Stochastic Population Growth Rate Research Articles

Life history costs drive the evolution of mycoheterotrophs: Increased sprouting and flowering in a strongly mycoheterotrophic Pyrola species

Abstract Land plants are typically photosynthetic, but some species have lost the ability to photosynthesize, instead relying on mycorrhizal fungi to obtain carbon. Increasing levels of partial mycoheterotrophy, in which seemingly autotrophic plants receive fungal carbon from their fungal partners, and reduced sprouting in concert with greater reproduction when sprouting, may be intermediate steps in the evolution of this trait. We studied the microevolutionary demography of Pyrola japonica and the closely related species P. subaphylla, which are currently considered sister species. While these species are both partially mycoheterotrophic, P. subaphylla is more strongly so, indicating that it may be closer to evolving full mycoheterotrophy. We tracked individuals in two nearby populations in Fukushima, Japan from 2015 to 2020. We analysed vital rates, population trends and long‐run population structure with historical function‐based matrix projection models. We assessed how shifts in P. subaphylla demography relate to fitness using a stochastic life table response experiment (SLTRE). P. subaphylla exhibited strong costs of growth to survival and of reproduction to sprouting, size and fecundity. It sprouted more than P. japonica but to smaller stages. P. subaphylla's flowering frequency was approximately four times that of P. japonica, but after flowering it was smaller and more likely to die. These life history costs appeared to drive differences in fitness, measured as stochastic population growth rate. Relative to P. japonica, shifts in fitness in P. subaphylla were due more to shifts in means than standard deviations of matrix elements. Shifts in growth transitions had the strongest negative impacts on fitness but also had the strongest positive impacts. Increased shifts to small non‐flowering stages were associated with increased fitness, while transitions involving dormancy were associated more strongly with drops in fitness. Synthesis. As far as we are aware, we found the first evidence that costs of growth and reproduction drive the evolution of increased sprouting and smaller size as a species becomes more intensely mycoheterotrophic. Surprisingly, vegetative dormancy did not evolve with mycoheterotrophy. We suggest further studies assessing whether these seemingly maladaptive trends result from genetic drift, genetic linkage or another mechanism.

Contrasting demographic responses under future climate for two populations of a montane amphibian

For species with complex life histories, climate change can have contrasting effects for different life stages within locally adapted populations and may result in responses counter to general climate change predictions. Using data from two, 14-year demographic studies for a North American montane amphibian, Cascades frog (Rana cascadae), we quantified how aspects of current climate influenced annual survival of larvae and adult stages and modeled the stochastic population growth rate (λs) of each population for current (1980–2006) and future periods (2080s). Climate drivers of survival for the populations were similar for larvae (i.e., decreases in precipitation lead to pond drying and mortality), but diverged for terrestrial stages where decreases in winter length and summer precipitation had opposite effects. By the 2080s, we predict one population will be in sharp decline (λs = 0.90), while the other population will remain nearly stable (λs = 0.99) in the absence of other stressors, such as mortality due to disease. Our case study demonstrates a result counter to many climate envelope predictions in that stage-specific responses to local climate and hydrology result in a higher extinction risk for the more northern population.

Consequences of cross-season demographic correlations for population viability.

Demographic correlations are pervasive in wildlife populations and can represent important secondary drivers of population growth. Empirical evidence suggests that correlations are in general positive for long-lived species, but little is known about the degree of variation among spatially segregated populations of the same species in relation to environmental conditions. We assessed the relative importance of two cross-season correlations in survival and productivity, for three Atlantic puffin (Fratercula arctica) populations with contrasting population trajectories and non-overlapping year-round distributions. The two correlations reflected either a relationship between adult survival prior to breeding on productivity, or a relationship between productivity and adult survival the subsequent year. Demographic rates and their correlations were estimated with an integrated population model, and their respective contributions to variation in population growth were calculated using a transient-life table response experiment. For all three populations, demographic correlations were positive at both time lags, although their strength differed. Given the different year-round distributions of these populations, this variation in the strength population-level demographic correlations points to environmental conditions as an important driver of demographic variation through life-history constraints. Consequently, the contributions of variances and correlations in demographic rates to population growth rates differed among puffin populations, which has implications for-particularly small-populations' viability under environmental change as positive correlations tend to reduce the stochastic population growth rate.

