Are functional traits good predictors of demographic rates? Evidence from five neotropical forests

A central goal of comparative plant ecology is to understand how functional traits vary among species and to what extent this variation has adaptive value. Here we evaluate relationships between four functional traits (seed volume, specific leaf area, wood density, and adult stature) and two demographic attributes (diameter growth and tree mortality) for large trees of 240 tree species from five Neotropical forests. We evaluate how these key functional traits are related to survival and growth and whether similar relationships between traits and demography hold across different tropical forests. There was a tendency for a trade-off between growth and survival across rain forest tree species. Wood density, seed volume, and adult stature were significant predictors of growth and/or mortality. Both growth and mortality rates declined with an increase in wood density. This is consistent with greater construction costs and greater resistance to stem damage for denser wood. Growth and mortality rates also declined as seed volume increased. This is consistent with an adaptive syndrome in which species tolerant of low resource availability (in this case shade-tolerant species) have large seeds to establish successfully and low inherent growth and mortality rates. Growth increased and mortality decreased with an increase in adult stature, because taller species have a greater access to light and longer life spans. Specific leaf area was, surprisingly, only modestly informative for the performance of large trees and had ambiguous relationships with growth and survival. Single traits accounted for 9-55% of the interspecific variation in growth and mortality rates at individual sites. Significant correlations with demographic rates tended to be similar across forests and for phylogenetically independent contrasts as well as for cross-species analyses that treated each species as an independent observation. In combination, the morphological traits explained 41% of the variation in growth rate and 54% of the variation in mortality rate, with wood density being the best predictor of growth and mortality. Relationships between functional traits and demographic rates were statistically similar across a wide range of Neotropical forests. The consistency of these results strongly suggests that tropical rain forest species face similar trade-offs in different sites and converge on similar sets of solutions.     

Genetic and color morph differentiation in the Caribbean sea anemone<Emphasis Type="Italic"> Condylactis gigantea</Emphasis>

The distribution of phenotypic and genetic variation across environments can provide insights into local adaptation. The tropical sea anemone Condylactis gigantea inhabits a broad spectrum of coral-reef habitats and displays a variety of phenotypes, particularly with respect to color. At the coast of Discovery Bay, Jamaica, individuals with either pink or green tentacle tips show distinct distributions. Pink morphs are more abundant in the lagoon and in deeper areas, while green morphs are more abundant in the forereef and in shallower areas. We use DNA sequence data (ITS1-5.8S) to investigate if variation in color is associated with genetic differentiation in lagoon and forereef habitats about 5 km apart. Population genetic analyses reveal two distinct ITS1-5.8S variants, which differ in relative frequency. The two variants are present in both habitats, but a dearth of intermediates suggests reduced gene flow. In the lagoon, but not the forereef, ITS variants show an association with color. In order to address the potential ecological significance of color, we study UV absorbance and UV acclimatization capacities of pink and green color morphs in the lagoon. Color morphs differed significantly in UV-B absorbance. These results suggest genetic and ecological differentiation in the face of gene flow over short distances.Communicated by O. Kinne, Oldendorf/Luhe     

Modelling environmental stochasticity in adult survival for a long-lived species

Stochastic matrix population models are often used to help guide the management of animal populations. For a long-lived species, environmental stochasticity in adult survival will play an important role in determining outcomes from the model. One of the most common methods for modelling such stochasticity is to randomly select the value of adult survival for each year from a distribution with a specified mean and standard deviation. We consider four distributions that can provide realistic models for stochasticity in adult survival. For values of the mean and standard deviation that cover the range we would expect for long-lived species, all four distributions have similar shapes, with small differences in their skewness and kurtosis. This suggests that many of the outcomes from a population model will be insensitive to the choice of distribution, assuming that distribution provides a realistic model for environmental stochasticity in adult survival. For a generic age-structured model, the estimate of the long-run stochastic growth rate is almost identical for the four distributions, across this range of values for the mean and standard deviation. Model outcomes based on short-term projections, such as the probability of a decline over a 20-year period, are more sensitive to the choice of distribution.     

How sensitive are elasticities of long-run stochastic growth to how environmental variability is modelled?

