Differences In Vital Rates Research Articles

Major axes of variation in tree demography across global forests

The future trajectory of global forests is closely intertwined with tree demography, and a major fundamental goal in ecology is to understand the key mechanisms governing spatio‐temporal patterns in tree population dynamics. While previous research has made substantial progress in identifying the mechanisms individually, their relative importance among forests remains unclear mainly due to practical limitations. One approach to overcome these limitations is to group mechanisms according to their shared effects on the variability of tree vital rates and quantify patterns therein. We developed a conceptual and statistical framework (variance partitioning of Bayesian multilevel models) that attributes the variability in tree growth, mortality, and recruitment to variation in species, space, and time, and their interactions – categories we refer to as organising principles (OPs). We applied the framework to data from 21 forest plots covering more than 2.9 million trees of approximately 6500 species. We found that differences among species, the species OP, proved a major source of variability in tree vital rates, explaining 28–33% of demographic variance alone, and 14–17% in interaction with space, totalling 40–43%. Our results support the hypothesis that the range of vital rates is similar across global forests. However, the average variability among species declined with species richness, indicating that diverse forests featured smaller interspecific differences in vital rates. Moreover, decomposing the variance in vital rates into the proposed OPs showed the importance of unexplained variability, which includes individual variation, in tree demography. A focus on how demographic variance is organized in forests can facilitate the construction of more targeted models with clearer expectations of which covariates might drive a vital rate. This study therefore highlights the most promising avenues for future research, both in terms of understanding the relative contributions of groups of mechanisms to forest demography and diversity, and for improving projections of forest ecosystems.

Sex-specific recruitment rates contribute to male-biased sex ratio in Adélie penguins.

Sex-related differences in vital rates that drive population change reflect the basic life history of a species. However, for visually monomorphic bird species, determining the effect of sex on demographics can be a challenge. In this study, we investigated the effect of sex on apparent survival, recruitment, and breeding propensity in the Adélie penguin (Pygoscelis adeliae), a monochromatic, slightly size dimorphic species with known age, known sex, and known breeding history data collected during 1996-2019 (n = 2127 birds) from three breeding colonies on Ross Island, Antarctica. Using a multistate capture-mark-recapture maximum-likelihood model, we estimated apparent survival (), recapture (resighting) probability (), and the probability of transitioning among breeding states and moving between colonies (; colony-specific non-juvenile pre-breeders, breeders, and non-breeders). Survival rate varied by breeding status and colony, but not sex, and pre-breeders had higher survival rates than breeders and non-breeders. Females had a higher probability of recruiting into the breeding population each year and may enter the breeding pool at younger ages. In contrast, both sexes had the same probability of breeding from year to year once they had recruited. Although we detected no direct sex effects on survival, the variation in recruitment probability and age-at-first reproduction, along with lower survival rates of breeders compared to pre-breeders, likely leads to shorter lifespans for females. This is supported by our findings of a male-biased mean adult sex ratio (ASR) of 1.4 males for every female ( proportion of males = 0.57, SD = 0.07) across all colonies and years in this metapopulation. Our study illustrates how important it can be to disentangle sex-related variation in population vital rates, particularly for species with complex life histories and demographic dynamics.

Extreme polygyny results in intersex differences in age-dependent survival of a highly dimorphic marine mammal.

Developmental differences in vital rates are especially profound in polygamous mating systems. Southern elephant seals (Mirounga leonina) are highly dimorphic and extremely polygynous marine mammals. A demographic model, supported by long-term capture-mark-recapture records, investigated the influence of sex and age on survival in this species. The study revealed clear differences between female and male age-dependent survival rates. Overall juvenile survival estimates were stable around 80-85% for both sexes. However, male survival estimates were 5-10% lower than females in the same age classes until 8 years of age. At this point, male survival decreased rapidly to 50% ± 10% while female estimates remained constant at 80% ± 5%. Different energetic requirements could underpin intersex differences in adult survival. However, the species' strong sexual dimorphism diverges during early juvenile development when sex-specific survival rates were less distinct. Maximizing growth is especially advantageous for males, with size being a major determinant of breeding probability. Maturing males may employ a high-risk high-reward foraging strategy to compensate for extensive sexual selection pressures and sex-specific energetic needs. Our findings suggest sex-specific adult survival is a result of in situ ecological interactions and evolutionary specialization associated with being a highly polygynous marine predator.

