Recruitment rates exhibit high elasticity and high temporal variation in populations of a short-lived perennial herb

Empirical studies for different life histories have shown an inverse relationship between elasticity (i.e. the proportional contribution to population growth rate) and temporal variation in vital rates. It is accepted that this relationship indicates the effect of selective pressures in reducing variation in those life‐history traits with a major impact on fitness. In this paper, we sought to determine whether changes in environmental conditions affect the relationship between elasticity of vital rates and their temporal variation, and whether vital rates with simultaneously large elasticity and temporal variation might represent a characteristic life‐history strategy. We used demographic data on 13 populations of the short‐lived Hypericum cumulicola over 5–6 years, in three time‐since‐fire classes. For each population of each time‐since‐fire, we computed the mean matrix over years and its respective elasticity matrix, and the coefficients of variation in matrix entries over study years as an estimate of temporal variability. We found that mean elasticity negatively significantly correlated with temporal variation in vital rates in populations (overall eight out of 13) included in each time‐since‐fire. However, seedling recruitment exhibited both high elasticity and high temporal variation in almost all study populations. These results indicated that (1) the general relationship between elasticity and temporal variation in vital rates was not modified by environmental changes due to time‐since‐fire, and (2) high elasticity and high temporal variation in seedling recruitment in H. cumulicola is a particular trait of the species' life history. After seed survival in the soil seed bank, seedling recruitment represents the most important life‐history trait influencing H. cumulicola population growth rate (and fitness). The high temporal variability in seedling recruitment suggests that this trait is determined by environmental cues, leading to an increase in population size and subsequent replenishment of the seed bank in favorable years.     

Population declines of tuna and relatives depend on their speed of life

Larger-bodied species in a wide range of taxonomic groups including mammals, fishes and birds tend to decline more steeply and are at greater risk of extinction. Yet, the diversity in life histories is governed not only by body size, but also by time-related traits. A key question is whether this size-dependency of vulnerability also holds, not just locally, but globally across a wider range of environments. We test the relative importance of size- and time-related life-history traits and fishing mortality in determining population declines and current exploitation status in tunas and their relatives. We use high-quality datasets of half a century of population trajectories combined with population-level fishing mortalities and life-history traits. Time-related traits (e.g. growth rate), rather than size-related traits (e.g. maximum size), better explain the extent and rate of declines and current exploitation status across tuna assemblages, after controlling for fishing mortality. Consequently, there is strong geographical patterning in population declines, such that populations with slower life histories (found at higher cooler latitudes) have declined most and more steeply and have a higher probability of being overfished than populations with faster life histories (found at tropical latitudes). Hence, the strong, temperature-driven, latitudinal gradients in life-history traits may underlie the global patterning of population declines, fisheries collapses and local extinctions.     

Are certain life histories particularly prone to local extinction?

Using stochastic simulations and elasticity analysis, we show that there are inherent differences in the risk of extinction between life histories with different demographies. Which life history is the most vulnerable depends on which vital rate varies. When juvenile survival varies semelparous organisms with delayed reproduction are the most vulnerable ones, while a varying developmental rate puts a greater threat to semelparous organisms with rapid development. Iteroparous organisms are the most vulnerable ones when adult survival varies. Generally, we do not expect to observe organisms in nature having variation in vital rates that produce a high risk of extinction. Given the results here we therefore predict that iteroparous organisms should show low variation in adult survival. Moreover, we predict that semelparous organisms should show low variation in juvenile survival and low variation in developmental rate. The effect of temporal correlation on extinction risk is most pronounced in organisms with semelparous life histories.     

Minimum viable population sizes and global extinction risk are unrelated

Theoretical and empirical work has shown that once reduced in size and geographical range, species face a considerably elevated risk of extinction. We predict minimum viable population sizes (MVP) for 1198 species based on long-term time-series data and model-averaged population dynamics simulations. The median MVP estimate was 1377 individuals (90% probability of persistence over 100 years) but the overall distribution was wide and strongly positively skewed. Factors commonly cited as correlating with extinction risk failed to predict MVP but were able to predict successfully the probability of World Conservation Union Listing. MVPs were most strongly related to local environmental variation rather than a species' intrinsic ecological and life history attributes. Further, the large variation in MVP across species is unrelated to (or at least dwarfed by) the anthropogenic threats that drive the global biodiversity crisis by causing once-abundant species to decline.     

