Vebjørn Veiberg

Temporal scales, trade-offs, and functional responses in red deer habitat selection

Animals selecting habitats often have to consider many factors, e.g., food and cover for safety. However, each habitat type often lacks an adequate mixture of these factors. Analyses of habitat selection using resource selection functions (RSFs) for animal radiotelemetry data typically ignore trade-offs, and the fact that these may change during an animals daily foraging and resting rhythm on a short-term basis. This may lead to changes in the relative use of habitat types if availability differs among individual home ranges, called functional responses in habitat selection. Here, we identify such functional responses and their underlying behavioral mechanisms by estimating RSFs through mixed-effects logistic regression of telemetry data on 62 female red deer (Cervus elaphus) in Norway. Habitat selection changed with time of day and activity, suggesting a trade-off in habitat selection related to forage quantity or quality vs. shelter. Red deer frequently used pastures offering abundant forage and little canopy cover during nighttime when actively foraging, while spending much of their time in forested habitats with less forage but more cover during daytime when they are more often inactive. Selection for pastures was higher when availability was low and decreased with increasing availability. Moreover, we show for the first time that in the real world with forest habitats also containing some forage, there was both increasing selection of pastures (i.e., not proportional use) and reduced time spent in pastures (i.e., not constant time use) with lowered availability of pastures within the home range. Our study demonstrates that landscape-level habitat composition modifies the trade-off between food and cover for large herbivorous mammals. Consequently, landscapes are likely to differ in their vulnerability to crop damage and threat to biodiversity from grazing.

Monitoring population size of red deer Cervus elaphus: an evaluation of two types of census data from Norway

Abstract Proper management of wildlife relies on metrics of population development. Typically, the best estimation techniques are too expensive for coarse-scale management. In marine fisheries, catch-per-unit effort is commonly used, but problems may arise due to changes in spatial harvest effort or in habitat use as density changes. Managers in Norway are in the early phases of implementing ′seen deer′ during harvesting and ′spring counts′ on farmland as a means of monitoring red deer Cervus elaphus populations. We provide a first evaluation of how suitable these methods are by comparing the results with population estimates obtained using cohort analysis, and by analysing the within-season variation in number of seen deer. ′Seen deer′ predicted annual increases in populations fairly well. Adjusting for harvesting effort provided less good estimates, due to a proportionally larger increase in effort relative to deer population size as population size increased. The number of seen deer per day decreased rapidly at the beginning of the season, and then levelled off or increased slightly during the rut, especially on farmland. The number of seen deer increased both with the number of harvesters and hours harvested, but at a diminishing rate. The current practice of ′spring counts′ was not successful in predicting population changes, probably due to a lack of replication. Indeed, date strongly affected the number of deer seen during spring counts. While ′seen deer′ seems to be a very promising tool for monitoring population size of red deer, there are some limitations to the practice as implemented for moose Alces alces in Scandinavia due to a more complex relationship with harvesting effort. Our study highlights that the large number of hours harvesters observe wildlife can provide a useful tool for population monitoring. However, the use of such indices may vary between species and according to harvest techniques and should thus be assessed with care before implementation.

Social rank, feeding and winter weight loss in red deer: any evidence of interference competition?

During winter at northern latitudes, large herbivores often exploit patches of concentrated, relatively high quality forage, which may lead to interference competition. The factors affecting success in contests and subsequent dominance rank, such as age and body weight, remain key issues in ungulate behavioural ecology. Maternal effects on offspring body weight are well known, but few studies have investigated if mother’s social rank influence offspring rank. Moreover, no study has related dominance rank in ungulates to weight loss during winter. Outcomes of social interactions (n=7,609), feeding time and spatial position in red deer (Cervus elaphus) hinds and calves, and weight loss of calves, were registered from 1981 to 1996 at six winter-feeding sites within the county of Sør-Trøndelag in Norway. The level of aggressiveness was higher among calves than among adult hinds, and the factors determining the outcome of contests also differed. The initiator won the majority of interactions (more than 90% in both hinds and calves). Social rank was related to both age and body weight in adult hinds, and related to body weight and mother rank in calves. The relationship between feeding time and rank was non-linear. Feeding time was correlated with rank only among high ranked hinds, while there was no such relationship among low ranked hinds or calves. There was no correlation between winter weight loss and social rank in calves. Our study therefore underlines that, although frequent aggression is observed at artificial feeding sites of northern herbivores, this is not necessarily sufficient to give rise to interference competition.

