Reidar Andersen

A slow life in hell or a fast life in heaven: demographic analyses of contrasting roe deer populations

1. Environmental conditions shape population growth through their impact on demographic parameters. While knowledge has accumulated concerning the effects of population density and climatic conditions, a topical question now concerns how predation and harvest influence demographic parameters and population growth (lambda). 2. We performed a comparative demographic analysis based on projection matrix models for female roe deer. Population-specific matrices were parameterized based on longitudinal data from five intensively monitored populations in Norway and France, spanning a large variability in environmental characteristics such as densities of large predators, hunter harvest and seasonality. 3. As expected for a large iteroparous vertebrate, temporal variation was invariably higher in recruitment than in adult survival, and the elasticity of adult survival was consistently higher than that of recruitment. However, the relative difference in elasticity of lambda to recruitment and adult survival varied strongly across populations, and was closely correlated with adult survival. 4. Different traits accounted for most of the variance in lambda in different ecological settings. Adult survival generally contributed more in populations with low mean adult survival and low mean growth rate during the study period. Hunters and predators (Eurasian lynx and red foxes) occurred in two of our study populations and contributed substantially to the variance in lambda, accounting for a total of 35% and 70% in the two populations respectively. 5. Across populations, we did not find any evidence that roe deer increased their reproductive output when faced with harsh conditions, resulting in some populations having negative growth rates. 6. Generation time, a measure of the speed of the life-history cycle, increased from less than 4 years in the most productive population (roe deer heaven) to more than 6 years in declining populations facing predation from lynx, red fox and hunters (roe deer hell), and was tightly and inversely correlated with lambda. Such a deceleration of the life cycle in declining populations might be a general feature in large herbivores. 7. Our results shows that the plethora of environmental conditions faced by populations of large herbivores also induce high intraspecific variation in their ranking along the fast-slow continuum of life-history tactics.

Climate, season, and social status modulate the functional response of an efficient stalking predator: the Eurasian lynx

1. Predation plays a major role in shaping the structure and dynamics of ecological communities, and the functional response of a predator is of crucial importance to the dynamics of any predator-prey system by linking the trophic levels. For large mammals, there is a dearth of field studies documenting functional responses, and observations at low prey density are particularly scarce. Furthermore, there is a lack of understanding about how variables such as season, social status and climate modulate the functional response curves. 2. We analysed kill rate data collected over a 10-year period based on radio-marked lynx (Lynx lynx) mainly preying on roe deer (Capreolus capreolus) along a steep prey density gradient in south-eastern Norway. 3. The asymptotic kill rate was reached at a very low prey density for both solitary individuals and family groups (i.e. females with their dependent kittens), indicative of an efficient predator. This highlights the importance of understanding the interplay between predator and prey at low prey densities. 4. A purely prey-dependent functional response was a poor descriptor of the data, as the curve was strongly modulated by season and differences between lynx of different social status. In addition, there was a clear effect of abiotic climatic factors (indexed by the North Atlantic Oscillation) on observed kill rates in the more snow-rich portion of our study area. 5. Our analysis suggests that simple functional response curves might be poor descriptors of predator consumption rates in complex natural system, and that auxiliary factors are likely to induce complexity into any predator-prey systems that would not be captured by simple deterministic approaches.

Habitat and roe deer fawn vulnerability to red fox predation

1. Notwithstanding the growing amount of literature emphasizing the link between habitat, life-history traits and behaviour, few empirical studies investigated the combined effect of these parameters on individual predation risk. We investigated direct and indirect consequences of habitat composition at multiple spatial scales on predation risk by red foxes on 151 radio-monitored roe deer fawns. We hypothesized that the higher resource availability in fragmented agricultural areas increased predation risk because of: (i) shorter prey movements, which may increase predictability; (ii) larger litter size and faster growth rates, which may increase detectability in species adopting a hiding neonatal anti-predator strategy. The sharing of risky habitat among littermates was expected to promote whole-litter losses as a result of predation. 2. The landscape-scale availability of agricultural areas negatively affected pre-weaning movements, but did not influence growth rates or litter size. Predation risk was best described by the interplay between movements and fine-scale habitat fragmentation: a higher mobility increased the encounter rate and predation risk in highly fragmented home ranges, while it reduced predation risk in forest-dominated areas with clumped resources because of decreased predictability. This is one of the first demonstrations that movement patterns can be an efficient anti-predator strategy when adjusted to local conditions. 3. In accordance with previous studies documenting the existence of family effects (i.e. non-independence among siblings) in survival, littermates survived or died together more often than expected by chance. In addition, our study specifically demonstrated the occurrence of behaviourally mediated family effects in predation risk: after a fox killed one fawn the probability of a sibling being killed within a few days rose from 20% to 47%, likely because of the win-stay strategy (i.e. return to a previously rewarding site) adopted by the predator. Hence, the predators hunting strategy has the potential to raise fawn mortality disproportionately to predator abundance. 4. There is increasing evidence that fawns inhabiting highly productive predator-free habitats are granted lifetime fitness benefits; these potential advantages, however, can be cancelled out when predation risk increases in the very same high-productivity areas, which might thus turn into attractive sinks.

