Comparison of budburst phenology trends and precision among participants in a citizen science program.

Quantifying shifts in plant phenology in response to climate change represents an ongoing challenge, particularly in mountain ecosystems. Because climate change and phenological responses vary in space and time, we need long-term observations collected at a broad spatial scale. While data collection by volunteers is a promising approach to achieve this goal, one major concern with citizen science programs is the quality and reliability of data. Using a citizen science program (Phenoclim) carried out in the western European Alps, the goals of this study were to analyze (1) factors influencing participant retention rates, (2) the efficacy of a citizen science program for detecting temporal changes in the phenology of mountain trees, (3) differences in budburst date trends among different observer categories, and (4) the precision of trends quantified by different categories of participants. We used 12 years of annual tree phenology measurements recorded by volunteers (schools and private individuals) and professionals within the Phenoclim program. We found decadal-scale shifts in budburst date consistent with the results from other studies, including significant advances in budburst date for the common birch and European ash (- 4.0 and - 6.5 days per decade respectively). In addition, for three of six species, volunteers and professionals detected consistent directional trends. Finally, we show how differences in precision among the categories of participants are determined by the number of years of participation in the program, the number of sites surveyed, and the variability in trends among sites. Overall, our results suggest that participants with a wide range of backgrounds are capable of collecting data that can significantly contribute to the study of the impacts of climate change on mountain plant phenology.

When relative allocation depends on total resource acquisition: implication for the analysis of trade-offs.

A central tenet of evolutionary biology states that life-history traits are linked via trade-offs, as classically exemplified by the van Noordwijk and de Jong model. This model, however, assumes that the relative resource allocation to a biological function varies independently of the total resource acquisition. Based on current empirical evidence, we first explored the dependency between the total resource acquisition and the relative resource allocation to reproduction and showed that such dependency is the rule rather than the exception. We then derived the expression of the covariance between traits when the assumption of independence is relaxed and used simulations to quantify the importance of such dependency on the detection of trade-offs between current reproduction and future survival. We found that the dependency between the total energy acquisition and the relative allocation to reproduction can influence the probability to detect trade-offs between survival and reproduction. As a general rule, a negative dependency between the total energy acquisition and the relative allocation to reproduction should lead to a higher probability of detecting a trade-off in species with a fast pace of life, whereas a positive dependency should lead to a higher probability of detecting a trade-off in species with a slow pace of life. In addition to confirming the importance of resource variation to reveal trade-offs, our finding demonstrates that the covariance between resource allocation and resource acquisition is generally not null and also plays a fundamental role in the detection of trade-offs.

DNA from soil mirrors plant taxonomic and growth form diversity.

Ecosystems across the globe are threatened by climate change and human activities. New rapid survey approaches for monitoring biodiversity would greatly advance assessment and understanding of these threats. Taking advantage of next-generation DNA sequencing, we tested an approach we call metabarcoding: high-throughput and simultaneous taxa identification based on a very short (usually <100 base pairs) but informative DNA fragment. Short DNA fragments allow the use of degraded DNA from environmental samples. All analyses included amplification using plant-specific versatile primers, sequencing and estimation of taxonomic diversity. We tested in three steps whether degraded DNA from dead material in soil has the potential of efficiently assessing biodiversity in different biomes. First, soil DNA from eight boreal plant communities located in two different vegetation types (meadow and heath) was amplified. Plant diversity detected from boreal soil was highly consistent with plant taxonomic and growth form diversity estimated from conventional above-ground surveys. Second, we assessed DNA persistence using samples from formerly cultivated soils in temperate environments. We found that the number of crop DNA sequences retrieved strongly varied with years since last cultivation, and crop sequences were absent from nearby, uncultivated plots. Third, we assessed the universal applicability of DNA metabarcoding using soil samples from tropical environments: a large proportion of species and families from the study site were efficiently recovered. The results open unprecedented opportunities for large-scale DNA-based biodiversity studies across a range of taxonomic groups using standardized metabarcoding approaches.

Echinococcus multilocularis in Svalbard, Norway: microsatellite genotyping to investigate the origin of a highly focal contamination.

