Assessing deer densities and impacts at the appropriate level for management: a review of methodologies for use beyond the site scale

• 1 Population density alone is unlikely to be a good predictor of the impacts of deer on their environment. The assessment of management requirements should therefore be based on assessment of deer impacts, alongside estimates of density.
• 2 Both density and impacts need to be monitored at a landscape scale, and there is a need to develop appropriate methodologies that allow managers to consider the current and likely future impact of deer.
• 3 The relevant scale for assessment (and management) varies both with deer species and context of impact, but should always encompass at least the estimated biological range of the population of deer present in an area. Some impacts (e.g. deer–vehicle collisions, and risks of disease transmission) may need to be assessed at a wider regional level.
• 4 In this review we consider various approaches available for assessing: absolute or relative animal abundance; impacts of ungulates on agriculture, forestry, amenity woodlands and other conservation sites; impacts on public safety (e.g. through road traffic accidents) and on humans or livestock through potential spread of disease.
• 5 In each case the advantages and disadvantages of a variety of methods are considered, before recommendations are made for methodologies which are sufficiently accurate, sufficiently robust and sufficiently practical to be favoured in a management context.
• 6 To address impacts at the landscape scale requires management policies that integrate information on both positive and negative impacts of deer in order to ensure appropriate and holistic management. We present a decision‐support framework suitable for use within the UK, using inputs from a variety of possible impact types to assist managers and forewarn of situations where current management may need to be modified.

     

Estimating abundance of introduced Chinese muntjac Muntiacus reevesi and native roe deer Capreolus capreolus using portable thermal imaging equipment

• 1 Managing the conservation impacts of deer requires knowledge of their numbers. However, estimating densities in forested areas is problematic, with pellet‐group‐based methods subject to error and uncertainty in estimating defecation and decay rates in addition to sampling variance. Use of thermal imaging equipment allows direct census by distance sampling.
• 2 Densities of introduced Chinese muntjac Muntiacus reevesi and native roe deer Capreolus capreolus were estimated in 12.8 km² of conifer forest in eastern England by thermal imaging distance transects. Estimated density of introduced muntjac (±95% confidence intervals [CI]: 20 ± 8 km⁻²) exceeded that of native roe deer (16 ± 6 km⁻²); 95% CI of c. 40% were achieved in 10 days of fieldwork.
• 3 Density estimates were not sensitive to the number of width bands applied during analysis. Detectability functions differed, with narrower effective strip widths (ESW) for muntjac (74 m) than roe deer (123 m). Thus, it is important to discriminate between species when censusing mixed assemblages. Detectability and ESW also differed among plantation growth stages. Stratification by habitat may therefore improve accuracy of density estimates but would require additional survey effort.

     

Temporal shifts of DNA‐microsatellite allele profiles in roe deer (Capreolus capreolus L.) within three decades

DNA‐microsatellite polymorphism (four loci) was studied in 56 male roe deer (Capreolus capreolus) from a 900‐ha hunting territory in the Vosges du Nord Mountains (France), culled over 34 years (1956–1990). Changed allele frequencies at two loci within this period, and increased allelic diversity, were traced to a phase of reduced population density and subsequent immigration. Decadic population samples collected within 900‐ha were distinguished by higher genetic variability measures than were certain geographical samples across Central Europe (4–900 km). On average, the decadic cohorts were distinguished by a gene diversity index of G_ST = 0.0286, and a genetic distance of D = 0.0938, which reflect 54% (G_ST) and 69% (D) of the respective geographic (350 km) differentiation indices of roe deer in Central Europe. The importance of demography and population ecology effects for microevolution in a large mammal is demonstrated, as is the risk of artefact by composing population samples of deer over several years. Population genetic screening should cover various demes of roe deer from the same general region, and be based on many unlinked polymorphic loci, to minimize the distorting effects of genetic dynamics at the small spatial scale.     

