Deer Habitat Use Research Articles

Novel insights into red deer (Cervus elaphus) habitat use and suitability in human-dominated versus near-natural landscapes

Abstract Red deer populations, once extirpated in large parts of Europe, have recovered over the last century in many regions. In Switzerland, primarily inhabiting mountain forests, they have recently started to expand into human-dominated lowland landscapes. To study their habitat selection, we analyzed day- and nighttime positions of 15 GPS-collared red deer. Our results show that during daytime, all animals generally preferred forests; open land was mainly used at night. In summer and fall, some lowland individuals deviated from this pattern and used agricultural crops as daytime retreats. Human settlements were often avoided in the lowlands. Day-site selection in the lowlands occurred irrespective of the distance to trails and the probability of being accessed by humans, in contrast to the pattern observed in the mountains. At night, red deer preferred forest edges and open land. In the lowlands, human settlements were avoided. Trails and transport infrastructures had a marginal influence at night. Our study confirmed a high behavioral plasticity of red deer. The use of daytime retreats close to trails and in high-standing crops represent major extensions of their behavior. Our habitat models indicate that there is ample suitable habitat in the lowlands, which is of major relevance for their management.

Temporal habitat use of mule deer in the Pueblo of Santa Ana, New Mexico

AbstractMule deer (Odocoileus hemionus) are important economically, culturally, and recreationally to the Pueblo of Santa Ana in central New Mexico, USA. Studies of habitat selection improve our understanding of mule deer ecology in central New Mexico and provide the Tribe with valuable information for management of mule deer. We used global positioning system telemetry‐collar data collected on mule deer around the Pueblo of Santa Ana to create resource selection functions from proximity‐based habitat predictors using a generalized linear mixed model. We created separate resource selection functions for females and males during summer and winter at different times of the day. Season generally had a greater effect on mule deer habitat use than the time of day. Female and male mule deer selected for similar habitats but were sexually segregated in their summer distributions. These findings are consistent with results from other locations where mule deer partitioned habitat similarly between seasons and sexes. Supported models reaffirm accepted patterns of habitat selection for mule deer to the Pueblo of Santa Ana where local results were lacking. Our results can help managers identify locations in and around the Pueblo of Santa Ana where future development such as highway expansion are likely to conflict with mule deer activity and locations where habitat enhancement projects such as adding water sources can have the greatest effect for the deer population.

Chronic wasting disease alters the movement behavior and habitat use of mule deer during clinical stages of infection.

Integrating host movement and pathogen data is a central issue in wildlife disease ecology that will allow for a better understanding of disease transmission. We examined how adult female mule deer (Odocoileus hemionus) responded behaviorally to infection with chronic wasting disease (CWD). We compared movement and habitat use of CWD-infected deer (n = 18) to those that succumbed to starvation (and were CWD-negative by ELISA and IHC; n = 8) and others in which CWD was not detected (n = 111, including animals that survived the duration of the study) using GPS collar data from two distinct populations collared in central Wyoming, USA during 2018-2022. CWD and predation were the leading causes of mortality during our study (32/91 deaths attributed to CWD and 27/91 deaths attributed to predation). Deer infected with CWD moved slower and used lower elevation areas closer to rivers in the months preceding death compared with uninfected deer that did not succumb to starvation. Although CWD-infected deer and those that died of starvation moved at similar speeds during the final months of life, CWD-infected deer used areas closer to streams with less herbaceous biomass than starved deer. These behavioral differences may allow for the development of predictive models of disease status from movement data, which will be useful to supplement field and laboratory diagnostics or when mortalities cannot be quickly retrieved to assess cause-specific mortality. Furthermore, identifying individuals who are sick before predation events could help to assess the extent to which disease mortality is compensatory with predation. Finally, infected animals began to slow down around 4 months prior to death from CWD. Our approach for detecting the timing of infection-induced shifts in movement behavior may be useful in application to other disease systems to better understand the response of wildlife to infectious disease.