Echinococcus multilocularis is a threatening cestode involved in the human alveolar echinococcosis. The parasite, mainly described in temperate regions of the Northern hemisphere was described for the first time in 1999 in the High Arctic Svalbard archipelago, Norway. The origin of this contamination could be due to an anthropogenic introduction from mainland Europe by domestic dogs or with the introduction of the sibling vole, perhaps from mainland Russia (St. Petersburg area), or with roaming Arctic foxes, known as the main definitive host of the parasite in Arctic regions. The genetic diversity of E. multilocularis in Svalbard was investigated here for the first time by genotyping using EmsB microsatellite and compared to other genotyped populations in the main worldwide endemic areas. We found low polymorphism amongst the 27 metacestode isolates from sibling voles trapped in the core of the distribution area of the vole on Svalbard. E. mutilocularis Arctic populations, using the Arctic fox as the definitive host, were genetically separated from European temperate populations that use the red fox, but closely related to St. Lawrence Island samples from Alaska. The result is inconsistent with the hypothesis of an anthropogenic introduction from mainland Europe, but can be seen as consistent with the hypothesis that Arctic foxes introduced E. multilocularis to Svalbard.

Intestinal parasites of the Arctic fox in relation to the abundance and distribution of intermediate hosts.

The intestinal parasite community of Arctic foxes (Vulpes lagopus) on the Svalbard archipelago in the High Arctic was investigated in relation to the abundance and distribution of intermediate hosts. Five species of cestodes (Echinococcus multilocularis, Taenia crassiceps, Taenia polyacantha, Taenia krabbei and Diphyllobothrium sp.), ascaridoid nematodes and one unidentified acanthocephalan species were found. The cestodes E. multilocularis, T. crassiceps and T. polyacantha all showed a decreasing prevalence in the fox population with increasing distance from their spatially restricted intermediate host population of sibling voles (Microtus levis). In addition, the prevalence of E. multilocularis in a sample from the vole population was directly related to the local vole abundance. The cestode T. krabbei uses reindeer as intermediate host, and its prevalence in female foxes was positively related to the density of reindeer (Rangifer tarandus platyrhyncus). Finally, the prevalence of the ascaridoid nematodes also decreased with increasing distance from the vole population, a finding that is consistent with the idea that voles are involved in transmission, most likely as paratenic hosts. The prevalence of the remaining species (Diphyllobothrium sp. and an unidentified acanthocephalan) was very low. We conclude that the distribution and abundance of intermediate host structure the gastrointestinal parasite community of the Arctic fox on the Svalbard archipelago.

Weak compensation of harvest despite strong density-dependent growth in willow ptarmigan.

Ptarmigan and grouse species (Lagopus spp.) are thought to be able to compensate for a modest harvest because there is a surplus of breeding birds that are prevented from breeding by territory holders. To estimate the degree of harvest-mortality compensation reliably we experimentally harvested 0%, 15% and 30% of the willow ptarmigan (Lagopus lagopus) on 13 estates ranging from 20 to 54 km2 in size during four hunting seasons in Norway according to a regional block design. Population overwinter growth rate was strongly negatively density dependent, but despite this, and contrary to earlier findings, only 33% of the harvest was compensated for. The lack of compensation was probably caused by long-distance juvenile dispersal that was unaffected by the harvest. The need for large-scale management experiments to detect the effects of harvest was clearly demonstrated: lack of compensation was found only when we used the whole dataset and not when the data were analysed by year or block. Our study shows that it is very difficult to demonstrate a population's lack of harvest compensation and warns against using small-scale, out-of-season or poorly replicated studies as a basis for future harvest-management decisions.

Consequences of inter-population crosses on developmental stability and canalization of floral traits in Dalechampia scandens (Euphorbiaceae).

Congruence between changes in phenotypic variance and developmental noise in inter-population hybrids was analysed to test whether environmental canalization and developmental stability were controlled by common genetic mechanisms. Developmental stability assessed by the level of fluctuating asymmetry (FA), and canalization by the within- and among-individual variance, were measured on several floral traits of Dalechampia scandens (Euphorbiaceae). Hybridization affected canalization. Both within- and among-individual phenotypic variance decreased in hybrids from populations of intermediate genetic distance, and strongly increased in hybrids from genetically distant populations. Mean-trait FA differed among cross-types, but hybrids were not consistently more or less asymmetric than parental lines across traits. We found no congruence between changes in FA and changes in phenotypic variance. These results suggest that developmental stability (measured by FA) and canalization are independently controlled. This study also confirms the weak relationship between FA and the breakdown of coadapted gene complexes following inter-population hybridization.

Demography and population dynamics of the mouse opossum (Thylamys elegans) in semi-arid Chile: seasonality, feedback structure and climate.