Predicting the status of wild deer as hosts of <Emphasis Type="Italic">Mycobacterium bovis</Emphasis> infection in Britain

Control of livestock diseases can become complicated when wild animals are involved. The Eurasian badger (Meles meles) is considered the principle wildlife host of Mycobacterium bovis (which causes bovine tuberculosis, bTB) in Great Britain and Ireland, but wild deer have also been implicated. Whether wild deer are likely to perpetuate bTB in cattle depends on the exposure risks they pose, the mode of pathogen transmission, the distances over which the disease can be transported and whether they can maintain infection within their own populations independently of other sources. We evaluated the likely host status of each of four species of wild British deer (red, roe, fallow and Reeves' muntjac) and the badger across a range of densities typically observed in Britain by manipulating the reproductive number equation proposed by Anderson and May (1991). We estimate that roe deer almost certainly act as spillover hosts at densities lower than 30 km⁻², red deer below 16 km⁻², muntjac below 6 km⁻², fallow below 4 km⁻² and the badger below 2 km⁻². We also estimate that muntjac will almost certainly act as maintenance hosts at densities above 56 km⁻², fallow above 47 km⁻² and badgers above 24 km⁻². For densities between these values, we cannot be certain of the host status of these species, and for red and roe deer we cannot be certain of host status under most natural conditions typically experienced in parts of Britain experiencing high incidence of bTB in cattle. However, enhanced transmission rates resulting from artificially high densities such as might be experienced at supplementary feeding sites may be sufficient to promote independent maintenance of infection. We were not able to confidently assign host status to any species over a wide range of densities, but conclude that this is likely to reflect reality, where host status may be affected as much by, for example, demographic fluctuations as it is by population density. Our results imply densities below which populations of wild deer inhabiting cattle bTB hotspots ought to be maintained in order to control the possibility of them perpetuating the cycle of intra- and interspecific M. bovis transmission.     

Roe and fallow deer: are they compatible neighbours?

The analysis of the relationships between population density and habitat features is important to evaluate the ecological needs of a species, its potential impact on ecosystems and its interspecific interactions. We analysed the spatial variation of roe deer Capreolus capreolus and fallow deer Dama dama densities in a Mediterranean area in summer 2007 and winter 2007/2008. Previous research has shown that fallow deer can actively displace and exclude roe deer from natural feeding sites. Here we show that both species have the greatest densities in ecotone habitats between wood and open fields (abandoned olive groves and pastures), but with contrasting geographic patterns. The fallow deer showed the greatest densities in the central northern part of the study area near to local historical release sites. The densities of roe deer were great where fallow deer were rare and low where fallow deer were abundant. Spatial overlap was great at the habitat scale, indicating a high potential for competition, but was low at the plot scale, suggesting that partitioning of space occurred at a fine scale. Supporting great numbers of deer, the ecotone areas are crucial for the management of ecosystems. We suggest that roe deer avoid areas with great densities of fallow deer and that interspecific interference from the latter affects the density and distribution of the former both at a fine and at a large scale.     

Regional-Scale Assessment of Deer Impacts on Vegetation Within Western Connecticut,USA

Abstract: High densities of white-tailed deer (Odocoileus virginianus) are believed to cause broad-scale forest regeneration failure and loss of plant diversity. But, the empirical basis for such presumptions is limited. We, therefore, conducted a survey in western Connecticut, USA, woodlots to examine how spatial variation in deer densities influences variation in impacts on plant species abundance, identity and diversity, and tree regeneration. We also used a Geographic Information System to quantify trends between land-cover type and deer density. Deer density was not correlated with any vegetation or land use variable. This suggests that deer density is not a leading factor determining variation in vegetation impacts across western Connecticut.     

