Spatial and temporal habitat use and selection by red deer: The use of direct and indirect methods

Given the importance of red deer Cervus elaphus for hunting and conservation purposes, understanding the interactions between this species and its habitats in the Mediterranean region is a crucial step for the sustainable management of this species. Aiming to compare pellet group counts and direct observations methods to study the habitat use and selection by red deer, the results obtained by both methods were compared, and their advantages and disadvantages were discussed. To understand the temporal patterns of habitat use and selection, the survey was conducted at three different seasons, birth period, rut season and winter. The habitat use and selection were studied in relation to land cover, watercourse, roads, ecotone zones and other topographic features (altitude, slope and aspect), using generalized linear models and selection ratios. The similarity of the results provided by pellet group counts and direct observations indicate that both methods may constitute useful tools to study the habitat use and selection by red deer. Globally, red deer seemed to select habitats that provide simultaneously food and some cover, as shown by its preference for shrublands, independently of the sampling season. The positive selection of ecotone zones embodies the need for open spaces. Males and females showed a similar use of shrubland, but selected patches with different characteristics therein. The spatial and temporal patterns exhibited by our results suggest that red deer balance their habitat requirements in respect to each phase of their reproductive cycle. Pellet group counts and direct observations seem to be useful methods to analyze habitat use and selection, and may provide helpful knowledge to the management and conservation of red deer.     

Solar Radiation Determines Site Occupancy of Coexisting Tropical and Temperate Deer Species Introduced to New Zealand Forests

Assemblages of introduced taxa provide an opportunity to understand how abiotic and biotic factors shape habitat use by coexisting species. We tested hypotheses about habitat selection by two deer species recently introduced to New Zealand’s temperate rainforests. We hypothesised that, due to different thermoregulatory abilities, rusa deer (Cervus timorensis; a tropical species) would prefer warmer locations in winter than red deer (Cervus elaphus scoticus; a temperate species). Since adult male rusa deer are aggressive in winter (the rut), we also hypothesised that rusa deer and red deer would not use the same winter locations. Finally, we hypothesised that in summer both species would prefer locations with fertile soils that supported more plant species preferred as food. We used a 250 × 250 m grid of 25 remote cameras to collect images in a 100-ha montane study area over two winters and summers. Plant composition, solar radiation, and soil fertility were also determined for each camera location. Multiseason occupancy models revealed that direct solar radiation was the best predictor of occupancy and detection probabilities for rusa deer in winter. Multistate, multiseason occupancy models provided strong evidence that the detection probability of adult male rusa deer was greater in winter and when other rusa deer were present at a location. Red deer mostly vacated the study area in winter. For the one season that had sufficient camera images of both species (summer 2011) to allow two-species occupancy models to be fitted, the detection probability of rusa deer also increased with solar radiation. Detection probability also varied with plant composition for both deer species. We conclude that habitat use by coexisting tropical and temperate deer species in New Zealand likely depends on the interplay between the thermoregulatory and behavioural traits of the deer and the abiotic and biotic features of the habitat.     

Habitat manipulation preferred by Eld’s Deer in Hainan Island,China

In order to improve the conservation status of the Eld’s Deer (Cervus eldii), a critically endangered species in China, a population in a nature reserve on Hainan Island was studied. Abundances and behaviours in two different habitat types, natural and manipulated, were recorded. We found that different habitats were preferred according to season. Both sexes were more abundant in the manipulated habitat from January to May, with males being more abundant in the natural habitat from September to October. However, females, were significantly more abundant from April and May in the manipulated habitat. The deer preferred the more open manipulated habitat for feeding, consistent with greater food availability. This study provides useful evidence for managers to create seasonally suitable habitat for the conservation of this species on Hainan Island. Similar methods could also be applied to other deer species in China, most of which are facing critical survival challenges.     