Raising offspring increases ageing: Differences in senescence among three populations of a long-lived seabird, the Atlantic puffin.

Actuarial senescence, the decline of survival with age, is well documented in the wild. Rates of senescence vary widely between taxa, to some extent also between sexes, with the fastest life histories showing the highest rates of senescence. Few studies have investigated differences in senescence among populations of the same species, although such variation is expected from population-level differences in environmental conditions, leading to differences in vital rates and thus life histories. We predict that, within species, populations differing in productivity (suggesting different paces of life) should experience different rates of senescence, but with little or no sexual difference in senescence within populations of monogamous, monomorphic species where the sexes share breeding duties. We compared rates of actuarial senescence among three contrasting populations of the Atlantic puffin Fratercula arctica. The dataset comprised 31 years (1990-2020) of parallel capture-mark-recapture data from three breeding colonies, Isle of May (North Sea), Røst (Norwegian Sea) and Hornøya (Barents Sea), showing contrasting productivities (i.e. annual breeding success) and population trends. We used time elapsed since first capture as a proxy for bird age, and productivity and the winter North Atlantic Oscillation Index (wNAO) as proxies for the environmental conditions experienced by the populations within and outside the breeding season, respectively. In accordance with our predictions, we found that senescence rates differed among the study populations, with no evidence for sexual differences. There was no evidence for an effect of wNAO, but the population with the lowest productivity, Røst, showed the lowest rate of senescence. As a consequence, the negative effect of senescence on the population growth rate (λ) was up to 3-5 times smaller on Røst (Δλ=-0.009) than on the two other colonies. Our findings suggest that environmentally induced differences in senescence rates among populations of a species should be accounted for when predicting effects of climate variation and change on species persistence. There is thus a need for more detailed information on how both actuarial and reproductive senescence influence vital rates of populations of the same species, calling for large-scale comparative studies.

A two‐sex integrated population model reveals intersexual differences in life history strategies in Cooper's hawks

AbstractIntegrated population models (IPMs) combine population counts with demographic data to improve the precision of estimates of population size and demographic rates. IPMs can also provide information on demographic parameters for which data are lacking (e.g., immigration) if other sufficient information is available. IPMs often use demographic data from one sex, usually females, implicitly assuming the chosen sex adequately represents the species' life history; for highly size‐dimorphic raptors, that assumption may not hold true. We developed a two‐sex IPM for a New Mexico population of the highly size‐dimorphic Cooper's hawk (Accipiter cooperii) from 2011 to 2020 and compared estimates of life history characteristics between sexes. Because we had data to directly estimate sex‐ and age‐specific probabilities of breeding, fecundity, survival, brood sex ratios, and emigration rates, we could indirectly estimate age‐specific immigration rates for both sexes. Our two‐sex IPM revealed that population growth was most strongly associated with increased immigration and decreased first‐year (FY) emigration in females and with after‐first‐year (AFY) survival in males. Most males that were recruited as new breeders in our study area were AFY residents, whereas most female recruits were AFY immigrants. All females that fledged in our study area and survived until their first breeding season bred in their first year, whereas only 3% of surviving FY males bred. We found evidence of density dependence in the survival of AFY males (r = −0.11, 95% credible interval = −0.36 to −0.03) and in AFY female immigration (r = −0.17, 95% credible interval = −0.27 to −0.12). Our findings reveal that male and female Cooper's hawks differ in their response to population density, and in how they contribute to population growth. Thus, sexual differences in vital rates can be an important consideration in raptor population models.

