Sheep grazing and rodent populations: evidence of negative interactions from a landscape scale experiment

Inter-specific competition, facilitation and predation influence herbivore assemblages, but no study has experimentally explored the interactions between large ungulates and small rodents. In a fully replicated, landscape scale experiment, we manipulated densities of domestic sheep in mountain pastures in Norway. We then determined population growth and densities of rodents by live trapping in each of the areas with different sheep densities. We found that the (summer) population growth rate and autumn density of the field vole (Microtus agrestis) was lower at high sheep density. This provides the first experimental evidence of negative interactions between an ungulate and small rodent species. There was no effect on the bank vole (Clethrionomys glareolus), whose diet differs from sheep. Sheep density, therefore, potentially alters the pattern of inter-specific population synchrony amongst voles. Our study shows that negative interactions between large ungulates and small rodents may be species-specific and negative population consequences for the rodent population appear above threshold ungulate densities.Electronic supplementary material is available for this article at     

The feeding habits of wolves in relation to large prey availability in northern Italy

We investigated wolf feeding habits in relation to the abundance of wild and domestic ungulates to test the hypothesis that large prey are preferred and that their abundance affects the use of other food categories and diet breadth. We determined diet composition by scat analysis from December 1987 to December 1992. The research was carried out in three study areas located in northern Italy and characterised by marked differences in wild and domestic ungulate abundance. In study area A (low wild and domestic ungulate availability) fruits, livestock, other vertebrates and wild ungulates made up the bulk of the diet (71% in volume). In area B (high availability of livestock) wolf diet was mainly based on sheep and wild boars (80% in volume). In study area C (high availability of wild ungulates) wild ungulates were the main food of wolves (90% in volume). Significant differences were found among study areas in the mean percentage volume of all food categories and in particular for wild ungulates, livestock, other vertebrates and fruits (p < 0.0001 in all cases). Diet breadth decreased in areas with high availability of large wild and domestic herbivores. The use of livestock species was lower where there was high abundance, richness and diversity of the wild ungulate guild. Selection for wild ungulate species was partially affected by their abundance: however other factors as prey social behaviour, adaptability to the habitat (for introduced species), and body size could have an important role in species selection by wolves. In particular in area C wild boars were selected for, roe and red deers avoided, and fallow deers and mouflons used as available. Livestock species were used in relation to their abundance and accessibility, in particular sheep were selected for and cattle avoided; but if calves bom in the pastures were considered as the only available cattle, they were selected for and sheep were used as available. Large and in particular wild herbivores were found to be of great importance for the wolf population maintenance in northern Italy, one of the most important recovery areas of Mediterranean wolves.     

Resource partitioning between ungulate populations in arid environments

Herbivores are major drivers of ecosystem structure, diversity, and function. Resilient ecosystems therefore require viable herbivore populations in a sustainable balance with environmental resource availability. This balance is becoming harder to achieve, with increasingly threatened species reliant on small protected areas in increasingly harsh and unpredictable environments. Arid environments in North Africa exemplify this situation, featuring a biologically distinct species assemblage exposed to extreme and volatile conditions, including habitat loss and climate change‐associated threats. Here, we implement an integrated likelihood approach to relate scimitar‐horned oryx (Oryx dammah) and dorcas gazelle (Gazella dorcas) density, via dung distance sampling, to habitat, predator, and geographic correlates in Dghoumes National Park, Tunisia. We show how two threatened sympatric ungulates partition resources on the habitat axis, exhibiting nonuniform responses to the same vegetation gradient. Scimitar‐horned oryx were positively associated with plant species richness, selecting for vegetated ephemeral watercourses (wadis) dominated by herbaceous cover. Conversely, dorcas gazelle were negatively associated with vegetation density (herbaceous height, litter cover, and herbaceous cover), selecting instead for rocky plains with sparse vegetation. We suggest that adequate plant species richness should be a prerequisite for areas proposed for future ungulate reintroductions in arid and semi‐arid environments. This evidence will inform adaptive management of reintroduced ungulates in protected environments, helping managers and planners design sustainable ecosystems and effective conservation programs.     

