Overabundant wild ungulate populations in Europe: management with consideration of socio-ecological consequences

1. Throughout recent years, ungulates have experienced significant increases in numbers and geographic range sizes in Europe, becoming locally overabundant populations in some regions. Changes in legislation regarding poaching, abandonment of land and re-naturalisation of habitats, and decreasing numbers of hunters, among other things, have led to alarming scenarios in wild ungulate biology. Although ungulates bring some financial benefits for ecosystems and society through tourism and hunting, the problems associated with populations that are no longer controlled can outweigh the advantages. Damage to forestry and agriculture, ungulate-vehicle collisions, and diseases are among the most concerning problems related to ungulate overabundance.
2. To deal with these problems and to decide on the best management strategy to apply, it is essential to have tools available to monitor populations with an integrative approach based on ecological change indicators, and to assess population and ecosystem status. Furthermore, in a globalised world, people’s opinions matter, and sociological studies regarding human perception of wild mammals must take place in order to allow proper management, including consideration of people’s expectations as well as animal and ecosystem needs. Successful and unsuccessful management strategies have already been attempted, and the knowledge of consequences over time enables an adaptive approach.
3. Management of ungulate populations is a complex subject, and each case should be studied, analysing the cost_performance balance of measures to be taken, and ensuring ongoing financial means to carry out and continue with successful ecosystem management strategies. Multidisciplinary teams should be built, including biologists, veterinarians, stakeholders, sociologists, and others, to deal with the management of European wild ungulate populations.

     

Insights into the Management of Large Carnivores for Profitable Wildlife-Based Land Uses in African Savannas

Large African predators, especially lions (Panthera leo) and leopards (Panthera pardus), are financially valuable for ecotourism and trophy hunting operations on land also utilized for the production of other wildlife species for the same purpose. Predation of ungulates used for trophy hunting can create conflict with landholders and trade off thus exists between the value of lions and leopards and their impact on ungulate populations. Therefore productionist and conservation trade-offs are complexly graded and difficult to resolve. We investigated this with a risk-benefit analysis on a large private wildlife production area in Zimbabwe. Our model showed that lions result in substantial financial costs through predation on wild ungulates that may not be offset by profits from hunting them, whereas the returns from trophy hunting of leopards are projected to exceed the costs due to leopard predation. In the absence of additional income derived from photo-tourism the number of lions may need to be managed to minimize their impact. Lions drive important ecological processes, but there is a need to balance ecological and financial imperatives on wildlife ranches, community wildlife lands and other categories of multiple use land used for wildlife production. This will ensure the competitiveness of wildlife based land uses relative to alternatives. Our findings may thus be limited to conservancies, community land-use areas and commercial game ranches, which are expansive in Africa, and should not necessarily applied to areas where biodiversity conservation is the primary objective, even if hunting is allowed there.     

“Ecology of fear” in ungulates: Opportunities for improving conservation

Because ungulates are important contributors to ecosystem function, understanding the “ecology of fear” could be important to the conservation of ecosystems. Although studying ungulate ecology of fear is common, knowledge from ungulate systems is highly contested among ecologists. Here, we review the available literature on the ecology of fear in ungulates to generalize our current knowledge and how we can leverage it for conservation. Four general focus areas emerged from the 275 papers included in our literature search (and some papers were included in multiple categories): behavioral responses to predation risk (79%), physiological responses to predation risk (15%), trophic cascades resulting from ungulate responses to predation risk (20%), and manipulation of predation risk (1%). Of papers focused on behavior, 75% were about movement and habitat selection. Studies were biased toward North America (53%), tended to be focused on elk (Cervus canadensis; 29%), and were dominated by gray wolves (40%) or humans (39%) as predators of interest. Emerging literature suggests that we can utilize predation risk for conservation with top‐down (i.e., increasing predation risk) and bottom‐up (i.e., manipulating landscape characteristics to increase risk or risk perception) approaches. It is less clear whether fear‐related changes in physiology have population‐level fitness consequences or cascading effects, which could be fruitful avenues for future research. Conflicting evidence of trait‐mediated trophic cascades might be improved with better replication across systems and accounting for confounding effects of ungulate density. Improving our understanding of mechanisms modulating the nature of trophic cascades likely is most important to ensure desirable conservation outcomes. We recommend future work embrace the complexity of natural systems by attempting to link together the focal areas of study identified herein.     

