The European bitterling: a model for oviposition decision

For oviparous species, oviposition decisions can have significant fitness consequences. The European bitterling ( Rhodeus sericeus ) is a cyprinid fish that lays its eggs on the gills of freshwater mussels. Because bitterling use a discrete spawning site that can be readily manipulated, they are unusually amenable to field and laboratory studies aimed at understanding the adaptiveness and consequences of oviposition decisions. Here I present data demonstrating the adaptiveness of oviposition decisions by female bitterling, and link these decisions, using game‐theory models, with bitterling population dynamics. I explain the proximate cues used by females in making oviposition choices, and how these respond to environmental variables. I also show how the oviposition decisions of males relate to the risk of sperm competition in mussels, and propose that oviposition choices may represent an inter‐sexual conflict in bitterling. Finally, I present behavioural and genetic data from mesocosm experiments to show the implications of global and local male densities for female oviposition opportunities, and consider the consequences for bitterling population dynamics.     

Density dependence in a recovering osprey population: demographic and behavioural processes

1. Understanding how density-dependent and independent processes influence demographic parameters, and hence regulate population size, is fundamental within population ecology. We investigated density dependence in growth rate and fecundity in a recovering population of a semicolonial raptor, the osprey Pandion haliaetus [Linnaeus, 1758], using 31 years of count and demographic data in Corsica. 2. The study population increased from three pairs in 1974 to an average of 22 pairs in the late 1990s, with two distinct phases during the recovery (increase followed by stability) and contrasted trends in breeding parameters in each phase. 3. We show density dependence in population growth rate in the second phase, indicating that the stabilized population was regulated. We also show density dependence in productivity (fledging success between years and hatching success within years). 4. Using long-term data on behavioural interactions at nest sites, and on diet and fish provisioning rate, we evaluated two possible mechanisms of density dependence in productivity, food depletion and behavioural interference. 5. As density increased, both provisioning rate and the size of prey increased, contrary to predictions of a food-depletion mechanism. In the time series, a reduction in fledging success coincided with an increase in the number of non-breeders. Hatching success decreased with increasing local density and frequency of interactions with conspecifics, suggesting that behavioural interference was influencing hatching success. 6. Our study shows that, taking into account the role of non-breeders, in particular in species or populations where there are many floaters and where competition for nest sites is intense, can improve our understanding of density-dependent processes and help conservation actions.     

On harvesting a structured ungulate population

Variation in demographic rates within a spatially structured population could have important consequences for management decisions, harvesting strategies and offtake rates. Although there is a growing body of evidence suggesting that demographic rates vary within populations over a range of spatial scales, there has been little research investigating the consequences of this variation for population management. In this paper, data on the dynamics of two female red deer sub-populations on Rum are analysed, and evidence is presented for differences between the fecundity and mortality rates of the two sub-populations. A simple harvesting model is developed to represent the dynamics of the two sub-populations, including density-independent migration between sub-populations and spatially correlated environmental variability. The highest monetary yield in the model is obtained by harvesting the more resilient sub-population at a higher rate. Surprisingly the losses involved in harvesting both sub-populations at the same rate are insignificant. However, if migration were density-dependent, the size of one sub-population would be more relevant to harvesting policy for the other sub-population. The results of this empirical study are compared to theoretical work on spatially structured populations; it is shown that when a species has complex age- and sex-structured population dynamics, previous theoretical results may not hold.     

The evolution of migration and some life history parameters in marine fishes

Synopsis Migration is a common phenomenon in marine fishes but the consequences with respect to life history variation have been little explored. Migration both influences the evolution of other traits and is contigent upon the evolution of other behavioural and demographic characters. The interaction between such factors is illustrated by considering the relationship between the cost of migration in relation to fecundity and the advantages and disadvantages of schooling, a phenomenon hypothesized to favour the evolution of migration. These analyses predict that migratory species will be relatively large, mature late and at a relatively large size and grow relatively fast. Data from north temperate marine fish support these predictions.     