Lack of feral livestock interference with native guanaco during the dry season in a South American desert

Analyzing coexistence of exotic and native ungulates in arid areas is important from both a theoretical and a species conservation perspective. We assess the habitat use patterns and possible interference between guanaco (Lama guanicoe) and feral livestock (donkey and cattle) in arid environments of South America. To determine habitat use and niche overlap between exotic and native ungulate species, ten sites with different habitats and six natural waterholes were selected. Plots (20 at each site, ten around each waterhole) were randomly set up and characterized by environmental variables and relative use by cattle, donkey and guanaco through faecal pellet counts. Aggregation, niche breadth and niche overlap of the three herbivores were analyzed at habitat level (mesoscale). A direct redundancy analysis was used to examine the relationships between abundance of herbivore faeces and environmental variables at microscale. Mesoscale analyses showed (i) an extensive use of the area by all three species, with guanaco having the highest niche breadth followed by donkey and cattle and (ii) a large, broad guanaco–donkey and donkey–cattle habitat overlap. However, results at a finer scale showed high spatial aggregation of feral livestock species and an independent use of territory by guanacos. This study is the first to provide information about habitat partitioning between guanacos and feral livestock in the hyper-arid Monte Desert biome and points to an apparent lack of negative effects on the native ungulate.     

Habitat niche relationships within an assemblage of ungulates in Bardia National Park,Nepal

Species co-exist in their individual niches by partitioning resources. The main prey of the big predators – wild boar, chital, and sambar are sympatric in tropical south-Asia. In order to obtain a better understanding of their habitat use and habitat niche relationships, we used an indirect sign (faecal pellets and rooting by wild boar) survey along line transects in Bardia National Park. Ground cover, in particular grass cover for wild boar and chital, and shrub cover for chital and sambar positively affected the occurrence of ungulate signs. We found a niche differentiation between wild boar, which preferred hill sal forest and grassland, and chital and sambar, which avoided hill sal forest and grassland. Habitat niche overlap between chital and sambar in open habitat was clearly evident during the dry season. It is suggested that the fine grained habitat mosaics in space are the underlying causes of the coexistence of the ungulates studied. Therefore, focus on maintaining the habitat mosaic throughout the regions of the species’ distribution may prove successful for effective conservation management.     

Shifts in native ungulate communities on a former cattle ranch in Tanzania

When an area is brought under protection, current animal populations and their habitat preferences need to be assessed to predict population trends and future habitat availability. Using data from walking transects, we estimated the size of native ungulate populations on an abandoned cattle ranch in a coastal savannah in Tanzania, now included in the new Saadani National Park. Data were analysed with distance sampling and conventional strip transect techniques and were compared with results of previous wildlife counts. Few individuals of mainly browsing species were present in former cattle grazing areas exhibiting high bush‐encroachment while a ten times higher biomass of browsers and grazers was found in the cattle‐unmodified savannah. Population sizes of some species increased twofold between 1991 and 2001 within the entire area but neither population size nor species richness increased in the abandoned rangeland during our 3‐year study period from 2001 to 2003. We conclude that the former ranch has potential for future recolonization by wild ungulates. Resettlement will take place gradually with ‘pioneer‐species’ facilitating the entry of more demanding species. Habitat restoration through wildlife can be observed and quantified on Mkwaja Ranch which will be of importance for future management of native ungulates reclaiming abandoned rangeland.     

Taxonomic homogenization in ungulates: patterns and mechanisms at local and global scales