Multi-Scale Foraging Habitat Use and Interactions by Sympatric Cervids in Northeastern China

Abstract: Moose (Alces alces) and roe deer (Capreolus pygargus) are sympatric in the forest region of northeastern China. Using univariate analyses of feeding sign data, we found the 2 species were positively associated, but there were distinctions between their use of forage resources across landscape, patch, and microhabitat scales. We used resource selection function models to predict the influence of environmental covariates on moose and roe deer foraging; we detected covariate effects at the landscape and microhabitat scales but not at the patch scale. Forage resources used by the 2 species were similar, but moose used wetter areas and more low-visibility habitats than did roe deer, which strongly avoided areas with sparse vegetation. Both species were influenced by forage abundance and distribution at the microhabitat scale but exhibited differences in intensity of use of plant species and microhabitats. Moose used areas with deeper snow and avoided hiding cover; roe deer avoided areas with higher total basal areas of tree stems and preferred areas with high plant species richness. For moose, there was a trade-off in the use of concealment cover between the landscape and microhabitat scales. We detected avoidance by moose of roads where roe deer occurred. Roe deer exhibited more capacity for coping with human disturbance and interspecific interaction. In areas similar to our study area, road closures and suppression of roe deer near roads within 3–5 years postlogging may benefit moose. Furthermore, a mosaic of areas with different logging intervals may contribute to spatial separation of moose and roe deer and promote their coexistence.     

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.     

Thirty‐four polymorphic microsatellites for European roe deer

The European roe deer (Capreolus capreolus) is an interesting model for molecular ecology studies because of its abundance and adaptability across a range of environments (including human‐modified habitats), and because of its increasing impacts on agricultural crops and on regenerating forests. We identify polymorphic microsatellites in two managed populations of roe deer in France by using cross‐species amplification of primers from other Cervids and from Bovids. Of the 62 primer pairs tested, 45 amplified microsatellites in roe deer, and 34 were polymorphic. Eleven primer pairs were selected for multiplex gel‐loading for routine genotyping of the studied populations.     

Achieving landscape-scale deer management for biodiversity conservation: The need to consider sources and sinks

Hyper-herbivory following predator removal is a global issue. Across North America and Europe, increasing deer numbers are affecting biodiversity and human epidemiology, but effectiveness of deer management in heterogeneous landscapes remains poorly understood. In forest habitats in Europe, deer numbers are rarely assessed and management is mainly based on impacts. Even where managed areas achieve stable or improving impact levels, the extent to which they act as sinks or persist as sources exporting deer to the wider landscape remains unknown. We present a framework to quantify effectiveness of deer management at the landscape scale. Applied across 234 km² of Eastern England, we assessed management of invasive Reeve's muntjac (Muntiacus reevesi) and native roe (Capreolus capreolus), measuring deer density (using thermal imaging distance transects 780 km/year), fertility, neonatal survival, and culling to quantify source-sink dynamics over 2008–2010. Despite management that removed 23–40% of the annual population, 1,287 (95% CI: 289–2,680) muntjac and 585 (454–1,533) roe deer dispersed annually into the wider landscape, consistent with their ongoing range expansion. For roe deer, culled individuals comprised fewer young deer than predicted by a Leslie matrix model assuming a closed population, consistent with age-dependent emigration. In this landscape, for roe and muntjac, an annual cull of at least 60% and 53%, respectively, is required to offset annual production. Failure to quantify deer numbers and productivity has allowed high density populations to persist as regional sources contributing to range expansion, despite deliberative management programs, and without recognition by managers who considered numbers and impacts to be stable. Reversing an unfavorable condition of woodland biodiversity requires appropriate culls across large contiguous areas, supported by knowledge of deer numbers and fertility. © 2013 The Wildlife Society.     

Distribution and range expansion of deer in Ireland

• 1 Throughout Europe, the range of many deer species is expanding. We provide current distribution maps for red deer Cervus elaphus, sika Cervus nippon, fallow deer Dama dama and muntjac deer Muntiacus sp. in Ireland, and estimates of range expansion rates for red deer, sika and fallow deer.
• 2 There was a considerable expansion in the ranges of red deer, sika and fallow deer between 1978 and 2008. The compound annual rate of expansion was 7% for red deer, 5% for sika and 3% for fallow deer. The total range increase was 565% for red deer, 353% for sika and 174% for fallow deer. The potential implications of these expansions are discussed.
• 3 There are unknown numbers of red‐sika hybrid deer in some parts of Ireland. Range expansion is likely to lead to further hybridizations with implications for the genetic integrity of deer stocks.
• 4 Sightings of free‐roaming muntjac deer were first recorded in 2007. The distribution of confirmed sightings of single and multiple animals in the eastern region of Ireland suggests multiple releases.
• 5 Deer are already impacting on both the economic and biodiversity values of habitats in Ireland, where, at present, no sustainable deer management policy exists.