Habitat associations of four ungulates in mountain forests of southwest China, based on camera trapping and dung counts data

The present study aimed to assess abundance indices and habitat associations of four sympatric ungulate species (alpine musk deer Moschus chrysogaster, tufted deer Elaphodus cephalophus, Chinese serow Capricornis milneedwardsii, and Chinese goral Naemorhedus griseus) in Baima Xueshan Nature Reserve of southwest China, using camera trapping and dung counts data. Camera traps were set along six dung transects in different habitats and explored habitat use of the sympatric ungulates using trapping rates. The results obtained revealed that Chinese serow showed a negative association with open canopy cover and low canopy cover. Alpine musk deer were associated with oak shrubs, oak forests and open canopy cover, while tufted deer avoided oak shrubs. Goral showed no significant associations with habitat variables. Alpine musk deer and tufted deer had considerable habitat overlap with Chinese serow. By finding a high correlation between indices, the study indicates that camera trapping may represent a valid index of relative abundance, matching results from other studies.     

Breeding success of an endangered island endemic kestrel increases with extent of invasion by an alien plant species

The Mauritius Kestrel Falco punctatus, once the rarest kestrel worldwide, became an icon of bird conservation after it recovered from four to six individuals in 1974 to some 800 by 2005 following intense conservation management. Its population however then halved within about a decade prompting a re-evaluation of the IUCN status and up listing of the species in 2014 and an increased conservation attention. Drivers of this new decline are unclear and the influence of habitat structure and diet on breeding success may be important contributors but have received relatively little attention, particularly in the way they may interact to influence production of new fledglings. We address this knowledge gap by studying whether breeding success is influenced by habitat structure (in terms of cover of the invasive Ravenala in native habitats, an alien plant causing strong structural shift in the forests that it invades, and extent of cleared area), diet composition and food pass frequency (as a proxy for food intake) and food quality at 28 nests of a re-introduced kestrel population in south east Mauritius during the 2015–2016 breeding season. The kestrel’s diet comprised native and alien birds, reptiles, insects, and small alien mammals, with a disproportionately high proportion of Phelsuma gecko. A higher frequency of food provisioning and percentage cover of Ravenala both contributed to higher breeding success. Ravenala may increase gecko density or increase gecko detectability and predation by the kestrel, or both, while changed land use (pasture and sugar cane fields) may increase prey diversity in the form of non-forest prey known to be eaten by Kestrels (e.g. alien agamids, small mammals and birds). These prey related influences on breeding suggest that the Bambou mountain range provides a human-generated novel ecosystem altering food availability and increasing the kestrel’s breeding success. However, Ravenala is an invasive alien species harmful to the wider forest biodiversity. Progressive weeding of Ravenala and concurrent re-introduction and augmentation of native palms and Pandanus species which geckos can use at comparable densities to Ravenala, is recommended. This would likely improve the kestrel’s hunting habitat quality and maintain high gecko density or detectability and the vegetation structure required for hunting manoeuvrability and prey availability without the negative consequences of invasive Ravenala.     

舟山群岛春秋季黄麂的栖息地特征及其利用的初步研究

栖息地选择是野生动物与其环境之间的基本关系,野生动物的栖息地特征研究对野生动物的保护和利用具有重要意义(骆颖等,2009).动物的食物资源具有季节性变化(蒋志刚,2004),在舟山群岛,春季有蹄类动物的可利用资源相对较为缺乏而到秋季相对丰富,有蹄类动物的栖息地特征由于不同季节间的环境可利用资源不同而产生差异(Sehmitz,1991).研究不同季节动物的柄息地特征及利用机制,可以深入了解动物是如何适应环境的(蒋志刚,2004).     

Do exotic pine plantations favour the spread of invasive herbivorous mammals in Patagonia?

Changes in land use patterns and vegetation can trigger ecological change in occupancy and community composition. Among the potential ecological consequences of land use change is altered susceptibility to occupancy by invasive species. We investigated the responses of three introduced mammals (red deer, Cervus elaphus; wild boar, Sus scrofa; and European hare, Lepus europaeus) to replacement of native vegetation by exotic pine plantations in the Patagonian forest‐steppe ecotone using camera‐trap surveys (8633 trap‐days). We used logistic regression models to relate species presence with habitat variables at stand and landscape scales. Red deer and wild boar used pine plantations significantly more frequently than native vegetation. In contrast, occurrence of European hares did not differ between pine plantations and native vegetation, although hares were recorded more frequently in firebreaks than in plantations or native vegetation. Presence of red deer and wild boar was positively associated with cover of pine plantations at the landscape scale, and negatively associated with mid‐storey cover and diversity at the stand scale. European hares preferred sites with low arboreal and mid‐storey cover. Our results suggest that pine plantations promote increased abundances of invasive species whose original distributions are associated with woodlands (red deer and wild boar), and could act as source or pathways for invasive species to new areas.     

