Pikas (Ochotona princeps : Lagomorpha) as allogenic engineers in an alpine ecosystem

Pikas (Ochotona princeps: Lagomorpha) build caches of vegetation (“haypiles”), which serve as a food source during winter in alpine and subalpine habitats. Haypiles appear to degrade over time and form patches of nutrient-rich soils in barren talus and scree areas. We sampled soils underneath and next to haypiles, and plants growing on and near haypiles in an alpine cirque in northwestern Wyoming, USA, to determine the effects of pika food caches on N, C, and C/N ratios in soils and plants. We found that (1) haypile soils had significantly higher carbon and nitrogen levels and lower C/N ratios than both adjacent soils and soils in the general study area, (2) two of three plant species tested (Polemonium viscosum and Oxyria digyna) had significantly higher levels of tissue percent N when growing on haypile soils, and (3) total standing plant biomass at the study site increased with soil percent N suggesting that vegetation was nitrogen limited. Pikas may therefore function as allogenic ecosystem engineers by modulating nutrient availability to plants. Received: 5 July 1997 / Accepted: 30 November 1997     

Herbivore Effects on Ecosystem Process Rates in a Low-Productive System

Ecosystems - Mammalian herbivores shape the structure and function of many nutrient-limited or low-productive terrestrial ecosystems through modification of plant communities and plant–soil...     

Long-term dynamics of voles and lemmings at the timberline and above the willow limit as a test of hypotheses on trophic interactions

We have monitored population fluctuations of microtine rodents since 1977 in two habitat complexes in Finnmark, northernmost Norway - a low arctic plains landscape, with patches of willow scrubland embedded in lichen-dwarf birch tundra, and in adjacent highlands, occupied by scrub-free heaths, snow-beds and bogs. In the plains landscape, voles were cyclic, with a period of five years, and with wave-like density fluctuations. This pattern is consistent with time trajectory of prey in a predator-prey limit cycle. Autoregression analysis implies that the prey pattern is cleanest in the most productive plains habitats, while dynamics in the prevailing heath and bog habitats are governed by two significant lags, implying that even vole-plant interactions count. In the highlands, lemmings had two outbreaks, characterized by J-shaped growth curves, and separated by long periods of low density. The fluctuation pattern of lemmings in highlands was consistent with the predicted time trajectory of a predator. The implications of time trajectories are corroborated by direct evidence on microtine impacts upon the vegetation and on spatial patterns in predator activity. Even the strong dispersal tendency of lemmings during population peaks is consistent with the conjecture that they are adapted to play the role of a predator in a sustained predator-prey cycle. As a whole, the pattern supports T. Oksanen's modification of the hypothesis of exploitation ecosystems, where both local productivity and the structure of the landscape are taken in account.     

The Logic and Realism of the Hypothesis of Exploitation Ecosystems

Hypotheses on trophic dynamics in terrestrial ecosystems fall into two major categories: those in which plants are assumed to be invulnerable to their consumers and those in which the build-up of plant biomass is assumed to require top-down control of folivores. The hypothesis of exploitation ecosystems (EEH) belongs to the latter category and focuses particularly on the consequences of the high energetic costs of maintenance of endotherms. Carnivorous endotherms require relatively high prey densities in order to break even. Moreover, they are dependent on folivorous prey during the limiting season, at least at high latitudes. The endotherm branch of the grazing web is thus predicted to collapse from three-link trophic dynamics (carnivores --> folivores --> plants --> inorganic resources) to two-link dynamics (folivores --> plants --> inorganic resources) along gradients of decreasing primary productivity. Consequently, the vegetation of cold and unproductive areas is predicted to be under intense winter grazing pressure, which prevents the accumulation of aboveground plant biomass and excludes erect woody plants. In the most extreme habitats (e.g., polar deserts and their high alpine counterparts), even folivorous endotherms are predicted to be absent, and the scanty vegetation is predicted to be structured by preemptive competition. Within temperature-determined productivity gradients, EEH is corroborated by biomass patterns, by patterns in the structure and dynamics of carnivore, folivore, and plant communities, and by experimental results. The general idea of top-down trophic dynamics is supported for other autotroph-based systems, too, but the relevance and sufficiency of the energy constraint in explaining patterns in trophic dynamics appears to be variable. Moreover, critical empirical evidence for or against the capacity of folivorous insects to regulate plant biomass has not yet been obtained. Another open question is the ability of boreal and temperate browsers, evolved in productive environments with intense predation pressure and abundance of forage, to prevent the regeneration of the least palatable tree species. There are, thus, many open questions waiting to be answered and many exciting experiments waiting to be conducted.     