Comparing patterns of human harvest and predation by Eurasian lynx Lynx lynx on European roe deer Capreolus capreolus in a temperate forest

Human harvest is the most important mortality factor for wild ungulates in Europe and can affect several aspects of ungulate biology. There is a growing concern about possible negative side effects of human harvest. To better understand the differences between human and natural mortality, we compared the extent, age and sex structure, nutritional condition, spatial and temporal distribution of human harvest, and natural predation by the Eurasian lynx Lynx lynx on the European roe deer Capreolus capreolus, the most abundant wild ungulate in Europe. Compared to the human harvest, lynx were less likely to kill fawns and yearlings than adults, and among adult deer, lynx were more likely to kill females. The proportion of roe deer with fat-depleted bone marrow was higher among lynx prey than among harvested animals. Average lynx kill rate was estimated to 47.8 roe deer per year, and lynx predation was considerably lower than the human harvest in the same area. While human harvest increased with higher roe deer density, lynx predation was similar across the gradient of roe deer densities. Comparison with other countries indicated that differences between human harvest and natural mortality of ungulates vary considerably in different parts of Europe. Variation in hunting practices and, even more importantly, carnivore predation may have an important role in buffering unwanted side effects of harvest of wild ungulates.     

Is hunting large carnivores different from hunting ungulates? Some judgments made by Norwegian hunters

The increase in large carnivore populations in a number of European countries causes numerous social conflicts and populations need to be kept at levels that are acceptable to the public. This may eventually require opening up or increasing public hunting of species like bear, wolf, wolverines and lynx as a management strategy. We surveyed a sample of 672 Norwegian hunters to examine how they judged a set of dilemmas associated with hunting carnivores versus ungulates. More than one-half of the sample would like to maintain or increase the current population sizes of the large carnivores (for wolverines: 57.5%; bears: 65.6%; wolves: 66.1%; lynx: 59.6%). A large majority of hunters (85.5%) emphasise not stressing the game over securing success in killing game, require adequate shooting skills (87.7%), linking harvest strategies to ecological principles (79.5%) rather than the level of conflict, and sustainable wildlife populations (95.1%) over optimum economic yield. Carnivore hunting is associated with a stronger preference for quotas based on science rather than local knowledge as well as paying more attention to the conflict level, compared to the judgments made for ungulate hunting. Positive attitudes toward maintaining or increasing carnivore populations are associated with a concern for animal welfare, and faith in scientific information, ecological values and sustainable wildlife populations.     

Standardizing the double-observer survey method for estimating mountain ungulate prey of the endangered snow leopard

Mountain ungulates around the world have been threatened by illegal hunting, habitat modification, increased livestock grazing, disease and development. Mountain ungulates play an important functional role in grasslands as primary consumers and as prey for wild carnivores, and monitoring of their populations is important for conservation purposes. However, most of the several currently available methods of estimating wild ungulate abundance are either difficult to implement or too expensive for mountainous terrain. A rigorous method of sampling ungulate abundance in mountainous areas that can allow for some measure of sampling error is therefore much needed. To this end, we used a combination of field data and computer simulations to test the critical assumptions associated with double-observer technique based on capture-recapture theory. The technique was modified and adapted to estimate the populations of bharal (Pseudois nayaur) and ibex (Capra sibirica) at five different sites. Conducting the two double-observer surveys simultaneously led to underestimation of the population by 15%. We therefore recommend separating the surveys in space or time. The overall detection probability for the two observers was 0.74 and 0.79. Our surveys estimated mountain ungulate populations (± 95% confidence interval) of 735 (± 44), 580 (± 46), 509 (± 53), 184 (± 40) and 30 (± 14) individuals at the five sites, respectively. A detection probability of 0.75 was found to be sufficient to detect a change of 20% in populations of >420 individuals. Based on these results, we believe that this method is sufficiently precise for scientific and conservation purposes and therefore recommend the use of the double-observer approach (with the two surveys separated in time or space) for the estimation and monitoring of mountain ungulate populations.     