Transient population dynamics drive the spread of invasive wild pigs and reveal impacts of management in North America

Invasion of nonindigenous species is considered one of the most urgent problems affecting native ecosystems and agricultural systems. Mechanistic models that account for short-term population dynamics can improve prediction because they incorporate differing demographic processes that link the environmental conditions of a spatial location explicitly with the invasion process. Yet short-term population dynamics are rarely accounted for in spatial models of invasive species spread.Accounting for transient (short-term) population dynamics that arise from the interaction of age structure and vital rates, we predict the stochastic population growth rate and establishment probability of wild pigs following introduction into any location in North America. Established ecological theory suggests that the rate of spatial spread is proportional to population growth rate. Using observed geographic distribution data for wild pigs we calculated geographic spread rates (watersheds/year) from 1982 to 2021. We investigated if observed spread rates increased in watersheds with higher stochastic population growth rates. Stochastic population growth rate and establishment probability of wild pigs increased with increasing initial population (propagule) size and length of establishment time. Areas along the Mississippi, Ohio, and lower portions of the Missouri river drainages had the highest probability of wild pig establishment with many regions having probabilities close to 1. Spread rates demonstrated strong spatial heterogeneity with the greatest rates of spread (5.8 watersheds/year) occurring from 2008 to 2013 prior to the establishment of a National wild pig control program in 2013. Spread rates declined 82% (0.57 watersheds/year) in the period from 2013 to 2021 compared to the period from 1982 to 2013. We found significant positive associations among stochastic population growth rate and observed geographic rates of spread. Stochastic population growth rate explained a large amount of variation (79.3–92.1%) in annual rate of watershed spread of wild pigs. Our predicted probabilities of establishment and population growth can be used to inform surveillance and control efforts to reduce the potential for establishment and spread of wild pigs.

Anther smut pathogens as important drivers of population dynamics of long-lived perennial plants: A case study of Dianthus carthusianorum

Pollinator-transmitted pathogens typically hinder sexual reproduction of their hosts and affect pollen flow among remaining healthy individuals in a population. The extent to which a pathogen also influences host’s population growth depends on the importance of sexual reproduction for the host’s life cycle. Such pathogen impact cannot be traced by measuring only the vital rates directly affected by the pathogen, and thus a study of the host’s entire life cycle is necessary. In this study, we aimed to quantify the effects of the pollinator-transmitted anther smut pathogen Microbotryum carthusianorum on population growth rate in three populations of the long-lived perennial Dianthus carthusianorum. We followed plant individuals over three years and measured their size, disease state, and reproduction. We then constructed an Integral Projection Model (IPM). To evaluate the pathogen impact, we performed a stochastic analysis of the IPM for real diseased populations as well as for simulated populations without the pathogen. As the populations also hosted predispersal seed predators, the same approach was used to evaluate their impact. Stochastic population growth rates indicated two of the real populations to be increasing, and one to be declining. Comparison with the simulated healthy populations showed that the pathogen impact on the growth rate was negative and relatively strong, because the growth rate was highly sensitive to changes in sexual reproduction. However, the pathogen did not appear to cause the decline in the one decreasing population, since the growth rate there was impaired more substantially by high rates of predispersal seed predation and low germination rates than by the castration of diseased flowers. Overall, our study suggests that D. carthusianorum is highly vulnerable to biotic interactions affecting sexual reproduction pathway. Additionally, our study illustrated several complexities in disease dynamics (e.g., occurrence of partially or fully asymptomatic plants) that need to be incorporated into the assessment of the impact of pollinator-transmitted pathogens on long-lived perennials.