Assessing extinction risks under the combined effects of climate change and human disturbance through the analysis of life-history plasticity

Plastic responses of species to unprecedented conditions and increased variability caused by climate change (CC) and anthropogenic disturbances are expected to play a major role determining populations’ extinction risk. We developed a method for assessing CC effects on population dynamics based on analyzing plastic life history responses to weather conditions within an ensemble forecasting framework. The method is illustrated using two threatened cactus species with contrasting distribution ranges. Demographic models were parameterized using 5 years of field data, from which the relationship between life-history traits and rainfall and temperature was estimated. These functions were used in Integral Projection Models to predict population growth under CC scenarios and different anthropogenic disturbance regimes. Both species were affected by CC and its interaction with disturbance. The most widespread species was less affected by CC, suggesting that past selection on plasticity allows it to survive under variable conditions. Managing disturbance appropriately lessened the impact of CC. Some directives for conservation under CC were identified based on projected elasticity values. Our procedure for modeling population dynamics as a function of climate may be used for designing management plans for conservation or sustainable use aimed at important plant sizes or life-history traits, predicting potential distributions, and identifying viable populations.     

Lizards from Urban Areas Are More Asymmetric: Using Fluctuating Asymmetry to Evaluate Environmental Disturbance

The increase in human activities that leads to wildlife decline and species extinction poses an urgent need for simple indicators of environmental stress in animal populations. Several studies have suggested that fluctuating asymmetry (FA) can be an easy, direct measure of developmental instability because it is associated to environmental stress and, as such, it can be a useful indicator of population disturbance. We examined three different morphological traits in urban and rural populations of the common wall lizard (Podarcis muralis) to test whether anthropogenic disturbance causes an increase in FA. Compared to rural populations, urban ones showed higher levels of FA in all analyzed traits, thus providing evidence that FA can respond to anthropogenic disturbance. However, we also found significant differences in FA among traits, where femoral pores and subdigital lamellae, traits with a functional relevance, were more stable developmentally compared to supracilliar granules which have no evident function. Unsigned FA [abs(right-left)] exhibited significant, but weak, positive correlations among traits, indicating that developmental noise does not have a uniform effect across characters and thus questioning the view of developmental stability as an organism-wide property. The degree of signed FA (right-left) was more similar between structurally associated traits, possibly as an outcome of morphological integration. In conclusion, our results demonstrate that FA can be a reliable indicator of disturbance provided that it is analyzed on multiple traits simultaneously and examined at the population level.     

Buffering and plasticity in vital rates of oldfield rodents

1. Under the hypothesis of environmental buffering, populations are expected to minimize the variance of the most influential vital rates; however, this may not be a universal principle. Species with a life span <1 year may be less likely to exhibit buffering because of temporal or seasonal variability in vital rate sensitivities. Further, plasticity in vital rates may be adaptive for species in a variable environment with reliable cues. 2. We tested for environmental buffering and plasticity in vital rates using stage-structured matrix models from long-term data sets in four species of grassland rodents. We used periodic matrices to estimate stochastic elasticity for each vital rate and then tested for correlations with a standardized coefficient of variation for each rate. 3. We calculated stochastic elasticities for individual months to test for an association between increased reproduction and the influence of reproduction, relative to survival, on the population growth rate. 4. All species showed some evidence of buffering. The elasticity of vital rates of Peromyscus leucopus (Rafinesque, 1818), Sigmodon hispidus Say & Ord, 1825 and Microtus ochrogaster (Wagner, 1842) was negatively related to vital rate CV. Elasticity and vital rate CV were negatively related in Peromyscus maniculatus (Wagner, 1845), but the relationship was not statistically significant. Peromyscus leucopus and M. ochrogaster showed plasticity in vital rates; reproduction was higher following months where elasticity for reproduction exceeded that of survival. 5. Our results suggest that buffering is common in species with fast life histories; however, some populations that exhibit buffering are capable of responding to short-term variability in environmental conditions through reproductive plasticity.     