Conservation in Saudi Arabia; moving from strategy to practice

Conservation in the Kingdom of Saudi Arabia is relatively young, yet have made considerable gains in conservation through strategic proclamation and reintroductions. Changes in land use, illegal hunting and competition with domestic stock has decimated the native ungulates, meaning that the survival of the native ungulate species is now completely dependent on protected area network. The challenge is to sustain this network to make meaningful conservation impact into the future. We review the status of ungulate conservation in Saudi Arabia and highlight that the conservation strategy is well developed. The major challenge faced in conservation in Saudi Arabia now is to implement what has been sanctioned.     

Identifying Conservation Successes,Failures and Future Opportunities; Assessing Recovery Potential of Wild Ungulates and Tigers in Eastern Cambodia

Conservation investment, particularly for charismatic and wide-ranging large mammal species, needs to be evidence-based. Despite the prevalence of this theme within the literature, examples of robust data being generated to guide conservation policy and funding decisions are rare. We present the first published case-study of tiger conservation in Indochina, from a site where an evidence-based approach has been implemented for this iconic predator and its prey. Despite the persistence of extensive areas of habitat, Indochina''s tiger and ungulate prey populations are widely supposed to have precipitously declined in recent decades. The Seima Protection Forest (SPF), and broader Eastern Plains Landscape, was identified in 2000 as representing Cambodia''s best hope for tiger recovery; reflected in its designation as a Global Priority Tiger Conservation Landscape. Since 2005 distance sampling, camera-trapping and detection-dog surveys have been employed to assess the recovery potential of ungulate and tiger populations in SPF. Our results show that while conservation efforts have ensured that small but regionally significant populations of larger ungulates persist, and density trends in smaller ungulates are stable, overall ungulate populations remain well below theoretical carrying capacity. Extensive field surveys failed to yield any evidence of tiger, and we contend that there is no longer a resident population within the SPF. This local extirpation is believed to be primarily attributable to two decades of intensive hunting; but importantly, prey densities are also currently below the level necessary to support a viable tiger population. Based on these results and similar findings from neighbouring sites, Eastern Cambodia does not currently constitute a Tiger Source Site nor meet the criteria of a Global Priority Tiger Landscape. However, SPF retains global importance for many other elements of biodiversity. It retains high regional importance for ungulate populations and potentially in the future for Indochinese tigers, given adequate prey and protection.     

Distribution pattern of the threatened Himalayan serow (Capricornis thar) in western midhills of Nepal: An insight for conservation along an altitudinal gradient

Almost nothing is known about the distribution and status of the Himalayan serow, an IUCN category “near threatened” ungulate, formerly widely distributed from subtropical to subalpine regions, in the Nepalese Himalaya. Heavy hunting, together with habitat loss and fragmentation are believed to have caused local extinctions in many sites within its geographical range. Distribution pattern of this dense forest dwelling threatened ungulate thus indicates conservation status of the whole unique ecosystems of this region. The objective of this study was therefore to assess distribution and status of serow in the Nepalese midhills. We conducted surveys, based on sightings and indirect presence signs, in western Nepal, between the subtropical Bardia National Park in the south and mountainous Shey Phoksundo National Park in the north, during 2008–2010. We recorded serow in the Bardia National Park. Further to the north, we recorded it only in highly rugged and inaccessible areas in the high altitudes, which suggests their possible extinction in the midhill region. The serow presence was strongly positively correlated with hunting intensity and forest intactness index. The populations of serow, scattered in the region, are small and therefore likely to be sensitive to demographic stochasticity. This all may consequently lead to future extinction of serow in the entire region. This information can be used to aid further research and decision-making processes for conservation planning along altitudinal gradient of Nepal's Himalaya.     

