Habitat selection and population regulation in temporally fluctuating environments

Understanding and predicting the distribution of organisms in heterogeneous environments lies at the heart of ecology, and the theory of density-dependent habitat selection (DDHS) provides ecologists with an inferential framework linking evolution and population dynamics. Current theory does not allow for temporal variation in habitat quality, a serious limitation when confronted with real ecological systems. We develop both a stochastic equivalent of the ideal free distribution to study how spatial patterns of habitat use depend on the magnitude and spatial correlation of environmental stochasticity and also a stochastic habitat selection rule. The emerging patterns are confronted with deterministic predictions based on isodar analysis, an established empirical approach to the analysis of habitat selection patterns. Our simulations highlight some consistent patterns of habitat use, indicating that it is possible to make inferences about the habitat selection process based on observed patterns of habitat use. However, isodar analysis gives results that are contingent on the magnitude and spatial correlation of environmental stochasticity. Hence, DDHS is better revealed by a measure of habitat selectivity than by empirical isodars. The detection of DDHS is but a small component of isodar theory, which remains an important conceptual framework for linking evolutionary strategies in behavior and population dynamics.     

A simple model for diffusion in independent, temporally fluctuating pores

A simple model is presented for one-dimensional diffusion in an ensemble of semi-infinite and finite pores or capillaries in which the boundary at one end of each capillary is allowed to fluctuate randomly between a perfectly reflecting barrier and a perfectly absorbing barrier. The model is independent of the spatial distribution of the capillaries; it is only assumed that there are a large number of them and that they are noninteracting. Exact solutions are possible and results are obtained, in terms of the fluctuation parameters, for the total amount per unit area of solute passed through the capillary system in the semi-infinite case, and for a permeability coefficient and time lag to steady state in the finite system. Applications of the model to diffusion in biological membranes are discussed.     

Bacterial exopolymer and PHB production in fluctuating ground-water habitats

Abstract The physiological response of a strain of ground-water bacteria to short-term fluctuations in the quality of percolating water was followed in a continuous-flow water-saturated soil column. The cells reduced their production of uronic acids-characterized extracellular polysaccharides by more than 50% when pure ground water was enriched with phosphate and glucose but resumed it gradually as the poor growth conditions returned. Bacteria that were sorbed to the mineral particles in the soil column produced more exopolymers-about six times more in pure ground water-than free-living bacteria, but the composition of the polymer was the same. Glucose, mannose and galacturonic acid accounted for about 90% of the carbohydrate. The cells stored about five times more poly-β-hydroxybutyric acid (PHB) in pure than in enriched ground water, and sorbed cells had five times higher concentrations than free-living cells. The quality variations of the ground water were also reflected in the population size of bacteria, which differed by two to four orders of magnitude between poor and enriched conditions.     

An experimental evolutionary study on adaptation to temporally fluctuating pH in Escherichia coli

In this study, we use the bacterium Escherichia coli to examine evolutionary responses to environmental acidity fluctuating temporally among pH 5.3, 6.3, 7.0, and 7.8 (5,000-15 nM [H(+)]). Two experimental protocols of temporal variation were used. One group (six replicate lines) of populations evolved for 2,000 generations during exposure to a cycled regime fluctuating daily between pH 5.3 and 7.8. The other group (also in six replicate lines) evolved during exposure for 2,000 generations to a randomly shifting regime fluctuating stochastically each day among pH 5.3, 6.3, 7.0, and 7.8. Adaptation to these fluctuating acidity regimes was measured as a change in fitness relative to the common ancestor by direct competition experiments in both constant and fluctuating pH regimes. For comparisons with constant pH evolution, a group evolved at a constant pH of 5.3 and another group evolved at pH 7.8 were also tested. This study initiated the first long-term laboratory natural selection experiment on adaptation to variable acidity and addressed key questions concerning patterns of adaptation (trade-offs, specialists, generalists, plasticity, transitions, and acclimation) in temporally fluctuating environments.     