The effects of temperature extremes on survival in two semi‐arid Australian bird communities over three decades, with predictions to 2104

AbstractAimOrganisms in arid and semi‐arid regions are frequently exposed to climatic extremes and accordingly among the most vulnerable to climate change. Studies of seasonal differences in vital rates, which mediate effects of climate on viability, are rare in arid species, limiting ability to project population trends. We quantified survival patterns for two bird communities as a function of exposure to temperature extremes in winter and summer, then projected survival patterns to 2104.LocationSemi‐arid eastern Australia.Time period1986–2016; 1986–2104.Major taxa studiedBirds.MethodsUsing mark–recapture time‐dependent Cormack–Jolly–Seber models and data for 37 species from two >30‐year ringing programmes, we tested for effects on 6‐monthly survival of exposure to temperatures >38 and <0°C. We then predicted future survival for different emission scenarios, testing whether changes in survival associated with warming winters would be sufficient to offset the effects of rising summer temperatures.ResultsSurvival probability declined strongly with increasing exposure to days >38°C and to a lesser extent to days <0°C, with temperature extremes explaining 43 and 13% of temporal variation in survival among years, respectively. Summer survival patterns were similar across avian guilds but only survival of nectarivores declined in winter. Our models predict that gains in winter survival will not offset reductions in summer survival. Annual survival is predicted to decline substantially by the end of the century: from .63 in 1986 to .43 in 2104 under an optimistic emission scenario and to .11 under a pessimistic scenario.Main conclusionsWe highlight the significance of temperature extremes for species' persistence in arid and semi‐arid regions, comprising 70% of Australia's landmass, and 40% globally. Our demography‐based results are consistent with physiological‐based projections evaluating avian survival in arid and semi‐arid regions globally and suggest rising summer temperatures pose a risk to population persistence in these regions.

Large herbivores in a partially migratory population search for the ideal free home.

Migration is a tactic used across taxa to access resources in temporally heterogenous landscapes. Populations that migrate can attain higher abundances because such movements allow access to higher quality resources, or reduction in predation risk resulting in increased fitness. However, most migratory species occur in partially migratory populations, a mix of migratory and non-migratory individuals. It is thought that the portion of migrants in a partial migration population is maintained either through (1) a population-level evolutionary stable state where counteracting density-dependent vital rates act on migrants and residents to balance fitness or (2) conditional migration, where the propensity to migrate is influenced by the individual's state. However, in many respects, migration is also a form of habitat selection and the proportion of migrants and residents may be the result of density-dependent habitat selection. Here, we test whether the theory of Ideal Free Distribution (IFD) can explain the coexistence of different migratory tactics in a partially migratory population. IFD predicts individuals exhibit density-dependent vital rates and select different migratory tactics to maximize individual fitness resulting in equal fitness (λ) between tactics. We tested the predictions of IFD in a partially migratory elk population that declined by 70% with 19 years of demographic data and migratory tactic switching rates from >300 individuals. We found evidence of density dependence for resident pregnancy and adult female survival providing a fitness incentive to switch tactics. Despite differences in vital rates between migratory tactics, mean λ (fitness) was equal. However, as predicted by the IFD, individuals switched tactics toward those of higher fitness. Our analysis reveals that partial migration may be driven by tactic selection that follows the ideal free distribution. These findings reinforce that migration across taxa may be a polymorphic behavior in large herbivores where migratory tactic selection is determined by differential costs and benefits, mediated by density dependence.

Relationships between vital rates and ecological traits in an avian community.

Comparative studies about the relationships between vital rates and ecological traits at the community level are conspicuously lacking for most taxa because estimating vital rates requires detailed demographic data. Identifying relationships between vital rates and ecological traits could help to better understand ecological and evolutionary demographic mechanisms that lead to interspecific differences in vital rates. We use novel dynamic N-mixture models for counts to achieve this for a whole avian community comprising 53 passerine species, while simultaneously accounting for density dependence and environmental stochasticity in recruitment and survival and, importantly, correcting our inferences for imperfect detection. Demographic stochasticity is taken into account in the form of the binomial and Poisson distributions describing survival events and number of recruits. We then explore relationships between estimated demographic parameters (i.e., vital rates) and ecological traits related to migration patterns, diet, habitat and nesting location of each species. The relative importance of recruitment and adult survival as contributors to population growth varied greatly among species, and interspecific differences in vital rates partly reflected differences in ecological traits. Migratory mode was associated with interspecific differences in population growth and density dependence. Resident species had higher population growth rates than long- and short-distance migrants. We found no relationships between diet and population growth rate. Habitat differences were associated with different growth rates: alpine, wetland and farmland species had lower population growth rates than forest species. Differences in population growth rates among nesting locations showed that breeding habitat is essential for population dynamics. Our study reveals relationships between ecological traits and contributions of vital rates to population growth and suggests ways in which patterns of population growth fluctuations in a community might be determined by life history.

