Long-term Population Growth Rates Research Articles

Unclear relationships between mean survival rate and its environmental variance in vertebrates.

Current environmental changes may increase temporal variability of life history traits of species thus affecting their long-term population growth rate and extinction risk. If there is a general relationship between environmental variances (EVs) and mean annual survival rates of species, that relationship could be used as a guideline for analyses of population growth and extinction risk for populations, where data on EVs are missing. For this purpose, we present a comprehensive compilation of 252 EV estimates from 89 species belonging to five vertebrate taxa (birds, mammals, reptiles, amphibians and fish) covering mean annual survival rates from 0.01 to 0.98. Since variances of survival rates are constrained by their means, particularly for low and high mean survival rates, we assessed whether any observed relationship persisted after applying two types of commonly used variance stabilizing transformations: relativized EVs (observed/mathematical maximum) and logit-scaled EVs. With raw EVs at the arithmetic scale, mean-variance relationships of annual survival rates were hump-shaped with small EVs at low and high mean survival rates and higher (and widely variable) EVs at intermediate mean survival rates. When mean annual survival rates were related to relativized EVs the hump-shaped pattern was less distinct than for raw EVs. When transforming EVs to logit scale the relationship between mean annual survival rates and EVs largely disappeared. The within-species juvenile-adult slopes were mainly positive at low (<0.5) and negative at high (>0.5) mean survival rates for raw and relativized variances while these patterns disappeared when EVs were logit transformed. Uncertainties in how to interpret the results of relativized and logit-scaled EVs, and the observed high variation in EV's for similar mean annual survival rates illustrates that extrapolations of observed EVs and tests of life history drivers of survival-EV relationships need to also acknowledge the large variation in these parameters.

Coevolution of Patch Selection in Stochastic Environments.

AbstractSpecies interact in landscapes where environmental conditions vary in time and space. This variability impacts how species select habitat patches. Under equilibrium conditions, evolution of this patch selection can result in ideal free distributions where per capita growth rates are zero in occupied patches and negative in unoccupied patches. These ideal free distributions, however, do not explain why species occupy sink patches, why competitors have overlapping spatial ranges, or why predators avoid highly productive patches. To understand these patterns, we solve for coevolutionarily stable strategies (coESSs) of patch selection for multispecies stochastic Lotka-Volterra models accounting for spatial and temporal heterogeneity. In occupied patches at the coESS, we show that the differences between the local contributions to the mean and the variance of the long-term population growth rate are equalized. Applying this characterization to models of antagonistic interactions reveals that environmental stochasticity can partially exorcize the ghost of competition past, select for new forms of enemy-free and victimless space, and generate hydra effects over evolutionary timescales. Viewing our results through the economic lens of modern portfolio theory highlights why the coESS for patch selection is often a bet-hedging strategy coupling stochastic sink populations. Our results highlight how environmental stochasticity can reverse or amplify evolutionary outcomes as a result of species interactions or spatial heterogeneity.

A demographic approach for predicting population responses to multifactorial stressors.

Populations face a suite of anthropogenic stressors acting simultaneously, which can combine additively or interact to have complex effects on population persistence. Yet we still know relatively little about the mechanisms underlying population-level responses to multifactorial combinations of stressors because multiple stressor impacts across organisms' life cycles have not been systematically considered in population models. Specifically, different anthropogenic stressors can have variable effects across an organism's life cycle, resulting in non-intuitive results for long-term population persistence. For example, synergistic or antagonistic interactions might exacerbate or alleviate the effects of stressors on population dynamics, and different life-history stages or vital rates might contribute unequally to long-term population growth rates. Demographic modelling provides a framework to incorporate individual vital rate responses to multiple stressors into estimates of population growth, which will allow us to make more informed predictions about population-level responses to novel combinations of anthropogenic change. Without integrating stressors' interactive effects across the entire life cycle on population persistence, we may over- or underestimate threats to biodiversity and risk missing conservation management actions that could reduce species' vulnerability to stress.

Optimal energy allocation trade-off driven by size-dependent physiological and demographic responses to warming.