Aim The aim of this paper is to examine taxonomic homogenization in ungulates globally and at the local scale in South Africa. Specifically, we aim to examine the roles of distance, scale, time, extinctions vs. introductions, and extralimital vs. extraregional introductions in the homogenization of ungulate biotas, and to determine pathways of introduction of ungulate species globally and the proximate explanatory variables of ungulate introductions in South Africa. Location Forty‐one countries globally and three spatial resolutions in South Africa. Methods Indigenous, extirpated and established introduced ungulate species data were obtained for countries globally, and at a quarter‐degree grid‐cell resolution in South Africa. Homogenization was calculated using Jaccard’s index of similarity (JI) for countries globally and for three spatial resolutions in South Africa. Zoo holdings and transfer data from the International Species Information System database were used to investigate the relationship between non‐indigenous ungulate species introductions and the number of non‐indigenous ungulate species in zoos. Relationships between JI and species richness, and between numbers of introductions and several environmental and social factors were examined using generalized linear models. Results Homogenization in ungulates was 2% for countries globally and 8% at the coarsest resolution in South Africa. Homogenization increased with increasing resolution and with time, but it decreased with increasing percentage change in species richness. Globally, introductions contributed more to homogenization than did extinctions. Within South Africa, extralimital introductions contributed more to the homogenization of ungulate assemblages than did extraregional ones, and ungulates were typically introduced to high‐income areas with high human population and livestock densities. The same was not true in the past, when ungulates were introduced to ungulate species‐poor areas. The number of non‐indigenous ungulate species established in a country is significantly related to the number of non‐indigenous ungulate species in zoos in the country, possibly owing to sales of surplus animals from zoos. Main conclusions Ungulate faunas are homogenized at both the global scale and in South Africa, with extralimital introductions being of considerable significance regionally. In consequence, increasing attention will have to be given to the conservation consequences of ungulate translocations, both within particular geopolitical regions and across the globe.     

Rewilding traditional grazing areas affects scavenger assemblages and carcass consumption patterns

The abandonment of traditional livestock farming systems in Mediterranean countries is triggering a large-scale habitat transformation, which, in general, consists of the replacement of open grazing areas by woodlands through non-managed regeneration. As a consequence, wild ungulates are occupying rapidly the empty niche left by domestic ungulates. Both types of ungulates represent the main trophic resource for large vertebrate scavengers. However, a comparison of how vertebrate scavengers consume ungulate carcasses in different habitats with different ungulate species composition is lacking. This knowledge is essential to forecast the possible consequences of the current farmland abandonment on scavenger species. Here, we compared the scavenging patterns of 24 wild and 24 domestic ungulate carcasses in a mountainous region of southern Spain monitored through camera trapping. Our results show that carcasses of domestic ungulates, which concentrate in large numbers in open pasturelands, were detected and consumed earlier than those of wild ungulate carcasses, which frequently occur in much lower densities at more heterogenous habitats such as shrublands and forest. Richness and abundance of scavengers were also higher at domestic ungulate carcasses in open habitats. Vultures, mainly griffons (Gyps fulvus), consumed most of the carcasses, although mammalian facultative scavengers, mainly wild boar (Sus scrofa) and red fox (Vulpes vulpes), also contributed importantly to the consumption of wild ungulate carcasses in areas with higher vegetation cover. Our findings evidence that the abandonment of traditional grazing may entail consequences for the scavenger community, which should be considered by ecologists and wildlife managers.     

Host preferences and behaviour of oxpeckers: co-existence of similar species in a fragmented landscape

The host preferences and behaviour of red-billed and yellow-billed oxpeckers (Buphagus erythrorynchus and Buphagus africanus) were studied at two locations in Kenya and combined with previously published data from East and South Africa. Red-billed oxpeckers are generally more common and have a greater niche breadth with respect to host preferences than yellow-billed oxpeckers, possibly due to the behavioural dominance of the latter. Otherwise, the host preferences are remarkably similar: both prefer larger species of ungulates and are found predominantly on species with manes. There are also no significant differences in the time–activity budgets of the two species controlling for site and host species, and differences in the proportion of time spent on different body parts of hosts are small in absolute magnitude even when statistically significant. Despite these and other basic ecological similarities, no evidence for competitive displacement was detected by comparing host preferences in areas of sympatry with areas of allopatry, even though each species appears to depress the population size of the other when sympatric. Traditional single-community competition theory cannot explain the geographical ecology of these two species. Instead, co-existence appears to be dependent on the patterns of extinction and colonization characterized by metapopulations distributed among a fragmented habitat.     

Ungulates increase forest plant species richness to the benefit of non‐forest specialists

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant‐plant interactions and through soil disturbance. In forest ecosystems, researchers’ attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation‐wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light‐demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non‐forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape‐level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.     