Individual heterogeneity determines sex differences in mortality in a monogamous bird with reversed sexual dimorphism.

Sex differences in mortality are pervasive in vertebrates, and usually result in shorter life spans in the larger sex, although the underlying mechanisms are still unclear. On the other hand, differences in frailty among individuals (i.e. individual heterogeneity), can play a major role in shaping demographic trajectories in wild populations. The link between these two processes has seldom been explored. We used Bayesian survival trajectory analysis to study age-specific mortality trajectories in the Eurasian sparrowhawk (Accipiter nisus), a monogamous raptor with reversed sexual size dimorphism. We tested the effect of individual heterogeneity on age-specific mortality, and the extent by which this heterogeneity was determined by average reproductive output and wing length as measures of an individual's frailty. We found that sex differences in age-specific mortality were primarily driven by the differences in individual heterogeneity between the two sexes. Females were more heterogeneous than males in their level of frailty. Thus, a larger number of females with low frailty are able to survive to older ages than males, with life expectancy for the least frail adult females reaching up to 4·23years, while for the least frail adult males it was of 2·68years. We found that 50% of this heterogeneity was determined by average reproductive output and wing length in both sexes. For both, individuals with high average reproductive output had also higher chances to survive. However, the effect of wing length was different between the two sexes. While larger females had higher survival, larger males had lower chances to survive. Our results contribute a novel perspective to the ongoing debate about the mechanisms that drive sex differences in vital rates in vertebrates. Although we found that variables that relate to the cost of reproduction and sexual dimorphism are at least partially involved in determining these sex differences, it is through their effect on the level of frailty that they affect age patterns of mortality. Therefore, our results raise the possibility that observed differences in age-specific demographic rates may in fact be driven by differences in individual heterogeneity.

Vitals rates and seasonal movements of two isolated greater sage-grouse populations in Utah's West Desert

Declines in greater sage-grouse (Centrocercus urophasianus; hereafter, sagegrouse) populations in Utah over the last century parallel range-wide trends. However, little is known about the ecology of sage-grouse populations that inhabit Utah’s naturally fragmented habitats. Utah’s West Desert sage-grouse populations occupy sagebrush (Artemisia spp.) habitats that are geographically separated by the Great Salt Lake, and largely confined to the Sheeprock and Deep Creek watersheds. From 2005 to 2006, we monitored sagegrouse that were radio-collared in each watershed to determine the factors affecting the vital rates in these isolated populations. Livestock grazing by domestic cattle was the dominate land use, and mammalian predator control for livestock protection was conducted in both watersheds. Corvid control was conducted only in the Sheeprock watershed. During the study, we identified 6 leks that had not been previously documented. Seasonal migration patterns for individual radio-collared sage-grouse in both watersheds varied across the sites. Habitat structure metrics were similar at brood-rearing and random sites for both areas. Nesting and brood success and the ratio of chicks per successful brood were higher for both populations in 2005 than 2006. We attributed these annual differences in vital rates to seasonal variation in precipitation. Spring precipitation in 2005 was twice the 30-year average following a 5 year drought. However, chick recruitment estimates for both populations regardless of year were lower than reported in the published literature. Adult sage-grouse survival rate estimates in Sheeprock and Deep Creek watersheds were lower and higher, respectively, than published reports indicated. These differences may reflect a difference in meso-predators communities. Sage-grouse conservation strategies in both areas should continue to emphasize protection of brood-rearing and seasonal habitat, but the risk of population extirpation as a consequence of extended droughts predicted by climate change models and the invasion of small mesopredators may remain problematic for these populations.

