Understanding how stochastic fluctuations in the environment influence population dynamics is crucial for sustainable management of biological diversity. However, because species do not live in ...isolation, this requires knowledge of how species interactions influence population dynamics. In addition, spatial processes play an important role in shaping population dynamics. It is therefore important to improve our understanding of how these different factors act together to shape patterns of abundance across space within and among species. Here, we present a new analytical model for understanding patterns of covariation in space between interacting species in a stochastic environment. We show that the correlation between two species in how they experience the same environmental conditions determines how correlated fluctuations in their densities would be in the absence of competition. In other words, without competition, synchrony between the species is driven by the environment, similar to the Moran effect within a species. Competition between the two species causes their abundances to become less positively or more negatively correlated. The same strength of competition has a greater negative effect on the correlation between species when one of them has a more variable growth rate than the other. In addition, dispersal or other movement weakens the effect of competition on the interspecific correlation. Finally, we show that movement increases the distance over which the species are (positively or negatively) correlated, an effect that is stronger when the species are competitors, and that there is a close connection between the spatial scaling of population synchrony within a species and between species. Our results show that the relationships between the different factors influencing interspecific correlations in abundance are not simple linear ones, but this model allows us to disentangle them and predict how they will affect population fluctuations in different situations.
•Understanding evolutionary responses to environmental changes requires ecological dynamics.•Ecology determines which measure evolution will maximize.•Environmental variability affects the rate of ...adaptive change.•Density-dependent selection can produce evolutionary stasis.
Fitness is a central concept in evolutionary biology, but there is no unified definition. We review recent theoretical developments showing that including fluctuating environments and density dependence has important implications for how differences among phenotypes in their contributions to future generations should be quantified. The rate of phenotypic evolution will vary through time because of environmental stochasticity. Density dependence may produce fluctuating selection for large growth rates at low densities but for larger carrying capacities when population sizes are large. In general, including ecologically realistic assumptions when defining the concept of fitness is crucial for estimating the potential of evolutionary rescue of populations affected by environmental perturbations such as climate change.
Extreme climate events often cause population crashes but are difficult to account for in population-dynamic studies. Especially in long-lived animals, density dependence and demography may induce ...lagged impacts of perturbations on population growth. In Arctic ungulates, extreme rain-on-snow and ice-locked pastures have led to severe population crashes, indicating that increasingly frequent rain-on-snow events could destabilize populations. Here, using empirically parameterized, stochastic population models for High-Arctic wild reindeer, we show that more frequent rain-on-snow events actually reduce extinction risk and stabilize population dynamics due to interactions with age structure and density dependence. Extreme rain-on-snow events mainly suppress vital rates of vulnerable ages at high population densities, resulting in a crash and a new population state with resilient ages and reduced population sensitivity to subsequent icy winters. Thus, observed responses to single extreme events are poor predictors of population dynamics and persistence because internal density-dependent feedbacks act as a buffer against more frequent events.
We propose that the ecological resilience of communities to permanent changes of the environment can be based on how variation in the overall abundance of individuals affects the number of species. ...Community sensitivity is defined as the ratio between the rate of change in the log expected number of species and the rate of change in the log expected number of individuals in the community. High community sensitivity means that small changes in the total abundance strongly impact the number of species. Community resistance is the proportional reduction in expected number of individuals that the community can sustain before expecting to lose one species. A small value of community resistance means that the community can only endure a small reduction in abundance before it is expected to lose one species. Based on long-term studies of four bird communities in European deciduous forests at different latitudes large differences were found in the resilience to environmental perturbations. Estimating the variance components of the species abundance distribution revealed how different processes contributed to the community sensitivity and resistance. Species heterogeneity in the population dynamics was the largest component, but its proportion varied among communities. Species-specific response to environmental fluctuations was the second major component of the variation in abundance. Estimates of community sensitivity and resistance based on data only from a single year were in general larger than those based on estimates from longer time series. Thus, our approach can provide rapid and conservative assessment of the resilience of communities to environmental changes also including only short-term data. This study shows that a general ecological mechanism, caused by increased strength of density dependence due to reduction in resource availability, can provide an intuitive measure of community resilience to environmental variation. Our analyses also illustrate the importance of including specific assumptions about how different processes affect community dynamics. For example, if stochastic fluctuations in the environment affect all species in a similar way, the sensitivity and resistance of the community to environmental changes will be different from communities in which all species show independent responses.