Congruent responses to weather variability in high arctic herbivores

Assessing the role of weather in the dynamics of wildlife populations is a pressing task in the face of rapid environmental change. Rodents and ruminants are abundant herbivore species in most Arctic ecosystems, many of which are experiencing particularly rapid climate change. Their different life-history characteristics, with the exception of their trophic position, suggest that they should show different responses to environmental variation. Here we show that the only mammalian herbivores on the Arctic islands of Svalbard, reindeer (Rangifer tarandus) and sibling voles (Microtus levis), exhibit strong synchrony in population parameters. This synchrony is due to rain-on-snow events that cause ground ice and demonstrates that climate impacts can be similarly integrated and expressed in species with highly contrasting life histories. The finding suggests that responses of wildlife populations to climate variability and change might be more consistent in Polar regions than elsewhere owing to the strength of the climate impact and the simplicity of the ecosystem.

Bigger teeth for longer life? Longevity and molar height in two roe deer populations

The role of tooth wear as a proximate cause of senescence in ruminants has recently been highlighted. There are two competing hypotheses to explain variation in tooth height and wear; the diet-quality hypothesis predicting increased wear in low-quality habitats, and the life-history hypothesis predicting molar height to be related to expected longevity. We compared tooth height and wear from roe deer of known age from two contrasting populations of roe deer (Capreolus capreolus) in France: Trois Fontaines (TF) with good habitat and shorter animal life expectancy and Chizé (CH) with poor habitat and longer animal life expectancy. There was no population difference in tooth wear, leading to rejection of the diet-quality hypothesis. However, despite their smaller body size, initial molar height for animals from CH was larger than for animals from TF. This provides the first evidence that variation in longevity between populations can lead to differences in molar height within a species.

Negative density-dependent emigration of males in an increasing red deer population

In species with polygynous mating systems, females are regarded as food-limited, while males are limited by access to mates. When local density increases, forage availability declines, while mate access for males may increase due to an increasingly female-biased sex ratio. Density dependence in emigration rates may consequently differ between sexes. Here, we investigate emigration using mark-recovery data from 468 young red deer Cervus elaphus marked in Snillfjord, Norway over a 20-year period when the population size has increased sixfold. We demonstrate a strong negative density-dependent emigration rate in males, while female emigration rates were lower and independent of density. Emigrating males leaving the natal range settled in areas with lower density than expected by chance. Dispersing males moved 42 per cent longer at high density in 1997 (37 km) than at low density in 1977 (26 km), possibly caused by increasing saturation of deer in areas surrounding the marking sites. Our study highlights that pattern of density dependence in dispersal rates may differ markedly between sexes in highly polygynous species. Contrasting patterns reported in small-scale studies are suggestive that spatial scale of density variation may affect the pattern of temporal density dependence in emigration rates and distances.

Spatial patterns of accumulated browsing and its relevance for management of red deer Cervus elaphus