Demographic consequences of management actions for the successful reintroduction of the White Stork Ciconia ciconia to the UK

Summary Species reintroductions can be used in conservation management to increase biodiversity and aid in restoring ecosystem function. For reintroductions to be successful, it is important to identify the conditions required to establish a viable population. We developed a demographic model using Vortex10, an individual-based simulation software, to assess the long-term consequences of different management interventions on the success of the recent reintroduction of the white stork, Ciconia ciconia, to the UK. Demographic data obtained from this reintroduced population were supplemented with information from western European populations to build the models. The impact of incorporating different management actions (e.g. supplementing with captive-bred juveniles, provision of nesting platforms, and habitat improvement/supplementary feeding) on the stochastic population growth rate was assessed. Survival rates also differ depending on an individual’s migratory strategy, hence we tested the impact of having different proportions of the population as residents or migrants. Our models showed that if the British stork population adopts a fully migratory strategy, with its associated higher mortality rates, i.e. all individuals migrating to southern Europe or northern Africa, increasing the supplementation rate would not lead to a positive population growth rate. However, management actions which increased the number of fledglings per nest generated a slight positive growth rate that led to a 54.3% increase in population size after 50 years and, when combining all three management options, the population grew by 378.3%. Alternatively, if a minimum of 9% of individuals overwintered in Britain, which is likely based on field observations and tracking data, a positive growth rate can be achieved without additional management due to this behaviour’s lower mortality rates. We conclude that the British white stork population will likely be viable in the long term, but these models and projections should be updated as more demographic data on this novel population become available.

Flexible demographic strategies promote the population persistence of a pioneer conifer tree (Pinus massoniana) in ecological restoration

Understanding how plant species are able to persist in temporally variable environments is a fundamental goal in ecology. According to the demographic buffering hypothesis (DBH), organisms persist by minimizing variation over time in those vital rates (survival, growth, and/or reproduction) which affect the population growth rate the most. Alternatively, the demographic lability hypothesis (DLH) posits that a population could instead track and benefit from environmental fluctuations, by adjusting one or more of its vital rates accordingly, to maximize fitness. Empirical evidence has been found for and against each hypothesis, yet rarely are both tested in tandem for pioneer tree species, which may be the most responsive or vulnerable to environmental stochasticity in the course of forest restoration. Here we evaluated the stochastic population growth rate (λs) of Pinus massoniana, a pioneer conifer tree, using integral projection models (IPMs) applied to five plots spanning distinctive stages along early ecological restoration in Southeast China. We assessed the demographic strategy operating in each plot through an elasticity analysis and uncovered the sources of variability in these demographic strategies across plots. We found that the ca. 40-year stand populations of P. massoniana were stable or growing (i.e., λs ≥ 1) in most plots, but strategies shifted between demographically labile and buffer across plots, which was predominately driven by among-plot variation in reproduction. Our findings suggest that diverging demographic strategies may promote the persistence of pioneer tree species of P. massoniana and others similar to it during the ecological restoration of subtropical forests.

Amplified Cyclicality in Mast Seeding Dynamics Positively Influences the Dynamics of a Seed Consumer Species.

AbstractTemporal autocorrelation in environmental conditions influences population dynamics through its effects on vital rates. However, a comprehensive understanding of how and to what extent temporal autocorrelation shapes population dynamics is still lacking because most empirical studies have unrealistically assumed that environmental conditions are temporally independent. Mast seeding is a biological event characterized by highly fluctuating and synchronized seed production at the tree population scale as well as a marked negative temporal autocorrelation. In the current context of global change, mast seeding events are expected to become more frequent, leading to strengthened negative temporal autocorrelations and thereby amplified cyclicality in mast seeding dynamics. Theory predicts that population growth rates are maximized when the environmental cyclicality of consumer resources and their generation times are closely matched. To test this prediction, we took advantage of the long-term monitoring of a wild boar population, a widespread seed consumer species characterized by a short generation time (∼2 years). As expected, simulations indicated that its stochastic population growth rate increased as mast seeding dynamics became more negatively autocorrelated. Our findings demonstrate that accounting for temporal autocorrelations in environmental conditions relative to the generation time of the focal population is required, especially under conditions of global warming, where the cyclicality in resource dynamics is likely to change.

