Covariation between mean vole density and variability drives the numerical response of storks to vole prey

Hušek et al. (Popul Ecol 55:363–375, 2013 ) showed that the numerical response of storks to vole prey was stronger in regions where variability in vole density was higher. This finding is, at first sight, in contradiction with the predictions of life-history theory in stochastic environments. Since the stork productivity-vole density relationship is concave, theory predicts a negative association between the temporal variability in vole density and stork productivity. Here, we illustrate this negative effect of vole variability on stork productivity with a simple mathematical model relating expected stork productivity to vole dynamics. When comparing model simulations to the observed mean density and variability of thirteen Czech and Polish vole populations, we find that the observed positive effect of vole variability on stork numerical response is most likely due to an unusual positive correlation between mean and variability of vole density.     

Responses of a specialist and a generalist seed predator to variation in their common resource

Fluctuations of resources in time and space will influence not only species abundance but also interactions among species. For plant–consumer interactions, the effects of resource variation have mostly been studied in systems with high resource variability. Systems with moderate variations are less studied, although ecological and evolutionary dynamics of plants and consumers are likely to be affected also by less extreme variability. The effects of variation in a particular resource should depend on consumer diet width.
We examined how spatial and temporal variation in seed production in the perennial herb Lathyrus vernus influenced population dynamics and resource utilization in two beetle pre-dispersal seed predators with different host ranges over six years. The monophagous Apion opeticum occupied fewer patches and had lower densities than the oligophagous Bruchus atomarius . The proportion of seeds attacked increased with increases in seed production between years for both seed predators. A possible explanation for these patterns is that population dynamics of beetles are driven largely by local factors and that the same factors influence both beetle performance and seed production. In B. atomarius , patterns may also be influenced by a more pronounced preference for L. vernus in years with a high seed production in L. vernus . We conclude that relatively modest variation in seed production may result in responses that differ in both direction and extent from those usually observed in systems with high variation in seed production.     

Can temporal covariation and autocorrelation in demographic rates affect population dynamics in a raptor species?

Theoretical studies suggest that temporal covariation among and temporal autocorrelation within demographic rates are important features of population dynamics. Yet, empirical studies have rarely focused on temporal covariation and autocorrelation limiting our understanding of these patterns in natural populations. This lack of knowledge restrains our ability to fully understand population dynamics and to make reliable population forecasts. In order to fill this gap, we used a long‐term monitoring (15 years) of a kestrel Falco tinnunculus population to investigate covariation and autocorrelation in survival and reproduction at the population level and their impact on population dynamics. Using Bayesian joint analyses, we found support for positive covariation between survival and reproduction, but weak autocorrelation through time. This positive covariation was stronger in juveniles compared with adults. As expected for a specialized predator, we found that the reproductive performance was strongly related to an index of vole abundance explaining 86% of the temporal variation. This very strong relationship suggests that the temporally variable prey abundance may drive the positive covariation between survival and reproduction in this kestrel population. Simulations suggested that the observed effect size of covariation could be strong enough to affect population dynamics. More generally, positive covariation and autocorrelation have a destabilizing effect increasing substantially the temporal variability of population size.     

Evolution of dispersal in a multi‐trophic community context

Much is known about the evolution of dispersal when species interact with their resources or natural enemies, but very little is known about dispersal evolution when species interact with both resources and natural enemies. Here I investigate how the dispersal of an intermediate consumer evolves in response to its interactions with a basal resource and top predator. I find that dispersal evolution is possible even when the consumer species is not directly affected by environmental variability, but rather experiences the consequences that such variability has on its resource and predator. Spatial variation in the consumer's fitness is driven by spatial heterogeneity in resource productivity, which determines whether a predator can colonize a resource‐consumer community. Temporal variation in the consumer's fitness is driven by random disturbances that cause periodic local extinctions of the predator, followed by recolonizations that lead to transient fluctuations in consumer abundance. When spatial variation in resource productivity is low and the predator can colonize all patches in the landscape, there is no spatial variation in consumer fitness but temporal variation in fitness favors the evolution of a dispersal monomorphism. When spatial variation in resource productivity is high and the predator cannot colonize many patches in the landscape, spatial variation in fitness selects against dispersal. In this case, temporal variation can promote the evolution of a dispersal polymorphism with sedentary and mobile phenotypes, but only for certain types of tri‐trophic interactions. This finding underscores the importance of indirect interactions in shaping the evolution of dispersal. While the ecological community can provide a strong selective environment for the evolution of dispersal, the nature of interactions between trophic levels can also impose constraints on evolution.     