Key factors determining the seasonal population growth rate of European wild rabbits and their implications for management

Recently, the European rabbit (Oryctolagus cuniculus) has received contrasting considerations due to its multidimensional role in the Mediterranean ecosystems. Within this framework, knowledge of factors determining its population size may have important consequences for designing an effective management plan. In this paper, we quantified the combined influence of the major demographic and mechanistic factors on seasonal population fluctuation of European rabbits on Lemnos Island (Greece), during 2007–2009. We developed a hypothetical model taking into account direct (productivity, predation, hunting pressure, food shortage, habitat treatment) and indirect factors (soil moisture, adverse weather conditions) using structural equation modeling. We tested for their influence on the seasonal population growth rate (spgr) as determined by line transects to estimate rabbit abundance. The productivity induced higher pgr. Food shortage, which is affected by low soil moisture during the late summer and early autumn, is demonstrated to be the most important negative factor followed by the hunting pressure and predation. Demographic and mechanistic factors highlighted in this analysis could be used either for conservation or for controlling populations of the species.     

青藏高原有蹄类动物多样性和特有性: 演化与保护

青藏高原是地球上一个独特的自然地理单元, 具有丰富的生境类型和生物种类, 是生物多样性与全球环境变化领域的热点研究区域。青藏高原发育着独特的动物区系, 尤其是有蹄类动物, 如藏羚(Pantholops hodgsonii)、野牦牛(Bos mutus)、藏原羚(Procapra picticaudata)、普氏原羚(P. przewalskii)、白唇鹿(Przewalskium albirostris)等青藏高原特有动物。在本文中, 我们探讨了如下问题: 青藏高原有多少种有蹄类动物?有多少种特有有蹄类动物?其分布格局如何?生存状况如何?保护现状如何?我们首先确定了青藏高原动物地理区的地理边界。发现青藏高原有28种有蹄类, 其中10种是青藏高原特有种。青藏高原有蹄类种数占中国有蹄类的42%, 单位面积上的有蹄类物种密度比中国现生有蹄类物种平均密度高62%。特有种比例高达36%。然而, 青藏高原有蹄类物种分布不均匀, 其丰富度呈东部高西部低格局, 而高原特有有蹄类则分布于高原腹地。青藏高原有蹄类动物生存受威胁比例高, 其中71%为受威胁物种, 54%被列入濒危野生动植物种国际贸易公约(CITES)附录I或附录II。野外调查发现大额牛(Bos frontalis)已经野外灭绝, 目前仅在高黎贡山有人工养殖的群体。青藏高原有蹄类物种红色名录指数从1998-2015年呈现持续下降趋势, 全球气候变化将加剧这一趋势。青藏高原有蹄类受威胁局面仍在继续, 物种生存状况持续恶化。虽然经过40年的自然保护区建设, 但一些青藏高原有蹄类的重要种群与栖息地并没有被自然保护区和新建的三江源国家公园所覆盖。因此, 在开发青藏高原、实现人类社会经济发展目标的同时, 应尽量保存高原野生动物种群与生境, 实现青藏高原草地生态系统的可持续利用与生物多样性保护的双赢目标。     

Evidence for top predator control of a grazing ecosystem

The importance of top predators in controlling ecological processes in large, intact ecosystems is unclear. In grasslands that support abundant ungulates, top–down control by predators may be particularly important, because of the tight biogeochemical linkages of ungulate prey with plants and soil microbes. Here, I examined the effects of the recent reintroduction of the gray wolf Canis lupus on ecosystem processes in Yellowstone National Park, where herds of grazing ungulates previously have been shown to stimulate several processes, including soil net nitrogen (N) mineralization. Rates of ungulate grazing intensity and soil net N mineralization were compared before and after wolf reintroduction in grasslands ranging five‐fold in aboveground production. Grazing intensity and grassland net N mineralization declined after wolf reintroduction, a likely partial function of fewer ungulates; wolf predation has been one of several factors implicated in causing the decline in Yellowstone ungulates. In addition, the spatial pattern of grazing and net N mineralization changed after reintroduction. A shift in the spatial patterns of grazer‐associated processes is consistent with a growing body of work indicating that wolves have changed habitat use patterns of ungulates in Yellowstone National Park. These findings suggest widespread wolf effects on ungulate prey, plants, and microbial activity that have spatially reorganized grassland energy and nutrient dynamics in Yellowstone Park.     