Body size-dependent physiological effects of temperature influence individual growth, reproduction, and survival, which govern animal population responses to global warming. Considerable knowledge has been established on how such effects can affect population growth and size structure, but less is known of their potential role in temperature-driven adaptation in life-history traits. In this study, we ask how warming affects the optimal allocation of energy between growth and reproduction and disentangle the underlying fitness trade-offs. To this end, we develop a novel dynamic energy budget integral projection model (DEB-IPM), linking individuals' size- and temperature-dependent consumption and maintenance via somatic growth, reproduction, and size-dependent energy allocation to emergent population responses. At the population level, we calculate the long-term population growth rate (fitness) and stable size structure emerging from demographic processes. Applying the model to an example of pike (Esox lucius), we find that optimal energy allocation to growth decreases with warming. Furthermore, we demonstrate how growth, fecundity, and survival contribute to this change in optimal allocation. Higher energy allocation to somatic growth at low temperatures increases fitness through survival of small individuals and through the reproduction of larger individuals. In contrast, at high temperatures, increased allocation to reproduction is favored because warming induces faster somatic growth of small individuals and increased fecundity but reduced growth and higher mortality of larger individuals. Reduced optimum allocation to growth leads to further reductions in body size and an increasingly truncated population size structure with warming. Our study demonstrates how, by incorporating general physiological mechanisms driving the temperature dependence of life-history traits, the DEB-IPM framework is useful for investigating the adaptation of size-structured organisms to warming.

Spatial consistency in drivers of population dynamics of a declining migratory bird.

Many migratory species are in decline across their geographical ranges. Single-population studies can provide important insights into drivers at a local scale, but effective conservation requires multi-population perspectives. This is challenging because relevant data are often hard to consolidate, and state-of-the-art analytical tools are typically tailored to specific datasets. We capitalized on a recent data harmonization initiative (SPI-Birds) and linked it to a generalized modelling framework to identify the demographic and environmental drivers of large-scale population decline in migratory pied flycatchers (Ficedula hypoleuca) breeding across Britain. We implemented a generalized integrated population model (IPM) to estimate age-specific vital rates, including their dependency on environmental conditions, and total and breeding population size of pied flycatchers using long-term (34-64 years) monitoring data from seven locations representative of the British breeding range. We then quantified the relative contributions of different vital rates and population structure to changes in short- and long-term population growth rate using transient life table response experiments (LTREs). Substantial covariation in population sizes across breeding locations suggested that change was the result of large-scale drivers. This was supported by LTRE analyses, which attributed past changes in short-term population growth rates and long-term population trends primarily to variation in annual survival and dispersal dynamics, which largely act during migration and/or nonbreeding season. Contributions of variation in local reproductive parameters were small in comparison, despite sensitivity to local temperature and rainfall within the breeding period. We show that both short- and long-term population changes of British breeding pied flycatchers are likely linked to factors acting during migration and in nonbreeding areas, where future research should be prioritized. We illustrate the potential of multi-population analyses for informing management at (inter)national scales and highlight the importance of data standardization, generalized and accessible analytical tools, and reproducible workflows to achieve them.

Application of modern coexistence theory to rare plant restoration provides early indication of restoration trajectories.

Restoration ecology commonly seeks to re-establish species of interest in degraded habitats. Despite a rich understanding of how succession influences re-establishment, there are several outstanding questions that remain unaddressed: are short-term abundances sufficient to determine long-term re-establishment success, and what factors contribute to unpredictable restorations outcomes? In other words, when restoration fails, is it because the restored habitat is substandard, because of strong competition with invasive species, or alternatively due to changing environmental conditions that would equally impact established populations? Here, we re-purpose tools developed from modern coexistence theory to address these questions, and apply them to an effort to restore the endangered Contra Costa goldfields (Lasthenia conjugens) in constructed ("restored") California vernal pools. Using 16 years of data, we construct a population model of L. conjugens, a species of conservation concern due primarily to habitat loss and invasion of exotic grasses. We show that initial, short-term appearances of restoration success from population abundances is misleading, as year-to-year fluctuations cause long-term population growth rates to fall below zero. The failure of constructed pools is driven by lower maximum growth rates compared with reference ("natural") pools, coupled with a stronger negative sensitivity to annual fluctuations in abiotic conditions that yield decreased maximum growth rates. Nonetheless, our modeling shows that fluctuations in competition (mainly with exotic grasses) benefit L. conjugens through periods of competitive release, especially in constructed pools of intermediate pool depth. We therefore show how reductions in invasives and seed addition in pools of particular depths could change the outcome of restoration for L. conjugens. By applying a largely theoretical framework to the urgent goal of ecological restoration, our study provides a blueprint for predicting restoration success, and identifies future actions to reverse species loss.

