows

Natural and sexual selection arise when individual fitness varies according to focal traits. Extra‐pair paternities (EPPs) can affect the intensity of selection by influencing variance in fitness among individuals. Studies of selection require that individual fitness is estimated using proxies of lifetime reproductive success (LRS). However, estimating LRS is difficult in large, open populations where EPPs cause reallocation of biological paternity. Here, we used extensive field sampling to estimate LRS in a population of barn swallows (Hirundo rustica) to estimate selection on lifespan and ornamental traits of males. We found selection on lifespan mediated both by within‐ and extra‐pair fertilization success and selection on tail length mediated by within‐ but not extra‐pair fertilization success. In addition, we found selection on tail white spots via extra‐pair fertilization success after controlling for selection on other traits. These results were not confounded by factors that hamper studies of LRS, including nonexhaustive sampling of offspring and biased sampling of males. Hence, natural and sexual selection mediated by LRS operates on lifespan, tail length, and size of the tail white spots in barn swallows.     

完达山林区森林采伐和非采伐区马鹿、狍子对冬季生境因子选择的比较

2003—2005年的冬季,在黑龙江省完达山地区的五泡林场,通过样线调查,利用逐步判别分析的方法,对冬季马鹿和狍子在森林采伐区和非采伐区生境选择的差异进行了比较研究,其结果表明,在该地区虽然两物种在生境选择上发生部分重叠,但在生境利用方式上均存在显著差异在采伐区马鹿主要利用中坡位或上坡位、半阴半阳或阳坡、人为干扰距离大于1000m、海拔相对较高、食物丰富度较高、乔木胸径较小、灌木密度较高的灌丛或杂木林生境;狍子主要利用中坡位或下坡位、阳坡、海拔相对较低、食物丰富度适中、乔木胸径较大、灌木密度较低、人为干扰距离要求不严格的杂木林或柞树林生境。在非采伐区,马鹿主要选择距人为干扰大于1000m、海拔较高、食物丰富度较高、乔木胸径较小的杂木林生境;狍子主要选择中坡位或下坡位、人为干扰距离要求不严格、海拔较低、食物丰富度较适中和乔木胸径较大的杂木林或杨-桦混交林生境。