Population Dynamic Responses Research Articles

Ecological success of no-take marine protected areas: Using population dynamics theory to inform a global meta-analysis.

Adaptively managing marine protected areas (MPAs) requires accurately assessing whether established MPAs are achieving their goals of protecting and conserving biomass, especially for harvested populations. Ecological MPA assessments commonly compare inside of the MPA to a reference point outside of and/or before implementation (i.e., calculating "response ratios"). Yet, MPAs are not simple ecological experiments; by design, protected populations interact with those outside, and population dynamic responses can be nonlinear. This complicates assessment interpretations. Here, we used a two-patch population model to explore how MPA response ratios (outside-inside, before-after, and before-after-control-impact [BACI]) for fished populations behave under different conditions, like whether the population is receiving a sustainable larval supply or if it is declining despite protection from harvest. We then conducted a Bayesian evaluation of MPA effects on fish and invertebrate populations based on data collected from 82 published studies on 264 no-take MPAs worldwide, using the results of an earlier global meta-analysis as priors. We considered the effects of calculating different summary metrics on these results, drawing on the theoretical insights from our population model as a comparative framework. We demonstrate that not all response ratio comparison types provide the same information: For example, outside-inside and BACI comparisons can fail to detect population decline within MPAs, whereas before-after comparisons likely detect that pattern. Considering these limitations, we nonetheless found that MPAs globally are producing positive outcomes, with on average greater biomass, density, and organism size within their boundaries than reference sites. However, only a small portion of studies (18 of 82) provided the temporal data necessary to determine that protection, on average, has led to increased abundance of populations within MPAs over time. These findings demonstrate the importance of considering the underlying system dynamics when assessing MPA effects. Assuming that large outside-inside or BACI response ratios always reflect large and net positive conservation effects may lead to misleading conclusions, we recommend that: (1) when assessing specific MPA effects, empirical findings be considered alongside theoretical knowledge relevant to that MPA system, and (2) management should respond to the local conditions and outcomes, rather than a blanket expectation for positive MPA effects.

Early-life variation in migration is subject to strong fluctuating survival selection in a partially migratory bird.

Population dynamic and eco-evolutionary responses to environmental variation and change fundamentally depend on combinations of within- and among-cohort variation in the phenotypic expression of key life-history traits, and on corresponding variation in selection on those traits. Specifically, in partially migratory populations, spatio-seasonal dynamics depend on the degree of adaptive phenotypic expression of seasonal migration versus residence, where more individuals migrate when selection favours migration. Opportunity for adaptive (or, conversely, maladaptive) expression could be particularly substantial in early life, through the initial development of migration versus residence. However, within- and among-cohort dynamics of early-life migration, and of associated survival selection, have not been quantified in any system, preventing any inference on adaptive early-life expression. Such analyses have been precluded because data on seasonal movements and survival of sufficient young individuals, across multiple cohorts, have not been collected. We undertook extensive year-round field resightings of 9359 colour-ringed juvenile European shags Gulosus aristotelis from 11 successive cohorts in a partially migratory population. We fitted Bayesian multi-state capture-mark-recapture models to quantify early-life variation in migration versus residence and associated survival across short temporal occasions through each cohort's first year from fledging, thereby quantifying the degree of adaptive phenotypic expression of migration within and across years. All cohorts were substantially partially migratory, but the degree and timing of migration varied considerably within and among cohorts. Episodes of strong survival selection on migration versus residence occurred both on short timeframes within years, and cumulatively across entire first years, generating instances of instantaneous and cumulative net selection that would be obscured at coarser temporal resolutions. Further, the magnitude and direction of selection varied among years, generating strong fluctuating survival selection on early-life migration across cohorts, as rarely evidenced in nature. Yet, the degree of migration did not strongly covary with the direction of selection, indicating limited early-life adaptive phenotypic expression. These results reveal how dynamic early-life expression of and selection on a key life-history trait, seasonal migration, can emerge across seasonal, annual, and multi-year timeframes, yet be substantially decoupled. This restricts the potential for adaptive phenotypic, microevolutionary, and population dynamic responses to changing seasonal environments.

A user-friendly guide to using distance measures to compare time series in ecology.