Effects of habitat disturbance and rainfall on a dominant medicinal dry forest tree

The interactive effects of anthropogenic disturbance and climatic variables can shape the demography of plant populations, but they remain poorly understood especially for long-lived species like tropical trees. Understanding these interactions is critical for designing forest management strategies in the face of both growing anthropogenic pressures and climate change. We explored the effects of habitat disturbance and interannual variation in rainfall on the population dynamics of Amphipterygium adstringens (Anacardiaceae), an ecologically dominant tree species endemic to the tropical deciduous forests of Mexico, that is heavily harvested for its medicinal bark. Specifically, we compared A. adstringens vital rates and long-term population growth rates of unharvested populations in a disturbed versus a protected tropical deciduous forest site; and explored the effects of inter-annual variation in annual rainfall on A. adstringens by comparing vital rates and population growth rates across three years with differing levels of rainfall. We established four plots in each of the two sites and monitored survival, growth and reproduction of 251 individuals over three years. We found that anthropogenic disturbance and rainfall differentially affected the vital rates of A. adstringens. Survival did not differ across sites or years. Growth tended to be lower and fecundity higher, in the protected site, but these effects varied across years and as a function of plant size. No seedlings were found in the disturbed site over the three years study period. Integral projection models showed that projected population growth rates (λ values) in the protected site were higher and much less variable over time than those of the disturbed site. Lambda values in the disturbed site were <1, indicating long-term population decline. In both sites, the rainiest year showed the highest population growth rate but the increase in λ was much higher in the disturbed site than in the protected site. These findings suggest that the population dynamics of A. adstringens are heavily affected by habitat disturbance forest management regime and that the impacts of interannual variability in rainfall or other environmental conditions may be exacerbated in disturbed conditions. The persistence of A. adstringens populations over time will require the exclusion of livestock as well as the implementation of silvicultural practices that will lead to seedling establishment and survival. This study provides key data for the management and conservation of A. adstringens, a highly valued medicinal species.

The Disconnect Between Short- and Long-Term Population Projections for Plant Reintroductions

The reintroduction of rare species in natural preserves is a commonly used restoration strategy to prevent species extinction. An essential first step in planning successful reintroductions is identifying which life stages (e.g., seeds or large adults) should be used to establish these new populations. Following this initial establishment phase, it is necessary to determine the level of survival, growth, and recruitment needed to maintain population persistence over time and identify management actions that will achieve these goals. In this 5-year study, we projected the short- and long-term population growth rates of a critically endangered long-lived shrub, Delissea waianaeensis. Using this model system, we show that reintroductions established with mature individuals have the lowest probability of quasi-population extinction (10 individuals) and the highest increase in population abundance. However, our results also demonstrate that short-term increases in population abundances are overly optimistic of long-term outcomes. Using long-term stochastic model simulations, we identified the level of natural seedling regeneration needed to maintain a positive population growth rate over time. These findings are relevant for planning future reintroduction efforts for long-lived species and illustrate the need to forecast short- and long-term population responses when evaluating restoration success.

Archaeology, demography and life history theory together can help us explain past and present population patterns.

Population matters. Demographic patterns are both a cause and a consequence of human behaviour in other important domains, such as subsistence, cooperation, politics and culture. Demographers interested in contemporary and recent historical populations have rich data at their fingertips; the importance of demography means many interested parties have gathered demographic data, much of which is now readily available for all to explore. Those interested in the demography of the distant past are not so fortunate, given the lack of written records. Nevertheless, the emergence in recent years of a new interest in the demography of ancient populations has seen the development of a range of new methods for piecing together archaeological, skeletal and DNA evidence to reconstruct past population patterns. These efforts have found evidence in support of the view that the relatively low long-term population growth rates of prehistoric human populations, albeit ultimately conditioned by carrying capacities, may have been owing to 'boom-bust' cycles at the regional level; rapid population growth, followed by population decline. In fact, this archaeological research may have come to the same conclusion as some contemporary demographers: that demography can be remarkably hard to predict, at least in the short term. It also fits with evidence from biology that primates, and particularly humans, may be adapted to environmental variability, leading to associated demographic stochasticity. This evidence of the fluctuating nature of human demographic patterns may be of considerable significance in understanding our species' evolution, and of understanding what our species future demographic trajectories might be. This article is part of the theme issue 'Cross-disciplinary approaches to prehistoric demography'.