Utilizing spatial demographic and life history variation to optimize sustainable yield of a temperate sex-changing fish

Fish populations vary geographically in demography and life history due to environmental and ecological processes and in response to exploitation. However, population dynamic models and stock assessments, used to manage fisheries, rarely explicitly incorporate spatial variation to inform management decisions. Here, we describe extensive geographic variation in several demographic and life history characteristics (e.g., size structure, growth, survivorship, maturation, and sex change) of California sheephead (Semicossyphus pulcher), a temperate rocky reef fish targeted by recreational and commercial fisheries. Fish were sampled from nine locations throughout southern California in 2007-2008. We developed a dynamic size and age-structured model, parameterized separately for each location, to assess the potential cost or benefit in terms of fisheries yield and conservation objectives of changing minimum size limits and/or fishing mortality rates (compared to the status quo). Results indicate that managing populations individually, with location-specific regulations, could increase yield by over 26% while maintaining conservative levels of spawning biomass. While this local management approach would be challenging to implement in practice, we found statistically similar increases in yield could be achieved by dividing southern California into two separate management regions, reflecting geographic similarities in demography. To maximize yield, size limits should be increased by 90 mm in the northern region and held at current levels in the south. We also found that managing the fishery as one single stock (the status quo), but with a size limit 50 mm greater than the current regulations, could increase overall fishery yield by 15%. Increases in size limits are predicted to enhance fishery yield and may also have important ecological consequences for the predatory role of sheephead in kelp forests. This framework for incorporating demographic variation into fisheries models can be exported generally to other species and may aid in identifying the appropriate spatial scales for fisheries management.     

Social consequences of rapid environmental change

While direct influences of the environment on population growth and resilience are well studied, indirect routes linking environmental changes to population consequences are less explored. We suggest that social behavior is key for understanding how anthropogenic environmental changes affect the resilience of animal populations. Social structures of animal groups are evolved and emergent phenotypes that often have demographic consequences for group members. Importantly, environmental drivers may directly influence the consequences of social structure or indirectly influence them through modifications to social interactions, group composition, or group size. We have developed a framework to study these demographic consequences. Estimating the strength of direct and indirect pathways will give us tools to understand, and potentially manage, the effect of human-induced rapid environmental changes.     

Evidence for selective angling of introduced trout and their hybrids in a stocked brown trout population

Comparisons of the genetic composition of brown trout Salmo trutta captured by anglers and by electrofishing based on three diagnostic microsatellite loci provided strong evidence that angling is selective in a stocked brown trout population. At two sites, anglers caught significantly younger trout and proportionally more introduced hatchery trout and hybrids than were observed in electrofishing surveys. Selective angling, in combination with a small legal catch size, may have considerably eliminated introduced trout and hybrids before spawning at the study sites, and thus may have reduced the introgression of alien genes into the local gene pool. Angling can be an important factor influencing the genetic structure of fish populations and should be taken into account in studies of introgressive hybridization in stocked fish populations and their management. In this study, demographic consequences of stocking were not assessed. Thus, even though the genetic consequences of stocking may be minimal or largely reversible through angling, resource competition between native and introduced trout, until they reach legal catch size, is expected to have a negative effect on the productivity of the indigenous trout population.     

Behavioural models of population growth rates: implications for conservation and prediction

Conservation biologists often wish to predict how vertebrate populations will respond to local or global changes in conditions such as those resulting from sea-level rise, deforestation, exploitation, genetically modified crops, global warming, human disturbance or from conservation activities. Predicting the consequences of such changes almost always requires understanding the population growth rate and the density dependence. Traditional means of directly measuring density dependence are often extremely difficult and have the problem that if the environment changes then it is necessary to remeasure the density dependence. We describe an alternative approach that does not require such long datasets and can be used to predict the density dependence under novel conditions. Game theory can be used to describe behavioural decisions that individuals make in response to interference, prey depletion, territorial behaviour or social dominance, and the resultant fitness consequences. It is then possible to predict how survival or reproductive output changes with population size. From this we can then make predictions about the responses of populations to environmental changes. We will illustrate how this can be applied to a range of species and a range of applied problems.     