The Temporal Spectrum of Adult Mosquito Population Fluctuations: Conceptual and Modeling Implications

An improved understanding of mosquito population dynamics under natural environmental forcing requires adequate field observations spanning the full range of temporal scales over which mosquito abundance fluctuates in natural conditions. Here we analyze a 9-year daily time series of uninterrupted observations of adult mosquito abundance for multiple mosquito species in North Carolina to identify characteristic scales of temporal variability, the processes generating them, and the representativeness of observations at different sampling resolutions. We focus in particular on Aedes vexans and Culiseta melanura and, using a combination of spectral analysis and modeling, we find significant population fluctuations with characteristic periodicity between 2 days and several years. Population dynamical modelling suggests that the observed fast fluctuations scales (2 days-weeks) are importantly affected by a varying mosquito activity in response to rapid changes in meteorological conditions, a process neglected in most representations of mosquito population dynamics. We further suggest that the range of time scales over which adult mosquito population variability takes place can be divided into three main parts. At small time scales (indicatively 2 days-1 month) observed population fluctuations are mainly driven by behavioral responses to rapid changes in weather conditions. At intermediate scales (1 to several month) environmentally-forced fluctuations in generation times, mortality rates, and density dependence determine the population characteristic response times. At longer scales (annual to multi-annual) mosquito populations follow seasonal and inter-annual environmental changes. We conclude that observations of adult mosquito populations should be based on a sub-weekly sampling frequency and that predictive models of mosquito abundance must include behavioral dynamics to separate the effects of a varying mosquito activity from actual changes in the abundance of the underlying population.     

Vole fluctuations,red fox responses,predation on fawns,and roe deer dynamics in a temperate latitude

The alternative prey hypothesis predicts that predators respond both functionally and numerically (with a time lag) to fluctuations in the main prey abundance, which affects the survival of alternative prey. This pattern was found in northern Europe in the community formed by voles (Microtidae), red foxes (Vulpes vulpes) and roe deer (Capreolus capreolus). We studied the same predator—prey community in a temperate latitude where, according to the predation hypothesis, only the functional response of predators to changes in main prey availability should occur. In the years 1997–2007, in western Poland, we estimated the index of common vole (Microtus arvalis) abundance (burrow counts), the density of foxes (spotlight counts), the young production in foxes (young/adult ratio), the index of fox predation on fawns (prey remains near dens) as well as the reproduction index (fawn/female ratio) and density of roe deer (total counts). The vole abundance fluctuated considerably, the young production in foxes did not correlate with the main prey availability, but the density of foxes showed direct numerical response. The index of fox predation on fawns decreased with the vole abundance and negatively affected the fawn/female ratio in roe deer. Thus, the relationships between voles and foxes were not fully consistent with the predation hypothesis. The direct numerical response of foxes should tend to stabilize this predator—prey community. It is suggested, however, that responses showed by vole-eating predators in temperate latitudes may sometimes affect their alternative prey, including animals with unfavourable conservation status.     

Climate and spatio-temporal variation in the population dynamics of a long distance migrant, the white stork

1. A central question in ecology is to separate the relative contribution of density dependence and stochastic influences to annual fluctuations in population size. Here we estimate the deterministic and stochastic components of the dynamics of different European populations of white stork Ciconia ciconia. We then examined whether annual changes in population size was related to the climate during the breeding period (the 'tap hypothesis' sensu Saether, Sutherland & Engen (2004, Advances in Ecological Research, 35, 185 209) or during the nonbreeding period, especially in the winter areas in Africa (the 'tube hypothesis'). 2. A general characteristic of the population dynamics of this long-distance migrant is small environmental stochasticity and strong density regulation around the carrying capacity with short return times to equilibrium. 3. Annual changes in the size of the eastern European populations were correlated by rainfall in the wintering areas in Africa as well as local weather in the breeding areas just before arrival and in the later part of the breeding season and regional climate variation (North Atlantic Oscillation). This indicates that weather influences the population fluctuations of white storks through losses of sexually mature individuals as well as through an effect on the number of individuals that manages to establish themselves in the breeding population. Thus, both the tap and tube hypothesis explains climate influences on white stork population dynamics. 4. The spatial scale of environmental noise after accounting for the local dynamics was 67 km, suggesting that the strong density dependence reduces the synchronizing effects of climate variation on the population dynamics of white stork. 5. Several climate variables reduced the synchrony of the residual variation in population size after accounting for density dependence and demographic stochasticity, indicating that these climate variables had a synchronizing effect on the population fluctuations. In contrast, other climatic variables acted as desynchronizing agents. 6. Our results illustrate that evaluating the effects of common environmental variables on the spatio-temporal variation in population dynamics require estimates and modelling of their influence on the local dynamics.     