Small-scale spatial and temporal variation in the demographic processes underlying the large-scale decline of eiders in the Baltic Sea

The application of uniform conservation schemes often fails to account for small-scale spatial variation in the drivers of population decline. Demographic comparisons of imperilled populations across locations are therefore crucial for successful conservation, but progress is hampered by lack of long-term data from more than a single population. The recent large-scale decline of eider ducks (Somateria mollissima) in the Baltic Sea is ideal for determining to what extent mechanisms underlying population decline can be extrapolated over larger areas. We utilized stochastic demographic methods incorporating both environmental and sampling variation to assess small-scale spatial and temporal variation in the population dynamics of eiders at Söderskär (eastern range-margin) and Tvärminne (core breeding area), situated 130 km apart. The stochastic growth rate models accurately predicted the observed differences in the rate of decline between sites and time periods. At Söderskär, established breeder survival had by far the greatest elasticity, whereas elasticity was more evenly distributed among vital rates at Tvärminne. Although the study sites showed the single largest difference in fecundity, stochastic life table response experiment analyses revealed that reduced adult female survival at Tvärminne mainly determined the observed difference in growth rates between sites. In contrast, reduced fecundity primarily differentiated the past population increase from the present population decline at Söderskär. Our results demonstrate that different mechanisms may be associated with population decline across adjacent geographic locations, and indicate that dispersal of first-time breeders may be important for population dynamics. Safeguarding adult female survival and/or fecundity should be prioritized in management efforts.     

Distribution, ecology, life history, genetic variation, and risk of extinction of nonhuman primates from Costa Rica

We examined the association between geographic distribution, ecological traits, life history, genetic diversity, and risk of extinction in nonhuman primate species from Costa Rica. All of the current nonhuman primate species from Costa Rica are included in the study; spider monkeys (Ateles geoffroyi), howling monkeys (Alouatta palliata), capuchins (Cebus capucinus), and squirrel monkeys (Saimiri oerstedii). Geographic distribution was characterized accessing existing databases. Data on ecology and life history traits were obtained through a literature review. Genetic diversity was characterized using isozyme electrophoresis. Risk of extinction was assessed from the literature. We found that species differed in all these traits. Using these data, we conducted a Pearson correlation between risk of extinction and ecological and life history traits, and genetic variation, for widely distributed species. We found a negative association between risk of extinction and population birth and growth rates; indicating that slower reproducing species had a greater risk of extinction. We found a positive association between genetic variation and risk of extinction; i.e., species showing higher genetic variation had a greater risk of extinction. The relevance of these traits for conservation efforts is discussed.     

Sheltered from the storm? Population viability analysis of a rare endemic under periodic catastrophe regimes

Rare species are important targets for biodiversity conservation efforts because rarity often equates to small populations and increased endangerment. Rare species are prone to stochastic extinction events and may be particularly susceptible to catastrophes. Therefore, understanding how rare species respond to disturbances is critical for evaluating extinction risk and guiding conservation managers. Population viability analyses (PVAs) are essential for assessing rare species' status yet they seldom consider catastrophic events. Accordingly, we present a PVA of a rare tropical epiphyte, Lepanthes caritensis (Orchidaceae), under simulated disturbance regimes to evaluate its demographics and extinction risk. We aimed to test how demographic models incorporating catastrophes affect population viability estimates. Our goal was to better guide management of these orchids and other rare plants. Results revealed L. caritensis numbers have declined recently, but projected growth rates indicated that most subpopulations should increase in size if undisturbed. Still, projection models show that moderate catastrophes reduce growth rates, increase stochasticity in subpopulation sizes, and elevate extinction risk. Severe catastrophes had a more pronounced effect in simulations; growth rates fell below replacement level, there was greater variation in projected population sizes, and extinction risk was significantly higher. PVAs incorporating periodic catastrophes indicate that rare species may have greater extinction probabilities than standard models suggest. Thus, precautionary conservation measures should be taken in disturbance prone settings and we encourage careful monitoring after environmental catastrophes. Future rare plant PVAs should incorporate catastrophes and aim to determine if rescue and reintroduction efforts are necessary after disturbances to insure long-term population viability.     

Interactive life‐history traits predict sensitivity of plants and animals to temporal autocorrelation

Temporal autocorrelation in demographic processes is an important aspect of population dynamics, but a comprehensive examination of its effects on different life‐history strategies is lacking. We use matrix population models from 454 plant and animal populations to simulate stochastic population growth rates (log λ_s) under different temporal autocorrelations in demographic rates , using simulated and observed covariation among rates. We then test for differences in sensitivities, or changes of log λ_s to changes in autocorrelation among two major axes of life‐history strategies, obtained from phylogenetically informed principal component analysis: the fast‐slow and reproductive‐strategy continua. Fast life histories exhibit highest sensitivities to simulated autocorrelation in demographic rates across reproductive strategies. Slow life histories are less sensitive to temporal autocorrelation, but their sensitivities increase among highly iteroparous species. We provide cross‐taxonomic evidence that changes in the autocorrelation of environmental variation may affect a wide range of species, depending on complex interactions of life‐history strategies.     