Are genetic variation and demographic performance linked?

Quantifying relationships between genetic variation and population viability is important from both basic biological and applied conservation perspectives, yet few populations have been monitored with both long-term demographic and population genetics approaches. To empirically test whether and how genetic variation and population dynamics are related, we present one such paired approach. First, we use eight years of historical demographic data from five populations of Boechera fecunda (Brassicaceae), a rare, self-compatible perennial plant endemic to Montana, USA, and use integral projection models to estimate the stochastic population growth rate (λ S) and extinction risk of each population. We then combine these demographic estimates with previously published metrics of genetic variation in the same populations to test whether genetic diversity within populations is linked to demographic performance. Our results show that in this predominantly inbred species, standing genetic variation and demography are weakly positively correlated. However, the inbreeding coefficient was not strongly correlated with demographic performance, suggesting that more inbred populations are not necessarily less viable or at higher extinction risk than less inbred populations. A contemporary re-census of these populations revealed that neither genetic nor demographic parameters were consistently strong predictors of current population density, although populations showing lower probabilities of extinction in demographic models had larger population sizes at present. In the absence of evidence for inbreeding depression decreasing population viability in this species, we recommend conservation of distinct, potentially locally adapted populations of B. fecunda rather than alternatives such as translocations or reintroductions.

Demography of cape petrels in response to environmental changes

AbstractPredicting the responses of populations in changing environments is an important task for ecologists. Understanding the population dynamics of high‐latitude breeding species is critical given the particularly rapid environmental changes that occur in these regions. Using long‐term mark–resighting data acquired over 53‐years in Pointe Géologie, Terre Adélie, Antarctica, we estimated age‐specific demographic parameters and evaluated the effect of the environment on survival of a poorly known species, the cape petrel Daption capense. We then modeled the dynamics of this population using a life‐history model and performed prospective and retrospective analyses to estimate the sensitivity of the population growth rate to demographic parameters, and to quantify their relative contribution. Survival of cape petrel increased with age, being 0.610 (±0.193) for juveniles, 0.739 (±0.158) for individuals from 2 to 4, and 0.920 (±0.031) for older individuals. Minimum age at first reproduction was 3 years old, the age at which all birds were recruited was 14 years, and mean age at first reproduction was 9.05 (±2.06) years. Adult survival increased over time and was positively correlated with the southern annular mode (SAM). The stochastic population growth rate was estimated at 1.019, and adult survival over age 5 made the largest contribution to variance of the population growth rate. Sensitivity analyses revealed that population regulation was mainly driven by the SAM. Our results suggest that despite the decrease in breeding success, the population of cape petrels at Pointe Géologie increased due to the increase in immature and adult survival.

Effects of climate variability on the demography of wild geladas.

Nonhuman primates are an essential part of tropical biodiversity and play key roles in many ecosystem functions, processes, and services. However, the impact of climate variability on nonhuman primates, whether anthropogenic or otherwise, remains poorly understood. In this study, we utilized age‐structured matrix population models to assess the population viability and demographic variability of a population of geladas (Theropithecus gelada) in the Simien Mountains, Ethiopia with the aim of revealing any underlying climatic influences. Using data from 2008 to 2019 we calculated annual, time‐averaged, and stochastic population growth rates (λ) and investigated relationships between vital rate variability and monthly cumulative rainfall and mean temperature. Our results showed that under the prevailing environmental conditions, the population will increase (λ s = 1.021). Significant effects from rainfall and/or temperature variability were widely detected across vital rates; only the first year of infant survival and the individual years of juvenile survival were definitively unaffected. Generally, the higher temperature in the hot‐dry season led to lower survival and higher fecundity, while higher rainfall in the hot‐dry season led to increased survival and fecundity. Overall, these results provide evidence of greater effects of climate variability across a wider range of vital rates than those found in previous primate demography studies. This highlights that although primates have often shown substantial resilience to the direct effects of climate change, their vulnerability may vary with habitat type and across populations.