Time series are a critical component of ecological analysis, used to track changes in biotic and abiotic variables. Information can be extracted from the properties of time series for tasks such as classification (e.g., assigning species to individual bird calls); clustering (e.g., clustering similar responses in population dynamics to abrupt changes in the environment or management interventions); prediction (e.g., accuracy of model predictions to original time series data); and anomaly detection (e.g., detecting possible catastrophic events from population time series). These common tasks in ecological research all rely on the notion of (dis-) similarity, which can be determined using distance measures. A plethora of distance measures have been described, predominantly in the computer and information sciences, but many have not been introduced to ecologists. Furthermore, little is known about how to select appropriate distance measures for time-series-related tasks. Therefore, many potential applications remain unexplored. Here, we describe 16 properties of distance measures that are likely to be of importance to a variety of ecological questions involving time series. We then test 42 distance measures for each property and use the results to develop an objective method to select appropriate distance measures for any task and ecological dataset. We demonstrate our selection method by applying it to a set of real-world data on breeding bird populations in the UK and discuss other potential applications for distance measures, along with associated technical issues common in ecology. Our real-world population trends exhibit a common challenge for time series comparisons: a high level of stochasticity. We demonstrate two different ways of overcoming this challenge, first by selecting distance measures with properties that make them well suited to comparing noisy time series and second by applying a smoothing algorithm before selecting appropriate distance measures. In both cases, the distance measures chosen through our selection method are not only fit-for-purpose but are consistent in their rankings of the population trends. The results of our study should lead to an improved understanding of, and greater scope for, the use of distance measures for comparing ecological time series and help us answer new ecological questions.

Properties of phenotypic plasticity in discrete threshold traits.

Forms of phenotypic plasticity in key traits, and forms of selection on and genetic variation in such plasticity, fundamentally underpin phenotypic, population dynamic, and evolutionary responses to environmental variation and directional change. Accordingly, numerous theoretical and empirical studies have examined properties and consequences of plasticity, primarily considering traits that are continuously distributed on observed phenotypic scales with linear reaction norms. However, many environmentally sensitive traits are expressed as discrete alternative phenotypes and are appropriately characterized as quantitative genetic threshold traits. Here, we highlight that forms of phenotypic plasticity, genetic variation, and inheritance in plasticity, and outcomes of selection on plasticity, could differ substantially between threshold traits and continuously distributed traits (as are typically considered). We thereby highlight theoretical developments that are required to rationalize and predict phenotypic and microevolutionary dynamics involving plastic threshold traits, and outline how intrinsic properties of such traits could provide relatively straightforward explanations for apparently idiosyncratic observed patterns of phenotypic variation. We summarize how key quantitative genetic parameters underlying threshold traits can be estimated, and thereby set the scene for embedding dynamic discrete traits into theoretical and empirical understanding of the role of plasticity in driving phenotypic, population, and evolutionary responses to environmental variation and change.

Ecosystem-specific growth responses to climate pattern by a temperate freshwater fish

Abstract Somatic growth patterns among animal populations are maintained through complex processes that vary among ecosystems. Changes in growth patterns may be concomitant with changes in climate; however, understanding how growth will manifest among ecosystems is limited. Information embedded within fish hard-parts (i.e., otoliths, spines, vertebrae) can account for variation in growth patterns resulting from changing climate conditions. Channel catfish Ictalurus punctatus is a freshwater fish species widely distributed across North America with limited information regarding climate influences on growth and differences in climate-growth relations among ecological systems. We assessed growth (total length) response to changing climate conditions for channel catfish among three waterbody types—pit lakes, irrigation and power-generation reservoirs, and flood-control reservoirs in Nebraska, USA. We used linear mixed-effect models and an information theoretic approach to assess the relative strengths among competing hypotheses. The most supported linear mixed-effect model of channel catfish growth was a function of fish age and an interaction between waterbody type and growing-degree-day (GDD). A positive trend existed in GDD from 1990 through 2008 whereby the predicted increase in GDD among waterbody types ranged from 182 GDD to 189 GDD. The predicted change in channel catfish growth resulting from increased GDD ranged from 1% to 39% among waterbody types. Channel catfish population rate functions, thus, may not respond similarly to climate conditions across ecosystem types. Changes in climate variables may contribute to system-specific responses in population dynamics for channel catfish as well as other similar freshwater species. The establishment of relations between climate and growth variables for a freshwater generalist with a plastic diet and broad temperature tolerance serves as an indication of the breadth of responses possible for freshwater fishes under global changes in climate conditions.