Trait-demography relationships underlying small mammal population fluctuations.

Large-scale fluctuations in abundance are a common feature of small mammal populations and have been the subject of extensive research. These demographic fluctuations are often associated with concurrent changes in the average body mass of individuals, sometimes referred to as the 'Chitty effect'. Despite the long-standing recognition of this phenomenon, an empirical investigation of the underlying coupled dynamics of body mass and population growth has been lacking. Using long-term life-history data combined with a trait-based demographic approach, we examined the relationship between body mass and demography in a small mammal population that exhibits non-cyclic, large-scale fluctuations in abundance. We used data from the male segment of a 25-year study of the monogamous prairie vole, Microtus ochrogaster, in Illinois, USA. Specifically, we investigated how trait-demography relationships and trait distributions changed between different phases of population fluctuations, and the consequences of these changes for both trait and population dynamics. We observed phase-specific changes in male adult body mass distribution in this population of prairie voles. Our analyses revealed that these changes were driven by variation in ontogenetic growth, rather than selection acting on the trait. The resulting changes in body mass influenced most life-history processes, and these effects varied among phases of population fluctuation. However, these changes did not propagate to affect the population growth rate due to the small effect of body mass on vital rates, compared to the overall differences in vital rates between phases. The increase phase of the fluctuations was initiated by enhanced survival, particularly of juveniles and fecundity, whereas the decline phase was driven by an overall reduction in fecundity, survival and maturation rates. Our study provides empirical support, as well as a potential mechanism, underlying the observed trait changes accompanying population fluctuations. Body size dynamics and population fluctuations resulted from different life-history processes. Therefore, we conclude that body size dynamics in our population do not drive the observed population dynamics. This more in-depth understanding of different components of small mammal population fluctuations will help us to better identify the mechanistic drivers of this interesting phenomenon.

Host genus and rainfall drive the population dynamics of a vascular epiphyte

AbstractVascular epiphytes constitute up to 25% of tropical plant diversity and play an important role in providing food, water, and shelter to many organisms. However, the factors that drive their population dynamics, including the influence of their host plants (phorophytes) and of climatic factors, are still poorly understood. We provide the first test of whether host tree genus can affect population dynamics of an epiphytic species, and assess the interactive effects of host tree and rainfall. We carried out a five‐year study of the demography of >1000 plants of the endemic bromeliad, Tillandsia macdougallii L. B. Sm, growing on pine and oak trees in a Mexican montane forest. We tested for differences in vital rates and used integral projection models to test whether these scaled up to differences in long‐term population growth rates between the two host genera and as a function of rainfall. T. macdougallii survival and growth were higher on pines than on oaks for larger plants but not for seedlings. The probability of producing capsules was higher for T. macdougallii on oaks than on pines. Increasing dry season rainfall was significantly correlated with increased survival of individuals, but not of ramets, growing on both oaks and pines. The probability of clonal reproduction increased significantly with increasing dry season rainfall for T. macdougallii growing on oaks, but not on pines. Projected long‐term population growth rates, λ, for T. macdougallii on pines were higher than on oaks. Lambda increased steeply as a function of dry season rainfall for T. macdougallii populations on both host tree genera, but the increase was steeper for populations on oaks, likely because the former are deciduous. Our results show that host tree genus can affect the long‐term dynamics of epiphyte populations and suggest that these effects may depend on rainfall. The high sensitivity of this species to changes in dry season rainfall suggests that a better understanding of how rainfall may drive vascular epiphyte populations will be critical for understanding the effects of climate change on species persistence.

Within- and among-population variation in vital rates and population dynamics in a variable environment.