Red deer <Emphasis Type="Italic">Cervus elaphus</Emphasis> resting place characteristics obtained from differential GPS data in a forest habitat

We recorded 30 24-h monitoring periods with 10-min sampling intervals on seven (three female; four male) Global-Positioning-System-collared adult free-ranging red deer (Cervus elaphus), from June 1999 to December 2000, in the Parc National des Cévennes, France. We observed the duration of resting bouts (n = 385) and then microhabitat variables (aspect, slope, presence of edge and litter, visibility, abundance of vegetation consumed or not) at 178 resting places. Resting bouts were shorter during the night than during the day from June to October but did not vary between sexes. Resting place visibility was lower during the day, especially in August. Daytime resting places generally offered more litter. Females used steeper slopes than males. We found higher variability in visibility and slope during the night. Aspect used did not vary from month to month or between day and night. Observed differences between day and night resting place characteristics suggest that red deer were probably facing a tradeoff between feeding and cover. Use of cover prevailed during the daytime whereas night resting place characteristics were more variable, indicating less constrained behaviour. Thus, cover (as a protection from disturbance), as well as food, is an important factor in red deer habitat use (at least during the day in disturbed areas) and should not be neglected in forest carrying capacity management.     

Landscape structure affects food quality of sika deer (<Emphasis Type="Italic">Cervus nippon</Emphasis>) evidenced by fecal nitrogen levels

Evaluating the quality of wildlife habitat is essential for understanding and predicting population dynamics in heterogeneous environments. We used fecal nitrogen levels as an indicator of habitat quality of sika deer (Cervus nippon) and explored important landscape elements influencing nitrogen levels, taking deer density into account. We established 92 plots differing in deer density and landscape structure on the Boso Peninsula, central Japan, and collected fecal samples along a 1-km transect at each plot. The regression models involving two independent variables, i.e., deer density and the length of forest edge within an area of 100 or 200 m from the transect, were selected based on the Akaike’s Information Criterion (AIC). Levels of fecal nitrogen were positively correlated with the length of the forest edge and negatively correlated with population density of deer. The area of 100 or 200 m from the transect most likely reflected the behavioral scale of the deer. Coverage of palatable understory vegetation increased with proximity to forest edge and decreased with deer density. Variability in the level of fecal nitrogen could thus be explained by food availability in the landscape. These results suggest that landscape alterations increase the carrying capacity of sika deer and thereby increase impacts upon the ecosystem.     

The Origin of Winter Visitors to the British Isles 7. Fieldfare (Turdus pilaris)

Capsule Large‐scale intensification of agricultural management during the past 50 years has resulted in a reduction of invertebrate abundance and higher and denser ground vegetation. Food availability for insectivorous birds foraging on the ground has been negatively affected, but the interactions between birds and their food availability are complex and often species‐specific. Populations of Wrynecks Jynx torquilla are declining all over Europe, possibly because of reduced accessibility to their main prey, ground‐dwelling ants, due to higher and denser ground vegetation. However, it is not clear which ground vegetation structures are tolerated by foraging Wrynecks and which habitats are preferred.

Aims To identify the optimal ground vegetation structure and the main habitat types in which Wrynecks search for food.

Method We radiotracked seven Wrynecks in high‐intensity farmland in Switzerland to study foraging habitat use during the reproduction season. Several habitat variables were mapped at each foraging location and compared with locations selected randomly within individual home ranges.

Results Wrynecks preferentially foraged at places with ≥50% bare ground. Vegetation height was not important. Older fruit tree plantations and fallow land were the preferred foraging habitats.

