Increased hormonal stress response of Apennine chamois induced by interspecific interactions and anthropogenic disturbance

Responses of animals to environmental changes and their interactions with other species play an important role in conservation. Sharing a common habitat may lead to interspecific competition for resources, but field assessment of these biological events is not always easily accomplished. By using a non-invasive method, we evaluated the physiological stress responses of Apennine chamois (Rupicapra pyrenaica ornata) to the presence of cattle, sheep and goat, red deer (Cervus elaphus), people (hikers), and predators to identify which factors may affect this endangered species. During September 2012, November 2012, and July 2013, a total of 318 faecal samples were collected in representative sites and analysed for faecal cortisol metabolites (FCM). FCM concentration was analysed through linear mixed-effect models. A significant increase in FCM values in Apennine chamois sharing their habitat with domestic animals was recorded during all study periods. On the contrary, stress responses to red deer and people were limited in time and emerged only during summer months, when hikers are more frequent and red deer extend their altitudinal range reaching chamois’ habitat. The observed effects of domestic animals, red deer, and hikers should be considered in future Apennine chamois management plans, which should include the regulation of pastured domestic livestock, anthropogenic disturbances, and possible interferences with other wild species within parks.     

Disentangling demographic effects of red deer on chamois population dynamics

Investigating the impact of ecological factors on sex‐ and age‐specific vital rates is essential to understand animal population dynamics and detect the potential for interactions between sympatric species. We used block count data and autoregressive linear models to investigate variation in birth rate, kid survival, female survival, and male survival in a population of Alpine chamois Rupicapra rupicapra rupicapra monitored over 27 years within the Stelvio National Park, Central Italian Alps, as function of climatic variables, density dependence, and interspecific competition with red deer Cervus elaphus. We also used path analysis to assess the indirect effect of deer abundance on chamois growth rate mediated by each demographic parameter. Based on previous findings, we predicted that birth rate at [t] would negatively relate to red deer abundance at year [t − 1]; survival rates between [t] and [t + 1] would negatively relate to red deer abundance at year [t − 1] and to the interactive effect of winter precipitation at [t + 1] and chamois density at [t]. Our results showed that birth rate was positively related to spring–summer precipitation in the previous year, but this effect was hampered by increasing red deer abundance. Kid and female survival rates were negatively related to the combined effect of chamois abundance and winter precipitation. Male and female survival rates were negatively related to lagged red deer abundance. The path analysis supported a negative indirect effect of red deer abundance on chamois growth rate mediated by birth rate and female survival. Our results suggest that chamois population dynamics was largely explained by the synergistic effect of density dependence and winter harshness, as well as by interspecific competition with red deer, whose effects were seemingly stronger on the kid–female segment of the population.     

Boreal forest titans do not clash: low overlap in winter habitat selection by moose (<Emphasis Type="Italic">Alces americanus</Emphasis>) and reintroduced bison (<Emphasis Type="Italic">Bison bison</Emphasis>)

Despite moose (Alces americanus) likely being both an ecological and cultural keystone species in the North American boreal forest, few studies have examined niche overlap between moose and other ungulates. In response to concerns from local people about the potential for competition for habitat between moose and reintroduced bison (Bison bison), and the potential for disturbance to moose by bison and bison hunters, we investigated overlap in winter habitat selection by these species in southwestern Yukon, Canada. We used available geo-referenced data of animal locations, and associated environmental covariates, to develop resource selection probability function (RSPF) models of early-winter and late-winter habitat selection by moose and bison. In early-winter, moose selected sub-alpine shrub-dominated habitats, whereas bison extensively selected wet sedge meadows in lowland valleys. Both species used a greater variety of habitats in late-winter, with moose selecting drainages above tree line or river valleys with deciduous forest cover, while bison selected either south-facing grassy slopes or wet sedge meadows. Given observed differences in seasonal habitat selection between moose and bison, these species are predicted to overlap on only 0.5 and 6.6% of our 12,818-km² study area during early-winter and late-winter, respectively. The lack of demonstrated winter habitat overlap between moose and reintroduced bison, coupled with low diet overlap, points to an overall low potential for competition for winter habitat between these species, or for disturbance to moose attributable to bison or bison hunters. Resource partitioning that facilitates coexistence on a shared landscape suggests that concern over the impact of reintroduced bison on resident populations of moose is likely unwarranted.     