When a generalist becomes a specialist: patterns of red fox predation on roe deer fawns under contrasting conditions

The red fox (Vulpes vulpes (L., 1758)) functional response to roe deer (Capreolus capreolus (L., 1758) fawn density was investigated in two Norwegian study areas characterized by a 26-fold differen...

Soil and vegetation nutrient response to bison carcasses in Bialowieza Primeval Forest, Poland

Ungulate carcasses can have important effects on the surrounding soil and vegetation. The impact of six carcasses of European bison (Bison bonasus) was investigated for the first time in a natural temperate forest (Białowieża, Poland) by measuring soil and plant nutrient concentrations along a gradient extending from the centre of each carcass. Calcium concentration and pH were found to be higher at the centre of the carcass, decreasing towards the periphery. This effect lasted up to 7xa0years after the death of the animal. The concentration of most nutrients in the soil and plants varied irregularly, suggesting an effect of the carcass at its centre but the absence of a clear pattern of variation along the gradient. Concentrations of NO3− in the soil differed only at the 1-year old carcass, suggesting a fast turnover of nitrate in temperate forests. Our results show that the effects of large herbivore carcasses on soil and plant nutrient concentrations are not easily detectable in a temperate forest as in more homogeneous habitats, such as tundra and prairie. This may be due to the high activity of scavengers and nutrient recycling in the study area, but it may also be a consequence of a more complex and patchy interaction between nutrient availability and other limiting factors in temperate forests.

Vulnerability of Domestic Sheep to Lynx Depredation in Relation to Roe Deer Density

Abstract The Eurasian lynx (Lynx lynx) causes large loss of free-ranging domestic sheep in Norway. We tested whether the observed higher kill rates by male lynx than female lynx were related to an association between the availability of the main natural prey, as measured by roe deer (Capreolus capreolus) habitat suitability, presence of sheep, and habitat selection of male and female lynx. We found that lynx selected areas with high roe deer suitability during summer and winter. Moreover, during summer, compared to male lynx, females had greater selection for roe deer areas and a stronger avoidance for sheep grazing areas, which suggests that previously observed differences in kill rates between male and female lynx can be attributed to sex-specific habitat use during summer. The connection between lynx habitat use and roe deer also was reflected in a positive relationship between the roe deer suitability of a sheep grazing area and the total loss of lambs, which suggests that livestock, rather than being actively selected, are mainly killed by lynx incidentally when encountered during other lynx activities (e.g., searching for natural prey species). Therefore, any management practice that separates lynx and sheep, such as concentrating livestock into small patches or less preferred habitats, may reduce depredation.

Selecting habitat to survive: The impact of road density on survival in a large carnivore

Habitat selection studies generally assume that animals select habitat and food resources at multiple scales to maximise their fitness. However, animals sometimes prefer habitats of apparently low quality, especially when considering the costs associated with spatially heterogeneous human disturbance. We used spatial variation in human disturbance, and its consequences on lynx survival, a direct fitness component, to test the Hierarchical Habitat Selection hypothesis from a population of Eurasian lynx Lynx lynx in southern Norway. Data from 46 lynx monitored with telemetry indicated that a high proportion of forest strongly reduced the risk of mortality from legal hunting at the home range scale, while increasing road density strongly increased such risk at the finer scale within the home range. We found hierarchical effects of the impact of human disturbance, with a higher road density at a large scale reinforcing its negative impact at a fine scale. Conversely, we demonstrated that lynx shifted their habitat selection to avoid areas with the highest road densities within their home ranges, thus supporting a compensatory mechanism at fine scale enabling lynx to mitigate the impact of large-scale disturbance. Human impact, positively associated with high road accessibility, was thus a stronger driver of lynx space use at a finer scale, with home range characteristics nevertheless constraining habitat selection. Our study demonstrates the truly hierarchical nature of habitat selection, which aims at maximising fitness by selecting against limiting factors at multiple spatial scales, and indicates that scale-specific heterogeneity of the environment is driving individual spatial behaviour, by means of trade-offs across spatial scales.