Conclusion Conservation measures should concentrate on preserving semi‐open agricultural landscape matrices with loose ground vegetation cover to provide suitable foraging conditions. This can be achieved even in intensively managed farmland as illustrated in this study.     

中国大陆鸟类栖息地选择研究十年

栖息地选择研究一直是鸟类生态学研究的重要内容之一。通过对2001年1月至2010年12月10年期间中国大陆鸟类学家在国内外期刊上发表的鸟类栖息地选择研究的论文进行分析,对我国鸟类栖息地选择研究提出展望。10年间,我国鸟类学家共发表有关鸟类栖息地选择或利用的论文170篇,共涉及到鸟类10目31科73种。在这10年里,中国大陆有关鸟类栖息地选择或利用的文章持续增长。然而,我国鸟类栖息地选择的研究也存在着如下问题:(1)存在栖息地选择和栖息地利用误用的现象,这种现象在10年内并无明显改善;(2)在选择研究方法时,很少考虑个体的可获得性、种群密度及抽样尺度对栖息地选择的影响;(3)大多数的栖息地选择的论文没有对鸟类的栖息地选择行为进行研究,也缺乏对其选择的适合度背景去进行研究。根据这些问题,对我国未来的鸟类栖息地选择研究提出了建议。     

带岭地区马鹿冬季对生境的选择性

本文研究马鹿冬季对生境的选择性,选择了10类与马鹿生存有关的生态因子(植被型、食物、雪深、隐蔽条件、坡向、坡位、坡度、水源、人为干扰、海拔)。根据这些生态因子的调查数据按一定要求分级,并转变为数量化方法来研究。经数量化理论Ⅰ的公式运算得出预测方程。筛选出影响马鹿选择生境的主要生态因子是植被型、坡向、雪深和人为干扰。用数量化理论Ⅱ分析,认为人工杨树幼林、天然杨桦幼林、阳坡、雪深25厘米、离人为干扰1000米以外是马鹿适宜生境。另外还调查了马鹿的卧迹,从直观上分析马鹿冬季选择卧地的生态条件。     

The effect of environmental gradients on the bed site selection of roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>)

A wildlife species’ selection of bedding sites is often characterised by strong trade-offs, as habitat quality, predator avoidance and foraging needs should be achieved simultaneously. Human activities often represent major threats in addition. In areas of intensive agriculture, e.g. mowing is one of the main causes of mortality of roe deer (Capreolus capreolus) fawns due their hiding strategy. For a species’ offspring, the selection of bedding sites is particularly crucial and thus, identifying how and when animals use such habitats is important for management. We used a long-term dataset of marked roe deer fawns in Switzerland (1971–2015) to reveal the characteristics of optimal bedding sites within the first two weeks of a fawn’s life in three contrasting landscapes and the potential trade-offs that may occur. We hypothesised that roe deer adjust the selection of bedding sites to current environmental conditions and available habitat to achieve sufficient levels of predator avoidance and thermoregulation necessary for the fawn’s survival, as well as the availability of sufficient food resources for the mother doe. We found that, in general, grassland habitats with medium vegetation height (20–50 cm) and habitats in close proximity to the edge of the forest were favoured to achieve those basic requirements. However, the use of bed site habitats differed between the three contrasting landscapes in dependence of elevation and hence vegetation phenology. Our results provide essential information to reduce mortality rates caused by mowing and improve the reproductive success of this species.     

Resource partitioning by ungulates on the Isle of Rhum

Summary This paper describes the seasonal changes in vegetation community use by red deer, cattle, goats and ponies on the Isle of Rhum, Scotland. During the winter, when food resources were of low abundance and digestibility, the ungulates showed extensive resource partitioning. During the summer, when resource availability and digestibility was high, the grazing species, red deer, cattle and ponies congregated on the vegetation communities which contained high biomasses of a high quality resource, mesotrophic graminoids and forbs. Goats, with a digestive system adapted to dealing with browse, foraged primarily on the communities dominated by dwarf shrubs. The patterns of resource use in this group of ungulates are discussed in relation to competition; species had relatively exclusive esource use during periods of low food availability during tye winter and had a high degree of resource use overlap when food was abundant during the summer. This suggests that there was little competition for food during the summer and that exploitative competition for the high quality foods led to resource partitioning during the winter. Senarios are described which predict the pattern of resource use between two species (one competitively superior to the other on the preferred resource) utilizing mutually or exclusively preferred resources. A model developed by Illius and Gordon (1987), based on the allometry of metabolic requirements and bite size, is used to provide a mechanistic explanation for the observation that the red deer were able to exploit the high quality plant communities during the winter, whereas the cattle moved off to feed on poorer quality communities at this time.     

