Rolf Langvatn

The Response of Terrestrial Ecosystems to Climate Variability Associated with the North Atlantic Oscillation

Climatic factors influence a variety of ecological processes determining patterns of species density and distribution in a wide range of terrestrial ecosystems. We review the effects of the NAO on processes and patterns of terrestrial ecosystems, including both plants and animals. In plants, the NAO index correlates with date of first flowering, tree ring growth and with quality of agricultural crops (wheat and wine grapes). Also, breeding dates are earlier after high NAO index winters for amphibians and birds in Europe. Population dynamical consequences of the NAO have also been reported for birds, and the differential impact of the NAO on two similar species may prevent competitive exclusion. Different effects of the NAO on large herbivore populations have been reported for different regions, depending on limiting factors and the correlation with local weather parameters. The NAO synchronizes population dynamics of lynx and some other carnivore populations in the eastern U.S. Most effects are on an ecological time scale; the evolutionary consequences of long term trends in the NAO are poorly documented. Important for predator and prey dynamics is (1) the disruption of phenology (the match-mismatch hypothesis), (2) that there may be delayed effects (cohort-effects), and (3) that effects of the NAO may interact with other factors such as density. We discuss the challenges related to nonlinearity, of using different climate indices, and how we can progress using these pattern-oriented NAO studies at coarse scales to conduct better process-oriented small-scale experiments.

The role of parasites in the dynamics of a reindeer population

Even though theoretical models show that parasites may regulate host population densities, few empirical studies have given support to this hypothesis. We present experimental and observational evidence for a host–parasite interaction where the parasite has sufficient impact on host population dynamics for regulation to occur. During a six year study of the Svalbard reindeer and its parasitic gastrointestinal nematode Ostertagia gruehneri we found that anthelminthic treatment in April–May increased the probability of a reindeer having a calf in the next year, compared with untreated controls. However, treatment did not influence the over–winter survival of the reindeer. The annual variation in the degree to which parasites depressed fecundity was positively related to the abundance of O. gruehneri infection the previous October, which in turn was related to host density two years earlier. In addition to the treatment effect, there was a strong negative effect of winter precipitation on the probability of female reindeer having a calf. A simple matrix model was parameterized using estimates from our experimental and observational data. This model shows that the parasite–mediated effect on fecundity was sufficient to regulate reindeer densities around observed host densities.

Climate, Plant Phenology and Variation in Age of First Reproduction in a Temperate Herbivore

Density-independent weather effects can have important consequences for the demography of terrestrial herbivores because precipitation, temperature and insolation influence plant phenology, forage quality and biomass production, which in turn affects the habitat carrying capacity. Since forage digestibility influences intake and weight gain, life-history traits of young, growing animals are likely to reflect variation in the prevailing weather. This paper specifically investigates spatial and temporal variation in age at maturation in female red deer (Cervus elaphus) in Norway in relation to climate variables known to influence primary production. Our findings are corroborated by analysing differences in age at maturation in 21 cohorts of red deer on the Isle of Rum, Scotland. In Norway the majority of females ovulated as yearlings and calved for the first time as 2-year-olds. The proportion calving for the first time at two years varied from 0.23 to 0.67 between regions and fluctuated from 0.46 to 0.76 between cohorts. On Rum, where age of maturation was delayed at least a year, the proportion calving for the first time as 3-year-olds varied between cohorts from 0.0 to 0.89. In a subset of yearlings culled in Norway at the time of conception, the spatial and temporal differences in ovulation rates were related to the geographical and annual variation in body weight. 5. Both the spatial and temporal variation in the proportion of 2-year-olds calving in Norway, and cohort differences in the proportion calving as 3-year-olds on Rum, were negatively related to variation in May-June degree days 12 months earlier. Although primary production on the preferred herb-rich Agrostis-Festuca grasslands was positively correlated with temperature in May and June on Rum, the proportion of females calving as three years old, was negatively correlated with annual differences in May June primary production. We argue that retarded phenological development, during periods of cooler weather, enhances diet quality because leaf :stem ratios and digestibility of plant parts decline more slowly. Thus, weight gain during the early summer growth spurt should be rapid during cool May-June weather, increasing the probability of conception in the autumn. Since density-independent variation in food availability also influences fitness components which commonly have a more pronounced influence on population demography, for example offspring survival, we argue that our results highlight the potential importance of variation in weather on herbivore abundance.