同域分布中华鬣羚与中华斑羚时空生态位特征

为了解同域分布有蹄类在环境复杂的山地森林生境中以何种方式维持种间关系以实现稳定共存,基于物种分布模型与日活动模式分析了四川省岷山、邛崃、大相岭、小相岭和凉山五大山系同域分布中华鬣羚(Capricornis milneedwardsii)与中华斑羚(Naemorhedus griseus)的时空生态位特征。结果显示:(1)在四川省五大山系,中华鬣羚的适宜栖息地面积为28006.07 km²,占研究区总面积的26.18%,其中高适宜栖息地面积为10015.90 km²,中华斑羚的适宜栖息地面积为21073.32 km²,占研究区总面积的19.71%,其中高适宜栖息地面积为8396.22 km²;(2)中华鬣羚与中华斑羚在生境因子选择上相似性高、栖息地重叠面积大,其空间生态位重叠度指数D=0.776,I=0.949,其适宜栖息地的主要重叠区域位于岷山和邛崃山系;(3)中华鬣羚与中华斑羚的日活动节律重叠指数为0.812;(4)中华鬣羚与中华斑羚属于同域分布的资源利用型竞争物种,中华鬣羚的存在会显著影响中华斑羚的日活动节律(P=0.016);二者同域分布时都会增加其昼间活动强度,并增加活动高峰期的强度及持续时间。本研究初步分析了中华鬣羚与中华斑羚的时空生态位特征,揭示了二者在空间、时间生态位上种群共存及种间竞争的耦合关系。研究有利于深入理解同域分布动物时空生态位特征、近缘物种的共存机制及种间竞争关系,为有蹄类等珍稀野生动物种群及栖息地的保护提供科学参考。     

Food competition between a large ruminant and a small hindgut fermentor: the case of the roe deer and mountain hare

In this study, we demonstrate that the mountain hare and roe deer compete with each other. This was determined using "natural experiments" of populations found in sympatry and allopatry on the islands along the west coast of Norway. We demonstrate that both species occupy the same habitats, share the same food resources and that food availability is limited. Two browsing species as different in size as roe deer and mountain hare might be expected to partition the available vegetation (e.g. woody scrub) in terms of height above ground level. However, from the evidence collected, the feeding-height-separation hypothesis must be rejected as an explanation for ecological separation between roe deer and mountain hares because there was extensive height overlap in resource utilisation by both species and neither species changed its feeding height in response to the presence of the other. Total browse utilisation did not increase when both species were together; rather, species-specific browse utilisation declined when the other species was present. However, the foraging behaviour of each herbivore varied significantly between the allopatric and sympatric sites. When both herbivores were present, the clip diameter of shoots browsed by mountain hares declined to match those selected by roe deer, while roe deer switched from a browse-dominated diet to a diet dominated by winter-green gramineae. The change to smaller-diameter shoots likely resulted in the hare increasing its intake of digestion-inhibiting or toxic secondary metabolites, while the alternative choice of digging through the snow like roe deer to reach the winter-green gramineae is a practice considered energetically too costly for hares. On this basis, we conclude that the enforced switch to a nutritionally inferior diet by mountain hares at the sympatric sites may result in changes to growth rate and body size which consequently impact on mortality and may explain the competitive superiority of the roe deer.     

Foraging of lynxes in a managed boreal-alpine environment

Foraging of Eurasian lynxes Lynx lynx was studied with telemetry and snow tracking in central Norway. In all habitats and at all seasons, medium-sized ungulates (roe deer Capreolus capreolus , reindeer Rangifer tarandus and domestic sheep Ovis aries) dominated the diet (81% of ingested biomass estimated from faeces). Mountain hares Lepus timidus and galliform birds comprised the remainder of the diet (15% and 3%, respectively). Lynxes with different life history status did not differ in prey choice, but adult males utilised carcasses of ungulate prey considerably less (16% of the edible parts) than did females with offspring (80%) and subadults (58%.). Forest habitats in lowlands and adjacent to cultivated fields were the most favourable foraging habitats (indexed as the prey encounter rate per km lynx track) primarily owing to the presence of roe deer. Two family groups tracked in winter killed 0.2 ungulate per day. The importance of agricultural land as a foraging habitat and the dominance of livestock in the diet in remoter areas indicate that the lynx has responded to agriculture and reindeer husbandry during the past century by switching from smaff game to ungulates.     