上海滨江森林公园重引入獐野放后的活动范围与栖息地选择

本文对重引入后野放于上海滨江森林公园的獐进行无线电遥测跟踪研究,确定其在公园内286个斑块内的有效活动位点,分析其活动范围,并通过Vanderloeg选择系数和Scavia选择指数分析其栖息地选择。结果显示,以最小凸多边形法得到的重引入獐的活动范围面积从 24.94 hm² 至 83.24 hm² 不等,均值为 58.74 hm²;以固定核空间法得到的面积从 11.23 hm² 至 41.52 hm² 不等,均值为 26.93 hm²。野放后的獐偏好选择面积为 1-2 hm² 的较大斑块和距离水源较远的栖息地。在植被因子的选择上,野放后的獐倾向选择野草地为主、乔木和灌木密度适中、乔木高度10-15m、草本高度 30cm 以上的栖息地,避免选择乔木高度低于 5m、草本高度过低（<5cm）和无草本植物分布的栖息地。建议在今后重引入项目实施中,在郊野公园或者其它城市绿林地内为獐营造植被盖度和密度适中、隐蔽性良好、斑块面积适中的栖息地,并补种高草本,促进物种重引入成功。     

海南坡鹿(Cervus eldi hainanus)的食性研究

1986年9月至1987年7月作者在海南省大田保护区面积约95公顷的封闭性围栏内,随机设立灌木及草本植物样方各39个。检查了296个坡鹿的采食场。结果表明坡鹿采食灌木31种,草本43种。x~2检验证明,坡鹿喜欢在中生性热带草食生境中采食,避开砂生灌丛林生境。坡鹿对植物种类及取食部位具有明显的选择,这种选择有季节变化。坡鹿的采食频率与喜食不完全一致。采食频率高的植物并不一定是坡鹿最喜食的,但这些植物对坡鹿的生存具有重要意义。     

Forest edge creates small-scale variation in reproductive rate of sika deer

Habitat edges are considered to have an important role in determining the abundance of deer in forest landscapes, but to our knowledge there are few lines of evidence indicating that forest edge enhances the vital rate of deer. We examined pregnancy of female sika deer in Boso peninsula, central Japan, and explored how forest edges, food availability in forests, and local population density influence the pregnancy rate of sika deer. Local deer density was estimated by the number of fecal pellets, and food availability in forests was estimated by combining GIS data of vegetation distribution and the relationship between vegetation biomass and local deer density. Forest edge length was also determined by GIS data. Model selection was performed with multiple logistic regression analyses using the AIC to find the best model for accounting for the observed variation in pregnancy rates of the deer. Multiple logistic regression analysis showed that the length of forest edge had a positive effect on the pregnancy rate of females, whereas food availability in forests and local deer density had little effect. This forest edge effect was detected in a 100–200-m radius from deer captured locations, indicating that deer pregnancy is primarily determined by habitat quality within a 10-ha area. This result was confirmed by tracking females with GPS telemetry, which found that the core areas of the home range were less than 12 ha. The positive effect of edges and the lack of density dependence could be a result of high plant productivity in open environments that produces forages not depleted by high deer densities. Our results support the view that land management is the cause of the current problem of deer overabundance. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

A Long-Term Assessment of the Variability in Winter Use of Dense Conifer Cover by Female White-Tailed Deer

Backgound

Long-term studies allow capture of a wide breadth of environmental variability and a broader context within which to maximize our understanding of relationships to specific aspects of wildlife behavior. The goal of our study was to improve our understanding of the biological value of dense conifer cover to deer on winter range relative to snow depth and ambient temperature.