Understanding the causes of within- and among-population differences in vital rates, life histories, and population dynamics is a central topic in ecology. To understand how within- and among-population variation emerges, we need long-term studies that include episodic events and contrasting environmental conditions, data to characterize individual and shared variation, and statistical models that can tease apart shared and individual contribution to the observed variation. We used long-term tag-recapture data to investigate and estimate within- and among-population differences in vital rates, life histories, and population dynamics of marble trout Salmo marmoratus, an endemic freshwater salmonid with a narrow range. Only ten populations of pure marble trout persist in headwaters of Alpine rivers in western Slovenia. Marble trout populations are also threatened by floods and landslides, which have already caused the extinction of two populations in recent years. We estimated and determined causes of variation in growth, survival, and recruitment both within and among populations, and evaluated trade-offs between them. Specifically, we estimated the responses of these traits to variation in water temperature, density, sex, early life conditions, and extreme events. We found that the effects of population density on traits were mostly limited to the early stages of life and that growth trajectories were established early in life. We found no clear effects of water temperature on vital rates. Population density varied over time, with flash floods and debris flows causing massive mortalities (>55% decrease in survival with respect to years with no floods) and threatening population persistence. Apart from flood events, variation in population density within streams was largely determined by variation in recruitment, with survival of older fish being relatively constant over time within populations, but substantially different among populations. Marble trout show a fast to slow continuum of life histories, with slow growth associated with higher survival at the population level, possibly determined by food conditions and age at maturity. Our work provides unprecedented insight into the causes of variation in vital rates, life histories, and population dynamics in an endemic species that is teetering on the edge of extinction.

Biocontrol insect impacts population growth of its target plant species but not an incidentally used nontarget

AbstractUnderstanding the impact of herbivory on plant populations is a fundamental goal of ecology. Damage to individual plants can be visually striking and affect the fates of individuals, but these impacts do not necessarily translate into population‐level differences in vital rates (survival, growth, or fecundity) or population growth rates. In biological control of weeds, quantitative assessments of population‐level impacts of released agents on both target invasive plants and native, nontarget plants are needed to inform evaluations of the benefits and risks of releasing agents into new regions. Here we present a 3‐yr experimental demographic field study using the European root‐feeding biocontrol weevil, Mogulones crucifer, first released in Canada in 1997 to control the invasive weed Cynoglossum officinale (Boraginaceae). Mogulones crucifer is an effective “search and destroy” agent in Canada, but sporadically feeds, oviposits, and develops on native nontarget Boraginaceae. We investigated the population‐level impacts of this biocontrol insect on its target weed and a native nontarget plant, Hackelia micrantha (Boraginaceae), by releasing large numbers of weevils into naturally occurring patches of H. micrantha growing isolated from or interspersed with C. officinale. We followed the fates of individual plants on release and nonrelease (control) sites for two transition years, developed matrix models to project population growth rates (λ) for each plant species, and examined the contributions from differences in vital rates to changes in λ using life table response experiments (LTRE). In contrast to studies of the insect–plant interaction in its native range, as a biocontrol agent, M. crucifer increased mortality of C. officinale rosettes in the year immediately following release, depressing the weed's λ to below the population replacement level. However, λ for H. micrantha was never depressed below the replacement level, and any differences between release and nonrelease sites in the nontarget could not be explained by significant contributions from vital rates in the LTRE. This study is the first to simultaneously and experimentally examine target and nontarget population‐level impacts of a weed biocontrol insect in the field, and supports the theoretical prediction that plant life history characteristics and uneven herbivore host preferences can interact to produce differences in population‐level impacts between target and nontarget plant species.

The role of demography, intra‐species variation, and species distribution models in species' projections under climate change

Organisms are projected to shift their distribution ranges under climate change. The typical way to assess range shifts is by species distribution models (SDMs), which predict species’ responses to climate based solely on projected climatic suitability. However, life history traits can impact species’ responses to shifting habitat suitability. Additionally, it remains unclear if differences in vital rates across populations within a species can offset or exacerbate the effects of predicted changes in climatic suitability on population viability. In order to obtain a fuller understanding of the response of one species to projected climatic changes, we coupled demographic processes with predicted changes in suitable habitat for the monocarpic thistle Carlina vulgaris across northern Europe. We first developed a life history model with species‐specific average fecundity and survival rates and linked it to a SDM that predicted changes in habitat suitability through time with changes in climatic variables. We then varied the demographic parameters based upon observed vital rates of local populations from a translocation experiment. Despite the fact that the SDM alone predicted C. vulgaris to be a climate ‘winner’ overall, coupling the model with changes in demography and small‐scale habitat suitability resulted in a matrix of stable, declining, and increasing patches. For populations predicted to experience declines or increases in abundance due to changes in habitat suitability, altered fecundity and survival rates can reverse projected population trends.