In a stable environment, evolution maximizes growth rates in populations that are not density regulated and the carrying capacity in the case of density regulation. In a fluctuating environment, ...evolution maximizes a function of growth rate, carrying capacity and environmental variance, tending to r-selection and K-selection under large and small environmental noise, respectively. Here we analyze a model in which birth and death rates depend on density through the same function but with independent strength of density dependence. As a special case, both functions may be linear, corresponding to logistic dynamics. It is shown that evolution maximizes a function of the deterministic growth rate r₀ and the lifetime reproductive success (LRS) R₀, both defined at small densities, as well as the environmental variance. Under large noise this function is dominated by r₀ and average lifetimes are small, whereas R₀ dominates and lifetimes are larger under small noise. Thus, K-selection is closely linked to selection for large R₀ so that evolution tends to maximize LRS in a stable environment. Consequently, different quantities (r₀ and R₀) tend to be maximized at low and high densities, respectively, favoring density-dependent changes in the optimal life history.
Understanding how environmental variation affects phenotypic evolution requires models based on ecologically realistic assumptions that include variation in population size and specific mechanisms by ...which environmental fluctuations affect selection. Here we generalize quantitative genetic theory for environmentally induced stochastic selection to include general forms of frequency-and density-dependent selection. We show how the relevant fitness measure under stochastic selection relates to Fisher’s fundamental theorem of natural selection, and present a general class of models in which density regulation acts through total use of resources rather than just population size. In this model, there is a constant adaptive topography for expected evolution, and the function maximized in the long run is the expected factor restricting population growth. This allows us to generalize several previous results and to explain why apparently “K-selected” species with slow life histories often have low carrying capacities. Our joint analysis of density-and frequency-dependent selection reveals more clearly the relationship between population dynamics and phenotypic evolution, enabling a broader range of eco-evolutionary analyses of some of the most interesting problems in evolution in the face of environmental variation.
The degree of spatial autocorrelation in population fluctuations increases with dispersal and geographical covariation in the environment, and decreases with strength of density dependence. Because ...the effects of these processes can vary throughout an individual’s lifespan, we studied how spatial autocorrelation in abundance changed with age in three marine fish species in the Barents Sea. We found large interspecific differences in age-dependent patterns of spatial autocorrelation in density. Spatial autocorrelation increased with age in cod, the reverse trend was found in beaked redfish, while it remained constant among age classes in haddock. We also accounted for the average effect of local cohort dynamics, i.e. the expected local density of an age class given last year’s local density of the cohort, with the goal of disentangling spatial autocorrelation patterns acting on an age class from those formed during younger age classes and being carried over. We found that the spatial autocorrelation pattern of older age classes became increasingly determined by the distribution of the cohort during the previous year. Lastly, we found high degrees of autocorrelation over long distances for the three species, suggesting the presence of far-reaching autocorrelating processes on these populations. We discuss how differences in the species’ life history strategies could cause the observed differences in age-specific variation in spatial autocorrelation. As spatial autocorrelation can differ among age classes, our study indicates that fluctuations in age structure can influence the spatio-temporal variation in abundance of marine fish populations.