High red deer density depresses body mass of roe deer fawns

Many previous studies have pointed out that, when resources are limited, the potential for competition should be high among sympatric species that display overlaps in habitat and nutritional niches. However, reliable evidence of competition between red deer (Cervus elaphus) and roe deer (Capreolus capreolus) has not been yet reported for life history traits directly measuring performance such as body mass, reproduction, or survival. From long-term monitoring of deer populations in the reserve of La Petite Pierre (France), we measured the sex-specific responses of roe deer fawn body mass to changes in red deer density after accounting for possible confounding effects of date of shooting, climatic conditions, and roe deer density. As expected under the hypothesis of competition, red deer density in a given year had a marked negative influence on body mass of roe deer fawns born the same year and the following year. Fawn mass of roe deer males and females responded in similar ways to changes in red deer density. Our study provides the first evidence of a negative response of roe deer performance to high red deer density.     

Toward an identification of resources influencing habitat use in a multi-specific context

Interactions between animal behaviour and the environment are both shaping observed habitat use. Despite the importance of inter-specific interactions on the habitat use performed by individuals, most previous analyses have focused on case studies of single species. By focusing on two sympatric populations of large herbivores with contrasting body size, we went one step beyond by studying variation in home range size and identifying the factors involved in such variation, to define how habitat features such as resource heterogeneity, resource quality, and openness created by hurricane or forest managers, and constraints may influence habitat use at the individual level. We found a large variability among individual's home range size in both species, particularly in summer. Season appeared as the most important factor accounting for observed variation in home range size. Regarding habitat features, we found that (i) the proportion of area damaged by the hurricane was the only habitat component that inversely influenced roe deer home range size, (ii) this habitat type also influenced both diurnal and nocturnal red deer home range sizes, (iii) home range size of red deer during the day was inversely influenced by the biomass of their preferred plants, as were both diurnal and nocturnal core areas of the red deer home range, and (iv) we do not find any effect of resource heterogeneity on home range size in any case. Our results suggest that a particular habitat type (i.e. areas damaged by hurricane) can be used by individuals of sympatric species because it brings both protected and dietary resources. Thus, it is necessary to maintain the openness of these areas and to keep animal density quite low as observed in these hunted populations to limit competition between these sympatric populations of herbivores.     

Food–niche relationships within a guild of alpine ungulates including an introduced species

To assess competition for trophic resources within a guild of ungulates, we studied the food niches of sympatric ungulates in an alpine area of the Italian Western Alps and evaluated the extent of their dietary overlap, especially with regard to the presence of the introduced mouflon Ovis musimon . The roe deer Capreolus capreolus had the most diverse diet including dicotyledons, young sprouts of trees and an unusually high percentage of graminoids. The red deer Cervus elaphus and chamois Rupicapra rupricapra mainly consumed graminoids, supplemented with dicotyledonous herbs and woody plants. The mouflon was a grass feeder, with graminoids representing the majority of items ingested in all seasons. Among the native species, we recorded a high overlap of feeding choices between red deer and chamois, which frequently used the same areas and are both intermediate feeders. The mouflon showed considerable dietary overlap with the chamois and red deer in all seasons, while the similarity with roe deer was low. High food–niche overlap can imply competition but only if resources are limited and the species use the same habitats.     

Red mason bees cannot compete with honey bees for floral resources in a cage experiment

Intensive beekeeping to mitigate crop pollination deficits and habitat loss may cause interspecific competition between bees. Studies show negative correlations between flower visitation of honey bees (Apis mellifera) and wild bees, but effects on the reproduction of wild bees were not proven. Likely reasons are that honey bees can hardly be excluded from controls and wild bee nests are generally difficult to detect in field experiments. The goal of this study was to investigate whether red mason bees (Osmia bicornis) compete with honey bees in cages in order to compare the reproduction of red mason bees under different honey bee densities. Three treatments were applied, each replicated in four cages of 18 m³ with 38 red mason bees in all treatments and 0, 100, and 300 honey bees per treatment with 10–20% being foragers. Within the cages, the flower visitation and interspecific displacements from flowers were observed. Niche breadths and resource overlaps of both bee species were calculated, and the reproduction of red mason bees was measured. Red mason bees visited fewer flowers when honey bees were present. Niche breadth of red mason bees decreased with increasing honey bee density while resource overlaps remained constant. The reproduction of red mason bees decreased in cages with honey bees. In conclusion, our experimental results show that in small and isolated flower patches, wild bees can temporarily suffer from competition with honey bees. Further research should aim to test for competition on small and isolated flower patches in real landscapes.     