Linking inter‐annual variation in environment, phenology, and abundance for a montane butterfly community

Climate change has caused widespread shifts in species’ phenology, but the consequences for population and community dynamics remain unclear because of uncertainty regarding the species‐specific drivers of phenology and abundance, and the implications for synchrony among interacting species. Here, we develop a statistical model to quantify inter‐annual variation in phenology and abundance over an environmental gradient, and use it to identify potential drivers of phenology and abundance in co‐occurring species. We fit the model to counts of 10 butterfly species with single annual generations over a mountain elevation gradient, as an exemplar system in which temporally limited availability of biotic resources and favorable abiotic conditions impose narrow windows of seasonal activity. We estimate parameters describing changes in abundance, and the peak time and duration of the flight period, over ten years (2004–2013) and across twenty sample locations (930–2,050 m) in central Spain. We also use the model outputs to investigate relationships of phenology and abundance with temperature and rainfall. Annual shifts in phenology were remarkably consistent among species, typically showing earlier flight periods during years with warm conditions in March or May–June. In contrast, inter‐annual variation in relative abundance was more variable among species, and generally less well associated with climatic conditions. Nevertheless, warmer temperatures in June were associated with increased relative population growth in three species, and five species had increased relative population growth in years with earlier flight periods. These results suggest that broadly coherent interspecific changes to phenology could help to maintain temporal synchrony in community dynamics under climate change, but that the relative composition of communities may vary due to interspecific inconsistency in population dynamic responses to climate change. However, it may still be possible to predict abundance change for species based on a robust understanding of relationships between their population dynamics and phenology, and the environmental drivers of both.

Contrasting consequences of climate change for migratory geese: Predation, density dependence and carryover effects offset benefits of high-arctic warming.

Climate change is most rapid in the Arctic, posing both benefits and challenges for migratory herbivores. However, population-dynamic responses to climate change are generally difficult to predict, due to concurrent changes in other trophic levels. Migratory species are also exposed to contrasting climate trends and density regimes over the annual cycle. Thus, determining how climate change impacts their population dynamics requires an understanding of how weather directly or indirectly (through trophic interactions and carryover effects) affects reproduction and survival across migratory stages, while accounting for density dependence. Here, we analyse the overall implications of climate change for a local non-hunted population of high-arctic Svalbard barnacle geese, Branta leucopsis, using 28years of individual-based data. By identifying the main drivers of reproductive stages (egg production, hatching and fledging) and age-specific survival rates, we quantify their impact on population growth. Recent climate change in Svalbard enhanced egg production and hatching success through positive effects of advanced spring onset (snow melt) and warmer summers (i.e. earlier vegetation green-up) respectively. Contrastingly, there was a strong temporal decline in fledging probability due to increased local abundance of the Arctic fox, the main predator. While weather during the non-breeding season influenced geese through a positive effect of temperature (UK wintering grounds) on adult survival and a positive carryover effect of rainfall (spring stopover site in Norway) on egg production, these covariates showed no temporal trends. However, density-dependent effects occurred throughout the annual cycle, and the steadily increasing total flyway population size caused negative trends in overwinter survival and carryover effects on egg production. The combination of density-dependent processes and direct and indirect climate change effects across life history stages appeared to stabilize local population size. Our study emphasizes the need for holistic approaches when studying population-dynamic responses to global change in migratory species.

When Do Shifts in Trait Dynamics Precede Population Declines?

Predicting population responses to environmental change is an ongoing challenge in ecology. Studies investigating the links between fitness-related phenotypic traits and demography have shown that trait dynamic responses to environmental change can sometimes precede population dynamic responses and thus can be used as an early warning signal. However, it is still unknown under which ecological and evolutionary circumstances shifts in fitness-related traits can precede population responses to environmental perturbation. Here, we take a trait-based demographic approach and investigate both trait and population dynamics in a density-regulated population in response to a gradual change in the environment. We explore the ecological and evolutionary constraints under which shifts in fitness-related traits precede a decline in population size. We show both analytically and with experimental data that under medium to slow rates of environmental change, shifts in a trait value can precede population decline. We further show the positive influence of environmental predictability, net reproductive rate, plasticity, and genetic variation on shifts in trait dynamics preceding potential population declines. These results still hold under nonconstant genetic variation and environmental stochasticity. Our study highlights ecological and evolutionary circumstances under which a fitness-related trait can be used as an early warning signal of an impending population decline.