Density dependence of daily activity in three ungulate species

Daily activity in herbivores reflects a balance between finding food and safety. The safety‐in‐numbers theory predicts that living in higher population densities increases safety, which should affect this balance. High‐density populations are thus expected to show a more even distribution of activity—that is, spread—and higher activity levels across the day. We tested these predictions for three ungulate species; red deer (Cervus elaphus), roe deer (Capreolus capreolus), and wild boar (Sus scrofa). We used camera traps to measure the level and spread of activity across ten forest sites at the Veluwe, the Netherlands, that widely range in ungulate density. Food availability and hunting levels were included as covariates. Daily activity was more evenly distributed when population density was higher for all three species. Both deer species showed relatively more feeding activity in broad daylight and wild boar during dusk. Activity level increased with population density only for wild boar. Food availability and hunting showed no correlation with activity patterns. These findings indicate that ungulate activity is to some degree density dependent. However, while these patterns might result from larger populations feeling safer as the safety‐in‐numbers theory states, we cannot rule out that they are the outcome of greater intraspecific competition for food, forcing animals to forage during suboptimal times of the day. Overall, this study demonstrates that wild ungulates adjust their activity spread and level based on their population size.     

Invasive exotic aoudad (Ammotragus lervia) as a major threat to native Iberian ibex (Capra pyrenaica): a habitat suitability model approach

The introduction of alien species to new environments is one of the main threats to the conservation of biodiversity. One particularly problematic example is that of wild ungulates which are increasingly being established in regions outside their natural distribution range due to human hunting interests. Unfortunately, we know little of the effects these large herbivores may have on the host ecosystems. This study deals with a first comparative analysis of the habitat requirements of two ungulate species that may be facing competition for resources in the south of Europe: the native Iberian ibex (Capra pyrenaica) and the exotic aoudad (Ammotragus lervia). The aoudad is a North African caprid introduced in 1970 as a game species in south‐eastern Spain. It has adapted well, and populations have been freely expanding since then. Ecological Niche Factor Analysis is used to describe the realized niche of both species where their distribution ranges merge. Both species occupy marginal areas of rugged terrain in the region. Marginality is higher for the Iberian ibex, which also presents a higher tolerance of secondary environmental gradients than the aoudad. Highly suitable areas for each species are secondarily suitable for the other. Reclassified and cross‐tabulated habitat suitability maps showing the areas of potential spatial coexistence and differences in ecological traits between both species are provided. The results obtained do not allow inferring resource competition between these species. However, current aoudad expansion could result in it invading the favoured habitats of the ibex. Inadequate hunting policy and monitoring, and increasing climatic resemblance of the study region to the native aoudad areas, due to a strong desertification process, are facilitating a high rate of expansion. We strongly recommend to eradicate or, at least, monitor these exotic populations, and promote active conservation practices, if one wants to preserve the unique natural resources present in this European region.     

Identification of ungulates used in a traditional Chinese medicine with DNA barcoding technology

Horns of Saiga antelope (Saiga tatarica) have always been an ingredient of “Lingyangjiao”, a traditional Chinese medicine (TCM). Persistent hunting for Saiga antelope has already threatened the survival of critical endangered populations in wild. To control the growing pressure, CITES and Chinese government have legislated for monitoring the trade of Saiga horns. However, similar ungulate horns are difficult to identify by their morphological characteristics, which has impeded the law enforcement. Besides Saiga antelope, other seven ungulate species which have similar horns are also sold and marked as “Lingyangjiao” in TCM markets to offset shortage of Saiga antelope horns. Such species are Gazella subgutturosa, Pantholops hodgsonii, Procapra picticaudata, Procapra gutturosa, Procapra przewalskii, Capra hircus, and Ovis aries. Our study aimed at implementing DNA barcoding technology to diagnose Saiga horns and the substitutes. We successfully extracted genomic DNA from horn samples. We recovered COI sequences of 644 bp with specific primers and 349 bp with nested PCR primers designed for degraded horn samples. The mean interspecific genetic distance of data set of the 644‐bp full barcodes and the 349‐bp mini‐barcodes was 14.96% and 15.38%, respectively, and the mean intraspecific distance was 0.24% and 0.20%, respectively. Each species formed independent clades in neighbor‐joining (NJ) phylogenetic tree of the two data sets with >99% supporting values, except P. gutturosa and P. przewalskii. The deep genetic distances gap and clear species clades in NJ tree of either full barcodes or mini‐barcodes suggest that barcoding technology is an effective tool to diagnose Saiga horns and their substitutes. Barcoding diagnosis protocol developed here will simplify diagnosis of “Lingyangjiao” species and will facilitate conservation of endangered ungulates involved in TCM “Lingyangjiao” markets, especially the Saiga antelope.     