Annual mowing has the potential to reduce the invasion of herbaceous Lupinus polyphyllus

In order to manage invasive plant species efficiently, it is necessary to have a thorough understanding of different strategies of population control, including the underlying mechanisms of action and the consequences for target populations. Here, I explored the effectiveness of biomass removal as a method of control for the invasive perennial herb Lupinus polyphyllus. More specifically, using seed material from 11 populations, I assessed among-population variation (if any) in plant compensatory growth as a response to annual biomass removal under standardised growing conditions over two consecutive years, and quantified the demographic effects of a single biomass-removal event. In all study populations, annual biomass removal reduced plant size, flowering probability, and shoot and root biomass. Biomass removal also reduced plant survival and the number of flowering shoots, but these effects were pronounced at certain time points only. A population-level demographic analysis revealed that a single biomass-removal event considerably decreased the long-term population growth rate (λ); this decline in λ was due to a reduction in plant fecundity followed by survival and growth. These findings suggest that annual mowing has the potential to curb invasions of L. polyphyllus because plants are not able to fully compensate for drastic biomass loss.

Apparent Survival and Long-Term Population Growth Rate of The Kentish Plover Charadrius alexandrinus in Mallorca, Archipelago, Spain

During the last few decades, many breeding waders have been declining worldwide, probably due to the reduction of suitable coastal habitats. Diagnosis of population parameters has become increasingly important for the conservation of waders. We used capture-recapture information of 214 adult Kentish Plovers Charadrius alexandrinus, marked between 1977 and 2015 at two breeding areas in Mallorca, Spain, to assess any possible temporal trend in the survival probability. Mean adult survival probability was 0.756 ± 0.05 with no differences between the two areas. The sparseness of the data did not permit robust estimates of yearly survival. However, models with a temporal trend in survival probability were not retained. An age-structured population model including demographic stochasticity suggested a stable population (λ = 1.024 ± 0.09). Perturbation analyses indicated that a 10% increase in mean breeding success (from 0.44 to 0.63), by reducing habitat loss or nest predation, would be sufficient to obtain positive population trajectories. —Garcias, P.J. & Tavecchia, G. (2018). Apparent survival and long-term population growth rate of the Kentish Plover Charadrius alexandrinus in Mallorca, Balearic Archipelago, Spain. Ardeola, 65: 59–68.

The generality of management recommendations across populations of an invasive perennial herb

Demographic models are widely used to produce management recommendations for different species. For invasive plants, current management recommendations to control local population growth are often based on data from a limited number of populations per species, and the assumption of stable population structure (asymptotic dynamics). However, spatial variation in population dynamics and deviation from a stable structure may affect these recommendations, calling into question their generality across populations of an invasive species. Here, I focused on intraspecific variation in population dynamics and investigated management recommendations generated by demographic models across 37 populations of a short-lived, invasive perennial herb (Lupinus polyphyllus). Models that relied on the proportional perturbations of vital rates (asymptotic elasticities) indicated an essential role for plant survival in long-term population dynamics. The rank order of elasticities for different vital rates (survival, growth, retrogression, fecundity) varied little among the 37 study populations regardless of population status (increasing or declining asymptotically). Summed elasticities for fecundity increased, while summed elasticities for survival decreased with increasing long-term population growth rate. Transient dynamics differed from asymptotic dynamics, but were qualitatively similar among populations, that is, depending on the initial size structure, populations tended to either increase or decline in density more rapidly than predicted by asymptotic growth rate. These findings indicate that although populations are likely to exhibit transient dynamics, management recommendations based on asymptotic elasticities for vital rates might be to some extent generalised across established populations of a given short-lived invasive plant species.