Wild ungulate effects on soil fungal diversity in Mediterranean mixed forests

The effect of wild ungulate density on ecosystems varies according to different factors: climate and physiography conditions, forest type, management history, and herbivore identity. In this study, we evaluated the effect of historically high densities of red deer (Cervus elaphus L.) on the soil fungal communities in Mediterranean ecosystems using 30 paired plots, open on the one hand and exclosure plots on the other one. Plots were established at the end of 2020 in a perimeter-fenced hunting estate of 6600 ha in Toledo, Spain. Three months after plots were established, we analysed fungal communities in 60 soil samples using Illumina 250-bp paired-end sequencing. We estimated changes in total fungal richness and in the richness of trophic groups through Linear Mixed Effects models using fenced/unfenced type and deer habitat use as fixed variables and the location of the plots and the main tree host species as random variables. Fungal composition was analysed using non-metric multidimensional scaling and permutational multivariate ANOVA; edaphic characteristics were incorporated to explain differences. Soil fungal communities were not differentially affected by excluding ungulates for three months. Areas with high deer densities had a richer saprotrophic community and where lowland environments were dominated by the main tree hosts Quercus faginea and Quercus ilex. Arbutus unedo was found in mountain areas where there was less herbivore pressure, a greater richness of ectomycorrhizal and lichenized fungi and soils positively associated to nitrogen, phosphorus, potassium and organic matter levels.

Fine-scale habitat characteristics influence the winter habitat use of swamp deer (Rucervus duvaucelii) in Shuklaphanta National Park, Nepal

Endemic to India and Nepal, the swamp deer (Rucervus duvaucelii) is crucial for restoring and sustaining populations of top predators like tigers. Despite recent signs of species recovery in protected areas, the species remains vulnerable to extinction due to increasing anthropogenic activities. Information on factors influencing the occurrence of swamp deer is essential to formulate suitable management plans that can help to maintain their viable population and protect their habitats. We recorded swamp deer presence via direct sightings and indirect signs (pellets and footprints) through line transect surveys. Two ecological drivers i.e., habitat variables and disturbance variables that included eleven predictor variables were measured in the field where swamp deer was present (“used plots”) and absent (“habitat availability plot”). Binary logistic regression was used to analyze the factors associated with the probability of detecting the swamp deer. The model's outcomes showed that an increase in road/path/fireline distance and settlement distance, mixed forest habitat, and depth of leaf litter were positively correlated with the probability of locating swamp deer, while the other significant predictor variables (canopy cover, distance from water source, and presence of predator) had inverse relationships. In summary, our study has provided crucial insights into the factors influencing the presence of swamp deer in their habitat. These findings are useful for local governments and protected area authorities by providing fine-scale field-based scientific information, that can benefit swamp deer and its habitat. We also recommend similar studies in species range locations considering other influencing factors such as season, climate, interaction with other herbivore species, and predators that might affect the habitat use of swamp deer.

Using a Cost-Distance Time-Geographic Approach to Identify Red Deer Habitat Use in Banff National Park, Alberta, Canada

Animal movements are realizations of complex spatiotemporal processes. Central to these processes are the varied environmental contexts in which animals move, which fundamentally impact the movement trajectories of individuals at fine spatial and temporal scales. An emerging perspective in time geography is the direct examination of the influence that varying contexts may have on observed movements. An approach that considers environmental context can yield actionable information for wildlife management, planning, and conservation; for instance, identifying areas of probable occupancy by an animal may improve the efficiency of fieldwork. This research develops the first known practical application of a new cost-distance-based, probabilistic voxel space–time prism (CDBPSTP) in efforts to more realistically characterize the unobserved habitat occupancies of animals occurring between known positions provided by location-aware technologies. The CDBPSTP method is applied to trajectory data collected for a group of red deer (Cervus elaphus) tracked near Banff National Park, Alberta, Canada. As a demonstration of the added value from examining how context influences movement, CDBPSTP habitat occupancy results are compared to the earlier PSTP method in context with empirical and theoretical understandings of red deer habitat preference and space-use behaviors. This comparison reveals that with CDBPSTP, variation present in the mover’s environment is explicitly considered as an influence on the mover’s probable path and occupancies between observations of its location. With the increasing availability of high-resolution geolocational and associated environmental data, this study highlights the potential for CDBPSTP to be leveraged as a broadly applicable tool in animal movement analysis.

Overlap in habitat use and activity patterns between sika deer (Cervus nippon) and Japanese serows (Capricornis crispus) in subalpine habitats: exploitative competition rather than direct interference?