The spatial scale of animal space use, e.g. measured as individual home range size, is a key trait with important implications for ecological and evolutionary processes as well as management and ...conservation of populations and ecosystems. Explaining variation in home range size has therefore received great attention in ecological research. However, few studies have examined multiple hypotheses simultaneously, which is important provided the complex interactions between life history, social system and behaviour. Here, we review previous studies on home range size in ungulates, supplementing with a meta-analysis, to assess how differences in habitat use and species characteristics affect the relationship between body mass and home range size. Habitat type was the main factor explaining interspecific differences in home range size after accounting for species body mass and group size. Species using open habitats had larger home ranges for a given body mass than species using closed habitats, whereas species in open habitats showed a much weaker allometric relationship compared with species living in closed habitats. We found no support for relationships between home range size and species diet or mating system, or any sexual differences. These patterns suggest that the spatial scale of animal movement mainly is a combined effect of body mass, group size and the landscape structure. Accordingly, landscape management must acknowledge the influence of spatial distribution of habitat types on animal behaviour to ensure natural processes affecting demography and viability of ungulate populations.
Telomere dynamics could underlie life‐history trade‐offs among growth, size and longevity, but our ability to quantify such processes in natural, unmanipulated populations is limited. We investigated ...how 4 years of artificial selection for either larger or smaller tarsus length, a proxy for body size, affected early‐life telomere length (TL) and several components of fitness in two insular populations of wild house sparrows over a study period of 11 years. The artificial selection was expected to shift the populations away from their optimal body size and increase the phenotypic variance in body size. Artificial selection for larger individuals caused TL to decrease, but there was little evidence that TL increased when selecting for smaller individuals. There was a negative correlation between nestling TL and tarsus length under both selection regimes. Males had longer telomeres than females and there was a negative effect of harsh weather on TL. We then investigated whether changes in TL might underpin fitness effects due to the deviation from the optimal body size. Mortality analyses indicated disruptive selection on TL because both short and long early‐life telomeres tended to be associated with the lowest mortality rates. In addition, there was a tendency for a negative association between TL and annual reproductive success, but only in the population where body size was increased experimentally. Our results suggest that natural selection for optimal body size in the wild may be associated with changes in TL during growth, which is known to be linked to longevity in some bird species.
Sammenfatning
Telomerdynamik kan ligge bag afvejninger mellem livshistorietræk såsom vækst, kropsstørrelse og livslængde, men vores evne til at kvantificere sådanne processer i naturlige, umanipulerede bestande er begrænset. Vi har undersøgt hvorledes 4 års kunstig selektion for enten større eller mindre tarsuslængde, et estimat for kropsstørrelse, påvirkede telomerlængden (TL) i det tidlige liv, samt overlevelses‐ og formeringsevner, i to øbestande af vilde gråspurve over en periode på 11 år. Vores forventning var, at den kunstige selektion ville skubbe bestandene væk fra deres optimale kropsstørrelse og øge den fænotypiske varians i kropsstørrelse. Kunstig selektion for større individer forårsagde en reduktion i TL, men der var begrænset evidens for en øgning i TL når vi selekterede for mindre individer. Der var en negativ korrelation mellem fugleungernes TL og tarsuslængde under begge selektionsregimer. Hanner havde længere telomerer end hunner og der var en negativ effekt af ugunstige vejrforhold på TL. Dernæst undersøgte vi om ændringer i TL kunne underbygge effekter på overlevelses‐ og formeringsevner som følge af afvigelsen fra den optimale kropsstørrelse. Analyser af dødeligheden indikerede disruptiv selektion på TL fordi både korte og lange telomerer i det tidlige liv viste tendens til at være associeret med de laveste dødelighedsrater. Derudover var der en tendens til en negativ sammenhæng mellem TL og årlig reproduktiv succes, men kun i bestanden hvor kropsstørrelse var øget eksperimentelt. Vores resultater antyder, at naturlig selektion for optimal kropsstørrelse i vildtlevende dyr kan være associeret med ændringer i TL (i løbet af vækstperioden), som er kendt for at være forbundet med levetid hos nogle fuglearter.
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