     

老爷岭南部狍冬季移动、卧息生境选择及其适宜性评价

在动物生境研究中,移动生境和卧息生境是生境研究的焦点。开展移动生境和卧息生境选择,并在此基础上进行生境评价,有利于深入了解动物对移动和卧息生境条件的需求,制定科学合理的栖息地保护计划。以东北虎（Panthera tigris altaica）的主要猎物物种之一-狍（Capreolus pygargus）为研究对象,于2017-2019年冬季积雪覆盖期在老爷岭南部通过随机布设28个大样方和84条用于足迹链跟踪的样线收集狍的移动点和卧息点信息,再结合近年来收集的东北虎出现点,利用广义可加模型（GAM）和最大熵模型（MaxEnt）进行狍移动、卧息生境选择及评价研究。移动生境选择研究表明,狍在移动的过程中偏好选择坡度小、距农田距离>500 m、远离道路、居民点和低海拔或较高海拔的区域;移动生境评价分析表明,移动适宜和次适宜生境面积之和为1318.16 km²,占研究区域面积的51.28%,当加入虎活动点影响因子后,狍移动适宜和次适宜生境面积之和为901.52 km²,适宜和次适宜生境面积之和减少了31.61%。狍卧息生境选择研究表明,水源、农田、道路和雪深是影响狍卧息的关键因素,其中雪深对狍卧息生境选择的贡献率达到70.13%;卧息生境评价表明,卧息适宜和次适宜生境面积之和为1243.77 km²,占研究区域面积的48.39%,当加入虎出现点因子后,适宜生境和次适宜生境面积之和减少了61.00%,仅为485.02 km²。研究认为,虎的出现对狍移动和卧息生境选择均产生影响,虎的活动及捕食行为可能会减少狍的活动范围和频次,狍远离虎活动区域卧息休息,压缩了狍适宜卧息的空间。     

黑龙江省胜山林场冬季驼鹿、马鹿和狍的种间关系

1987—1989年的两个冬季在黑龙江省黑河市胜山林场,对驼鹿、马鹿和狍的食性、采食高度和栖息地进行了研究。驼鹿的主要食物是柳(22.4％)、榛(16.0％)、红松(14.3％)和桦(13.8％);马鹿为杨(36.0％)、桦(22.7％)和红松(17.3％);狍为桦(26.5％)、柳(19.0％)、杨(12.7％)。三种动物采食高度的频次为正态分布。驼鹿的采食高度(标准差)为137.4(±39.0)厘米;马鹿为136.2(±37.0)厘米;狍为84.2(±29.2)厘米。驼鹿和马鹿对红松林的利用程度最高,而狍主要利用沼泽植物和红松林。驼鹿与狍在食物维和栖息地维上的生态位重叠最大,驼鹿与马鹿在采食高度维上的重叠最大。三种动物在食物、采食高度和栖息地三维生态位上的重叠大小的顺序为:驼鹿和马鹿>驼鹿和狍>马鹿和狍。     

The relative densities of red (Cervus elaphus) and roe (Capreolus capreolus) deer and their relationship in Scottish plantation forests

Population densities of red and roe deer were estimated in 20 conifer plantations throughout central and northern Scotland. Ten forests were selected on the basis of culling records that were considered likely to cover the range of possible densities. Ten further forests were selected at random to investigate the relationship between red and roe deer densities. Density assessment was by standing-crop dung-group counts, on 1.5 x 100 m transects. Dung decay rates were estimated in four areas and used to convert the values to deer densities. Red deer densities were estimated to range from 0.3-35 km⁻²; roe deer densities ranged from 0.5-25 km⁻². Preferences for different structural habitats within individual forests (i.e. blocks with different age classes of tree) were inferred from dung densities. Red deer preferred unplanted ground, open-thicket, and prethicket; roe deer likewise preferred prethicket (highly) and open-thicket. Both species avoided closed-canopy habitats. Red and roe deer densities were negatively correlated and the relationship was significant when the data were log transformed ( P < 0.05).     