Nonlinear effects of large-scale climatic variability on wild and domestic herbivores.

Large-scale climatic fluctuations, such as the North Atlantic Oscillation (NAO), have been shown to affect many ecological processes. Such effects have been typically assumed to be linear. Only one study has reported a nonlinear relation; however, that nonlinear relation was monotonic (that is, no reversal). Here we show that there is a strong nonlinear and non-monotonic (that is, reversed) effect of the NAO on body weight during the subsequent autumn for 23,838 individual wild red deer (Cervus elaphus) and 139,485 individual domestic sheep (Ovis aries) sampled over several decades on the west coast of Norway. These relationships are, at least in part, explained by comparable nonlinear and non-monotonic relations between the NAO and local climatic variables (temperature, precipitation and snow depth). The similar patterns observed for red deer and sheep, the latter of which live indoors during winter and so experience a stable energy supply in winter, suggest that the (winter) climatic variability (for which the index is a proxy) must influence the summer foraging conditions directly or indirectly.

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.

Age- and density-dependent reproductive effort in male red deer.

Reproductive effort in female ungulates originates from gestation and lactation and has been studied extensively; however, no comparable studies of reproductive effort in males (due to fighting for access to mates) have, to our knowledge, previously been reported. Here, we report on weight loss of male red deer during the annual mating season—a direct measure of male reproductive effort (or somatic reproductive costs). The ‘terminal investmen’ hypothesis predicts that reproductive effort should increase with age, given that costs remain stable. We also propose the ‘mating strategy–effort’ hypothesis, which predicts that reproductive effort peaks in prime–aged males, since they are most often the harem holders. Consistent with the mating strategy–effort hypothesis, relative weight loss during the rutting season peaked at prime age and was lower in younger and senescent males. Weight loss during the rut was relatively smaller as density increased and more so for older males. This is probably primarily due to males (particularly senescent males) starting their rut in poorer condition at high density. The pattern of reproductive effort in males with regard to age and density therefore differs markedly from the pattern reported for females.

Trading forage quality for quantity? Plant phenology and patch choice by Svalbard reindeer

Abstract Plant phenology of Luzula heathland plots in Spitsbergen (78°N) was manipulated by adding or removing snow, which altered the time for plots (2 m×2 m; n=10) to become snow-free. A 2-week difference in snowmelt, equivalent to approximately one-sixth of the growing season, was achieved between advanced (first to be snow-free) and delayed (last to be snow-free) treatments, which influenced plant biomass and plant quality. Nitrogen content of the forage species decreased with time after snowmelt, whereas C:N ratio increased. Manipulation of snowmelt led to a shift in ”phenological time”, without altering these plant quality parameters as such. Early in the growing season, Svalbard reindeer (Rangifer tarandus platyrhynchus) selected the advanced plots which had been snow-free for longest, presumably because of the greater biomass of both Luzula confusa and Salix polaris, major components of reindeer diet at that time of the year. Moreover, the proportion of live Luzula leaves was highest in advanced plots, relative to both unmanipulated control and delayed plots. In contrast, plant quality, measured as nitrogen content and C:N ratio of leaves, was lowest in the preferred plots. Phenolic content did not differ among treatments, and is therefore unlikely to play a role in reindeer selection for plots with early snowmelt. Unlike in temperate regions, where selection for plant quality seems to be of major importance, selection for plant quantity might be an outcome of generally low levels of plant biomass and high forage quality during the growing season in the high Arctic. Reindeer selection for high plant biomass is likely to lead to a more favourable nitrogen and energy return than selection for high plant quality.