From inducible defences to population dynamics: modelling refuge use and life history changes in Daphnia

We investigated the relative importance of a behavioural defence (refuge use through diel vertical migration) and a life history change (a reduced size at first reproduction) that are used by daphnids to decrease the risk of predation by visually hunting fish. We used an individual based model of a Daphnia population in a stratified lake to quantify the effects of these inducible defences on Daphnia predation-mortality and the resulting Daphnia population dynamics. Our analysis shows that diel vertical migration (DVM) confers a much stronger protection against fish predation than a reduced size at first reproduction (SFR). DVM allows daphnids to withstand a higher predation pressure in the epilimnion and it decelerates a Daphnia population decline more strongly than a reduced SFR. DVM effectively reduces the (P/B) flow of carbon from daphnids to fish.
Many theoretical studies have only considered the fitness benefits of DVM above 'staying up' in the epilimnion of a lake. Our results suggest that 'staying down' in the hypolimnion would confer an even stronger fitness benefit to Daphnia than DVM at times of peak predation risk. Daphnids that remain in the hypolimnion avoid the predation suffered by migrating daphnids around dusk and dawn. Staying down could prevent a Daphnia population decline, while DVM and a reduced SFR can only decelerate the decrease of Daphnia population densities under heavy fish predation. Staying down at high concentrations of fish infochemicals has in fact been observed within a variety of Daphnia clones and species, both in the laboratory and in stratified lakes.     

Coexistence of behavioural types in an aquatic top predator: a response to resource limitation?

Intra-population variation in behaviour unrelated to sex, size or age exists in a variety of species. The mechanisms behind behavioural diversification have only been partly understood, but density-dependent resource availability may play a crucial role. To explore the potential coexistence of different behavioural types within a natural fish population, we conducted a radio telemetry study, measuring habitat use and swimming activity patterns of pike (Esox lucius), a sit-and-wait predatory fish. Three behavioural types co-occurred in the study lake. While two types of fish only selected vegetated littoral habitats, the third type opportunistically used all habitats and increased its pelagic occurrence in response to decreasing resource biomasses. There were no differences in size, age or lifetime growth between the three behavioural types. However, habitat-opportunistic pike were substantially more active than the other two behavioural types, which is energetically costly. The identical growth rates exhibited by all behavioural types indicate that these higher activity costs of opportunistic behaviour were compensated for by increased prey consumption in the less favourable pelagic habitat resulting in approximately equal fitness of all pike groups. We conclude that behavioural diversification in habitat use and activity reduces intraspecific competition in preferred littoral habitats. This may facilitate the emergence of an ideal free distribution of pike along resource gradients.     

Indirect effects of human-induced environmental change on offspring production mediated by behavioural responses

Human-induced rapid environmental changes often cause behavioural alterations in animals. The consequences that these alterations in turn have for the viability of populations are, however, poorly known. We used a population of threespine sticklebacks Gasterosteus aculeatus in the Baltic Sea to investigate the consequences of behavioural responses to human-induced eutrophication for offspring production. The investigated population has been growing during the last decades, and one cause could be increased offspring production. We combined field-based surveys with laboratory-based experiments, and found that an enhanced growth of macroalgae relaxed agonistic interactions among males. This allowed more males to nest, improved hatching success, and increased the number of reproductive cycles that males completed. Thus, the behavioural responses were adaptive at the individual level and increased offspring production. However, a larger proportion of small males of low competitive ability reproduced in dense vegetation. As male size and dominance are heritable, this could influence the genetic composition of the offspring. Together with a higher number of offspring produced, this could influence natural selection and the rate of adaptation to the changing environment. Thus, behavioural responses to a rapid human-induced environmental change can influence offspring production, with potential consequences for population dynamics and evolutionary processes.     

Population genetic patterns suggest a behavioural change in wild common frogs (Rana temporaria) following disease outbreaks (Ranavirus)

We use 14 microsatellite loci to investigate the impact of a viral disease ( Ranavirus ) on the population genetic structure of wild common frogs ( Rana temporaria ). Populations with a history of Ranavirus mortalities (and 83% declines in the number of frogs) were compared with populations with no history of infection. Infected ponds showed significantly elevated F _IS (homozygote excess), significantly reduced relatedness, and no detectable effect on allelic richness. We hypothesize that the elevated F _IS and reduced relatedness are consequences of assortative mating, and that allelic richness is maintained by immigration from nearby populations. Simulations indicate that the elevated F _IS cannot be explained by population size reductions, but can indeed be explained by assortative mating (even if a mate choice locus is unlinked to the genetic markers). While the majority of studies consider demographic outcomes following disease outbreaks, our results indicate that emerging infectious diseases could also result in behavioural changes.     