Species richness,relative abundances and habitat use in local assemblages ofSorex shrews in Eurasian boreal forests

Using a large body of observational data on the occurrence ofSorex shrews in boreal forests, we test two models that predict the structure of small mammal communities along a gradient of increasing habitat productivity. Tilman’s (1982) model predicts a humped curve of species richness along productivity gradients. In contrast, we found a linear increase in species richness with increasing logarithm of the pooled density of shrews, which we use as a measure of habitat productivity for shrews. The model of Hanski and Kaikusalo (1989) assumes a trade-off between exploitative and interference competitive abilities, and it predicts that the size structure of small mammal communities should shift from the dominance of small species (superior in exploitative competition) in unproductive habitats to the dominance of large species (superior in interference competition) in productive habitats. Shrew assemblages show such a shift. Though it is not possible to draw definite conclusions about the role of interspecific competition from our observational data, the changing size structure of local shrew assemblages with increasing habitat productivity is a predictable feature of their community structure.     

Spatial distribution and habitat selection in coexisting species of mountain ungulates

One of the main objectives of community ecology is to understand the conditions allowing species to coexist, which requires identifying how co‐occurring species use and share space and resources. Species of the same trophic level, such as large herbivores, are of fundamental interest in that context because competition for resources is likely. Segregation in space or on some axes of the ecological niche are processes allowing coexistence, yet, both are seldom studied jointly. Based on annual spring censuses collected for 11 yr, we analysed the degree of overlap in spatial distribution among chamois Rupicapra rupicapra and mouflon Ovis gmelini musimon, 2 species of similar size that coexist on the same alpine pastures. We further investigated whether they differed in terms of habitat selection processes, and identified which environmental factors led to species being aggregated or segregated. The areas of intensive use for 2 species were more spatially aggregated than expected by chance. Habitat selection was studied using multivariate methods based on the niche concept, considering the presence of 1 species as an environmental variable for the other. Despite a large overlap in niches (88%), segregation was significant as chamois preferred meadows dominated by Sesleria and Carex sempervirens while mouflon selected meadows dominated by Carex ferruginea and avoided being close to areas affected by human activities. Importantly, habitat selection within each species was not affected by the presence of the other species. Coexistence between these 2 species and spatial overlap may be permitted because resource partitioning occurs at a fine temporal and/or spatial scale. We underscore that joint approaches of spatial and ecological processes are necessary to disclose the type of interaction (neutral, facilitation or competition) at play within a community.     

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.     

The effect of agricultural land use practice on habitat selection of red deer

The population of red deer (Cervus elaphus) has increased substantially in many western European countries during recent decades. Simultaneously, agricultural practices have undergone major changes. Observations suggest that meadows and pastures are important sources of nutrients for deer, but there are few studies quantifying the selection of agricultural land by deer in general. Here, we study red deer selection of various types of agricultural land and the history of land use (times of fertilisation and since renewal) in Norway. We used data from 14 female and 12 male red deer with GPS collars during the years 2007–2010. Our study design was to compare pairs of agricultural fields that had received “low” and “high” use by a given individual red deer and data were analysed by using case–control logistic regression. Our results showed that both sexes selected meadows over other types of agricultural land. Females selected intensively fertilised meadows and meadows of intermediate age, while male selection did not depend on meadow age (time since renewal) or fertilisation. The smaller females thus selected meadows of high quality, while the larger males continued to use old meadows of lower quality but rich in biomass. Our analysis suggests that a decreasing supply of meadows in the future due to lower production of livestock fodder may also affect management of red deer, and that a future change in the intensity of agricultural practices may affect sexes of wild large herbivores differently.     

Spatial variation in antler investment of Apennine red deer

Heterogeneity in resource availability and quality can trigger spatial patterns in the expression of sexually selected traits such as body mass and weaponry. While relationships between habitat features and phenotypic quality are well established at broad geographical scales, information is poor on spatial patterns at finer, intrapopulation scales. We analyzed biometric data collected on 1965 red deer Cervus elaphus males over 20 years from a nonmigratory population living on two sides of a mountainous ridge, with substantial differences in land cover and habitat quality but similar climate and population density. We investigate spatial patterns in (i) body mass, (ii) antler mass, and (iii) antler investment. We also tested for site‐ and age‐specific patterns in allometric relationship between body mass and antler mass. Statistically significant fine‐scale spatial variations in body mass, antler mass, and, to a lesser extent, antler allocation matched spatial differences in land cover. All three traits were greater in the northern slope, characterized by higher habitat heterogeneity and greater availability of open habitats, than in the southern slope. Moreover, the allometric relationship between body mass and antler mass differed among age‐classes, in a pattern that was consistent between the two mountain slopes. Our results support the occurrence of spatial patterns in the expression of individual attributes also at a fine, intrapopulation scale. Our findings emphasize the role of environmental heterogeneity in shaping spatial variations of key life‐history traits, with potential consequences for reproductive success.     