Examining multiple niche relationships, such as habitat use and activity patterns, between sympatric species contributes to an understanding of the mechanisms of coexistence and competition. Sika deer ( Cervus nippon Temminck, 1838) and Japanese serows ( Capricornis crispus (Temminck, 1836)) often show different habitat use, and diversity in vegetation and topography seems to facilitate their coexistence. Conversely, their habitat use may overlap in areas with low-diversity habitats; however, this remains unstudied. Moreover, whether they spatiotemporally avoid each other to reduce direct interference is unclear. We examined the fine-scale patterns of habitat use and activity of deer and serows in the subalpine forests of Mount Fuji, which have low-diversity habitats, using camera trapping data over 3 years. Deer and serow habitat niches largely overlapped, especially in summer (92%–94%) and autumn (89%–91%), suggesting that low-diversity habitats facilitate overlapping habitat use. Both species selected areas frequently used by the other during spring to autumn and their temporal niches largely overlapped, especially in summer (88%) and autumn (83%), suggesting that they do not spatiotemporally avoid each other (i.e., there is no direct interference). Rapid range expansion of deer into these subalpine habitats may exclude native serows through resource exploitative competition rather than interference competition.

Impacts of the novel coronavirus SARS-CoV-2 on wildlife behaviour via human activities.

Severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2) caused the pandemic of the coronavirus disease 2019 (COVID-19), resulting in a global lockdown in 2020. This stagnation in human activities ('anthropause') has been reported to affect the behaviour of wildlife in various ways. The sika deer Cervus nippon in Nara Park, central Japan, has had a unique relationship with humans, especially tourists, in which the deer bow to receive food and sometimes attack if they do not receive it. We investigated how a decrease and subsequent increase in the number of tourists visiting Nara Park affects the number of deer observed in the park and their behaviour (bows and attacks against humans). Compared with the pre-pandemic years, the number of deer in the study site decreased from an average of 167 deer in 2019 to 65 (39%) in 2020 during the pandemic period. Likewise, the number of deer bows decreased from 10.2 per deer in 2016-2017 to 6.4 (62%) in 2020-2021, whereas the proportion of deer showing aggressive behaviour did not change significantly. Moreover, the monthly numbers of deer and their bows both corresponded with the fluctuation in the number of tourists during the pandemic period of 2020 and 2021, whereas the number of attacks did not. Thus, the anthropause caused by the coronavirus altered the habitat use and behaviour of deer that have continuous interactions with humans.

Forage, forest structure or landscape: What drives roe deer habitat use in a fragmented multiple-use forest ecosystem?

Browsing damages to young trees can have lasting impacts on forest structure. Roe deer (Capreolus capreolus), the most common and widespread large herbivore in central Europe, create a vast majority of this damage. To lessen the impact, it is important to understand the relationship between roe deer and the landscape matrix, and which factors such as food availability and cover will drive the use of habitat by roe deer. In this study, we explored how small scale-food availability (5 × 5 m2), forest structure (100 × 100 m2) and landscape heterogeneity (500 m radius) influenced the use of habitat by roe deer in an intensively managed temperate mountainous mixed forest with implemented retention forestry practices. Using camera-trap detections of roe deer from 130 study plots in the southern Black Forest, monitored for 2.5 years, we found that local forest structure had the strongest influence on roe deer habitat use. Contrary to our expectations, landscape features, such as edge density between forest and non-forest, did not affect roe deer detections, probably because overall anthropogenic pressure is high and homogenous throughout our study system. Small-scale food availability also had little influence, which is likely due to widespread availability throughout the study area. Roe deer were also detected less where there were higher amounts of lying deadwood in autumn, indicating that retention forestry methods may have a negative impact on roe deer habitat use. Since forest structure was the strongest driver of roe deer habitat use, this study supports earlier claims that forests may be managed by affecting roe deer habitat use, thereby browsing damage intensity, through manipulation of food availability and cover.