A variable environment leaves a signature in a population's dynamics. Deriving statistical and mathematical models of how environmental variability affects population projections has - in the wake of reports of substantial climatic fluctuations - received much recent attention. If the model changes, then so too does the population projection. This is because a different model of environmental variability changes estimates of long-run stochastic growth, which is a function of demographic rates and their temporal sequence. Decomposing elasticities of long-run stochastic growth into constituent parts can assess the relative influence of different components. Here, we investigate the consequences of changing the environmental state definition, and therefore altering the shape of demographic rate distributions and their temporal sequence, by using age-structured matrix models to project vertebrate populations into the future under a range of environmental scenarios. The identity of the most influential demographic rate was consistent among all approaches that perturbed only the mean, but was not when only the variance was perturbed. Furthermore, the influence of each demographic rate fluctuated among projections by up to factors of six and two for changes to the variance and mean, respectively. These changes in influence depend in part upon how environmental variability - in particular, the color of environmental noise - is incorporated. In the light of predictions of increasing climatic variability in the future, these results suggest caution when drawing quantitative conclusions from stochastic population projections.     

Iteroparity in the variable environment of the salamander Ambystoma tigrinum

Simultaneous estimation of survival, reproduction, and movement is essential to understanding how species maximize lifetime reproduction in environments that vary across space and time. We conducted a four-year, capture-recapture study of three populations of eastern tiger salamanders (Ambystoma tigrinum tigrinum) and used multistate mark-recapture statistical methods to estimate the manner in which movement, survival, and breeding probabilities vary under different environmental conditions across years and among populations and habitats. We inferred how individuals may mitigate risks of mortality and reproductive failure by deferring breeding or by moving among populations. Movement probabilities among populations were extremely low despite high spatiotemporal variation in reproductive success and survival, suggesting possible costs to movements among breeding ponds. Breeding probabilities varied between wet and dry years and according to whether or not breeding was attempted in the previous year. Estimates of survival in the nonbreeding, forest habitat varied among populations but were consistent across time. Survival in breeding ponds was generally high in years with average or high precipitation, except for males in an especially ephemeral pond. A drought year incurred severe survival costs in all ponds to animals that attempted breeding. Female salamanders appear to defer these episodic survival costs of breeding by choosing not to breed in years when the risk of adult mortality is high. Using stochastic simulations of survival and breeding under historical climate conditions, we found that an interaction between breeding probabilities and mortality limits the probability of multiple breeding attempts differently between the sexes and among populations.     

Mate choice games, context-dependent good genes, and genetic cycles in the side-blotched lizard, Uta stansburiana

According to mate choice models, a female should prefer males with traits that are reliable indicators of genetic quality which the sire can pass on to their progeny. However, good genes may depend on the social environment, and female choice for good genes should be context dependent. The side-blotched lizard, Uta stansburiana, exhibits genetically based throat colors (orange, blue, or yellow) that could be used as a sexually selected signal since they reliably predict the genetic quality of mates. The frequencies of male and female morphs cycle between years, and both male and female morphs have an advantage when rare; thus genetic quality will depend on morph frequency. A female should choose a sire that maximizes the reproductive success of both male and female progeny. We examine a game theoretical model that predicts female mate choice as a function of morph frequency and population density. The model predicts the following flexible mate choice rule: both female morphs should prefer rare males in ’boom years’ of the female cycle (e.g., ’rarest-of-N rule’), but prefer orange males in ’crash years’ of the female cycle (’orange-male rule’). Cues from the current social environment should be used by females to choose a mate that maximizes the future reproductive success of progeny, given the social environment of the next generation. We predict that the cue is the density of aggressive orange females. In the side-blotched lizard, cycling mate choice games and context-dependent mate choice are predicted to maintain genetic variation in the presence of choice for good genes. Received: 8 March 2000 / Revised: 26 August 2000 / Accepted: 4 September 2000     

Seed availability and insect herbivory limit recruitment and adult density of native tall thistle