Changes over time in the spatiotemporal dynamics of cyclic populations of field voles (Microtus agrestis L.)

We demonstrate changes over time in the spatial and temporal dynamics of an herbivorous small rodent by analyzing time series of population densities obtained at 21 locations on clear cuts within a coniferous forest in Britain from 1984 to 2004. Changes had taken place in the amplitude, periodicity, and synchrony of cycles and density-dependent feedback on population growth rates. Evidence for the presence of a unidirectional traveling wave in rodent abundance was strong near the beginning of the study but had disappeared near the end. This study provides empirical support for the hypothesis that the temporal (such as delayed density dependence structure) and spatial (such as traveling waves) dynamics of cyclic populations are closely linked. The changes in dynamics were markedly season specific, and changes in overwintering dynamics were most pronounced. Climatic changes, resulting in a less seasonal environment with shorter winters near the end of the study, are likely to have caused the changes in vole dynamics. Similar changes in rodent dynamics and the climate as reported from Fennoscandia indicate the involvement of large-scale climatic variables.     

Barn Owl Productivity Response to Variability of Vole Populations

We studied the response of the barn owl annual productivity to the common vole population numbers and variability to test the effects of environmental stochasticity on their life histories. Current theory predicts that temporal environmental variability can affect long-term nonlinear responses (e.g., production of young) both positively and negatively, depending on the shape of the relationship between the response and environmental variables. At the level of the Czech Republic, we examined the shape of the relationship between the annual sum of fledglings (annual productivity) and vole numbers in both non-detrended and detrended data. At the districts’ level, we explored whether the degree of synchrony (measured by the correlation coefficient) and the strength of the productivity response increase (measured by the regression coefficient) in areas with higher vole population variability measured by the s-index. We found that the owls’ annual productivity increased linearly with vole numbers in the Czech Republic. Furthermore, based on district data, we also found that synchrony between dynamics in owls’ reproductive output and vole numbers increased with vole population variability. However, the strength of the response was not affected by the vole population variability. Additionally, we have shown that detrending remarkably increases the Taylor’s exponent b relating variance to mean in vole time series, thereby reversing the relationship between the coefficient of variation and the mean. This shift was not responsible for the increased synchrony with vole population variability. Instead, we suggest that higher synchrony could result from high food specialization of owls on the common vole in areas with highly fluctuating vole populations.     

Is the heart of Fennoscandian rodent cycle still beating? A 14‐year study of small mammals and Tengmalm's owls in northern Norway

Analyses of spatial and temporal patterns of the small mammal cycle in Fennoscandia have led to two main conclusions: a south-north geographical gradient in the strength of density dependence and period length, and a change in temporal dynamics with less clear periodicity and lower amplitude of fluctuations in recent years. Fourteen years (1985–1998) of small mammals trapping and Tengmalm's owls monitoring data provided clear evidence for 3-yr cycles of two vole species in inner Troms, north Norway, at the same latitude and only 80 km from Kilpisjärvi, north Finland, where 4–5-yr cycles have been the norm until the end of 1980s. The response of Tengmalm's owls to variation in small rodents abundance was not delayed, as was observed in central Finland. Common shrews appeared to have more stable dynamics. The features of the cycle, and a reassessment of the previously described patterns, show that the dynamics may be different over short distances. Furthermore, snow cover and habitat fragmentation should explicitly be taken into account in any understanding of the Fennoscandian gradient, and surrogate variables such as latitude should be avoided. We stress the need for a more extensive coverage of spatial and temporal patterns in cyclicity with respect to assumed ecological causes.     

Age-related environmental sensitivity and weather mediated nestling mortality in white storks Ciconia ciconia

We studied environmental sensitivity and mortality related to weather inclemency in white stork nestlings Ciconia ciconia in their southern European boundary (Doñana, SW Spain). The study of homeothermy acquisition and fault bars (i.e. a measure of stress on feathers) revealed that stork nestlings were specially sensitive to environmental conditions occurring before 20 d of age. Accordingly, most of nestling mortality concentrated during this sensitive period: 91% of deaths corresponded to nestlings younger than 20 d, 73% concentrating on nestlings up to 10 d-old. Nestling mortality and total breeding failure were highly variable among years, being especially high when rainy periods coincided with the early live of nestlings (between 1 April and 15 May). Maximum temperatures had a positive correlation with breeding success and nestling survival but this effect disappeared when controlling for rainfall. Our results are in agreement with previous studies conducted in other white stork populations in other latitudes. We suggest that this could be the result of a low homeothermy capacity of young nestlings jointly with an early breeding phenology that expose white storks to rain, but not to high temperatures. In the context of global climate change we suggest that the current decrease on spring rainfall could increase nestling survival while punctual rainy springs could have a negative effect on the reproduction of white storks.     