Population effects of increased climate variation

Global circulation models predict and numerous observations confirm that anthropogenic climate change has altered high-frequency climate variability. However, it is not yet well understood how changing patterns of environmental variation will affect wildlife population dynamics and other ecological processes. Theory predicts that a population's long-run growth rate is diminished and the chance of population extinction is increased as environmental variation increases. This results from the fact that population growth is a multiplicative process and that long-run population growth rate is the geometric mean of growth rates over time, which is always less than the arithmetic mean. However, when population growth rates for unstructured populations are related nonlinearly to environmental drivers, increasing environmental variation can increase a population's long-run growth rate. This suggests that patterns of environmental variation associated with different aspects of climate change may affect population dynamics in different ways. Specifically, increasing variation in rainfall might result in diminished long-run growth rates for many animal species while increasing variation in temperature might result in increased long-run growth rates. While the effect of rainfall is theoretically well understood and supported by data, the hypothesized effect of temperature is not. Here, I analyse two datasets to study the effect of fluctuating temperatures on growth rates of zooplankton. Results are consistent with the prediction that fluctuating temperatures should increase long-run growth rates and the frequency of extreme demographic events.     

Geographical variation in predictors of mammalian extinction risk: big is bad, but only in the tropics

Whereas previous studies have investigated correlates of extinction risk either at global or regional scales, our study explicitly models regional effects of anthropogenic threats and biological traits across the globe. Using phylogenetic comparative methods with a newly-updated supertree of 5020 extant mammals, we investigate the impact of species traits on extinction risk within each WWF ecoregion. Our analyses reveal strong geographical variation in the influence of traits on risk: notably, larger species are at higher risk only in tropical regions. We then relate these patterns to current and recent-historical human impacts across ecoregions using spatial modelling. The body–mass results apparently reflect historical declines of large species outside the tropics due to large-scale land conversion. Narrow-ranged and rare species tend to be at high risk in areas of high current human impacts. The interactions we describe between biological traits and anthropogenic threats increase understanding of the processes determining extinction risk.     

Relating life‐history traits,environmental constraints and local extinctions in river fish

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Human Population Density and Extinction Risk in the World's Carnivores

Understanding why some species are at high risk of extinction, while others remain relatively safe, is central to the development of a predictive conservation science. Recent studies have shown that a species' extinction risk may be determined by two types of factors: intrinsic biological traits and exposure to external anthropogenic threats. However, little is known about the relative and interacting effects of intrinsic and external variables on extinction risk. Using phylogenetic comparative methods, we show that extinction risk in the mammal order Carnivora is predicted more strongly by biology than exposure to high-density human populations. However, biology interacts with human population density to determine extinction risk: biological traits explain 80% of variation in risk for carnivore species with high levels of exposure to human populations, compared to 45% for carnivores generally. The results suggest that biology will become a more critical determinant of risk as human populations expand. We demonstrate how a model predicting extinction risk from biology can be combined with projected human population density to identify species likely to move most rapidly towards extinction by the year 2030. African viverrid species are particularly likely to become threatened, even though most are currently considered relatively safe. We suggest that a preemptive approach to species conservation is needed to identify and protect species that may not be threatened at present but may become so in the near future.     

Human Population Density and Extinction Risk in the World's Carnivores

Understanding why some species are at high risk of extinction, while others remain relatively safe, is central to the development of a predictive conservation science. Recent studies have shown that a species' extinction risk may be determined by two types of factors: intrinsic biological traits and exposure to external anthropogenic threats. However, little is known about the relative and interacting effects of intrinsic and external variables on extinction risk. Using phylogenetic comparative methods, we show that extinction risk in the mammal order Carnivora is predicted more strongly by biology than exposure to high-density human populations. However, biology interacts with human population density to determine extinction risk: biological traits explain 80% of variation in risk for carnivore species with high levels of exposure to human populations, compared to 45% for carnivores generally. The results suggest that biology will become a more critical determinant of risk as human populations expand. We demonstrate how a model predicting extinction risk from biology can be combined with projected human population density to identify species likely to move most rapidly towards extinction by the year 2030. African viverrid species are particularly likely to become threatened, even though most are currently considered relatively safe. We suggest that a preemptive approach to species conservation is needed to identify and protect species that may not be threatened at present but may become so in the near future.     