Understanding spatial and temporal variation in factors influencing plant regeneration is critical to predicting plant population growth. We experimentally evaluated seed limitation, insect herbivory, and their interaction in the regeneration and density of tall thistle (Cirsium altissimum) across a topographic ecosystem productivity gradient in tallgrass prairie over two years. On ridges and in valleys, we used a factorial experiment manipulating seed availability and insect herbivory to quantify effects of: seed input on seedling density, insect herbivory on juvenile density, and cumulative impacts of both seed input and herbivory on reproductive adult density. Seed addition increased seedling densities at three of five sites in 2006 and all five sites in 2007. Insect herbivory reduced seedling survival across all sites in both years, as well as rosette survival from the previous year's seedlings. In both years, insecticide treatment of seed addition plots led to greater adult tall thistle densities in the following year, reflecting the increase in juvenile thistle densities in the experimental year. Seedling survival was not density dependent. Our analytical projection model predicts a significant long-term increase in adult densities from seed input, with a greater increase under experimentally reduced insect herbivory. While plant community biomass and water stress varied significantly between ridges and valleys, the effects of seed addition and insect herbivory did not vary with gradient position. These results support conceptual models that predict seedling and adult densities of short-lived monocarpic perennial plants should be seed limited. Further, the experiment demonstrates that even at high juvenile plant densities, at which density dependence potentially could have overridden herbivore effects on plant survival, insect herbivory strongly affected juvenile thistle performance and adult densities of this native prairie species.     

Genetic and environmental effects on the growth rate of Littorina saxatilis

Transfer experiments with two morphs of Littorina saxatilis Olivi (=L. rudis) typically inhabiting exposed and sheltered localities, showed a between-morph difference in shell growth in the same type of habitat, and a withinmorph difference between exposed and sheltered environments. The former indicates a genetic difference between the two morphs, although growth rate has an environmental component as shown by the latter. Juvenile snails of the exposed morph were on average slightly larger than sheltered morph juveniles on hatching, but at 20 wk, when raised in identical environments, the sheltered morph juveniles had grown significantly larger than the exposed ones. A rise in temperature from 5° to 10°C enhanced growth rate for snails raised in the laboratory. Temperature alone could not however explain increased growth during the spring and summer in natural populations.     

Differential breeding investment in bridled and non-bridled common guillemots (Uria aalge): morph of the partner matters

Polymorphism is the coexistence of two or more phenotypically distinct and genetically determinate forms in a population and implies a selective balance between the alternative morphs to permanently exist. Common guillemots Uria aalge occur in two genetically distinct morphs—a non-bridled and a bridled—the latter with white eye rings and a well-defined stripe behind the eyes. In this study, we investigated differences between the morphs with regard to reproductive parameters. We used a detailed family-based sample providing data on mother, father, and chick over three breeding seasons. The mating between morphs was random but pure non-bridled and pure bridled pairs produced smaller chicks at age 15 days than the two mixed pair compositions. Body mass of the adults showed much the same pattern; pure pairs having lower body mass than mixed pairs. There were no differences between the two morphs in reproductive parameters without considering the morph of the partner. This suggests that reproductive decisions in some way not only depend on the morph but also on the tactic of the partner. Different reproductive strategies between morph family groups as found in this study may contribute to the understanding of the existence of a balanced polymorphism in common guillemots. The overall breeding conditions during the years of this study were good. However, over time in a variable environment, we suggest that tactics of different family groups may have different success stabilizing the frequency of non-bridled and bridled birds over time.     

The triple helix of Plantago lanceolata: genetics and the environment interact to determine population dynamics

The theory of evolution via natural selection predicts that the genetic composition of wild populations changes over time in response to the environment. Different genotypes should exhibit different demographic patterns, but genetic variation in demography is often impossible to separate from environmental variation. Here, we asked if genetic variation is important in determining demographic patterns. We answer this question using a long-term field experiment combined with general linear modeling of deterministic population growth rates (lambda), deterministic life table response experiment (LTRE) analysis, and stochastic simulation of demography by paternal lineage in a short-lived perennial plant, Plantago lanceolata, in which we replicated genotypes across four cohorts using a standard breeding design. General linear modeling showed that growth rate varied significantly with year, spatial block, and sire. In LTRE analysis of all cohorts, the strongest influences on growth rate were from year x spatial block, and cohort x year x spatial block interactions. In analysis of genetics vs. temporal environmental variation, the strongest impacts on growth rate were from year and year x sire. Finally, stochastic simulation suggested different genetic composition among cohorts after 100 years, and different population growth rates when genetic differences were accounted for than when they were not. We argue that genetic variation, genotype x environment interactions, natural selection, and cohort effects should be better integrated into population ecological studies, as these processes should result in deviations from projected deterministic and stochastic population parameters.     