Lack of habitat segregation and no interspecific territoriality in three syntopic cryptic species of the golden‐spectacled warblers Phylloscopus (Seicercus) burkii complex

It is widely accepted that exploitative competition prevents the coexistence of any two or more closely‐related species unless differences exists in their ecological niches and resource use. In sibling bird species, exploitative competition is reduced mainly by spatial segregation of competing species. Spatial segregation can be achieved in two basic ways: by using different habitats or microhabitats that each species is predominately exploiting, or by interspecific territoriality. To our knowledge, either habitat segregation or interspecific territoriality or both have been found in all dyads and groups of sympartic sibling bird species studied so far. In this study, we investigated the coexistence of three cryptic sibling species of the golden‐spectacled warblers Phylloscopus burkii complex in Hunan Province, China: Ph. tephrocephalus, Ph. omeiensis and Ph. valentini. We analyzed their habitat preferences, and spatial interrelations between the species. In order to analyze either presence or absence of interspecific territoriality, we additionally performed con‐ and heterospecific playback experiments. Contrary to expectations, we found no evidences for either habitat segregation or interspecific territoriality in these three species.     

Feeding habits,forage selection,and diet overlap in Alpine chamois (<Emphasis Type="Italic">Rupicapra rupicapra</Emphasis> L.) and domestic sheep

Fecal microhistological analyses have been used to assess the summer diet of chamois (Rupicapra rupicapra L., 1758) and domestic sheep and to evaluate diet overlap. The aim of our work was to investigate interspecific interactions and to assess the effects of sheep grazing on the summer feeding habits of chamois. A high dietary overlap (Pianka’s index = 0.93–0.99) was found, despite differences in the use of some plant groups. Sheep presence strongly affected chamois feeding habits: a reduction of highly digestible forbs was observed in the chamois diet during August, when both species grazed in the same range. As a consequence of sheep grazing, chamois may have been forced to reduce niche breadth and to change their food habits, increasing percentages of monocotyledons in the diet and feeding mainly on Cyperaceae. The diet overlap between sheep and chamois suggests that resource-mediated interactions as well as direct competition must be taken into account when evaluating chamois and sheep compatibility on Alpine meadows.     

Influence of digestive morphology on resource partitioning in Amazonian ungulates

Summary Resource partitioning of diet and habitat use was studied in the entire Amazonian ungulate community of Northeastern Peru, which comprises the red brocket deer (Mazama americana), grey brocket deer (M. gouazoubira), collared peccary (Tayassu tajacu), white-lipped peccary (T. pecari), and lowland tapir (Tapirus terrestris). Each ungulate species partitioned at least one type of resource from every other species. Digestive morphology had a greater influence on resource partitioning of diet than body size. Neither digestive morphology nor body size were related to segregation of habitats. However, species with similar diets partitioned habitats, whereas species with different diets often used the same type of forest. Increases in habitat breadth of ungulates were positively correlated with increases in dietary breadth.     

Sexual segregation in ungulates: individual behaviour and the missing link

Previous "explanations" of sexual segregation in ungulates establish no more than a prerequisite for habitat segregation because they do not include a model of competitive habitat selection. Here we provide one based on the ideal free distributions of mutually competing, optimally foraging, individual deer. We parameterised our model using field data collected from a population of fallow deer (Dama dama) in a Mediterranean forest. The predictions of the inter-sex competition model were in full agreement with observational data, but those of single sex distributions (conventional theory) were not. The "conventional" hypothesis, that segregation arises simply from sex differences, predicted no more than moderate (20–40%) levels of segregation, even in optimal conditions. By introducing inter-sex resource competition, the predicted segregation can generally more than double and full segregation becomes possible in some circumstances. The modelling showed segregation to be density-dependent, varying in complicated ways with season and animal density. Sensitivity analysis showed the vulnerability of the "conventional" understanding of environmental variation and uncertainty. Using our competition model we show that as diet difference increases, direct competition between the sexes declines, so that as males increasingly differ from females, segregation declines and the two sexes are more likely to be found mixed (as long as the chosen food is available to both in the same area). Conversely, small differences among male and female deer are amplified by both food depletion and inter-sex competition to give substantial levels of segregation. The theoretical framework on which our model is built strongly suggests that sexual dimorphism in the context of scramble competition may be the fundamental cause of sexual habitat-segregation among ungulates.     