Diet,Prey Selection,and Predation Impact of Black-Backed Jackals in South Africa

ABSTRACT To investigate the role of black-backed jackals (Canis mesomelas) as predators, we studied diet, prey selection, and predation impact of jackals on 2 game ranches in South Africa that differed in ungulate diversity and biomass. Results showed that large (>15 kg) ungulate species dominated jackal diets throughout the year on both the less diverse (range of ingested biomass across seasons = 39–78%) and more diverse (26–69%) game ranch. Other important food items included medium-sized mammals (1–3 kg; 1–26%) and fruit (2–69%), whereas small mammals comprised 3–11% of ingested biomass across seasons on both sites. Jackals were not random in consumption of ungulates, and consumption patterns suggested jackals actively hunted certain species rather than consumed them as carrion. During ungulate birthing periods, jackals consumed almost exclusively those ungulate species that were hiders (i.e., fawns were hidden in tall vegetation away from herd) regardless of ungulate densities, suggesting that primarily fawns were preyed upon. Among hiders, there was a negative relationship (P = 0.01) between body size and percent of population consumed by jackals, indicating smaller species were more susceptible than larger species to jackal predation. Consequently, springbok (Antidorcas marsupialis) were always selected over other ungulate species on both sites, and this species was the most impacted by jackal predation. In contrast, ungulate species that were followers (i.e., fawns immediately followed mothers within protection of the herd) were scarcely or not at all consumed by jackals, regardless of body size or density. Medium-sized mammals were selectively consumed over ungulates, and there was a negative relationship (P < 0.01) between consumption of berries and ungulates, indicating alternative food resources influenced consumption of ungulates on our study sites. Our results will help wildlife managers in Africa identify ungulate species susceptible to jackal predation, and can be used to develop management strategies for reducing jackal predation in areas where it is problematic.     

Cougar Kill Rate and Prey Composition in a Multiprey System

Abstract: Assessing the impact of large carnivores on ungulate prey has been challenging in part because even basic components of predation are difficult to measure. For cougars (Puma concolor), limited field data are available concerning fundamental aspects of predation, such as kill rate, or the influence of season, cougar demography, or prey vulnerability on predation, leading to uncertainty over how best to predict or interpret cougar-ungulate dynamics. Global Positioning System (GPS) telemetry used to locate predation events in the field is an efficient way to monitor large numbers of cougars over long periods in all seasons. We applied GPS telemetry techniques combined with occasional snow-tracking to locate 1,509 predation events for 53 marked and an unknown number of unmarked cougars and amassed 9,543 days of continuous predation monitoring for a subset of 42 GPS-collared cougars in west-central Alberta, Canada. Cougars killed ungulates at rates near the upper end of the previously recorded range, and demography substantially influenced annual kill rate in terms of both number of ungulates (subad F [SAF] = 24, subad M [SAM] = 31, ad M = 35, ad F = 42, ad F with kittens <6 months = 47, ad F with kittens <6 months = 67) and kg of prey (SAF = 1,441, SAM = 2,051, ad M = 4,708, ad F = 2,423, ad F with kittens <6 months = 2,794, ad F with kittens >6 months = 4,280). Demography also influenced prey composition; adult females subsisted primarily on deer (Odocoileus spp.), whereas adult males killed more large ungulates (e.g., moose [Alces alces]), and subadults incorporated the highest proportion of nonungulate prey. Predation patterns varied by season and cougars killed ungulates 1.5 times more frequently in summer when juveniles dominated the diet. Higher kill rate in summer appeared to be driven primarily by greater vulnerability of juvenile prey and secondarily by reduced handling time for smaller prey. Moreover, in accordance with predictions of the reproductive vulnerability hypothesis, female ungulates made up a higher proportion of cougar diet in spring just prior to and during the birthing period, whereas the proportion of males increased dramatically in autumn during the rut, supporting the notion that prey vulnerability influences cougar predation. Our results have implications for the impact cougars have on ungulate populations and have application for using cougar harvest to manage ungulates.     