Variations in bark-stripping by red deer Cervus elaphus across Europe

• 1 The literature on bark‐stripping by red deer Cervus elaphus in Europe is reviewed to reveal quantitative variation in this behaviour and relate it to deer density and local characteristics such as dominant tree species, occurrence of artificial feeding, altitude, region and size of the study site. We also review the importance of bark in red deer diets over the seasons and discuss the causes of bark‐stripping, focusing on the significance of bark as food.
• 2 Over the 36 sites examined, the rate of bark‐stripping was highly variable (from 0 to 84% of susceptible trees debarked), with less damage in Scotland than in other European sites for which bark‐stripping rates were higher at high red deer density. Altitude, the size of the study site, the number of dominant tree species and the occurrence of artificial feeding do not significantly relate to the rate of bark‐stripping.
• 3 Bark sometimes made up a large proportion of red deer diet (> 10%), especially in areas with severe winters (high levels of snow), whereas in study sites with mild winters, bark was practically not eaten at all.
• 4 These results suggest that severe bark‐stripping could be related to a reduction in food resource availability. This food availability hypothesis needs to be better documented, dealing particularly with the possible interaction between food availability and red deer density.

     

Estimation of roe deerCapreolus capreolus and mouflonOvis aries densities, abundance and habitat use in a mountainous Mediterranean area

The abundance, density, and habitat use of roe deerCapreolus capreolus (Linnaeus, 1758) and mouflonOvis aries Linnaeus, 1758 were studied in a confined Mediterranean area in Greece with a dung survey based on the faecal accumulation rate (FAR) technique. Estimated density was modelled with generalized additive models using altitude, habitat type, and slope as potential covariates. Model selection among the set of candidate models was conducted based on their generalized cross-validation score. Roe deer had an estimated mean density of 13.9 ind./km² and the best model included slope and habitat type as covariates. The mean density of mouflon in the study area was 22.1 ind./km² and the best model used altitude and habitat type as covariates. For both species, the highest densities were encountered in abandoned cultivations and glades, followed by conifer forests, while the lowest densities were observed in maquis. However, use of open habitats by mouflon was much greater than it was for roe deer. The strong preference of mouflon (a grazer species) for open habitats that were abundant with grasses probably reflected food availability and contrasted with the more diverse habitat use by roe deer (a selective browser).     

Predation has a greater impact in less productive environments: variation in roe deer, Capreolus capreolus, population density across Europe

Aim We aimed to describe the large‐scale patterns in population density of roe deer Caprelous capreolus in Europe and to determine the factors shaping variation in their abundance. Location Europe. Methods We collated data on roe deer population density from 72 localities spanning 25° latitude and 48° longitude and analysed them in relation to a range of environmental factors: vegetation productivity (approximated by the fraction of photosynthetically active radiation) and forest cover as proxies for food supply, winter severity, summer drought and presence or absence of large predators (wolf, Canis lupus, and Eurasian lynx, Lynx lynx), hunter harvest and a competitor (red deer, Cervus elaphus). Results Roe deer abundance increased with the overall productivity of vegetation cover and with lower forest cover (sparser forest cover means that a higher proportion of overall plant productivity is allocated to ground vegetation and thus is available to roe deer). The effect of large predators was relatively weak in highly productive environments and in regions with mild climate, but increased markedly in regions with low vegetation productivity and harsh winters. Other potentially limiting factors (hunting, summer drought and competition with red deer) had no significant impact on roe deer abundance. Main conclusions The analyses revealed the combined effect of bottom‐up and top‐down control on roe deer: on a biogeographical scale, population abundance of roe deer has been shaped by food‐related factors and large predators, with additive effects of the two species of predators. The results have implications for management of roe deer populations in Europe. First, an increase in roe deer abundance can be expected as environmental productivity increases due to climate change. Secondly, recovery plans for large carnivores should take environmental productivity and winter severity into account when predicting their impact on prey.     