Importance of climatological downscaling and plant phenology for red deer in heterogeneous landscapes

Understanding how climate influences ecosystems represents a challenge in ecology and natural resource management. Although we know that climate affects plant phenology and herbivore performances at any single site, no study has directly coupled the topography–climate interaction (i.e. the climatological downscaling process) with large-scale vegetation dynamics and animal performances. Here we show how climatic variability (measured by the North Atlantic oscillation ‘NAO’) interacts with local topography in determining the vegetative greenness (as measured by the normalized difference vegetation index ‘NDVI’) and the body masses and seasonal movements of red deer (Cervus elaphus) in Norway. Warm springs induced an earlier onset of vegetation, resulting in earlier migration and higher body masses. Increasing values of the winter-NAO corresponded to less snow at low altitude (warmer, more precipitation results in more rain), but more snow at high altitude (colder, more precipitation corresponds to more snow) relative to winters with low winter-NAO. An increasing NAO thus results in a spatially more variable phenology, offering migrating deer an extended period with access to high-quality forage leading to increased body mass. Our results emphasize the importance of incorporating spring as well as the interaction between winter climate and topography when aiming at understanding how plant and animal respond to climate change.

Effects of age, sex and density on body weight of Norwegian red deer: evidence of density-dependent senescence

There are only a few recent studies that have demonstrated senescence in ungulates and nothing is known regarding how patterns of senescence may vary as a function of density. Senescence in general is linked to the cost of reproduction, which probably increases with density in ungulates and may differ between the sexes. Further, senescence in ungulates is also regarded to be a function of tooth wear rates. As density dependence and sexual differences in food choice have been well documented, this may lead to different tooth wear rates and, thus, possibly density–dependent and sex–specific patterns of senescence. We therefore investigated the effects of age, sex, density and their possible interactions on the variability of body weight in 29 047 red deer harvested during 1965–1998 from Norway, out of which 380 males and 1452 females were eight years or older.There was clear evidence that spatio–temporal variation in density correlated negatively with body weights. In addition, there was evidence of senescence in both male and female red deer. Age at onset of senescence in females was after 20 years of age and independent of population density. In males, the onset and rate of senescence increased with increasing population density. The onset of senescence for males was at ca. 12 years of age at low density, but decreased to approximately ten years of age at high density. The pattern of density–dependent senescence in males, but not that in females, can be explained if (i) the cost of reproduction increases with density more strongly in male than in female red deer, and/or (ii) tooth wear rates are density dependent in males, but not in females. We discuss the ability of these two different, not mutually exclusive hypotheses in explaining the observed pattern of senescence.

Feeding-patch choice by red deer in relation to foraging efficiency

We tested the idea that ruminants allocate their feeding time to habitat patches in relation to foraging efficiency. We used five tame red deer (Cervus elaphus) in an enclosure planted with four treatment of timothy grass (Phleum pratense) differing in their stage of growth. Older swards offered higher biomass but lower nutritional quality than younger swards. We observed time spent feeding in each treatment during each of seven trials. We measured goodness-of-fit between observed times and predictions from two alternative hypotheses differing in optimization strategy (maximizing versus matching), and a third, null hypothesis. We tested the hypotheses using two alternative currecies: digestible protein, and digestible dry matter or energy. Although digestible protein concentration and dry-matter digestibility were highly correlated (r=0.763, P<0.001), the wider range of digestible protein made it the much more sensitive measure of forage quality. Distributions of feeding time closely matched estimated intake rates of digestible protein (RinfPredsup2=0.899) across all animals and trials. The other hypotheses were rejected. The results have important ecological implications in showing the underlying role of food in the selection of habitat by ruminants, and that simple, mechanistic models of forage intake and digestion can be scaled up to the level of animal behavioural choices.