Abstract The substantial increase of deer populations in Europe and North America in recent decades has led to concerns regarding the long-term sustainability of current management from an ecosystem perspective. Key questions to be answered are how herbivore density relates to available resource levels, e.g. food availability and browsing pressure, and how this relation should be monitored and included in management. In Norway, the harvest of red deer Cervus elaphus has increased from 1,479 to 35,700 during 1960-2008. Current monitoring programmes focus on deer body mass and number of seen deer rather than on the state of the vegetation communities. In our study, we quantify browsing frequency on common shrubs and trees in two municipalities, Gloppen and Flora, in the county of Sogn and Fjordane, Norway, to document current browsing pressure levels and to evaluate the potential of using browsing frequency of indicator species as a tool in the monitoring of red deer populations. We found that several species were heavily browsed (median browsing frequency > 60%). Due to their wide availability, we analysed the spatial patterns of accumulated browsing of the highly selected rowan Sorbus aucuparia compared to the less selected bilberry Vaccinium myrtillus and birch Betula sp. We predicted less spatial variation in browsing frequency of the highly selected rowan. However, we found that the best model predicting browsing on rowan (forest type, habitat productivity, canopy cover, aspect, slope and distance to arable land) based on habitat variables was more complicated than for birch (habitat productivity and altitude) and bilberry (forest type, habitat productivity, altitude and distance from the coast). This suggests large spatial variation in browsing frequency of rowan even though the average browsing frequency of rowan was higher than for bilberry and birch. Browsing frequency for all species was positively correlated with faeces counts, but only bilberry showed additional correlation with red deer (harvest) density at the local management unit scale. Due to its wide distribution and promising link to local red deer density, bilberry stands out as the most promising species to monitor among winter browse species. However, browsing frequency on different species was not always correlated. It remains to be determined to which extent browsing on bilberry can be used as an indicator for the browsing pressure on the whole vegetation community. The documentation of high browsing frequencies clearly suggests that further focus on these issues is warranted.

Increased effect of harsh climate in red deer with a poor set of teeth

Teeth are vital for mammal performance and especially in ungulates relying on mechanical decomposition of plant material for effective microbial digestion and energy uptake. The main focus of the role of teeth in ungulate life histories has been on tooth wear, while no one has addressed to what extent deviation from the natural set of teeth (maldentition) causes variation in individual fitness components. Based on mandibles from 41,066 individual red deer (Cervus elaphus L.) collected from 1969 to 2001, we tested whether maldentition had an effect on individual body condition and whether this effect depended on environmental harshness. Females with maldentition (0.6% of the population) were in a poorer condition than individuals without tooth anomalies and the effect increased during unfavorable climatic conditions. The effect of maldentition in males was less clear. This study indicates that a well-functioning set of teeth is essential for mammal performance, and that selection pressure against (dental) anomalies is more pronounced when climate is unfavorable.

The influence of weather conditions during gestation on life histories in a wild Arctic ungulate

The internal predictive adaptive response (internal PAR) hypothesis predicts that individuals born in poor conditions should start to reproduce earlier if they are likely to have reduced performance in later life. However, whether this is the case remains unexplored in wild populations. Here, we use longitudinal data from a long-term study of Svalbard reindeer to examine age-related changes in adult female life-history responses to environmental conditions experienced in utero as indexed by rain-on-snow (ROSutero). We show that females experiencing high ROSutero had reduced reproductive success only from 7 years of age, independent of early reproduction. These individuals were able to maintain the same annual reproductive success between 2 and 6 years as phenotypically superior conspecifics that experienced low ROSutero. Young females born after high ROSutero engage in reproductive events at lower body mass (about 2.5 kg less) than those born after low ROSutero. The mean fitness of females that experienced poor environmental conditions in early life was comparable with that of females exposed to good environmental conditions in early life. These results are consistent with the idea of internal PAR and suggest that the life-history responses to early-life conditions can buffer the delayed effects of weather on population dynamics.

Dental microwear textural analyses to track feeding ecology of reindeer: a comparison of two contrasting populations in Norway

Using dental microwear textural analysis, we aim to explore variations in feeding habits within and between wild reindeer populations (Rangifer tarandus) in Norway. Specimens from the two wild reindeer areas of Knutshø and Hardangervidda were hunted between late August and the end of September 2014, and found to share very similar topological and ecological characteristics (continental alpine tundra). Despite the homogenisation of plant resources at the end of the snow-free period, dental microwear textural analysis on cheek teeth indicates differences in feeding behaviors between the two populations, in that reindeer from Knutshø might display a more “grazing” signal than the ones from Hardangervidda. Such differences in dental microwear textures reflect the differences in feeding habits that could be linked with differences in population density of reindeer. The Knutshø reindeer density is lower than at Hardangervidda, where primary food resources like grasses, sedges, and lichens are less abundant. Analyses also show the differences in gender for the population of Knutshø, but more studies are necessary for the clearest interpretation of data. Our results emphasize the need to explore local variations at a monthly time frame in order to better assess the versatility of dental microwear textures before applying this kind of analysis to either extant or extinct multi-population scales.