Integral projection models perform better for small demographic data sets than matrix population models: a case study of two perennial herbs

1. Matrix population models are widely used to describe population dynamics, conduct population viability analyses and derive management recommendations for plant populations. For endangered or invasive species, management decisions are often based on small demographic data sets. Hence, there is a need for population models which accurately assess population performance from such small data sets.
2. We used demographic data on two perennial herbs with different life histories to compare the accuracy and precision of the traditional matrix population model and the recently developed integral projection model (IPM) in relation to the amount of data.
3. For large data sets both matrix models and IPMs produced identical estimates of population growth rate (λ). However, for small data sets containing fewer than 300 individuals, IPMs often produced smaller bias and variance for λ than matrix models despite different matrix structures and sampling techniques used to construct the matrix population models.
4. Synthesis and applications . Our results suggest that the smaller bias and variance of λ estimates make IPMs preferable to matrix population models for small demographic data sets with a few hundred individuals. These results are likely to be applicable to a wide range of herbaceous, perennial plant species where demographic fate can be modelled as a function of a continuous state variable such as size. We recommend the use of IPMs to assess population performance and management strategies particularly for endangered or invasive perennial herbs where little demographic data are available.     

Temperature-driven regime shifts in the dynamics of size-structured populations

Global warming impacts virtually all biota and ecosystems. Many of these impacts are mediated through direct effects of temperature on individual vital rates. Yet how this translates from the individual to the population level is still poorly understood, hampering the assessment of global warming impacts on population structure and dynamics. Here, we study the effects of temperature on intraspecific competition and cannibalism and the population dynamical consequences in a size-structured fish population. We use a physiologically structured consumer-resource model in which we explicitly model the temperature dependencies of the consumer vital rates and the resource population growth rate. Our model predicts that increased temperature decreases resource density despite higher resource growth rates, reflecting stronger intraspecific competition among consumers. At a critical temperature, the consumer population dynamics destabilize and shift from a stable equilibrium to competition-driven generation cycles that are dominated by recruits. As a consequence, maximum age decreases and the proportion of younger and smaller-sized fish increases. These model predictions support the hypothesis of decreasing mean body sizes due to increased temperatures. We conclude that in size-structured fish populations, global warming may increase competition, favor smaller size classes, and induce regime shifts that destabilize population and community dynamics.     

Stocking of captive-bred fish can cause long-term population decline and gene pool replacement: predictions from a population dynamics model incorporating density-dependent mortality

Releasing captive-bred fish into natural environments (stocking) is common in fisheries worldwide. Although stocking is believed to have a positive effect on fish abundance over the short term, little is known about the long-term consequences of recurrent stocking and its influence on natural populations. In fact, there are growing concerns that genetically maladapted captive-bred fish can eventually reduce the abundance of natural population. In this study, we develop a simple model to quantitatively investigate the condition under which recurrent stocking has long-term effects on the natural population. Using a population dynamics model that takes into account a density-dependent recruitment, a gene responsible for the fitness difference between wild and captive-bred fish, and hybridization between them, we show that there is little or no contribution of recurrent stocking to the stock enhancement without a replacement of the wild gene pool by the captive-bred gene pool. The model further predicted that stocking of an intermediate level causes a reduction, rather than enhancement, of population size over the long term. The population decline due to stocking was attributed to the fitness disadvantage of captive-bred fish and strong overcompensation at recruitment stage. These results suggest that it would be difficult to simultaneously attain population size recovery and conservation of the local gene pool when captive-bred fish have fitness disadvantage in the wild, although caution is needed when applying the predictions from the simplified model to a specific species or population.     

Effects of branching spatial structure and life history on the asymptotic growth rate of a population

The dendritic structure of a river network creates directional dispersal and a hierarchical arrangement of habitats. These two features have important consequences for the ecological dynamics of species living within the network. We apply matrix population models to a stage-structured population in a network of habitat patches connected in a dendritic arrangement. By considering a range of life histories and dispersal patterns, both constant in time and seasonal, we illustrate how spatial structure, directional dispersal, survival, and reproduction interact to determine population growth rate and distribution. We investigate the sensitivity of the asymptotic growth rate to the demographic parameters of the model, the system size, and the connections between the patches. Although some general patterns emerge, we find that a species’ modes of reproduction and dispersal are quite important in its response to changes in its life history parameters or in the spatial structure. The framework we use here can be customized to incorporate a wide range of demographic and dispersal scenarios.     