Evidence of demographic buffering in an endangered great ape: Social buffering on immature survival and the role of refined sex‐age classes on population growth rate

Theoretical and empirical research has shown that increased variability in demographic rates often results in a decline in the population growth rate. In order to reduce the adverse effects of increased variability, life-history theory predicts that demographic rates that contribute disproportionately to population growth should be buffered against environmental variation. To date, evidence of demographic buffering is still equivocal and limited to analyses on a reduced number of age classes (e.g. juveniles and adults), and on single sex models. Here we used Bayesian inference models for age-specific survival and fecundity on a long-term dataset of wild mountain gorillas. We used these estimates to parameterize two-sex, age-specific stochastic population projection models that accounted for the yearly covariation between demographic rates. We estimated the sensitivity of the long-run stochastic population growth rate to reductions in survival and fecundity on ages belonging to nine sex-age classes for survival and three age classes for female fecundity. We found a statistically significant negative linear relationship between the sensitivities and variances of demographic rates, with strong demographic buffering on young adult female survival and low buffering on older female and silverback survival and female fecundity. We found moderate buffering on all immature stages and on prime-age females. Previous research on long-lived slow species has found high buffering of prime-age female survival and low buffering on immature survival and fecundity. Our results suggest that the moderate buffering of the immature stages can be partially due to the mountain gorilla social system and the relative stability of their environment. Our results provide clear support for the demographic buffering hypothesis and its predicted effects on species at the slow end of the slow-fast life-history continuum, but with the surprising outcome of moderate social buffering on the survival of immature stages. We also demonstrate how increasing the number of sex-age classes can greatly improve the detection of demographic buffering in wild populations.

Unusually Paced Life History Strategies of Marine Megafauna Drive Atypical Sensitivities to Environmental Variability

Understanding why different life history strategies respond differently to changes in environmental variability is necessary to be able to predict eco-evolutionary population responses to change. Marine megafauna display unusual combinations of life history traits. For example, rays, sharks and turtles are all long-lived, characteristic of slow life histories. However, turtles also have very high reproduction rates and juvenile mortality, characteristic of fast life histories. Sharks and rays, in contrast, produce a few live-born young, which have low mortality rates, characteristic of slow life histories. This raises the question if marine megafaunal responses to environmental variability follow conventional life history patterns, including the pattern that fast life histories are more sensitive to environmental autocorrelation than slow life histories. To answer this question, we used a functional trait approach to quantify for different species of mobulid rays, cheloniid sea turtles and carcharhinid sharks – all inhabitants or visitors of (human-dominated) coastalscapes – how their life history, average size and log stochastic population growth rate, log(λs), respond to changes in environmental autocorrelation and in the frequency of favorable environmental conditions. The faster life histories were more sensitive to temporal frequency of favourable environmental conditions, but both faster and slower life histories were equally sensitive, although of opposite sign, to environmental autocorrelation. These patterns are atypical, likely following from the unusual life history traits that the megafauna display, as responses were linked to variation in mortality, growth and reproduction rates. Our findings signify the importance of understanding how life history traits and population responses to environmental change are linked. Such understanding is a basis for accurate predictions of marine megafauna population responses to environmental perturbations like (over)fishing, and to shifts in the autocorrelation of environmental variables, ultimately contributing toward bending the curve on marine biodiversity loss.