Impacts of dispersal on rapid adaptation and dynamic stability of Daphnia in fluctuating environments

Prior ecological research has shown that spatial processes can enhance the temporal stability of populations in fluctuating environments. Less explored is the effect of dispersal on rapid adaptation and its concomitant impact on population dynamics. For asexually reproducing populations, theory predicts that dispersal in fluctuating environments can facilitate asynchrony among clones and enhance stability by reducing temporal variability of total population abundance. This effect is predicted when clones exhibit heritable variation in environmental optima and when fluctuations occur asynchronously among patches. We tested this in the field using artificial ponds and metapopulations composed of a diverse assemblage of Daphnia pulex clones. We directly manipulated dispersal presence/absence and environmental fluctuations in the form of nutrient pulses. Consistent with predictions, dispersal enhanced temporal asynchrony among clones in the presence of nutrient pulses; this in turn stabilized population dynamics. This effect only emerged when patches experienced spatially asynchronous nutrient pulses (dispersal had no effect when patches were synchronously pulsed). Clonal asynchrony was driven by strong positive selection for a single clone that exhibited a performance advantage under conditions of low resource availability. Our work highlights the importance of dispersal as a driver of eco-evolutionary dynamics and population stability in variable environments.     

Population dynamics in a cyclic environment: consequences of cyclic food abundance on tawny owl reproduction and survival

1. Understanding which factors regulate population dynamics may help us to understand how a population would respond to environmental change, and why some populations are declining.
2. In southern Finland, vole abundance shows a three-phased cycle of low, increase and decrease phases, but these have been fading out in recent years. During five such cycles (1981–1995), all tawny owls Strix aluco were censused in a 250-km² study area, and their reproduction and survival were monitored.
3. Males and females showed similar dynamics, but experienced breeders recruited more offspring and had higher survival than first breeders. Offspring recruitment, but not survival of breeding individuals varied in accordance with vole abundance.
4. The population's numerical response to prey abundance was primarily due to first-breeding individuals entering the population in the increase phase when immigration was the highest. First-breeding birds were younger, but experienced breeders were older in more favourable vole years.
5. A stage-specific matrix population model integrating survival and fecundity showed that, despite obvious variation in fecundity between vole cycle phases, this variation had limited importance for overall tawny owl population dynamics, but that the survival of experienced breeders during the low phase is most important for population growth.
6. Model and data agreed that the vole cycle drives the dynamics of this avian predator by limiting the recruitment of new breeders during the low phase. Population dynamics hence differ not only from the classic example of the species in a more temperate region in the UK where the number of territories is stable across years, but also from the dynamics of other avian vole predators in Fennoscandia where the recurring crash in vole abundance drastically lowers adult survival thereby creating vacancies.     

The influence of spatio-temporal resource fluctuations on insular rat population dynamics

Local spatio-temporal resource variations can strongly influence the population dynamics of small mammals. This is particularly true on islands which are bottom-up driven systems, lacking higher order predators and with high variability in resource subsidies. The influence of resource fluctuations on animal survival may be mediated by individual movement among habitat patches, but simultaneously analysing survival, resource availability and habitat selection requires sophisticated analytical methods. We use a Bayesian multi-state capture-recapture model to estimate survival and movement probabilities of non-native black rats (Rattus rattus) across three habitats seasonally varying in resource availability. We find that survival varies most strongly with temporal rainfall patterns, overwhelming minor spatial variation among habitats. Surprisingly for a generalist forager, movement between habitats was rare, suggesting individuals do not opportunistically respond to spatial resource subsidy variations. Climate is probably the main driver of rodent population dynamics on islands, and even substantial habitat and seasonal spatial subsidies are overwhelmed in magnitude by predictable annual patterns in resource pulses. Marked variation in survival and capture has important implications for the timing of rat control.     