Collapsing population cycles

During the past two decades population cycles in voles, grouse and insects have been fading out in Europe. Here, we discuss the cause and implication of these changes. Several lines of evidence now point to climate forcing as the general underlying cause. However, how climate interacts with demography to induce regime shifts in population dynamics is likely to differ among species and ecosystems. Herbivores with high-amplitude population cycles, such as voles, lemmings, snowshoe hares and forest Lepidoptera, form the heart of terrestrial food web dynamics. Thus, collapses of these cycles are also expected to imply collapses of important ecosystem functions, such as the pulsed flows of resources and disturbances.     

Breeding dispersal of Eurasian kestrels Falco tinnunculus under temporally fluctuating food abundance

Costs and benefits of dispersal can vary in space and time, depending on environmental factors and individual state. Plastic, condition‐dependent dispersal strategies, in which individuals rely on external cues such as food abundance to adjust their dispersal distances, are therefore expected to evolve in temporally fluctuating environments. We examined factors affecting breeding dispersal distances in Eurasian kestrels Falco tinnunculus subsisting on multi‐annually and cyclically fluctuating voles as their main food. We attempted to avoid traditional bias in dispersal studies by having large study areas and by taking detection probabilities into account. We observed 320 dispersal events of male and 215 events of female kestrels from our study areas in western Finland during 24 yr. After correcting for distance‐specific detection probability, the estimates of mean dispersal distances increased two‐fold being still clearly higher for females than males. Vole abundance in the spring of settlement was more important in determining average dispersal distances than vole abundance in the previous autumn. At the population level (cross‐sectional model), both males and females dispersed longer distances when the spring abundance of their main food (voles) was low compared to when it was abundant, as predicted by the food depletion hypothesis. At the individual level (longitudinal model), only females responded to the food situation by dispersing more when food abundance was low in the spring of settlement. Females also dispersed longer when vole abundance in the previous autumn had been high. Individual males did not respond to vole abundance, which implies that the population level response in males might have been caused by long‐distance dispersers, which breed in the study area only in good vole years. Our results show that the dispersal distances of kestrels at northern latitudes depend both on individual properties (gender, age, and possibly individual tendency to disperse) and environmental conditions (temporal variation in main food abundance).     

Phase switching in population cycles

Oscillatory populations may exhibit a phase change in which, for example, a high–low periodic pattern switches to a low–high pattern. We propose that phase shifts correspond to stochastic jumps between basins of attraction in an appropriate phase space which associates the different phases of a periodic cycle with distinct attractors. This mechanism accounts for two-cycle phase shifts and the occurrence of asynchronous replicates in experimental cultures of Tribolium.     

Site fidelity in temporally correlated environments enhances population persistence

Site fidelity, the phenomenon of remaining faithful to sites, often where an individual has bred successfully in the past, has important consequences for population dynamics. Previous results have shown that site fidelity results in a positive correlation between population density and fitness. Here, I build on this theme by incorporating site fidelity using the win‐stay : lose‐switch rule often seen among birds, i.e. individuals return to sites were they bred successfully in the past and vacate those where they have not. Results demonstrate that the combination of site fidelity and temporal autocorrelation in site quality can enhance the persistence of population networks, whereas either factor acting alone has little or no influence. Moreover, there is an abrupt threshold at moderate levels of temporal autocorrelation, ρ_time > 0.35–0.4, beyond which persistence time and the probability of surviving >500 years is greatly accelerated. These results suggest that temporal autocorrelation combined with appropriate behavioural responses may enhance population persistence.     

Small-scale spatial dynamics in a fluctuating ungulate population

1. The scale at which population dynamics are analysed is important, as results from analyses at different spatial scales can differ and affect interpretation.
2. In this study, detailed census data collected over a 10-year period from a population of Soay sheep ( Ovis aries L.) on the Island of Hirta in the St Kilda archipelago, Scotland, is used, together with cluster analysis, to distinguish a temporally stable spatial substructure.
3. Structured demographic accounting of the variance in population change (SDA) is also used to analyse the dynamics of the whole population treated as (a) one unit; (b) one unit subdivided into three subunits; and (c) three independent units.
4. Differences in survival, recruitment and dispersal rates are demonstrated between divisions of the population, which are probably associated with variation in grazing quality.
5. If these groups were not coupled by dispersal and density-independent entrainment, the population dynamics of the three groups would diverge, however, the dynamics of the three subunits are strongly correlated.     