Age-size plasticity for reproduction in monocarpic plants

Empirical and theoretical investigations of monocarpy have usually addressed the question of minimum or threshold sizes for reproduction. However, the range of flowering sizes observed in many monocarpic species is extraordinarily large (well beyond what can be called a "threshold"), and the sizes of flowering and nonflowering plants may overlap greatly. We attempt to explain these reproductive patterns in terms of optimal reaction norms predicted by simple deterministic life history models. We assume that individuals differ in their growth trajectories due to the heterogeneous quality of microsites and ask how the optimal age and size at flowering varies with environmental variation in growth and for different assumptions about fecundity and mortality. Under two very different growth functions (one with no age- or size-related decline in growth rate and another with such a decline as size approaches an asymptote), the optimal reaction norms imply considerable plasticity for size at reproduction, particularly when poor growth is associated with higher mortality or lower asymptotic size. Deterministic models such as these may be more applicable to long-lived than to short-lived monocarps, because fitness potential should be less affected by stochastic variability in yearly growing condition in the former than in the latter. We consider the case of a tropical monocarpic and masting tree species, Cerberiopsis candelabra (Apocynaceae), and show that our model results can account for wide ranges of reproductive size and overlap in size of flowering and nonflowering plants, in accord with observation. We suggest that empirical attention to norms of reaction across growth environments will be a more profitable approach than investigation of size thresholds per se.     

Social,environmental and genetic factors in the ontogeny of phenotypic differentiation in a lizard with alternative male reproductive strategies

Summary Adult male tree lizards, Urosaurus ornatus, practise alternative (territorial or sneaker/satellite) reproductive strategies that are correlated with differences in throat color and body size. In this study we raised tree lizards from hatching in the laboratory to examine the question of whether the phenotypic expression of secondary sex coloration and body size can be facultatively influenced by social or abiotic environmental factors. We compared males reared in the laboratory under different social and environmental conditions to males in the field and found no effect of different conditions on phenotypic differentiation (Figs. 2–4). Thus, phenotypic differences between morphs probably result largely from nonfacultative expression of different genotypes. This suggests that alternative male morphs practise a mixed evolutionary stable strategy (ESS) rather than one morph making the best of a bad situation. However, in the context of ESS theory it is difficult to explain our further result that the nonterritorial morph in this species grows faster and reaches a larger adult body size than the territorial morph (Fig. 5).     

Allozyme electrophoresis demonstrates that the scleractinian coral Montipora digitata is two species

Allozyme electrophoresis at five loci was used to clarify the species status of two morphs of the scleractinian coral Montipora digitata (Dana, 1846), using specimens collected from three locations off the Queensland coast between October 1991 and April 1993. The two morphs, occurring sympatrically, were distinguished by one fixed gene difference at Locus LT-2 and frequency differences at three other loci, indicating reproductive isolation between them. Populations of both morphs were generally in Hardy-Weinberg equilibrium, suggesting that sexual reproduction does occur. Despite sampling designed to reduce the chance of collecting clonemates, the genetic diversity ratio (G _O:G_E) indicated that asexual reproduction also occurs. This is in agreement with the known modes of reproduction for this species. There was significant genetic differentiation between populations of one of the morphs of M. digitata. This could have been influenced by greater degrees of asexual reproduction in this morph, selection within sites, or variation in larval survival and recruitment patterns. Relatively low F _st (differentiation among populations) values found for a broadcast spawning species during this study are in accordance with the hypothesis that brooded planulae are adapted for rapid settlement, whereas planulae produced by broadcast spawners are adapted for widespread dispersal.     

Interactive effects of prey and p,p'-DDE on burrowing owl population dynamics.

We used population models to explore the effects of the organochlorine contaminant p,p'-DDE and fluctuations in vole availability on the population dynamics of Burrowing Owls (Athene cunicularia). Previous work indicated an interaction between low biomass of voles in the diet and moderate levels of p,p'-DDE in Burrowing Owl eggs that led to reproductive impairment. We constructed periodic and stochastic matrix models that incorporated three vole population states observed in the field: average, peak, and crash years. We modeled varying frequencies of vole crash years and a range of impairment of owl demographic rates in vole crash years. Vole availability had a greater impact on owl population growth rate than did reproductive impairment if vole populations peaked and crashed frequently. However, this difference disappeared as the frequency of vole crash years declined to once per decade. Fecundity, the demographic rate most affected by p,p'-DDE, had less impact on population growth rate than adult or juvenile survival. A life table response experiment of time-invariant matrices for average, peak, and crash vole conditions showed that low population growth under vole crash conditions was due to low adult and juvenile survival rates, whereas the extremely high population growth under vole peak conditions was due to increased fecundity. Our results suggest that even simple models can provide useful insights into complex ecological interactions. This is particularly valuable when temporal or spatial scales preclude manipulative experimental work in the field or laboratory.     