Trawl ban in a heavily exploited marine environment: Responses in population dynamics of four stomatopod species

Intensive trawling activities in Hong Kong waters have seriously depleted fishery resources and damaged marine benthic habitats over the last four decades. To minimize further destruction and rehabilitate fishery resources, the Hong Kong Government implemented a permanent territory-wide trawling closure on 31 December 2012. Such a trawl ban creates a unique opportunity to investigate recoveries in ecosystem structure and function following a major shift in disturbance regime by removing impacts from a major gear. This study was designed to test the hypothesis that dominant predatory mantis shrimps, including Harpiosquilla harpax, Miyakella nepa, Oratosquillina interrupta, and Oratosquilla oratoria would show signs of recovery following the trawl ban. Their population dynamics were investigated before and after the trawl ban. The results showed that their mean weight, mean carapace length and proportion of large-sized individuals increased significantly 3.5 years after the trawl ban, whilst their abundance, biomass and maximum length remained unchanged. This study suggests that the stomatopod assemblage in the human-dominated Hong Kong waters shows some initial signs of possible recovery following the trawl ban but also highlights the complexity of implementing fishery management and detecting changes resulted from management measures in a heavily urbanized seascape where many biotic and abiotic factors can influence their population dynamics.

Population and evolutionary dynamics in spatially structured seasonally varying environments.

ABSTRACTIncreasingly imperative objectives in ecology are to understand and forecast population dynamic and evolutionary responses to seasonal environmental variation and change. Such population and evolutionary dynamics result from immediate and lagged responses of all key life‐history traits, and resulting demographic rates that affect population growth rate, to seasonal environmental conditions and population density. However, existing population dynamic and eco‐evolutionary theory and models have not yet fully encompassed within‐individual and among‐individual variation, covariation, structure and heterogeneity, and ongoing evolution, in a critical life‐history trait that allows individuals to respond to seasonal environmental conditions: seasonal migration. Meanwhile, empirical studies aided by new animal‐tracking technologies are increasingly demonstrating substantial within‐population variation in the occurrence and form of migration versus year‐round residence, generating diverse forms of ‘partial migration’ spanning diverse species, habitats and spatial scales. Such partially migratory systems form a continuum between the extreme scenarios of full migration and full year‐round residence, and are commonplace in nature.Here, we first review basic scenarios of partial migration and associated models designed to identify conditions that facilitate the maintenance of migratory polymorphism. We highlight that such models have been fundamental to the development of partial migration theory, but are spatially and demographically simplistic compared to the rich bodies of population dynamic theory and models that consider spatially structured populations with dispersal but no migration, or consider populations experiencing strong seasonality and full obligate migration. Second, to provide an overarching conceptual framework for spatio‐temporal population dynamics, we define a ‘partially migratory meta‐population’ system as a spatially structured set of locations that can be occupied by different sets of resident and migrant individuals in different seasons, and where locations that can support reproduction can also be linked by dispersal. We outline key forms of within‐individual and among‐individual variation and structure in migration that could arise within such systems and interact with variation in individual survival, reproduction and dispersal to create complex population dynamics and evolutionary responses across locations, seasons, years and generations. Third, we review approaches by which population dynamic and eco‐evolutionary models could be developed to test hypotheses regarding the dynamics and persistence of partially migratory meta‐populations given diverse forms of seasonal environmental variation and change, and to forecast system‐specific dynamics. To demonstrate one such approach, we use an evolutionary individual‐based model to illustrate that multiple forms of partial migration can readily co‐exist in a simple spatially structured landscape. Finally, we summarise recent empirical studies that demonstrate key components of demographic structure in partial migration, and demonstrate diverse associations with reproduction and survival. We thereby identify key theoretical and empirical knowledge gaps that remain, and consider multiple complementary approaches by which these gaps can be filled in order to elucidate population dynamic and eco‐evolutionary responses to spatio‐temporal seasonal environmental variation and change.