Here we present, to the authors' knowledge for the very first time for a small marsupial, a thorough analysis of the demography and population dynamics of the mouse opossum (Thylamys elegans) in western South America. We test the relative importance of feedback structure and climatic factors (rainfall and the Southern Oscillation Index) in explaining the temporal variation in the demography of the mouse opossum. The demographic information was incorporated into a stage-structured population dynamics model and the model's predictions were compared with observed patterns. The mouse opossum's capture rates showed seasonal (within-year) and between-year variability, with individuals having higher capture rates during late summer and autumn and lower capture rates during winter and spring. There was also a strong between-year effect on capture probabilities. The reproductive (the fraction of reproductively active individuals) and recruitment rates showed a clear seasonal and a between-year pattern of variation with the peak of reproductive activity occuring during winter and early spring. In addition, the fraction of reproductive individuals was positively related to annual rainfall, while population density and annual rainfall positively influenced the recruitment rate. The survival rates were negatively related to annual rainfall. The average finite population growth rate during the study period was estimated to be 1.011 +/- 0.0019 from capture-recapture estimates. While the annual growth rate estimated from the seasonal linear matrix models was 1.026, the subadult and adult survival and maturation rates represent between 54% (winter) and 81% (summer) of the impact on the annual growth rate.

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.

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.

Echinococcus multilocularis on Svalbard: introduction of an intermediate host has enabled the local life-cycle.

The taeniid tapeworm Echinococcus multilocularis is here reported for the first time at the Svalbard Archipelago in the Norwegian Arctic. This new finding is interesting because the establishment of E. multilocularis is due to a recent anthropogenic introduction of its intermediate host--the sibling vole Microtus rossiaemeridionalis at Svalbard. The parasite itself has probably become naturally transferred to Svalbard due to migratory movements of its final host--the arctic fox Alopex lagopus between source areas for E. multilocularis in Siberia and Svalbard. We report macroscopically determined prevalence of E. multilocularis from a sample of 224 voles trapped in August in 1999 and 2000. The prevalence was among the highest ever recorded in intermediate hosts and was dependent on age and sex of the hosts approaching 100% in overwintered males. The high prevalence and the simplicity of the vole-arctic fox-E. multilocularis system at Svalbard makes it an eminent model system for further epidemiological studies.

The effects of spatial habitat configuration on recruitment, growth and population structure in arctic Collembola.

The population density and demography of five species of arctic Collembola were studied in a naturally patchy habitat, consisting of Carex ursinae tussocks with varying degrees of isolation. Focal predictor variables were those describing the spatial configuration of tussocks, including tussock size and isolation and the amount of habitat (cover) at a 1-m2 scale surrounding each tussock population. The Collembola populations were heavily influenced by environmental stochasticity in the form of winter mortality and summer drought, and the influence of patchiness on population characteristics was evaluated in this context. The five species showed very different responses to the structuring effect of the habitat, depending on life history characteristics, mobility and habitat requirements. Population density was highly variable in both time and space. Spring densities indicated larger winter mortality compared to observations from a previous study, and the snow- and ice-free season from June to August only resulted in population growth for Folsomia sexoculata. In the other species, adult mortality must have been high as there was no net population growth despite observed reproduction. The exception was Hypogastrura viatica, whose population decline was more likely to have been the result of migration out of the study area. Cover was the most important variable explaining density. No pure area or isolation effects at the tussock level were detected, even in areas with very low habitat cover. Drought was probably an important mortality factor, as July was particularly warm and dry. Due to qualitative differences in the tussocks and the matrix substrate, desiccation risk would be higher during dispersal between tussocks. We suggest that increased dispersal mortality gave the observed pattern of increased density in relation to cover, both in general and in F. quadrioculata, an opportunistic species otherwise known for rapid population growth. Onychiurus groenlandicus, which had a similar density response to cover, may also be influenced by a rescue effect sustaining densities in areas with high cover. The cover effect can be viewed as a large-scale factor which encompasses the general spatial neighbourhood of each tussock, where inter-population processes are important, as opposed to internal patch dynamics.

From patterns to processes: phase and density dependencies in the Canadian lynx cycle.

Across the boreal forest of North America, lynx populations undergo 10-year cycles. Analysis of 21 time series from 1821 to the present demonstrates that these fluctuations are generated by nonlinear processes with regulatory delays. Trophic interactions between lynx and hares cause delayed density-dependent regulation of lynx population growth. The nonlinearity, in contrast, appears to arise from phase dependencies in hunting success by lynx through the cycle. Using a combined approach of empirical, statistical, and mathematical modeling, we highlight how shifts in trophic interactions between the lynx and the hare generate the nonlinear process primarily by shifting functional response curves during the increase and the decrease phases.