Components of breeding performance in two competing species: habitat heterogeneity, individual quality and density-dependence

Density‐dependent breeding performance due to habitat heterogeneity has been shown to regulate populations of territorial species, since the progressive occupation of low quality territories as breeding density increases may cause a decline in the mean per capita fecundity of a population while variation in fecundity increases. Although the preemptive use of sites may relegate low quality individuals to sites of progressively lower suitability, few studies on density dependence have tried to separate the effects of territory quality from individual quality, and none have simultaneously considered the effects of heterospecific competitors. Using two long‐term monitored populations, we assessed the relative contribution of habitat heterogeneity and bird quality (in terms of age) on the productivity of sympatric golden Aquila chrysaetos and Bonelli's eagles Hieraaetus fasciatus under different scenarios of intra‐ and inter‐specific competition. Productivity (number of offspring fledged) varied among territories and average annual productivity was negatively related to its variability in both species and populations, thus giving some support to the habitat heterogeneity hypothesis. However, the effect of habitat heterogeneity on productivity became non‐significant when parental age and local density estimators were included in multivariate analyses. Therefore, temporal changes in bird quality (age) combined with intra‐ and interspecific competition explained variability in territory productivity rather than habitat heterogeneity among territories per se. The recruitment of subadult breeders, a surrogate of mortality in eagles, strongly varied among territories. Habitat heterogeneity in productivity may thus arise not because sites differ in suitability for reproduction but because of differences in factors affecting survival. Territories associated with high mortality risks have a higher probability of being occupied by young birds, whose lower quality, interacting with the density competitors, leads to a reduction of productivity. Site‐dependent variability in adult survival and interspecific competition may be extensive, but so far largely overlooked, factors to be seriously considered for the site‐dependent population regulation framework.     

Habitat Interference by Axis Deer on White-Tailed Deer

Abstract: Many studies of interactions between exotic and native ungulates have not had temporal and spatial controls nor have they considered the types of competitive interactions that would allow coexistence. For exotic axis deer (Axis axis) and native white-tailed deer (Odocoileus virginianus) to coexist one species should be superior at interference competition and the other species should be superior at exploitative competition. We generated and tested predictions, based on body size and diet breadth, about habitat selection by white-tailed deer in the presence and absence of axis deer, dominance relationships, and time at sites provisioned with high quality forage. We conducted our study in treatment (axis and white-tailed deer) and control (white-tailed deer only) areas when both species were present and after axis deer were removed. We conducted vehicle surveys to determine habitat use of both species. At provisioned feeding sites we recorded aggressive behaviors and amount of time species spent at feeding sites alone and together. In the treatment area white-tailed deer selection for wooded habitat increased 2.1 times after axis deer were removed, whereas habitat selection by white-tailed deer was constant in the control area over the same time. At feeding sites axis deer were dominant to white-tailed deer; both species spent a significantly greater amount of time alone than at feeders together, but amount of time that individuals of each species spent at feeders did not differ. Axis deer were superior at interference competition, but white-tailed deer were not superior at exploitative competition; thus, species coexistence is unlikely. Whether white-tailed deer are negatively impacted by axis deer at spatial scales larger than our experiment probably depends on abundance of axis deer at larger spatial scales. Experiments of species interactions with temporal and spatial controls that consider types of competitive interactions increase a manager's understanding of when and how native ungulates may be negatively impacted by exotic ungulates.     

An allometric model of home range formation explains the structuring of animal communities exploiting heterogeneous resources

Understanding and predicting the composition and spatial structure of communities is a central challenge in ecology. An important structural property of animal communities is the distribution of individual home ranges. Home range formation is controlled by resource heterogeneity, the physiology and behaviour of individual animals, and their intra‐ and interspecific interactions. However, a quantitative mechanistic understanding of how home range formation influences community composition is still lacking. To explore the link between home range formation and community composition in heterogeneous landscapes we combine allometric relationships for physiological properties with an algorithm that selects optimal home ranges given locomotion costs, resource depletion and competition in a spatially‐explicit individual‐based modelling framework. From a spatial distribution of resources and an input distribution of animal body mass, our model predicts the size and location of individual home ranges as well as the individual size distribution (ISD) in an animal community. For a broad range of body mass input distributions, including empirical body mass distributions of North American and Australian mammals, our model predictions agree with independent data on the body mass scaling of home range size and individual abundance in terrestrial mammals. Model predictions are also robust against variation in habitat productivity and landscape heterogeneity. The combination of allometric relationships for locomotion costs and resource needs with resource competition in an optimal foraging framework enables us to scale from individual properties to the structure of animal communities in heterogeneous landscapes. The proposed spatially‐explicit modelling concept not only allows for detailed investigation of landscape effects on animal communities, but also provides novel insights into the mechanisms by which resource competition in space shapes animal communities.