Quantifying long-term population growth rates of threatened bull trout: challenges, lessons learned, and opportunities

Temporal symmetry models (TSM) represent advances in the analytical application of mark–recapture data to population status assessments. For a population of char, we employed 10 years of active and passive mark–recapture data to quantify population growth rates using different data sources and analytical approaches. Estimates of adult population growth rate were 1.01 (95% confidence interval = 0.84–1.20) using a temporal symmetry model (λTSM), 0.96 (0.68–1.34) based on logistic regressions of annual snorkel data (λA), and 0.92 (0.77–1.11) from redd counts (λR). Top-performing TSMs included an increasing time trend in recruitment (f) and changes in capture probability (p). There was only a 1% chance the population decreased ≥50%, and a 10% chance it decreased ≥30% (λMCMC; based on Bayesian Markov chain Monte Carlo procedure). Size structure was stable; however, the adult population was dominated by small adults, and over the study period there was a decline in the contribution of large adults to total biomass. Juvenile condition decreased with increasing adult densities. Utilization of these different information sources provided a robust weight-of-evidence approach to identifying population status and potential mechanisms driving changes in population growth rates.

Continent-scale global change attribution in European birds - combining annual and decadal time scales.

Species attributes are commonly used to infer impacts of environmental change on multiyear species trends, e.g. decadal changes in population size. However, by themselves attributes are of limited value in global change attribution since they do not measure the changing environment. A broader foundation for attributing species responses to global change may be achieved by complementing an attributes-based approach by one estimating the relationship between repeated measures of organismal and environmental changes over short time scales. To assess the benefit of this multiscale perspective, we investigate the recent impact of multiple environmental changes on European farmland birds, here focusing on climate change and land use change. We analyze more than 800 time series from 18 countries spanning the past two decades. Analysis of long-term population growth rates documents simultaneous responses that can be attributed to both climate change and land-use change, including long-term increases in populations of hot-dwelling species and declines in long-distance migrants and farmland specialists. In contrast, analysis of annual growth rates yield novel insights into the potential mechanisms driving long-term climate induced change. In particular, we find that birds are affected by winter, spring, and summer conditions depending on the distinct breeding phenology that corresponds to their migratory strategy. Birds in general benefit from higher temperatures or higher primary productivity early on or in the peak of the breeding season with the largest effect sizes observed in cooler parts of species' climatic ranges. Our results document the potential of combining time scales and integrating both species attributes and environmental variables for global change attribution. We suggest such an approach will be of general use when high-resolution time series are available in large-scale biodiversity surveys.

Genetic variation facilitates seedling establishment but not population growth rate of a perennial invader.

Assessing the demographic consequences of genetic variation is fundamental to invasion biology. However, genetic and demographic approaches are rarely combined to explore the effects of genetic variation on invasive populations in natural environments. This study combined population genetics, demographic data and a greenhouse experiment to investigate the consequences of genetic variation for the population fitness of the perennial, invasive herb Lupinus polyphyllus. Genetic and demographic data were collected from 37 L. polyphyllus populations representing different latitudes in Finland, and genetic variation was characterized based on 13 microsatellite loci. Associations between genetic variation and population size, population density, latitude and habitat were investigated. Genetic variation was then explored in relation to four fitness components (establishment, survival, growth, fecundity) measured at the population level, and the long-term population growth rate (λ). For a subset of populations genetic variation was also examined in relation to the temporal variability of λ. A further assessment was made of the role of natural selection in the observed variation of certain fitness components among populations under greenhouse conditions. It was found that genetic variation correlated positively with population size, particularly at higher latitudes, and differed among habitat types. Average seedling establishment per population increased with genetic variation in the field, but not under greenhouse conditions. Quantitative genetic divergence (Q(ST)) based on seedling establishment in the greenhouse was smaller than allelic genetic divergence (F'(ST)), indicating that unifying selection has a prominent role in this fitness component. Genetic variation was not associated with average survival, growth or fecundity measured at the population level, λ or its variability. The study suggests that although genetic variation may facilitate plant invasions by increasing seedling establishment, it may not necessarily affect the long-term population growth rate. Therefore, established invasions may be able to grow equally well regardless of their genetic diversity.

Jensen's Inequality and the Impact of Short-Term Environmental Variability on Long-Term Population Growth Rates.

It is well established in theory that short-term environmental fluctuations could affect the long-term growth rates of wildlife populations, but this theory has rarely been tested and there remains little empirical evidence that the effect is actually important in practice. Here we develop models to quantify the effects of daily, seasonal, and yearly temperature fluctuations on the average population growth rates, and we apply them to long-term data on the endangered Black-faced Spoonbill (Platalea minor); an endothermic species whose population growth rates follow a concave relationship with temperature. We demonstrate for the first time that the current levels of temperature variability, particularly seasonal variability, are already large enough to substantially reduce long-term population growth rates. As the climate changes, our results highlight the importance of considering the ecological effects of climate variability and not just average conditions.