Adaptation of timing of life history traits and population dynamic responses to climate change in spatially structured populations

Changes in the seasonal timing of life history events are documented effects of climate change. We used a general model to study how dispersal and competitive interactions affect eco-evolutionary responses to changes in the temporal distribution of resources over the season. Specifically, we modeled adaptation of the timing of reproduction and population dynamic responses in two competing populations that disperse between two habitats characterized by an early and late resource peak. We investigated three scenarios of environmental change: (1) food peaks advance in both habitats, (2) in the late habitat only and (3) in the early habitat only. At low dispersal rates the evolutionarily stable timing of reproduction closely matched the local resource peak and the environmental change typically caused population decline. Larger dispersal rates rendered less intuitive eco-evolutionary population responses. First, dispersal caused mismatch between evolutionarily stable timing of reproduction and local resource peaks and as a result, reproductive output for subpopulations could increase as well as decrease when resource availability underwent temporal shifts. Second, population responses were contingent on competition between populations. This could accelerate population declines and cause extinctions or even reverse population trends from negative to positive compared to the low dispersal case. When dispersal rate was large and the early resource peak was advanced available niche space was reduced. Hence, even when a population survived the environmental change and obtained positive equilibrium population density, subsequent adaptation of competing populations could drive it to extinction due to convergent evolution and competitive exclusion. These results shed new light on the role of competition and dispersal for the evolution of timing of life history events and provide guidelines for understanding short and long-term population response to climate change.

Spatial and temporal variation in amphibian metacommunity structure in Chiapas, Mexico

Abstract:Amphibians are known to be sensitive to environmental change but their responses at the level of metacommunities to short-term environmental variability are poorly understood. We used field data from two protected areas, La Pera and Nahá (Chiapas, Mexico) to test for variation in metacommunity properties for two consecutive years (2009 and 2010). Amphibians and accompanying environmental data were recorded to a standardized protocol within each landscape, in four or five transects of 50 × 2-m per patch, for 30 and 31 patches, respectively. We found 23 species in La Pera and 30 in Nahá (21 species shared). Metacommunity structure was analysed using reciprocal averaging (RA) ordination by means of metrics for coherence, turnover and boundary clumping, with Spearman rank correlation used to examine relationships with environmental variables. The metacommunity structure varied differentially among the landscapes between years, being classed as quasi-Gleasonian in La Pera in both years, but Clementsian for Nahá in 2009 and Gleasonian for Nahá in 2010. In further illustration of variation between years, in 2009 the principal community gradient (RA axis 1) for La Pera was significantly positively correlated with altitude (r = 0.36), forest disturbance status (r = 0.78), mean canopy cover (r = 0.79) and mean litter depth (r = 0.67), while in 2010 it was correlated with latitude (r = 0.38), mean grass-layer height (r = 0.38), incidence of rainfall prior to sampling (r = 0.35) and presence of temporary ponds (r = 0.45). Our findings support the notion that amphibians respond to short-term environmental changes by individualistic movement within the landscape as well as via population dynamic responses.

Long-term responses in population dynamics and diversity of small mammals in riparian and upland habitats within an agricultural landscape

Riparian zones in agricultural landscapes provide linear non-crop habitats for a variety of plant and mammal species, and hence are an important component of biodiversity. To date, variable responses of abundance, species richness, and species diversity of small mammals have been recorded in riparian and upland habitats. To address this variability, we provide a detailed analysis of seasonal changes in abundance and diversity of terrestrial small-mammal communities over a 7-year period within an agricultural landscape in south-central British Columbia, Canada. We tested the hypotheses (H) that abundance, species richness, and species diversity of communities of small mammals (H1), and demographic parameters of reproduction, recruitment, and survival of the major species: deer mouse (Peromyscus maniculatus) and montane vole (Microtus montanus) (H2), would be higher in riparian than upland habitats. Mean total abundance of small mammals was higher in summer and winter, and species richness higher in summer, in riparian than hedgerow habitats. Winter population data supported the total and species abundance patterns for small mammals, but species richness was similar, and diversity lower, in riparian than hedgerow sites during winter periods. Deer mice were the dominant species in terms of abundance and reproductive output for pregnancies and recruitment, but not survival, in riparian sites. Montane voles were similar in abundance and demographic parameters in the two habitats. House mice (Mus musculus) preferred hedgerows and wandering shrews (Sorex vagrans) riparian sites. Demographic parameters for deer mice and montane voles indicated that both riparian and hedgerow sites were “source” rather than “sink” habitats, and likely contribute to maintenance of mammal diversity in agricultural landscapes.