Elevated ozone modifies the feeding behaviour of the common leaf weevil on hybrid aspen through shifts in developmental, chemical, and structural properties of leaves

In this study, we tested the impact of moderately elevated ozone (O₃) – 1.5 × ambient, equivalent to predicted near-future ozone concentrations – on the feeding behaviour of the common leaf weevil Phyllobius pyri L. (Coleoptera: Curculionidae), on two hybrid aspen [ Populus tremula  × Populus tremuloides (Salicaceae)] clones (clones 55 and 110) differing in ozone sensitivity using the open-air ozone exposure site in Kuopio, Finland. Three host-selection tests (test between treatments, test between clones, and test between treatments* clones) with common leaf weevil females were carried out in the laboratory in the 2nd year of ozone exposure. The beetles were offered two (four for the tests between treatments and clones) freshly cut leaf discs from first flush leaves. After 24 h, the beetles were removed and the leaf disc area consumed was measured. In the field, the unfolding of the buds was followed and samples were taken for anatomical and chemical (salicylates, condensed tannins, nitrogen, and water content) leaf analyses. Phyllobius pyri significantly preferred leaves from clone 55 to those from clone 110 in the ambient air treatment, whereas this preference was less evident under elevated ozone. Leaves from ozone-exposed trees were significantly preferred to leaves grown in ambient air. Our results suggest that the preference of clone 55 and of ozone-exposed leaves can be explained by phenotypic properties of the plant and prevailing ozone concentration through shifts in leaf development process, phenolic composition, and leaf thickness.     

Optimization of reproductive effort and foraging time in mammals: The influence of resource level and predation risk

Summary The trade-off between fitness benefits from foraging and associated costs in terms of predation risk is analysed by a simple model which takes into account the differential predation risk for reproducing and non-reproducing individuals. The currency that animals are assumed to maximize is their expected absolute fitness (probability of survival plus half of the expected litter size) after a potential reproductive period. Depending on resource levels and predation risk, this maximization can be achieved by (1) opting for individual survival and behaving as a strict time minimizer, (2) by reproducing at the maximal rate and behaving as a strict energy maximizer or (3) by submaximal reproductive effort and a behaviour intermediate between time minimization and energy maximization. Small changes in the availability of food or cover or in the density of predators can shift the optimum from one strategy to another. The shift is particularly abrupt, if predation pressure increases and the availability of resources remains high. This could explain the spatial and temporal variation in the reproductive effort and body weight observed in boreal small mammals with sustained, multiannual population fluctuations.     

Predation on two vole species by a shared predator: antipredatory response and prey preference

In prey communities with shared predators, variation in prey vulnerability is a key factor in shaping community dynamics. Conversely, the hunting efficiency of a predator depends on the prey community structure, preferences of the predator and antipredatory behavioural traits of the prey. We studied experimentally, under seminatural field conditions, the preferences of a predator and the antipredatory responses of prey in a system consisting of two Myodes species of voles, the grey-sided vole (M. rufocanus Sund.) and the bank vole (M. glareolus Schreb.), and their specialist predator, the least weasel (Mustela nivalis nivalis L.). To quantify the preference of the weasels, we developed a new modelling framework that can be used for unbalanced data. The two vole species were hypothesised to have different habitat-dependent vulnerabilities. We created two habitats, open and forest, to provide different escape possibilities for the voles. We found a weak general preference of the weasels for the grey-sided voles over the bank voles, and a somewhat stronger preference specifically in open habitats. The weasels clearly preferred male grey-sided voles over females, whereas in bank voles, there was no difference. The activity of voles changed over time, so that voles increased their movements immediately after weasel introduction, but later adjusted their movements to times of lowered predation risk. Females that were more active had an elevated mortality risk, whereas in the case of males, the result was the opposite. We conclude that, in vulnerability to predation, the species- or habitat-specific characteristics of these prey species are playing a minor role compared to sex-specific characteristics.     