Well-temperate phage: optimal bet-hedging against local environmental collapses.

Upon infection of their bacterial hosts temperate phages must chose between lysogenic and lytic developmental strategies. Here we apply the game-theoretic bet-hedging strategy introduced by Kelly to derive the optimal lysogenic fraction of the total population of phages as a function of frequency and intensity of environmental downturns affecting the lytic subpopulation. “Well-temperate” phage from our title is characterized by the best long-term population growth rate. We show that it is realized when the lysogenization frequency is approximately equal to the probability of lytic population collapse. We further predict the existence of sharp boundaries in system’s environmental, ecological, and biophysical parameters separating the regions where this temperate strategy is optimal from those dominated by purely virulent or dormant (purely lysogenic) strategies. We show that the virulent strategy works best for phages with large diversity of hosts, and access to multiple independent environments reachable by diffusion. Conversely, progressively more temperate or even dormant strategies are favored in the environments, that are subject to frequent and severe temporal downturns.

Environmental variation and population responses to global change

Species' responses to environmental changes such as global warming are affected not only by trends in mean conditions, but also by natural and human-induced environmental fluctuations. Methods are needed to predict how such environmental variation affects ecological and evolutionary processes, in order to design effective strategies to conserve biodiversity under global change. Here, we review recent theoretical and empirical studies to assess: (1) how populations respond to changes in environmental variance, and (2) how environmental variance affects population responses to changes in mean conditions. Contrary to frequent claims, empirical studies show that increases in environmental variance can increase as well as decrease long-term population growth rates. Moreover, environmental variance can alter and even reverse the effects of changes in the mean environment, such that even if environmental variance remains constant, omitting it from population models compromises their ability to predict species' responses to changes in mean conditions. Drawing on theory relating these effects of environmental variance to the curvatures of population growth responses to the environment, we outline how species' traits such as phylogenetic history and body mass could be used to predict their responses to global change under future environmental variability.

Homing fidelity and reproductive rate for migratory populations

Short-term and long-term population growth rates can differ considerably. While changes in growth rates can be driven by external factors, we consider another source for changes in growth rate. That is, changes are generated internally by gradual modification of population structure. Such a modification of population structure may take many generations, particularly when the populations are distributed spatially in heterogeneous environments. Here, the net reproductive rate R0 is not sufficient to char- acterize short-term growth. Indeed, a population with net reproductiverate greater than one couldinitially declinepre- cipitously, or a population with net reproductive rate less than one could initially grow substantially. Thus, we aug- ment the net reproductive rate with lower and upper bounds for the transient reproductive rate, Rl and Ru. We apply these measures to the study of spatially structured salmon populations and show the effect of variable homing fidelity on short-term and long-term generational growth rates.

Frequency-dependent population dynamics: Effect of sex ratio and mating system on the elasticity of population growth rate

When vital rates depend on population structure (e.g., relative frequencies of males or females), an important question is how the long-term population growth rate λ responds to changes in rates. For instance, availability of mates may depend on the sex ratio of the population and hence reproductive rates could be frequency-dependent. In such cases change in any vital rate alters the structure, which in turn, affect frequency-dependent rates. We show that the elasticity of λ to a rate is the sum of (i) the effect of the linear change in the rate and (ii) the effect of nonlinear changes in frequency-dependent rates. The first component is always positive and is the classical elasticity in density-independent models obtained directly from the population projection matrix. The second component can be positive or negative and is absent in density-independent models. We explicitly express each component of the elasticity as a function of vital rates, eigenvalues and eigenvectors of the population projection matrix. We apply this result to a two-sex model, where male and female fertilities depend on adult sex ratio α (ratio of females to males) and the mating system (e.g., polygyny) through a harmonic mating function. We show that the nonlinear component of elasticity to a survival rate is negligible only when the average number of mates (per male) is close to α. In a strictly monogamous species, elasticity to female survival is larger than elasticity to male survival when α<1 (less females). In a polygynous species, elasticity to female survival can be larger than that of male survival even when sex ratio is female biased. Our results show how demography and mating system together determine the response to selection on sex-specific vital rates.