Bottom-up and top-down processes in African ungulate communities: resources and predation acting on the relative abundance of zebra and grazing bovids

African ungulate populations appear to be limited principally by their food resources. Within ungulate communities, plains zebras coexist with grazing bovids of similar body size, but rarely are the dominant species. Given the highly effective nutritional strategy of the equids and the resistance of zebras to drought, this is unexpected and suggests that zebra populations may commonly be limited by other mechanisms. Long-term research in the Serengeti ecosystem and in the Kruger National Park suggests that zebra could be less sensitive to food shortage, and more sensitive to predation, than grazing bovids: if this is a general principle, then, at a larger scale, resource availability should have a weaker effect on the abundance of zebra than on grazing ruminants of similar body size (wildebeest and buffalo), and zebras should be relatively more abundant in ecosystems where predators are rare or absent. We test these expectations using data on 23 near-natural ecosystems in east and southern Africa. The abundance of wildebeest is more closely related to resources than is that of zebra; buffalo are intermediate. We show that hyena densities are closely correlated with those of lions, and use the abundance of lions as an index of predation by large predators. The numerical response of lions to increases in the abundance of their prey was linear for mesoherbivores, and apparently so for the three species alone. Finally, the abundance of zebra relative to grazing bovids is lower in ecosystems with high biomasses of lions. These results indicate that zebras may commonly be more sensitive to top-down processes than grazing bovids: the mechanism(s) have not been demonstrated, but predation could play a role. If it is true, then when numbers of the large mammalian predators decline, zebra populations should increase faster than buffalo and wildebeest.     

Evaluation of multiple behavioral responses of sika deer to human hunting pressures

Several ungulate populations have become overabundant worldwide, resulting in ecological and social effects. Wildlife managers must establish effective population control programs to mitigate these negative influences. Hunting is a significant mortality factor for ungulates, which exhibit sensitive behavioral responses toward hunting activities. Wildlife managers need a comprehensive understanding of the ungulate responses to different hunting pressures across seasons. We developed a conceptual method to evaluate multiple behavioral responses to human hunting pressures and applied this to sika deer (Cervus nippon) in Hokkaido, Japan, during August and November in 2015. We measured flight behavior, spatial avoidance, and temporal activity shift during non-hunting and hunting seasons in 4 regions with different hunting pressures. Although we did not observe a clear difference in flight initiation distance among study areas or seasons, sika deer showed significant spatial avoidance during the hunting season in areas with high hunting pressure. Furthermore, sika deer were more active at night in areas with higher hunting pressures, regardless of the season. Collectively, these results suggest that sika deer responded to hunting pressure by changing diel activity to nocturnal throughout the year and avoiding risky areas (i.e., around hunting roads) during hunting season, rather than by increasing flight distance when they encounter humans. Our method can be used for a more efficient population control program for wildlife managers, as it considers the spatiotemporal variations of flexible ungulate behaviors in response to hunting pressures.     

Resource selection,utilization and seasons influence spatial distribution of ungulates in the western Serengeti National Park

Understanding herbivore selection and utilization of vegetation types is fundamental to conservation of multispecies communities. We tested three hypotheses for how ungulate species select their habitats and how this changes with season: first, resources are distributed as a mosaic of patches so that ungulates are also distributed patchily; this distribution reflects habitat selection, which changes with season, the different ungulates behaving differently. Second, resources become scarcer in the dry season relative to those in the wet season. If interspecific competition prevails, then all species should show a contraction of habitats chosen. Third, if predation is limiting, competition will be minimal, and hence, habitat selection by herbivores will not differ between seasons. We used frequencies of occurrence in four common vegetation types in western Serengeti National Park to determine selection coefficients and utilization patterns and Chi‐square analysis to test the hypotheses. The results showed that selection changes differently in each species, agreeing with the first hypothesis. Herbivores did not all become more selective, as predicted by the competition hypothesis, nor did selection remain the same across seasons, as predicted by the predation hypothesis. These results can be useful in constructing habitat suitability maps for ungulate species with special conservation needs.