Simulated Indigenous fire stewardship increases the population growth rate of an understorey herb

Abstract Understanding how plant populations respond to multiple drivers is increasingly critical for biodiversity conservation under global change. Indigenous knowledge can provide guidance for sustainable management, but the outcome of its application in novel ecosystems is rarely known. Simulating the re‐introduction of Indigenous stewardship in contemporary contexts with population models allows for the comparison of different management scenarios and the elucidation of the mechanisms driving population outcomes. Beargrass Xerophyllum tenax is an ecologically and culturally important understorey plant managed through fire and leaf harvest by Native Americans. We collected demographic and abiotic data on beargrass over 3 years across fire severities in nine populations and conducted an experiment to simulate Native American leaf gathering. These data were used to build integral projections models (IPMs) with soil moisture and light availability as covariates. With these IPMs, we simulated stochastic population growth rates across future fire and leaf harvest scenarios. We then decomposed our simulation results using stochastic life table response experiments (SLTREs). The ‘no fire’ and ‘business as usual’ (180‐year fire return interval, 58% probability of high‐severity fire) scenarios resulted in lower population growth rates than ‘Indigenous fire stewardship’ (10‐year fire return interval, 10% chance of high‐severity fire). SLTREs revealed that Indigenous stewardship led to higher beargrass population growth rates due to greater fire frequency, higher adult survival and increased vegetative reproduction. Fire also interacted with harvest in the simulations; leaf harvest increased population growth rate only in combination with Indigenous fire stewardship. Synthesis. Stochastic and retrospective population dynamics tools combined with an understanding of Indigenous management practices allow for the comparison of future socio‐ecological scenarios as well as mechanistic understanding of differences between scenarios. Simulated Indigenous stewardship supported the long‐term persistence of X. tenax populations while business as usual and no fire did not. The benefits of Indigenous stewardship to population dynamics, and the complexity of interactive effects of multiple drivers, provide further impetus for collaboration across Indigenous and western knowledge systems. Xerophyllum tenax is presented as a model system to explore the influence of Indigenous stewardship, or its absence, on population dynamics.

Fire effects on the vital rates and stochastic population growth rate of the rare shrub Lindera subcoriacea Wofford

Understanding demographic vital rates and the factors that affect those rates are key components of successful conservation strategies for many threatened and endangered rare plant species. Lindera subcoriacea is a rare dioecious shrub that occupies isolated wetland habitats in a small number of locations in the southeastern United States. The species faces a number of threats to its continued persistence, including habitat destruction, invasive species, and population isolation. From 2011 to 2019, we collected demographic information from 290 L. subcoriacea individuals within 28 populations on Fort Bragg, North Carolina and used the data to estimate demographic vital rates in unburned populations and after being exposed to prescribed fire. We then constructed population matrices and estimated population growth rates under a 3-, 5-, and 10-year return interval. Results indicated that L. subcoriacea individuals have high survivorship in both burned and unburned populations, seed production was reduced 1- and 2-year post-fire, seed production was highly uneven across individuals, seedling recruitment was extremely low, and simulated population growth rates were only above 1.0 under the 10-year fire return interval. Taken together, these results indicate that (1) L. subcoriacea populations are persisting with population growth rates close to one, (2) the short-term impacts of fire on the overall population growth rate of L. subcoriacea, while only 2–3% may determine long-term population viability, and (3) extremely uneven seed production and limited recruitment of seedlings into larger size classes make L. subcoriacea populations vulnerable to stochastic demographic processes.

Demography of the giant monocarpic herb Rheum nobile in the Himalayas and the effect of disturbances by grazing.