Evidence for transient dynamics in plant populations based on long‐term demographic data

Summary Recent interest in the importance of short‐term, transient responses in population dynamics has prompted the development of many new methods for describing transient responses in population models. Many of these methods focus on the largest possible responses for a model, thus providing a description of the ‘transient potential’ of a model that does not depend on knowledge of initial conditions in the population. This study uses long‐term demographic data from five perennial plant species to evaluate how well measures of transient potential reflect actual transient responses in the populations. I used previously published matrix population models to create a set of initial conditions derived from empirically‐based simulations for each population. Maximum transient responses within this set of initial conditions were then compared with responses used in describing transient potential. Transient responses based on empirical modeling results were smaller than responses used for measures of transient potential. Realized responses were strongly correlated with the distance between the current demographic structure in the population and the asymptotically stable stage structure of the model (SSD). Theoretical transient potential was predictive of the amount of variation in distance from SSD observed in each population in the simulations. Synthesis. Methods based on largest possible responses tend to overemphasize the role of transient dynamics. These results suggest that traditional, asymptotic analyses may be appropriate in many cases. Measures of transient potential can be helpful for identifying species and situations that may be prone to larger transient responses, but do not necessarily indicate that transient dynamics are more important in those systems.

Direct and indirect effects of temperature on the population dynamics and ecosystem functioning of aquatic microbial ecosystems

1. While much is known about the direct effect that temperature can have on aquatic communities, less is known about its indirect effect via the temperature dependence of viscosity and temperature-dependent trophic interactions. 2. We manipulated the temperature (5-20 °C) and the viscosity (equivalent to 5-20 °C) of water in laboratory-based bacteria-protist communities. Communities contained food chains with one, two or three trophic levels. Responses measured were population dynamics (consumer carrying capacity and growth rate, average species population density, and the coefficient of variation of population density through time) and ecosystem function (decomposition). 3. Temperature, viscosity and food chain length produced significant responses in population dynamics. Temperature-dependent viscosity had a significant effect on the carrying capacity and growth rates of consumers, as well as the average density of the top predator. Overall, indirect effects of temperature via changes in viscosity were subtle in comparison to the indirect effect of temperature via trophic interactions. 4. Our results highlight the importance of direct and indirect effects of temperature, mediated through trophic interactions and physical changes in the environment, both for population dynamics and ecosystem processes. Future mechanistic modelling of effects of environmental change on species will benefit from distinguishing the different mechanisms of the overall effect of temperature.

Overcompensatory population dynamic responses to environmental stochasticity

1. To quantify the interactions between density-dependent, population regulation and density-independent limitation, we studied the time-series dynamics of an experimental laboratory insect microcosm system in which both environmental noise and resource limitation were manipulated. 2. A hierarchical Bayesian state-space approach is presented through which it is feasible to capture all sources of uncertainty, including observation error to accurately quantify the density dependence operating on the dynamics. 3. The regulatory processes underpinning the dynamics of two different bruchid beetles (Callosobruchus maculatus and Callosobruchus chinensis) are principally determined by environmental conditions, with fluctuations in abundance explained in terms of changes in overcompensatory dynamics and stochastic processes. 4. A general, stochastic population model is developed to explore the link between abundance fluctuations and the interaction between density dependence and noise. Taking account of time-lags in population regulation can substantially increase predicted population fluctuations resulting from underlying noise processes.

Linking growth strategies to long-term population dynamics in a guild of desert annuals

Summary 1 Combining long-term observational studies with comparative physiological ecology can yield a deeper understanding of the contribution of individual function to population and community dynamics. Sonoran Desert winter annuals exhibit striking year-to-year variation in population dynamics that is driven by variable precipitation, but species differ in the strength of demographic response to precipitation and hence in the degree of temporal variance in population dynamics. To understand the physiological mechanisms of differing population dynamic responses to environmental variation, we investigated interspecific differences in functional traits that mediate responsiveness to precipitation. 2 We conducted sequential harvests throughout the growing season to examine relative growth rate and biomass allocation patterns. We then related growth parameters to leaf-level carbon isotope discrimination (a time-integrated measure of water-use efficiency) and long-term demographic variation. 3 We hypothesized that water-use efficiency should trade-off with rapid growth rates. Furthermore, we hypothesized that species having efficient water use should have buffered population dynamics in dry years but sacrifice high growth and fecundity in wet years, resulting in low long-term variance in demographic success. Conversely, species with high growth capacity should be very responsive to infrequent periods of high precipitation and thus exhibit high temporal variance. 4 Species differed in seasonal relative growth rate and allocation patterns. Species with the highest relative growth rates rapidly deployed large leaf area displays following mid-season rainfall. Species with intermediate relative growth rates exhibited high biomass assimilation rates per unit leaf area. Species with low relative growth rates exhibited low leaf area ratios and low assimilation rates per unit leaf area. 5 Relative growth rate was positively related to leaf carbon isotope discrimination, consistent with a trade-off between growth rate and water-use efficiency. 6 Seasonal relative growth rate did not predict long-term demographic variance. However, leaf area plasticity in response to precipitation was positively related to long-term demographic variance. Our results illustrate how morphological and physiological traits influence demographic tracking of environmental variability and demonstrate how species differences in functional strategies determine population and community dynamics.