Temporal effects of relative hunter activity on adult male white-tailed deer habitat use

Context Predation risk, whether from large carnivores or humans, is a primary driver of movement and habitat selection for ungulates. Throughout much of the geographical range of white-tailed deer, predation risk during autumn and winter is driven by hunters; however, information about how deer respond to hunting pressure during a long hunting season is lacking. Aims We aimed to determine how differences in daily temporal exposure to hunting risk affected habitat selection of adult male white-tailed deer. Methods We deployed GPS collars on 42 adult male white-tailed deer during the 2017–2018 and 2018–2019 firearm hunting seasons in Mississippi, USA. GPS collars provided fine-scale spatial and temporal information regarding habitat use of our study area. Key results We showed a mismatch between what hunters and deer selected; landscape characteristics selected most by hunters were least selected by deer during the day, which we believe has never been documented. Fine-scale temporal analysis showed that deer selected for some areas (food plots) as much as five times more often during the night, when no hunting risk was present, than during the day, when risk was present. Conclusions Habitat selection by deer varies depending on the amount of risk that is present. Even in a high-risk heavily hunted area, adult male white-tailed deer can recognise external cues on the landscape, and shift habitat selection both spatially and temporally to favour areas that present less risk. Implications This study, to our knowledge, provides the first evidence that white-tailed deer actively selected for habitat characteristics that hunters were not utilising. To minimise the effectiveness of risk-avoidance behaviours used by deer, we suggest hunters moderate temporal hunting effort across time to decrease the risk to which deer are exposed, thereby minimising the response by deer.

Responses to changes in precipitation of plant species vary by functional groups on understories of temperate forests in central Japan

Climate change has one of the most dramatic impacts on plant species diversity at the global scale. However, compared to global warming, there are few reports of the effects of changes in long-term precipitation patterns on plant community diversity in areas other than moisture-limited areas, for example temperate areas. In temperate forests, understory plant communities constitute a large proportion of plant diversity. It is crucial to investigate effects of change in long-term precipitation patterns and the other important environmental changes on plant species composition of understories in temperate forests for making the conservation strategy. The important changes that may have particularly affected plant composition in recent years include the followings; temperature rise, increase of large herbivores, change of habitat area, and changes in the cover of canopy and understory-abundant species according to the land use change. Vulnerability to the above environmental changes may be higher for species groups that are less adapted to their current habitat and have shrinking population sizes (e.g., grassland species on understories in abandoned secondary forests). The aim of this study was to estimate the effects of precipitation and the above-mentioned factors on changes in cover of individual species on understories in temperate forests from the 1980s to the 2000s. In the analysis, we took into account the differences in the functional groups of the species (grassland herbaceous species, forest herbaceous species, and woody species). Grassland herbaceous species responded more sensitively to precipitation, temperature, cover of dwarf bamboo, and deer habitat use than other functional groups. In the 1980s, the cover of grassland herbaceous species tended to be higher in places with lower precipitation, higher temperatures, and lower cover of dwarf bamboo and canopy, in association with climatic conditions and past land-use history. Over the last 20 years, increased precipitation and deer habitat use had particularly strong effects on the decline of grassland herbaceous species. This study found that precipitation variability is becoming a driving factor for biodiversity change through decline of vulnerable species groups in temperate areas. We should continue to monitor the precipitation effects on the ecosystems.

Impacts of human disturbance on flight response and habitat use of red deer

AbstractMany wild animals perceive humans as predators, and human disturbance, especially in the form of hunting, triggers antipredatory behavior among prey. Yet, knowledge of how game species react to different types of human disturbance and adapt to repeated disturbances is limited. We investigated how disturbance in the form of a solitary human approacher (stalker) impacted behavior (flight response and short‐term habitat use) of 28 GPS‐collared red deer (Cervus elaphus) in two populations with contrasting population densities in Norway. We studied how the behavioral response differed: (1) with season (pre‐hunting vs. hunting); (2) by consecutive approaches within a day; (3) among replicated experiments within the same season; and (4) between two regions with contrasting densities of red deer. The average flight initiation distance (FID) increased by 15% during the hunting season, and consecutive approaches within the same day caused the red deer to move 49% longer distances. Flight initiation distance was longer in the high‐density population, while escape distance was longer in the low‐density population. Red deer moved out of their weekly home range after 52% approaches, and after the onset of hunting season, time spent outside the home range increased by 89%. Red deer preferred denser resting sites after the disturbance and animal sites had shorter sighting distance and higher canopy cover than control plots. Tree density and canopy cover at animal sites increased at the onset of hunting season, from first to second approach within day, and after replicated experiments within season. Our results suggest that red deer preferred dense resting sites, especially in the hunting season. However, these animal sites had the same amount of the favorable forage plant bilberry (Vaccinium myrtillus), indicating no clear food–cover trade‐off in selection of habitat. Our study showed that onset of hunting initiates stronger fear responses in red deer, which may in turn affect red deer distribution and harvesting efficiency.