Low recruitment not mortality limits growth of peripheral populations of Silene spaldingii

Populations at the periphery of a species range are thought to be less viable than those in the center because they are at the limit of their ecological tolerances. Plant population viability is determined primarily by mortality and recruitment, thus knowing differences in vital rates between central and peripheral populations is key to understanding range limits. Silene spaldingii S. Watson is a long-lived iteroparous plant that occurs throughout the eastern Columbia Plateau region of eastern Washington and adjacent Idaho and Oregon and is disjunct in northwest Montana. I recorded the fate of mapped S. spaldingii plants annually for 10 years in four populations, two from eastern Washington and two from Montana. Recruitment averaged three times higher and relatively constant at two central populations in Washington compared with two peripheral sites in Montana. Mortality was three times higher at one of the central populations compared with the remaining three sites due to vole predation. Vole activity was observed at the second central population but came too late in the study to be certain that it resulted in mortality. Vole predation was not observed in the peripheral populations. My results suggest that peripheral populations of S. spaldingii may only be able to persist as long as predation or other sources of mortality remain low and that intrinsic low and variable recruitment rates coupled with predation may help define the eastern range margin of this species.

Pair‐wise analyses of the effects of demographic processes, vital rates, and life stages on the spatiotemporal variation in the population dynamics of the riparian tree Aesculus turbinata Blume

AbstractPopulation growth rates (λ) of the riparian tree Aesculus turbinata varied from 0.9988 to 1.0524 spatiotemporally. We conducted a series of pair‐wise demographic and matrix analyses, including randomization tests, three types of life table response experiments (LTREs), analysis of variance and χ2 tests, to test which life stages had the greatest effect on this variation in λ. Randomization tests detected significant variations in λ between plots affected or not by typhoons in three habitats and between periods with high and low recruitment in one habitat. Mixed‐level LTREs identified that the demographic processes and life stages that had the strongest effect on the actual variation in λ were: (1) progressions of small and intermediate juveniles and (2) founding process from seeds to 1‐year‐old seedlings. These juvenile stages had medium sensitivities and variances that explained high upper‐level LTRE contributions. Lower‐level LTREs showed that the vital rates contributing the most were the growth rates of these juvenile stages. These findings demonstrate that progression from one stage to the next, growth rates of 1‐year‐old seedlings, and stunted aging juveniles are the most important stages in the population dynamics of this long‐lived primary tree species. Transition matrix elements with high elasticities had little effect on the variation in λ, indicating that high‐elasticity vital rates do not necessarily drive variation in population growth. As compared with the results of randomization tests, significant differences in vital rates examined using ANOVA or χ2 tests showed that typhoon disturbance had the greatest effect on the demographic parameters of individual trees.

Variation in stochastic demography between and within central and peripheral regions in a widespread short‐lived herb

Analyzing intraspecific variation in population dynamics in relation to environmental factors is crucial to understand the current and future distributions of plant species. Across ranges, peripheral populations are often expected to show lower and more temporally variable vital rates than central populations, although it remains unclear how much any differences in vital rates actually contribute to differences in population growth rates. Moreover, few demographic studies accounting for environmental stochasticity have been carried out both at continental and regional scales. In this study we calculated stochastic growth rates in five central and six northern peripheral populations of the widespread shortlived herb Plantago coronopus along the Atlantic Coast in Europe. To evaluate at two spatial scales how mean values and variability of vital rates (i.e., fecundity, recruitment, survival, growth, and shrinkage) contributed to the differences in stochastic growth rates, we performed Stochastic Life Table Response Experiment (SLTRE) analyses between and within central and peripheral regions. Additionally, we searched for correlations between vital rate contributions and local environmental conditions. Lower mean values and greater variability for some vital rates in peripheral than in central populations had an overall negative but nonsignificant effect on the stochastic growth rates in the periphery. Different life cycle components accounted for differences in population growth depending on spatial scale, although recruitment was the vital rate with the highest influence both between and within regions. Interestingly, the same pattern of differentiation among populations was found within central and peripheral areas: in both regions, one group of populations displayed positive contributions of growth and shrinkage and negative contributions of recruitment and survival; the opposite pattern was found in the remaining populations. These differences in vital rate contributions among populations within regions were correlated with precipitation regime, whereas at the continental scale, differences in contribution patterns were related to temperature. Altogether, our results show how populations of P. coronopus exhibit life cycle differences that may enable the species to persist in locations with widely varying environmental conditions. This demographic flexibility may help to explain the success of widespread plants across large and heterogeneous ranges.