Understanding the multiple meanings of ‘inbreeding’ and ‘effective size’ for genetic management of African rhinoceros populations

Although some African rhinoceros populations are currently increasing, others are critically endangered. Even healthy populations are extensively managed in the wild and in captivity. While political and demographic considerations are of primary concern, many decisions are made in the name of genetic management. Such decisions should be informed by a full understanding of the multiple meanings of inbreeding and effective population size. In this essay, we examine inbreeding and effective size of wild and captive populations of African rhinoceroses. We conclude by showing how misunderstanding of effective size and Franklin’s 50/500 rule can make a crucial difference in informing management decisions.     

A general model of site-dependent population regulation: population-level regulation without individual-level interactions

In this paper, we use a modeling approach to explore the population regulatory consequences of individual choices for where to breed in heterogeneous environments. In contrast to standard models, we focus on individuals that interact only indirectly through their choices of breeding sites (i.e., individuals preempt the occupation of a breeding site by others when they choose to breed there). We consider the consequences of individuals choosing breeding sites either randomly or sequentially from best to worst. Our analysis shows that average per-capita fecundity of the population is independent of the number of occupied breeding sites if individuals choose sites at random and that variation in average per-capita fecundity increases as population size declines. In contrast, if individuals choose breeding sites sequentially from highest to lowest quality, then as population size increases average per-capita fecundity declines and variation in average per-capita fecundity increases. Consequently, aggregate population-level demographic rates can change in ways that generate population regulation, even when change in population size does not change the demographic performance of any individual on any particular breeding site. However, such regulation occurs only when individuals make adaptive choices of where to breed. Because variation in average per-capita fecundity decreases when population size declines, populations regulated in a site-dependent manner should be much less susceptible to the vicissitudes of small population size than those which choose breeding sites at random.     

When phenology matters: age–size truncation alters population response to trophic mismatch

Climate-induced shifts in the timing of life-history events are a worldwide phenomenon, and these shifts can de-synchronize species interactions such as predator–prey relationships. In order to understand the ecological implications of altered seasonality, we need to consider how shifts in phenology interact with other agents of environmental change such as exploitation and disease spread, which commonly act to erode the demographic structure of wild populations. Using long-term observational data on the phenology and dynamics of a model predator–prey system (fish and zooplankton in Windermere, UK), we show that age–size truncation of the predator population alters the consequences of phenological mismatch for offspring survival and population abundance. Specifically, age–size truncation reduces intraspecific density regulation due to competition and cannibalism, and thereby amplifies the population sensitivity to climate-induced predator–prey asynchrony, which increases variability in predator abundance. High population variability poses major ecological and economic challenges as it can diminish sustainable harvest rates and increase the risk of population collapse. Our results stress the importance of maintaining within-population age–size diversity in order to buffer populations against phenological asynchrony, and highlight the need to consider interactive effects of environmental impacts if we are to understand and project complex ecological outcomes.     

Temperature‐dependent body size effects determine population responses to climate warming

Current understanding of animal population responses to rising temperatures is based on the assumption that biological rates such as metabolism, which governs fundamental ecological processes, scale independently with body size and temperature, despite empirical evidence for interactive effects. Here, we investigate the consequences of interactive temperature‐ and size scaling of vital rates for the dynamics of populations experiencing warming using a stage‐structured consumer‐resource model. We show that interactive scaling alters population and stage‐specific responses to rising temperatures, such that warming can induce shifts in population regulation and stage‐structure, influence community structure and govern population responses to mortality. Analysing experimental data for 20 fish species, we found size–temperature interactions in intraspecific scaling of metabolic rate to be common. Given the evidence for size–temperature interactions and the ubiquity of size structure in animal populations, we argue that accounting for size‐specific temperature effects is pivotal for understanding how warming affects animal populations and communities.