Occupancy modeling of habitat use by white‐tailed deer after more than a decade of exclusion in the boreal forest

The exclusion of herbivores in forest areas is a strategy used to reduce the impact of selective browsing and increase the regeneration of desired plant species. On Anticosti Island (Québec, Canada), selective browsing by white‐tailed deer prevents the regeneration of balsam fir – white birch forests leading to their conversion into white spruce forests. Large deer exclosures were established for ca 10–12 years in clear‐cuts with patches of residual forest from 2001 to 2006 to assist in the natural regeneration of fir stands and to provide shelter and food resources for deer. Our objective was to assess how deer use exclosures after the removal of fences according to their spatial configuration and habitat composition. We randomly distributed automatic cameras for periods of 14 days during summer in six exclosures ranging from 3.1 to 11.2 km2 (n = 25 cameras per exclosure) from which deer were reduced for 10–12 years. We compared candidate occupancy models that included spatial configuration and food resource variables while simultaneously controlling for variables affecting detection probability. We obtained weak evidence that deer habitat use increased by 19% when forage resources, represented by the cover of Cornus canadensis, increased from 0 to 100%. None of the other variables (distance between the border of exclosures and cameras and distance between forest patches and cameras) was retained, suggesting that the use of regenerating forests by deer in summer after a period of exclusion is related to forage availability and therefore, any forest management that improves food production during summer should help maintain or increase habitat use by deer.

Reduction in local white-tailed deer abundance allows the positive response of northern white cedar regeneration to gap dynamics

Gap dynamics play a crucial role in forest regeneration by creating favourable regeneration and survival niches for some plant species. Nonetheless, potentially overriding factors, such as ungulate browsing, could limit or eliminate this gap dynamic-related regeneration. The deleterious effects of browsing may be exacerbated for slow-growing species such as northern white cedar ( Thuja occidentalis L.), a tree highly selected by white-tailed deer ( Odocoileus virginianus Zimmerman). We therefore aimed to understand how deer browsing and gap dynamics interact to affect cedar regeneration and hypothesized that cedar regeneration benefits from natural gaps but that deer browsing could override this effect. We inventoried natural canopy gaps along a spatiotemporal gradient of deer habitat use. We evaluated cedar regeneration abundance, tree height, and various gap, stand, and competition metrics. We found that deer browsing pressure greatly limited cedar regeneration; however, when deer populations decreased, cedar regeneration abundance increased within a decade, even after prolonged browsing pressure, and increased further over time. Our study illustrates that cedar regeneration can be favoured by gap creation and deer population control.

Ecological Factors Determining Barking Deer Distribution and Habitat Use in the Mid-Hills of Nepal

Barking deer is found in dense tropical and subtropical forests of Asia. It is listed as “least concerned” by the International Union of Conservation of Nature and as “vulnerable” in Nepal, where it is also protected. Due to the habitat loss and fragmentation by human activities, barking deer abundance is decreasing, which may even ultimately lead to its extinction. This in turn might negatively affect local ecosystem such as the abundance of the endangered common leopard, for which barking deer is the main prey species in the mid-hills of Nepal. We therefore need to know factors affecting barking deer abundance and its habitat preferences. To determine these factors, we recorded barking deer either by direct sighting or by any evidence of its indirect presence observed through transect surveys in January and February, 2019. To analyze habitat preference, the presence of barking deer was set to 1 if the barking deer or any sign of its presence were observed “used plots,” or to 0 if the barking deer or any sign of its presence were not observed (“habitat availability plot”). We measured main four ecological drivers such as forest management regime, microclimate, disturbance and food resources, which include 11 habitat characteristics in spots where barking deer was present, and in randomly selected spots. We found that elevation, slope, distance from settlement, presence of tree species, depth of leaf litter and percentage cover of leaf litter were most significantly affecting its presence. These results can serve as guidelines for local authorities to prevent decline in abundance of barking deer.