Perennity of giant rosette species in combination with a single 'big bang' reproduction followed by death of the genetic individual is relatively rare among plants. Such long-lived monocarpic plants are usually slow growing and can be found in deserts, bogs or in alpine regions of the tropics or sub-tropics. Due to their longevity, monocarpic perennials risk losing everything before reproduction, which make them particularly susceptible to disturbances. Because of the inherent difficulties in assessing whether long-lived populations are growing or declining, usually neither their demography nor the consequences of increasing grazing pressure are known. We used integral projection modelling (IPM) to measure the growth rate and passage time to flowering of Rheum nobile, a monocarpic perennial, and one of the most striking alpine plants from the high Himalayas. Rosettes which were no longer found due to disturbances or grazing by yaks were either treated as missing or as dead in two series of analysis, thereby simulating demography with and without the impact of grazing cattle. Data were collected from plants at 4500 m a.s.l. in Shangri-la County, Yunnan Province, south-west China. In four consecutive years (2011-2014) and in two populations, 372 and 369 individuals were measured, respectively, and size-dependent growth, survival and fecundity parameters were estimated. In addition, germination percentage, seedling survival and establishment probability were assessed. The probability of survival, flowering and fecundity were strongly size dependent. Time to reach flowering size was 33.5 years [95 % confidence interval (CI) 21.9-43.3, stochastic estimate from pooled transitions and populations]. The stochastic population growth rate (λs) of Rheum nobile was 1.013 (95 % CI 1.010-1.017). When disturbance by grazing cattle (yaks) was accounted for in the model, λs dropped to values <1 (0.940, 95 % CI 0.938-0.943). We conclude that natural populations of this unique species are viable, but that conservation efforts should be made to minimize disturbances by grazing and to protect this slow-growing flagship plant from the high Himalayas.

Using statistics to design and estimate vital rates in matrix population models for a perennial herb

AbstractMatrix population models are widely used to assess population status and to inform management decisions. Despite existing theories for building such models, model construction is often partially based on expert opinion. So far, model structure has received relatively little attention, although it may affect estimates of population dynamics. Here, we assessed the consequences of two published matrix structures (a 4 × 4 matrix based on expert opinion and a 10 × 10 matrix based on statistical modeling) for estimates of vital rates and stochastic population dynamics of the long‐lived herb Astragalus scaphoides. We explored the ways in which choice of model structure alters the accuracy (i.e., mean) and precision (i.e., variance) of predicted population dynamics. We found that model structure had a negligible effect on the accuracy and precision of vital rates and stochastic stage distribution. However, the 10 × 10 matrix produced lower estimates of stochastic population growth rates than the 4 × 4 matrix, and more accurately predicted the observed trends in population abundance for three out of four study populations. Moreover, estimates of realized variation in population growth rate due to fluctuations in population stage structure over time were occasionally sensitive to matrix structure, suggesting differential roles of transient dynamics. Our study indicates that statistical modeling for choosing categories in matrix models might be preferable over expert opinion to accurately predict population trends and can provide a more objective way for model construction when the biological knowledge of the species is limited.

Rainfall and temperature change drive Arnica montana population dynamics at the Northern distribution edge.

Plant species of semi-natural grasslands are threatened by several simultaneous global change drivers, most notably land-use and climate change. In this study, we explore spatiotemporal variation and changes in deterministic (λ) and stochastic population growth rates (λs), and the underlying vital rates of eight populations of Arnica montana at the species' north-western range margin in Norway. We assess to what extent variation in the demographic rates could be attributed to environmental correlates of the key global change drivers likely to operate at the range edge, including population size, surrogates of habitat quality, temperature and precipitation. We found no relationship between λ and population size or habitat quality, but λ declined in response to both increasing precipitation and increasing temperature. Life-table response experiments revealed that the temporal variability was driven by survival and clonality, whereas the spatial variation was driven by clonality. Our results suggest that A. montana has a threshold response to increasing precipitation, likely due to adaptations to local climatic conditions. Growth and flowering were both negatively affected by increasing temperature, but these effects had a low influence on the spatiotemporal variability in λ. In contrast, the stochastic growth rate was negatively influenced by climate change, indicating an increased extinction risk for marginal populations, possibly leading to range contraction of A. montana as climate change proceeds. Altogether, our study illustrates how the fates of peripheral populations, which are critically important in species range dynamics, may be affected by both deterministic and stochastic effects of multiple coinciding global change drivers.