Effective size of a fluctuating age-structured population

Previous theories on the effective size of age-structured populations assumed a constant environment and, usually, a constant population size and age structure. We derive formulas for the variance effective size of populations subject to fluctuations in age structure and total population size produced by a combination of demographic and environmental stochasticity. Haploid and monoecious or dioecious diploid populations are analyzed. Recent results from stochastic demography are employed to derive a two-dimensional diffusion approximation for the joint dynamics of the total population size, N, and the frequency of a selectively neutral allele, p. The infinitesimal variance for p, multiplied by the generation time, yields an expression for the effective population size per generation. This depends on the current value of N, the generation time, demographic stochasticity, and genetic stochasticity due to Mendelian segregation, but is independent of environmental stochasticity. A formula for the effective population size over longer time intervals incorporates deterministic growth and environmental stochasticity to account for changes in N.     

Adaptive topography of fluctuating selection in a Mendelian population

An adaptive topography is derived for a large randomly mating diploid population under weak density-independent selection in a fluctuating environment. Assuming a stationary distribution of environmental states with no temporal autocorrelation, a diffusion approximation for population size and allele frequency, p, reveals that the expected change in p involves the gradient with respect to p of the stochastic intrinsic rate of increase (the density-independent long-run growth rate), r = r - sigma 2 e/2, where r is the mean Malthusian fitness in the average environment and is the environmental variance in population growth rate. The expected relative fitness of a genotype is its Malthusian fitness in the average environment minus the covariance of its fitness with population growth rate. The influence of fitness correlation between genotypes is illustrated by an analysis of the Haldane-Jayakar model of fluctuating selection on a single diallelic locus, and on two loci with additive effects on a quantitative character.     

Dip listening and the cocktail party problem in grey treefrogs: Signal recognition in temporally fluctuating noise

Dip listening refers to our ability to catch brief "acoustic glimpses" of speech and other sounds when fluctuating background noise levels momentarily decrease. Exploiting dips in natural fluctuations of noise contributes to our ability to overcome the "cocktail party problem" of understanding speech in multi-talker social environments. We presently know little about how nonhuman animals solve analogous communication problems. Here, we asked whether female grey treefrogs (Hyla chrysoscelis) might benefit from dip listening in selecting a mate in the noisy social setting of a breeding chorus. Consistent with a dip listening hypothesis, subjects recognized conspecific calls at lower thresholds when the dips in a chorus-like noise masker were long enough to allow glimpses of nine or more consecutive pulses. No benefits of dip listening were observed when dips were shorter and included five or fewer pulses. Recognition thresholds were higher when the noise fluctuated at a rate similar to the pulse rate of the call. In a second experiment, advertisement calls comprising six to nine pulses were necessary to elicit responses under quiet conditions. Together, these results suggest that in frogs, the benefits of dip listening are constrained by neural mechanisms underlying temporal pattern recognition. These constraints have important implications for the evolution of male signalling strategies in noisy social environments.     

Life cycles and habitats of wisconsin heptageniidae (ephemeroptera)

Detailed studies were made of the life cycles of Heptageniidae known to occur in Wisconsin. 19 species had univoltine cycles while two and possibly a third had bivoltine cycles. Three univoltine species developed in late spring and early summer while the other univoltine species developed in fall, winter and early spring. For three of the univoltine species, eggs hatched both in fall and the following spring. Diagrams of the life cycles of 15 Wisconsin heptageniids are presented, illustrating the different types of life cycles. Also presented are observation on the food and habitats of the nymphs and flight periods of the adults.     

Limit cycles in periodically perturbed population systems

A perturbation method is proposed to calculate approximately the limit cycle type nonequilibrium steady-state resulting from periodic perturbation of coefficients of stable population systems; the two species Lotka-Volterra competition system is explicity studied and the results are formulated for general multi-species population systems. Avoidance of competitive or other types of exclusion of species in a periodic environment is indicated.     