Hypoderma actaeon: an emerging myiasis in roe deer (Capreolus capreolus)

Subcutaneous larvae of Hypoderma spp. (Diptera: Oestridae) were detected in the dorsal region in 10 roe deer, Capreolus capreolus (Artiodactyla: Cervidae), hunted in central Spain between January and March 2016. All larvae were found in the inner side of the hide during the skinning of the animals. The study of the morphological features of eight larvae of different stages collected from two animals allowed the identification of Hypoderma actaeon Brauer. The small size (4–5 mm) of some of the first instars suggests that the internal lifecycle of H. actaeon may be exclusively subcutaneous. This is the first confirmation of H. actaeon in roe deer; however, further studies to assess the spread of the parasite and to follow the evolution of this myiasis in roe deer are needed.     

Quaternary history of the European roe deer Capreolus capreolus

• 1 The European roe deer Capreolus capreolus is a typical faunal element of the Holocene. It was already present in Europe at least 600 000 years ago and it has been known from both glacial and interglacial phases since then. With nearly 3000 fossil and subfossil records, it is one of the most frequent mammals in the Late Quaternary.
• 2 During the Middle and Late Weichselian Pleniglacial, the distribution of the roe deer was not restricted to the Mediterranean peninsulas but repeatedly reached regions of central Europe. In contrast to that, roe deer records from the Last Glacial Maximum (LGM, 21.0–14.5 ka ¹⁴C BP) are largely confined to the Mediterranean peninsulas – with the exception of south‐western France and the surroundings of the Carpathians where several records attest to its occurrence during the LGM.
• 3 During the Greenland Interstadial 1 (12.5–10.8 ka ¹⁴C BP), the species' distribution extended further north and the roe deer appeared north of the Alps and reached regions of central Germany. This seems to be correlated with the abrupt change to more favourable environmental conditions during this period. It is very likely that the roe deer disappeared north of the Alps during the Younger Dryas cooling (10.8–10.0 ka¹⁴C BP). The northern regions of the central European lowlands were recolonized by roe deer during the late Preboreal 9.7–9.5 ka ¹⁴C BP for the first time since the Weichselian Glacial.
• 4 The combined pattern of genetic data and fossil records of European roe deer suggests several regions in the Iberian peninsula, southern France, Italy and the Balkans as well as in the Carpathians and/or eastern Europe as glacial refugia. It further suggests that C. capreolus might have recolonized most parts of central‐northern Europe out of one or more eastern European (not Balkan) and/or Carpathian refugia. This recolonization wave might have blocked immigration from the traditional Mediterranean areas.

     

Indirect effects of deer on insect pests and soybean plants

1. White-tailed deer (Odocoileus virginianus Zimmermann) and insect pests negatively affect soybean production; however, little is known about how these herbivores potentially interact to affect soybean yield. Previous studies have shown deer browse on non-crop plants affects insect density and insect-mediated leaf damage, which together reduce plant reproductive output. In soybeans, reproductive output is influenced by direct and indirect interactions of different herbivores.
2. Here, we quantified indirect interactions between two groups of herbivores (mammals and insects) and their effects on soybean growth and yield. We examined responses of insect pest communities along a gradient of deer herbivory (29% to 49% browsed stems) in soybean monocultures.
3. Structural equation models showed that deer browse had direct negative effects on soybean plant height and yield. Deer browse indirectly decreased insect-mediated leaf damage by reducing plant height. Deer browse also indirectly increased pest insect abundance through reductions in plant height. Similarly, deer herbivory had an indirect positive effect on leaf carbon: nitrogen ratios through changes in plant height, thereby decreasing leaf nutrition.
4. These results suggest that pest insect abundance may be greater on soybean plants in areas of higher deer browse, but deer browse may reduce insect herbivory through reduced leaf nutrition.