A functional trait approach to identifying life history patterns in stochastic environments.

Temporal variation in demographic processes can greatly impact population dynamics. Perturbations of statistical coefficients that describe demographic rates within matrix models have, for example, revealed that stochastic population growth rates (log(λ s)) of fast life histories are more sensitive to temporal autocorrelation of environmental conditions than those of slow life histories. Yet, we know little about the mechanisms that drive such patterns. Here, we used a mechanistic, functional trait approach to examine the functional pathways by which a typical fast life history species, the macrodetrivore Orchestia gammarellus, and a typical slow life history species, the reef manta ray Manta alfredi, differ in their sensitivity to environmental autocorrelation if (a) growth and reproduction are described mechanistically by functional traits that adhere to the principle of energy conservation, and if (b) demographic variation is determined by temporal autocorrelation in food conditions. Opposite to previous findings, we found that O. gammarellus log(λ s) was most sensitive to the frequency of good food conditions, likely because reproduction traits, which directly impact population growth, were most influential to log(λ s). Manta alfredi log(λs) was instead most sensitive to temporal autocorrelation, likely because growth parameters, which impact population growth indirectly, were most influential to log(λ s). This differential sensitivity to functional traits likely also explains why we found that O. gammarellus mean body size decreased (due to increased reproduction) but M. alfredi mean body size increased (due to increased individual growth) as food conditions became more favorable. Increasing demographic stochasticity under constant food conditions decreased O. gammarellus mean body size and increased log(λ s) due to increased reproduction, whereas M. alfredi mean body and log(λ s) decreased, likely due to decreased individual growth. Our findings signify the importance of integrating functional traits into demographic models as this provides mechanistic understanding of how environmental and demographic stochasticity affects population dynamics in stochastic environments.

Evaluating the efficacy of independent versus simultaneous management strategies to address ecological and genetic threats to population viability

Abstract Small, declining populations can face simultaneous, interacting, ecological and genetic threats to viability. Conservation management strategies designed to tackle such threats independently may then prove ineffective. Population viability analyses that evaluate the efficacy of management strategies implemented independently versus simultaneously are then essential to the design of effective management plans, yet such quantitative evaluations are typically lacking. We used stochastic individual‐based models, parameterised with high‐quality multi‐year demographic and genetic data, to evaluate the efficacy of independent or simultaneous ecological (supplementary feeding) and genetic (translocations to alleviate inbreeding) management strategies for a red‐billed chough (Pyrrhocorax pyrrhocorax) population of major conservation concern. This population is experiencing ecological threats from food limitation and genetic threats from escalating inbreeding. Conservation managers therefore face a dilemma: supplementary feeding may be ineffective if inbreeding is limiting stochastic population growth rate (λs), while translocations may be ineffective if food is limiting. Model simulations suggested that the focal population will decline to extinction relatively rapidly with no conservation management (mean λs ≈ 0.86) and with genetic management alone (λs ≈ 0.90). Ecological management alone reduced, but did not halt the population decline (λs ≈ 0.93). However, simultaneous genetic and ecological management yielded population stability (λs ≈ 1), with genetic rescue lasting ~25 years. These outcomes arose because the capacity for translocations to alleviate inbreeding depression is limited by food availability, while supplementary feeding cannot achieve population viability in the presence of accumulating inbreeding. However, supplementary feeding improved environmental quality enough to allow expression of variance in fitness and thus inbreeding depression, meaning that reductions in inbreeding following translocations can increase λs. Synthesis and applications. Our analyses suggest that simultaneous management of ecological and genetic threats will be critical to ensuring viability of Scotland's chough population; neither strategy independently is likely to achieve population persistence and may consequently waste conservation resources. Managers of other resource‐limited, inbred populations should consider that the efficacy of strategies designed to alleviate ecological and genetic threats may be interdependent, such that holistic management is essential to ensure population viability.