Methodology/Principal Findings

We examined variation among deer in their use of dense conifer cover during a 12-year study period as potentially influenced by winter severity and cover availability. Female deer were fitted with a mixture of very high frequency (VHF, n = 267) and Global Positioning System (GPS, n = 24) collars for monitoring use of specific cover types at the population and individual levels, respectively. We developed habitat composites for four study sites. We fit multinomial response models to VHF (daytime) data to describe population-level use patterns as a function of snow depth, ambient temperature, and cover availability. To develop alternative hypotheses regarding expected spatio-temporal patterns in the use of dense conifer cover, we considered two sets of competing sub-hypotheses. The first set addressed whether or not dense conifer cover was limiting on the four study sites. The second set considered four alternative sub-hypotheses regarding the potential influence of snow depth and ambient temperature on space use patterns. Deer use of dense conifer cover increased the most with increasing snow depth and most abruptly on the two sites where it was most available, suggestive of an energy conservation strategy. Deer use of dense cover decreased the most with decreasing temperatures on the sites where it was most available. At all four sites deer made greater daytime use (55 to >80% probability of use) of open vegetation types at the lowest daily minimum temperatures indicating the importance of thermal benefits afforded from increased exposure to solar radiation. Date-time plots of GPS data (24 hr) allowed us to explore individual diurnal and seasonal patterns of habitat use relative to changes in snow depth. There was significant among-animal variability in their propensity to be found in three density classes of conifer cover and other open types, but little difference between diurnal and nocturnal patterns of habitat use.

Conclusions/Significance

Consistent with our findings reported elsewhere that snow depth has a greater impact on deer survival than ambient temperature, herein our population-level results highlight the importance of dense conifer cover as snow shelter rather than thermal cover. Collectively, our findings suggest that maximizing availability of dense conifer cover in an energetically beneficial arrangement with quality feeding sites should be a prominent component of habitat management for deer.     

Migratory strategies, fawn recruitment, and winter habitat use by urban and rural mule deer (Odocoileus hemionus)

We examined the migratory strategies of mule deer (Odocoileus hemionus) using adjacent urban and rural winter ranges in northern Utah in relation to deer demography and patterns of habitat use. Urban deer were more likely to be migratory than rural deer, even though migratory animals from the two herds intermixed on a common, high-elevation summer range. Urban deer exhibited lower fawn recruitment than rural deer; but within each herd, demographic characteristics of migratory and nonmigratory animals suggested that game theory explained the ratios of deer adopting each behavior. Estimates of animal numbers and available habitat did not reveal clearly whether deer densities differed on the two winter ranges. However, patterns of habitat use by urban deer were so clustered around areas of concealment vegetation that animals probably experienced higher local densities than rural animals. In addition, the clustered patterns of habitat use by urban deer resulted in incomplete use of available forage. This may have contributed to the relatively poor fawn recruitment by urban deer, a phenomenon that appeared to be perpetuated by their strong fidelity to winter range.     

Ecological effects of fear: How spatiotemporal heterogeneity in predation risk influences mule deer access to forage in a sky‐island system

Forage availability and predation risk interact to affect habitat use of ungulates across many biomes. Within sky‐island habitats of the Mojave Desert, increased availability of diverse forage and cover may provide ungulates with unique opportunities to extend nutrient uptake and/or to mitigate predation risk. We addressed whether habitat use and foraging patterns of female mule deer (Odocoileus hemionus) responded to normalized difference vegetation index (NDVI), NDVI rate of change (green‐up), or the occurrence of cougars (Puma concolor). Female mule deer used available green‐up primarily in spring, although growing vegetation was available during other seasons. Mule deer and cougar shared similar habitat all year, and our models indicated cougars had a consistent, negative effect on mule deer access to growing vegetation, particularly in summer when cougar occurrence became concentrated at higher elevations. A seemingly late parturition date coincided with diminishing NDVI during the lactation period. Sky‐island populations, rarely studied, provide the opportunity to determine how mule deer respond to growing foliage along steep elevation and vegetation gradients when trapped with their predators and seasonally limited by aridity. Our findings indicate that fear of predation may restrict access to the forage resources found in sky islands.