Maintaining heritable variation via sex-limited temporally fluctuating selection: a phenotypic model accommodating non-Mendelian epigenetic effects

Using a phenotypic model, we show that significant heritable variation can be maintained in a population subjected to temporally fluctuating selection if only one sex is subject to selection. In fact, more variation is maintained with sex-limited selection at a given selection intensity than if both sexes are subject to half that selection intensity. This result is commensurate with existing population genetic models. However, genetic models may be inappropriate for sexually selected traits because many of them may be of non-genetic origin, such as maternal effects or – more likely –epigenetic effects. Phenotypic models obviate this problem by accommodating both genetic and epigenetic effects, as well as maternaleffects. Our phenotypic model of sex-limited temporally fluctuating selection shows that substantial heritable variation can be maintained and therebyprovides impetus to develop population epigenetic models.     

Conservation effort and assessment of population size in fluctuating environments

We consider optimal conservation strategies for endangered populations. We assume that the survival of the population is affected by unpredictable environmental fluctuation and can be improved by conservation effort. Furthermore, the exact value of the initial population size is assumed to be unknown. The conservation strategy involves two aspects: investment of assessment effort, to improve the estimate of the initial population size and investment of conservation effort. Both types of effort imply economic costs. The optimal management strategy is assumed to minimize the weighted sum of extinction probability and the economic cost of the conservation and the assessment effort. (1) We first analyse the optimal conservation effort when the current population size is known accurately. (2) Next, we consider the situation in which there is limited information (i.e. a cue) on population size. (3) We subsequently discuss the cases where the cue accuracy can be improved by assessment of the population. We study the optimal level of the assessment effort and discuss its dependence on various parameters.     

Age variation in a fluctuating population of the common vole

We analysed variation in age in a fluctuating population of the common vole (Microtus arvalis) in southern Moravia, Czech Republic, to test the assumption of the senescence hypothesis that the age of voles increases with increasing population density. Between 1996 and 1998, we monitored the demographic changes by snap-trapping and live-trapping in a field population passing through the increase, peak and decline phase of the population cycle. We used the eye lens mass method to determine the age of snap-trapped animals and those that died in live-traps. The average age of winter males was clearly higher after the peak phase breeding season than before it. No such phase-dependent shift in age, however, was observed in the female component. Male age continued to increase from autumn to spring over the pre-peak winter, and the highest age was in spring of the peak phase year. However, after the peak phase breeding season the highest age was achieved in winter, with the decline phase males during the next spring tending to be younger. The average age of females in spring populations was always lower than in winter populations. The average age of voles from live-traps was always higher than voles from snap-traps, particularly in winter and spring populations, suggesting the presence of senescent animals. Although the density-dependent changes in age are consistent with those observed for other voles, they provide only weak evidence that population cycles in the common vole are accompanied by pronounced shifts in individual age, particularly in female voles.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

The Wright-Fisher model with temporally varying selection and population size

The Wright-Fisher model is considered in the case where the population size is random and the magnitude of the selective advantage of one of the alleles varies with time. The central question addressed is the possibility of ultimate genetic polymorphism. Partial results are obtained in the general case and complete results in the case where the population size and selective advantage are not density dependent. Bounds on the fixation probability are obtained when the selective advantage is constant.     

Variation in pea aphid population development in three different habitats

Abstract.

• 1 Seasonal population growth rates for the pea aphid, Acyrthosiphon pisum Harris, were determined in three different host plant habitats; alfalfa, Medicago sativa (L.), clover, Trifolium pratense (L.), and peas, Pisum sativum (L.); over four years and eight places. It was possible to estimate a common intrinsic rate of increase for each host plant habitat.
• 2 An analysis of the relative influence of temporal, spatial and host plant habitat variation showed that the host plant habitat was most important in determining the growth rates of the populations, both in rate of build-up and decline.
• 3 Patterns of alate production in the three different habitats differed substantially between the annual peas and the two perennial legumes. During the summer, alate production was large and rapid in peas and remained low and constant in clover and alfalfa
• 4 Parasitism was highest in peas. The species composition of parasitoids differed between crops.
• 5 Aphids in annual peas had a higher intrinsic rate of increase and a faster rate of decline than in the two perennial legumes. This explains the presence of both migratory and sedentary forms among pea aphids.