Long distance pollen transport cause problems for determining the timing of birch pollen season in Fennoscandia by using phenological observations

The male flowering and leaf bud burst of birch take place almost simultaneously, suggesting that the observations of leaf bud burst could be used to determine the timing of birch pollen release. However, long‐distance transport of birch pollen before the onset of local flowering may complicate the utilization of phenological observations in pollen forecasting.

We compared the timing of leaf bud burst of silver birch with the timing of the stages of birch pollen season during an eight year period (1997–2004) at five sites in Finland. The stages of the birch pollen season were defined using four different thresholds: 1) the first date of the earliest three‐day period with airborne birch pollen counts exceeding 10 grains m^?3 air; and the dates when the accumulated pollen sum reaches 2) 5%; 3) 50% and 4) 95% of the annual total. Atmospheric modelling was used to determine the source areas for the observed long‐distance transported pollen, and the exploitability of phenological observations in pollen forecasting was evaluated.

Pair‐wise comparisons of means indicate that the timing of leaf bud burst fell closest to the date when the accumulated pollen sum reached 5% of the annual total, and did not differ significantly from it at any site (p<0.05; Student‐Newman‐Keuls test). It was found that the timing of leaf bud burst of silver birch overlaps with the first half of the main birch pollen season. However, phenological observations alone do not suffice to determine the timing of the main birch pollen season because of long‐distance transport of birch pollen.     

Signal reliability compromised by genotype-by-environment interaction and potential mechanisms for its preservation

Sexual selection based on signaling requires that signals used by females in mate choice are reliable indicators of a male's heritable total fitness. A signal and the preference for it are expected to be heritable, resulting in the maintenance of genetic covariance between these two traits. However, a recent article has proposed that signals may quickly become unreliable in the presence of both environmental variation and genotype-by-environment interaction (G x E) with crossing reaction norms, potentially compromising the mechanisms of sexual selection. Here we examine the heritability and plasticity of a male dominance advertisement in the bank vole, Clethrionomys glareolus, in stable and changing rearing environments from father to son. The bank vole is naturally exposed to considerable sources of spatial and temporal environmental variation and male reproductive success is determined by both intra- (male-male competition) and inter- (females prefer to mate with dominant males) sexual selection. Significant G x E for male dominance was found with crossing reaction norms. Plasma testosterone level (T), rather than condition, determined a male's dominance and T also showed a significant G x E. Dominance showed a considerable plasticity across environments, but was only heritable under stable conditions. We document a negative between-environments correlation of male dominance, suggesting that when the environment changes between father and son, the dominance signal is unreliable to females and sexual selection may be compromised. We discuss how G x E and environmental variation interacting with other mechanisms may preserve the reliability of signals and thus the mechanism of sexual selection itself.     

Enhanced leptin sensitivity and attenuation of diet-induced obesity in mice with haploinsufficiency of Socs3

Leptin is an adipocyte-derived hormone that regulates energy balance and neuroendocrine function primarily by acting on specific hypothalamic pathways. Resistance to the weight reducing effects of leptin is a feature of most cases of human and rodent obesity, yet the molecular basis of leptin resistance is poorly understood. We have previously identified suppressor of cytokine signaling-3 (Socs3) as a leptin-induced negative regulator of leptin receptor signaling and potential mediator of leptin resistance. However, due to the non-viability of mice with targeted disruption of Socs3 (ref. 6), the importance of Socs3 in leptin action in vivo was unclear. To determine the functional significance of Socs3 in energy balance in vivo we undertook studies in mice with heterozygous Socs3 deficiency (Socs3(+/-)). We report here that Socs3(+/-) mice display greater leptin sensitivity than wild-type control mice: Socs3(+/-) mice show both enhanced weight loss and increased hypothalamic leptin receptor signaling in response to exogenous leptin administration. Furthermore, Socs3(+/-) mice are significantly protected against the development of diet-induced obesity and associated metabolic complications. The level of Socs3 expression is thus a critical determinant of leptin sensitivity and obesity susceptibility in vivo and this molecule is a potential target for therapeutic intervention.