Correction to: Population density, habitat use and activity patterns of endangered hog deer in Cambodia

Correction to: Population density, habitat use and activity patterns of endangered hog deer in Cambodia

Population density, habitat use and activity patterns of endangered hog deer in Cambodia

Population density, habitat use and activity patterns of endangered hog deer in Cambodia

Grasslands half‐full: investigating drivers of spatial heterogeneity in ungulate occurrence in Indian Terai

AbstractGrassland ecosystems have declined in their extent globally, driving declines in wild ungulate populations. Even within the remnant grasslands, ungulate distribution is highly heterogeneous for reasons that are not well understood. This in turn undermines both local and landscape level conservation efforts for these often‐neglected ecosystems and the herbivores they support. We investigated grassland‐ungulate relationships in the Terai region of North India, where wild ungulates are patchily distributed across alluvial grasslands, a besieged ecosystem. Specifically, we posited that spatial variation in grassland habitat use by swamp deer (Rucervus duvaucelii duvaucelii), hog deer (Axis porcinus) and spotted deer (Axis axis) would be explained by community composition, palatability of key grasses, and fire and flooding regimes in grasslands. Hierarchical clustering and multivariate ordination revealed that three dominant tall‐grass communities and the narrowly distributed and more palatable short grass Cynodon‐Oryza community were only marginally influenced by fire and flood histories. Ungulate habitat use, assessed using occupancy models for spatially correlated data, indicated that hog deer were ubiquitously distributed across the grasslands in the park (habitat use probability, ᴪ = 0.92, se = 0.05) utilizing a diverse range of grassland types, while spotted deer (ᴪ = 0.80, se = 0.17) and swamp deer (ᴪ = 0.45, se = 0.10) occurrence within grasslands was relatively lower. Grass height and grassland extent positively influenced habitat use of swamp deer and spotted deer, respectively. However, grassland community composition, fire history, flood history and palatability were relatively uninformative predictors of fine‐scale habitat use. Our study provides a robust baseline for future monitoring of grasslands and ungulates as well as insights for the design and implementation of grassland management interventions.

There is only one winner: The negative impact of red deer density on roe deer numbers and distribution in the Słowiński National Park and its vicinity.

Red and roe deer are the most numerous cervids in Europe, and they occur in sympatry in most regions. Roe deer were considered to be an inferior competitor in studies in which they co‐occurred with fallow deer or muntjac. Despite the remarkable overlap of their ranges, there are few studies on the competition between the red and roe deer. Since interspecific interactions among ungulates are often related to their mutual densities, the current study focused on the effects of high red deer density on the roe deer numbers and spatial distribution in the unhunted Słowiński National Park (SNP) in northern Poland and forest districts open to hunting bordering the park. Using fecal pellet group counts, it was found that in the forest districts (where red deer densities were 2–3 times lower than in the SNP), roe deer densities were significantly higher than in the park. The red‐to‐roe deer density ratio was 10.8 and 2.7, in the SNP and the surrounding forest districts, respectively. Moreover, in the SNP, the roe deer distribution was negatively affected by the red deer habitat use, while in the hunting areas, such an effect was not recorded. The negative influence of the red deer on the roe deer population in the park was most probably due to the red deer impact on food availability. The biomass of the plant groups forming the staple food of the roe deer (Rubus spp., forbs, dwarf shrubs) was significantly higher in the fenced plots than in the unfenced ones. Lack of hunting in the protected areas may benefit only some species in ungulate assemblages which, in turn, may contradict one of their objectives—to maintain viable and ecologically functional populations.

Diets and habitat selection of takhi and red deer in Hustai National Park, Mongolia

Hustai National Park in Mongolia became a refuge for takhi Equus ferus przewalskii, a wild horse native to central Asia, in 1992. Wild takhi became extinct in the wild in the 1960s, although there was a small captive population in European zoos that was used to repopulate Hustai National Park. This park is also inhabited by red deer Cervus elaphus, which may compete with takhi for food. We analyzed the fecal composition and habitat use of takhi and red deer. Takhi mainly foraged on grasses in the steppe, while red deer foraged on grasses and dicots in the forest. The percentage similarity (PS) of foods was 65% in summer and 58% in winter. There does not appear to be competition for food or habitat between these species because both diets and the habitat selection were different. We discuss some management implications and emphasize the importance of integrated management of the refuge.