Roe Deer Population Growth and Lynx Predation along a Gradient of Environmental Productivity and Climate in Norway

Abstract: n The extent to which large carnivores compete with hunters for harvestable populations of wild ungulates is a topic of widespread controversy in many areas of the world where carnivore populations are recovering or are reintroduced. Theory predicts that predation impacts should vary with prey density and environmental conditions. To test this prediction, we analyzed trends in an index of population abundance of roe deer (Capreolus capreolus) over 9 y in 144 Norwegian municipalities. The municipalities span a wide range of landscapes and climatic conditions and were associated with a varying degree of Eurasian lynx (Lynx lynx) presence. There was a wide variation in trends of roe deer abundance (estimated long-term average &lgr; ranging from 0.69 to 1.23) among municipalities. Roe deer population growth rates were lower in the municipalities with lynx and harsh climatic conditions than in municipalities with mild climatic conditions and/or without lynx. Thus, lynx presence appears to be having a negative impact on roe deer populations; this was especially evident in areas with unfavourable environmental conditions. Our finding that estimated long-term average values of &lgr; were less than 1 in many municipalities indicates that roe deer populations in Norway may not be able to sustain current combined mortality from hunters and lynx, especially in marginal areas.

Reconstructing the history of a fragmented and heavily exploited red deer population using ancient and contemporary DNA

BackgroundRed deer (Cervus elaphus) have been an important human resource for millennia, experiencing intensive human influence through habitat alterations, hunting and translocation of animals. In this study we investigate a time series of ancient and contemporary DNA from Norwegian red deer spanning about 7,000 years. Our main aim was to investigate how increasing agricultural land use, hunting pressure and possibly human mediated translocation of animals have affected the genetic diversity on a long-term scale.ResultsWe obtained mtDNA (D-loop) sequences from 73 ancient specimens. These show higher genetic diversity in ancient compared to extant samples, with the highest diversity preceding the onset of agricultural intensification in the Early Iron Age. Using standard diversity indices, Bayesian skyline plot and approximate Bayesian computation, we detected a population reduction which was more prolonged than, but not as severe as, historic documents indicate. There are signs of substantial changes in haplotype frequencies primarily due to loss of haplotypes through genetic drift. There is no indication of human mediated translocations into the Norwegian population. All the Norwegian sequences show a western European origin, from which the Norwegian lineage diverged approximately 15,000 years ago.ConclusionsOur results provide direct insight into the effects of increasing habitat fragmentation and human hunting pressure on genetic diversity and structure of red deer populations. They also shed light on the northward post-glacial colonisation process of red deer in Europe and suggest increased precision in inferring past demographic events when including both ancient and contemporary DNA.

Trade-offs between maternal foraging and fawn predation risk in an income breeder

The choice of neonatal hiding place is critical for ungulates adopting hiding anti-predator strategies, but the consequences of different decisions have rarely been evaluated with respect to offspring survival. First, we investigated how landscape-scale choices made by roe deer fawns and their mothers affected predation risk by red foxes in a forest–farmland mosaic in southeastern Norway. After, we examined the effect of site-specific characteristics and behaviour (i.e. visibility, mother–fawn distance and abundance of the predator’s main prey item—small rodents) on predation risk. The study of habitat use, selection and habitat-specific mortality revealed that roe deer utilised the landscape matrix in a functional way, with different habitats used for feeding, providing maternal care and as refugia from predation. Mothers faced a trade-off between foraging and offspring survival. At the landscape-scale decisions were primarily determined by maternal energetic constraints and only secondarily by risk avoidance. Indeed, forage-rich habitats were strongly selected notwithstanding the exceptionally high densities of rodents which increased fawn predation. At fine spatial scales, a high visibility of the mother was the major factor determining predation risk; however, mothers adjusted their behaviour to the level of risk at the bed site to minimise predation. Fawns selected both landscape-scale refugia and concealed bed sites, but failure to segregate from the main prey of red foxes led to higher predation. This study provides evidence for the occurrence of spatial heterogeneity in predation risk and shows that energetically stressed individuals can tackle the foraging-safety trade-off by adopting scale-dependent anti-predator responses.