Predicting extinction risk in declining species

What biological attributes predispose species to the risk of extinction? There are many hypotheses but so far there has been no systematic analysis for discriminating between them. Using complete phylogenies of contemporary carnivores and primates, we present, to our knowledge, the first comparative test showing that high trophic level, low population density slow life history and, in particular, small geographical range size are all significantly and independently associated with a high extinction risk in declining species. These traits together explain nearly 50% of the total between-species variation in extinction risk. Much of the remaining variation can be accounted for by external anthropogenic factors that affect species irrespective of their biology.     

High variability in patterns of population decline: the importance of local processes in species extinctions

A fundamental goal of conservation science is to improve conservation practice. Understanding species extinction patterns has been a central approach towards this objective. However, uncertainty remains about the extent to which species-level patterns reliably indicate population phenomena at the scale of local sites, where conservation ultimately takes place. Here, we explore the importance of both species- and site-specific components of variation in local population declines following habitat disturbance, and test a suite of hypotheses about their intrinsic and extrinsic drivers. To achieve these goals, we analyse an unusually detailed global dataset for species responses to habitat disturbance, namely primates in timber extraction systems, using cross-classified generalized linear mixed models. We show that while there are consistent differences in the severity of local population decline between species, an equal amount of variation also occurs between sites. The tests of our hypotheses further indicate that a combination of biological traits at the species level, and environmental factors at the site level, can help to explain these patterns. Specifically, primate populations show a more marked decline when the species is characterized by slow reproduction, high ecological requirements, low ecological flexibility and small body size; and when the local environment has had less time for recovery following disturbance. Our results demonstrate that individual species show a highly heterogeneous, yet explicable, pattern of decline. The increased recognition and elucidation of local-scale processes in species declines will improve our ability to conserve biodiversity in the future.     

A trait‐based approach for predicting species responses to environmental change from sparse data: how well might terrestrial mammals track climate change?

Estimating population spread rates across multiple species is vital for projecting biodiversity responses to climate change. A major challenge is to parameterise spread models for many species. We introduce an approach that addresses this challenge, coupling a trait‐based analysis with spatial population modelling to project spread rates for 15 000 virtual mammals with life histories that reflect those seen in the real world. Covariances among life‐history traits are estimated from an extensive terrestrial mammal data set using Bayesian inference. We elucidate the relative roles of different life‐history traits in driving modelled spread rates, demonstrating that any one alone will be a poor predictor. We also estimate that around 30% of mammal species have potential spread rates slower than the global mean velocity of climate change. This novel trait‐space‐demographic modelling approach has broad applicability for tackling many key ecological questions for which we have the models but are hindered by data availability.     

Stochastic demography and population dynamics in the red kangaroo Macropus rufus

1.  Many organisms inhabit strongly fluctuating environments but their demography and population dynamics are often analysed using deterministic models and elasticity analysis, where elasticity is defined as the proportional change in population growth rate caused by a proportional change in a vital rate. Deterministic analyses may not necessarily be informative because large variation in a vital rate with a small deterministic elasticity may affect the population growth rate more than a small change in a less variable vital rate having high deterministic elasticity.
2.  We analyse a stochastic environment model of the red kangaroo ( Macropus rufus ), a species inhabiting an environment characterized by unpredictable and highly variable rainfall, and calculate the elasticity of the stochastic growth rate with respect to the mean and variability in vital rates.
3.  Juvenile survival is the most variable vital rate but a proportional change in the mean adult survival rate has a much stronger effect on the stochastic growth rate.
4.  Even if changes in average rainfall have a larger impact on population growth rate, increased variability in rainfall may still be important also in long-lived species. The elasticity with respect to the standard deviation of rainfall is comparable to the mean elasticities of all vital rates but the survival in age class 3 because increased variation in rainfall affects both the mean and variability of vital rates.
5.  Red kangaroos are harvested and, under the current rainfall pattern, an annual harvest fraction of c . 20% would yield a stochastic growth rate about unity. However, if average rainfall drops by more than c . 10%, any level of harvesting may be unsustainable, emphasizing the need for integrating climate change predictions in population management and increase our understanding of how environmental stochasticity translates into population growth rate.