At-risk individuals in Feline Immunodeficiency Virus epidemiology: evidence from a multivariate approach in a natural population of domestic cats (Felis catus).

Prevalence of Feline Immunodeficiency Virus (FIV) infection was measured during 6 consecutive years in a natural rural population of domestic cats. Sex, age, weight, origin, group size and presence of antibodies to FIV were recorded for each sampled cat. Logistic regressions were used to estimate the influence of the recorded parameters on infection. FIV prevalence rates are as high as 19.6% in the total population, and do not statistically change between years, after controlling for changes in samples' age structure. FIV infection is characterized by risk factors linked to aggressive behaviour: old mature male adults having dispersed are more likely to be infected. A study of the cats group size and of the spatial distribution of infected individuals indicates the absence of infection clusters in males, and suggests the importance of roaming in the spreading of FIV. In conclusion, FIV infection spreads, with low contagiousness, mainly between particularly aggressive individuals, and the virus is endemic in this population.

Retroviruses and sexual size dimorphism in domestic cats (Felis catus L.).

Hochberg and co-workers have predicted that an increase in host adult mortality due to parasites is balanced by an earlier age at first reproduction. In polygynous species we hypothesize that such a pattern would lead to diverging selection pressure on body size between sexes and increased sexual size dimorphism. In polygynous mammals, male body size is considered to be an important factor for reproductive success. Thus, under the pressure of a virulent infection, males should be selected for rapid growth and/or higher body size to be able to compete successfully as soon as possible with opponents. In contrast, under the same selection pressure, females should be selected for lighter adult body size or rapid growth to reach sexual maturity earlier. We investigated this hypothesis in the domestic cat Felis catus. Orange cats have greater body size dimorphism than non-orange cats. Orange females are lighter than non-orange females, and orange males are heavier than non-orange males. Here, we report the extent to which orange and non-orange individuals differ in infection prevelance for two retroviruses, feline immunodeficiency virus (FIV) and feline leukaemia virus (FeLV). FIV is thought to be transmitted almost exclusively through aggressive contacts between individuals, whereas FeLV transmission occurs mainly through social contacts. The pattern of infection of both diseases is consistent with the higher aggressiveness of orange cats. In both sexes, orange cats are significantly more infected by FIV, and tend to be less infected by FeLV than other cats. The pattern of infection is also consistent with an earlier age at first reproduction in orange than in non-orange cats, at least for females. These results suggest that microparasitism may have played an important role in the evolution of sexual size dimorphism of domestic cats.

Population dynamics of large herbivores: variable recruitment with constant adult survival.

Recent long-term studies of population ecology of large herbivorous mammals suggest that survival of prime-aged females varies little from year to year and across populations. Juvenile survival, on the other hand, varies considerably from year to year. The pattern of high and stable adult survival and variable juvenile survival is observed in contrasting environments, independently of the main proximal sources of mortality and regardless of whether mortality is stochastic or density-dependent. High yearly variability in juvenile survival may play a predominant role in population dynamics.

Optimal foraging: the importance of environmental stochasticity and accuracy in parameter estimation.

The Engen and Stenseth theorem is applied to a situation where the expected energy content and handling time of encountered food items are estimated with some stochastic error. We extended the Engen-Stenseth approach by incorporating in the model an explicit relationship between what the forager actually observes and what the situation actually is like. By such a formulation, we are able to evaluate the changes in the optimal foraging tactic and the cost of incomplete information; this cost is defined as the reduction in the optimal reward rate that results from the incomplete knowledge about the environmental situation. The approach presented in this article is general and makes it possible to study the problem of incomplete information in a variety of situations. In this article, however, we give some numerical examples that provide insights into the effect of incomplete information. For instance, we demonstrate that the optimal reward rate may attain a minimum for intermediate values of environmental uncertainty. However, since the location of this minimum depends on other parameters of the model, an increase in environmental uncertainty cannot in general be concluded to be favorable to the forager. In order to draw such conclusions, a thorough knowledge of both the environmental features entering the calculation of the optimal reward rate and also the intrinsic properties of the food items (measured by the energy content and the handling time) is necessary. We suggest how results like those derived in this article can be evaluated experimentally.