METHODOLOGICAL INSIGHT: Linking management changes to population dynamic responses: the transfer function of a projection matrix perturbation

Summary An important task in applied population ecology is to understand how changes to individual life‐history parameters, such as survival, growth and fecundity, affect population dynamics. Parameter changes, or perturbations, may be caused by deliberate attempts to manage populations (e.g. in pest control, harvesting or conservation) or they may be side‐effects of pollution, genetic modification and climate change. For organisms with complicated life cycles, links between individual life histories and population dynamics are made using population projection matrix (PPM) modelling. Changes to individual, or groups of, life‐history transition rates within a PPM have a nonlinear impact on the resulting eigenvalues. Conventional sensitivity analysis calculates the derivative of the perturbation‐eigenvalue curve to provide tangential linear extrapolation. Until now, only the simulation of perturbed PPMs has captured nonlinear perturbation effects. Here we describe the transfer function of a matrix perturbation. The transfer function captures analytically the true relationship between perturbation magnitude and PPM eigenvalues. This analytical link extends easily to multi‐transition and multiple perturbations, promotes an understanding of matrix properties, and provides a simple method to predict the perturbation required to achieve a desired population rate of increase. We use the transfer function approach to analyse a PPM for the desert tortoise Gopherus agasizzii Cooper, in the context of conservation management decisions. Synthesis and applications. The transfer function offers a novel and powerful framework for the analysis of population projection matrices (PPMs), giving precise predictive power and analytical understanding of population‐level responses to life‐history perturbations, for example in the design of conservation, pest control and population harvesting strategies, prediction of population effects of pollution in ecotoxicology, and in ecological risk assessment. A useful focus is to set a target for the desired rate of increase (or decline) of a population, and use the transfer function to determine how best to achieve this rate.

Why ecology fails at application: should we consider variability more than regularity?

Inability to solve applied ecological problems is argued to depend on inherent variability in intra‐ and interspecific interactions caused by individual utilisation of a variety of local environments, and on individual counter‐adaptations towards limiting or regulating factors. Such biological diversification is largely a response to environmental heterogeneity. Empirical and theoretical analyses of ecological mechanisms in various gradients of interaction and adaptation should result in improved use of ecological insight. Due to the many gradients, to the locally determined processes and to covert genetic variation very precise predictions of the outcome of ecological interactions will never be possible. Instead of focussing on possible general laws, ecology may try to establish common responses in population and community dynamics to main environmental gradients, particularly with regard to habitats and ecosystems.

Demographic dynamics of a neotropical small rodent (Phyllotis darwini): feedback structure, predation and climatic factors

Summary The leaf‐eared mouse (Phyllotis darwini) exhibits large numerical fluctuations associated with high‐ and low‐rainfall years in semi‐arid Chile. Using capture–mark–recapture (CMR) statistical modelling, we provide a detailed description of the demographic variation in this species. We studied between‐year and seasonal variation of demography, and tested for the relative importance of endogenous and exogenous factors as covariates of survival, recruitment, maturation and reproduction over a 12‐year period. The demographic variables studied show larger between‐year than within‐year (seasonal) variation, emphasizing the importance of this source of external variability. We found that both feedback structure and exogenous factors operate together on different demographic processes of this species. We found positive direct feedback effects on survival rate and negative feedback on recruitment, fraction of reproductive individuals and female maturation rates. The ratio between barn and magellanic horned owls showed an important effect on most of the demographic variables of leaf‐eared mice, in particular by influencing negatively survival rates and the fraction of reproductive individuals, thus suggesting the importance of non‐lethal predation. Hence, feedback intrinsic mechanisms (density‐dependence), owls and climate (rainfall and Southern Oscillation Index (SOI)) operate simultaneously in determining the demography of leaf‐eared mice. We hypothesize that trophic interactions with owls (and perhaps other predators) may cause second‐order feedback responses in demography and population dynamics of this small rodent species.