Orchids do not pay costs at emergence for prolonged dormancy

In plants, prolonged dormancy is often considered a response to resource depletion or environmental stress that comes at a fitness cost. However, apparent costs of dormancy could reflect the state in which plants entered dormancy, rather than effects of dormancy per se. We tested this hypothesis for a terrestrial orchid, Epipactis atrorubens, by analyzing differences in vital rates of dormant and emergent plants using generalized linear mixed models, applied to eight years of demographic data. Dormant E. atrorubens plants did not form one homogeneous stage class. Instead, the vital rates of dormant plants mirrored performance of plants in their life stage before dormancy. Plants emerging from dormancy were slightly (albeit only marginally statistically significantly) larger than plants transitioning from the matching aboveground stage class, especially for smaller and younger stage classes. Because small plants were most likely to go dormant, plants emerging from dormancy were also smaller than average, if one were to compare all previously dormant plants to all previously emergent plants. Therefore, misclassifying all dormant plants into a single stage class changes whether we view dormancy as intrinsically costly, in terms of future performance upon emergence. We suggest that prolonged dormancy may be a form of phenotypic plasticity in which plants distribute their performance and reproductive effort through time, rather than a simple stress response.

Size‐dependent flowering and costs of reproduction affect population dynamics in a tuberous perennial woodland orchid

Summary 1. In woodland herbs, the probability of flowering and costs associated with reproduction may strongly depend on environmental context (shade vs. light habitats) and on plant size. This may be particularly true for tuberous orchids that inhabit woodlands, as the amount of incoming radiation and total leaf area strongly determine photosynthetic capacity and hence the amount of carbohydrates that can be relocated to below‐ground storage organs that form next year’s rosette and flowering stalk. 2. To fully comprehend the impact of size‐dependent reproduction on population dynamics under varying light conditions, life cycle models should therefore include plant size in a continuous manner. In this study, annual changes in plant size and demographic behaviour of the tuberous perennial orchid Orchis purpurea were monitored during seven consecutive years (2003–2009) in open and shaded woodland. Integral projection models (IPMs) and life table response experiments (LTRE) were used to investigate the extent to which variation in plant size affected the overall population dynamics of this species and to decompose differences in population growth rates between populations of open and shaded woodland into contributions from growth, survival and reproduction. 3. Both plants in shaded and light environments needed to be a certain size to initiate flowering, but this threshold size was almost three times as large in shaded environments as in light environments. Plants in open woodlands flowered more frequently over the years, showed less size regression after flowering and produced significantly more fruits than plants in shaded environments, resulting in significantly larger population growth rates. 4. Our life cycle models revealed that costs of reproduction, measured at the population‐level, were small in the light environment, and more than buffered by the increase in survival of flowering plants compared to non‐flowering plants. In the shade environment, however, the costs of reproduction were significant and made the difference between a stable (current) and a growing (without reproduction costs) population. 5. Synthesis. Light penetration to the soil is a key variable determining population dynamics of woodland orchids. Our analyses show that differences in vital rates related to size‐dependent reproduction (flowering) and growth are essential drivers of changes in orchid population dynamics in different light environments. The combination of IPMs and LTREs thus proves to be very useful when studying the fitness consequences of size‐ and context‐dependent phenomena like flowering strategies and costs of reproduction.