Interannual bumble bee abundance is driven by indirect climate effects on floral resource phenology

Climate change can influence consumer populations both directly, by affecting survival and reproduction, and indirectly, by altering resources. However, little is known about the relative importance of direct and indirect effects, particularly for species important to ecosystem functioning, like pollinators. We used structural equation modelling to test the importance of direct and indirect (via floral resources) climate effects on the interannual abundance of three subalpine bumble bee species. In addition, we used long‐term data to examine how climate and floral resources have changed over time. Over 8 years, bee abundances were driven primarily by the indirect effects of climate on the temporal distribution of floral resources. Over 43 years, aspects of floral phenology changed in ways that indicate species‐specific effects on bees. Our study suggests that climate‐driven alterations in floral resource phenology can play a critical role in governing bee population responses to global change.     

Temporal resource variability and the habitat-matching rule

Summary The ideal free distribution of competitors in a heterogeneous environment often predicts habitat matching, where the equilibrium number of consumers in a patch is proportional to resource abundance in that patch. We model the interaction between habitat matching and temporal variation in resource abundance. In one patch the rate of resource input follows a Markov chain; a second patch does not vary temporally. We predict patch use by scaling transition rates in the variable patch to the time that consumers require to respond to changes in rates of resource input. If consumers respond very quickly, habitat matching tracks temporal variability. If resource input fluctuates faster than consumers respond, habitat matching averages over the equilibrium of the Markov chain. Tracking and averaging produce the same mean resource consumption for individuals, but long-term mean occupation of the patches differs. When habitat matching tracks temporal variability in resources, consumer density in the variable patch has a lower mean and a higher variance than when habitat matching reflects only average rates of resource input.We tested our model by feeding free-living mallard ducks (Anas platyrynchos) at two artificial patches. The foragers' behavior satisfied the quantitative predictions of the model in each of two experiments.     

Resource-dependent reproductive adjustment and the stability of consumer-resource dynamics

This study explored a consumer-resource model including reproductive and nonreproductive subpopulations of the consumer to consider whether resource-dependent reproductive adjustment by the consumer would stabilize consumer-resource dynamics. The model assumed that decreasing (increasing) resource availability caused reproductive suppression (facilitation), and that the reproductive consumer had a higher mortality rate than the nonreproductive one (i.e., a trade-off between reproduction and survival). The model predicted that the variability would be reduced when the consumer had a strong tendency to suppress reproduction in response to low resource availability or when the cost of reproduction was high, although consumer extinction became more likely. Furthermore, when the consumer-resource dynamics converged to limit cycles, reproductive adjustment enhanced the long-term average of the consumer density. It was also predicted that if reproductive suppression enhanced resource consumption efficiency (i.e., a trade-off between reproduction and foraging), then it would destabilize the system by canceling the stabilizing effect of the reproductive adjustment itself. These results suggest that it is necessary not only to identify the costs of reproduction, but also to quantify the changes in individual-level performances due to reproduction in order to understand the ecological consequences of reproductive adjustment.     

Survival and Population Dynamics of the Marabou Stork in an Isolated Population,Swaziland

Investigating the ecology of long lived birds is particularly challenging owing to the time scales involved. Here an analysis is presented of a long term study of the survival and population dynamics of the marabou stork (Leptoptilos crumeniferus), a wide ranging scavenging bird from Sub-Saharan Africa. Using resightings data of tagged nestlings and free flying birds we show that the stork population can be divided into three general life stages with unique survival probabilities and fecundities. Fecundity of the storks is inversely related to rainfall during their breeding season. Corroborative evidence for a metapopulation structure is discussed highlighting the impact of the Swaziland birds on the ecology of the species in the broader region. The importance of tag loss or illegibility over time is highlighted. Clearly, any attempt at conserving a species will require a detailed understanding of its population structure, of the sort examined here.     

The numerical response: rate of increase and food limitation in herbivores and predators

Two types of numerical response function have evolved since Solomon first introduced the term to generalize features of Lotka-Volterra predator-prey models: (i) the demographic numerical response, which links change in consumer demographic rates to food availability; and (ii) the isocline numerical response, which links consumer abundance per se to food availability. These numerical responses are interchangeable because both recognize negative feedback loops between consumer and food abundance resulting in population regulation. We review how demographic and isocline numerical responses have been used to enhance our understanding of population regulation of kangaroos and possums, and argue that their utility may be increased by explicitly accounting for non-equilibrium dynamics (due to environmental variability and/or biological interactions) and the existence of multiple limiting factors. Interferential numerical response functions may help bridge three major historical dichotomies in population ecology (equilibrium versus non-equilibrium dynamics, extrinsic versus intrinsic regulation and demographic versus isocline numerical responses).