Niche compression: Two tests of an hypothesis using narrowly sympatric predator species

The compression hypothesis predicts that when two species occur together in narrow sympatry, individuals in the overlap zone will use a smaller range of habitats and a larger or unchanged range of prey than individuals in allopatry. This report attempts to validate the assumptions underlying this hypothesis, and describes two tests of the predicted resource shifts using narrowly sympatric species of predatory marsupials (Antechinus spp.). In the first test, using A. stuartii and A. flavipes, there was little correspondence between observed and predicted changes in diet and habitat use (as measured by indices of niche breadth) for populations of either species. One index of niche breadth (Levins' B) revealed no trends in resource use, but a proportional similarity index relating resource use to resource abundance consistently indicated that the niche breadths of each species were greater in allopatry than in narrow sympatry. In the second test, using A. stuartii where it occurred alone and in narrow sympatry with A. swainsonii, there was again little correspondence between observed and predicted changes in resource use for seasonal or overall data. Differences in magnitudes of niche breadths between allopatric and narrowly sympatric A. stuartii were smaller in this test than in the first. While these findings provided no support for the compression hypothesis, they suggested three situations in which the predictions of the hypothesis may generally not apply. These are:

• (i) when the predators are active, dietary generalists (diet cannot expand in sympatry);
• (ii) when one species is a markedly superior competitor to the other (the dominant species suffers little or no niche compression in sympatry); and
• (iii) when the species compete by interference rather than exploitation (one species is evicted from sympatry or both show compression in habitat use and diet).

     

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.     

贺兰山同域分布岩羊和马鹿的夏季食性

2008年78月贺兰山对同域分布的岩羊和马鹿粪便进行采集,利用粪便显微分析法对二者夏季食性进行了研究。从10个沟系中一共收集了297堆岩羊和305堆马鹿粪便,每堆中捡拾2粒粪粒,组成40个复合样本进行分析。结果表明,夏季在岩羊采食的19科35种(属)植物中,针茅(29.74%)、早熟禾(18.82%)、内蒙古邪蒿(10.30%)、灰榆(7.76%)、冰草(7.47%)是其主要食物,针茅是岩羊夏季的大宗食物;夏季在马鹿取食的11科18种(属)植物中,小红柳(36.26%)、山杨(23.10%)、灰榆(16.84%)等是其主要食物,小红柳为马鹿夏季的大宗食物。夏季岩羊采食植物中,乔木占8.67%,灌木占3.97%,禾本科草本占69.43%,非禾本科草本占17.94%;马鹿采食植物中,乔木占89.61%,灌木占5.38%,禾本科草本占1.03%,非禾本科草本占3.99%。从食物多样性和食物生态位宽度来看,岩羊的Shannon-Wiener指数高于马鹿,而马鹿的Pielou均匀性指数和食物生态位宽度指数高于岩羊,岩羊和马鹿共有9种食物重叠,生态位重叠指数为65.17%。     

Trophic niche overlap and wild ungulate consumption by red fox and wolf in a mountain area in Italy

Red fox (Vulpes vulpes) and wolf (Canis lupus) are two widespread opportunistic predators living in simpatry in many areas. Nonetheless, scarce information are available on their trophic interactions. We investigated food habits of these two carnivores in a mountain area in Italy and assessed the extent of their trophic niche overlap, focusing on the consumption of wild ungulates. Thereby we analyzed the content of 669 red fox scats and 253 wolf scats collected between May 2008 and April 2009. Red foxes resulted to have a more than three times higher niche breadth than wolves. Vegetables, small mammals, wild ungulates, and invertebrates were major items (altogether 92% of volume) of the red fox annual diet. On the contrary wolf annual diet relied on wild ungulates (94% of volume) with wild boar (Sus scrofa) being the main food item. The degree of trophic niche overlap between the two species was found to be low (Pianka's O = 0.356). Diet variation between the warm and the cold seasons was limited in both species, and higher in red fox than in wolf. The two canids appeared to use wild ungulates unevenly being the former more selective for younger preys, smaller in size (newborn piglets and roe deer Capreolus capreolus fawns), whereas the latter exhibited a preference for medium-sized and large ungulates (10–35 kg wild boar and adult roe deer). Even if wild ungulates represent the main shared food category, the different use of age/weight classes by the two predators, together with their possible consumption as carrions by red fox, suggests a very limited trophic competition between wolf and red fox.This study represents a contribution to the knowledge of trophic interaction in predator–prey systems where sympatric carnivores are present.