Social structures,genetic structures and dispersal strategies in Australian rabbit (<Emphasis Type="Italic">Oryctolagus cuniculus</Emphasis>) populations

In this study, the pattern of movement of young male and female rabbits and the genetic structures present in adult male and female populations in four habitats was examined. The level of philopatry in young animals was found to vary between 18-90% for males and 32-95% for females in different populations. It was skewed, with more males dispersing than females in some but not all populations. Analysis of allozyme data using spatial autocorrelation showed that adult females from the same social group, unlike males, were significantly related in four of the five populations studied. Changes in genetic structure and rate of dispersal were measured before and during the recovery of a population that was artificially reduced in size. There were changes in the rate and distance of dispersal with density and sex. Subadults of both sexes moved further in the first year post crash (low density) than in the following years. While the level of dispersal for females was lower than that of the males for the first 3 years, thereafter (high density) both sexes showed similar, low levels of dispersal (20%). The density at which young animals switch behaviour between dispersal and philopatry differed for males and females. The level of genetic structuring in adult females was high in the precrash population, reduced in the first year post crash and undetectable in the second year. Dispersal behaviour of rabbits both affects the genetic structure of the population and changes with conditions. Over a wide range of levels of philopatry, genetic structuring is present in the adult female , but not the male population. Consequently, though genetic structuring is present, it does not lead to inbreeding. More long-distance movements are found in low-density populations, even though vacant warrens are available near birth warrens. The distances moved decreased as density increased. Calculation of the effective population size (N_e) shows that changes in dispersal distance offset changes in density, so that N_e remains constant.     

Environmental Surrogates for Predicting and Conserving Adaptive Genetic Variability in Tree Species

Abstract: Adaptive genetic variability within species is an essential component of biodiversity but has been largely ignored in studies aimed at assessing and predicting biodiversity of the forest environment. We used factorial regression and structuring models to test easily measured surrogates, such as ecological attributes, as predictors of adaptive genetic variation between populations of a tree species ( Eucalyptus delegatensis ). Adaptive variability was defined in terms of variation in average growth performance of 68 populations and of population-by-environment interaction across seven different environments. The best surrogates of genetic variability were measures of solar radiation and temperature range, each predicting more than 50% of the genetic variability within the species. Rock and understory types, when used either alone or in combination with other covariates, also were very efficient in discriminating between populations in groups showing similar adaptation. Significant relationships between particular surrogates and growth patterns of variation were attributed to effects of natural selection that had occurred in the population source locations. We recommend the development of studies focusing on the population level of biodiversity to improve the conservation of forest ecosystems in Australia.     

Genetic mechanisms of pollution resistance in a marine invertebrate.

Pollution is a common stress in the marine environment and one of today's most powerful agents of selection, yet we have little understanding of how anthropogenic toxicants influence mechanisms of adaptation in marine populations. Due to their life history strategies, marine invertebrates are unable to avoid stress and must adapt to variable environments. We examined the genetic basis of pollution resistance across multiple environments using the marine invertebrate, Styela plicata. Gametes were crossed in a quantitative genetic breeding design to enable partitioning of additive genetic variance across a concentration gradient of a common marine pollutant, copper. Hatching success was scored as a measure of stress resistance in copper concentrations of 0, 75, 150, and 350 microg/L. There was a significant genotype x environment interaction in hatching success across copper concentrations. Further analysis using factor analytic modeling confirmed a significant dimension of across-environment genetic variation where the genetic basis of resistance to stress in the first three environments differed from that in the environment of highest copper concentration. A second genetic dimension further differentiated between the genetic basis of resistance to low and high stress environments. These results suggest that marine organisms use different genetic mechanisms to adapt to different levels of pollution and that the level of genetic variation to adapt to intense pollution stresses may be limited.     

Environment-specific elasticity and sensitivity analysis of the stochastic growth rate

For matrix population models, analyses of how sensitive the population growth rate is to changes in vital rates (i.e. perturbations) are important for studies of life history evolution as well as for management and conservation of threatened species. There are two types of sensitivity analyses corresponding to absolute (sensitivity) or relative (elasticity) changes in the vital rates and both types can be applied to both deterministic and stochastic matrix population models. To date, most empirical studies of elasticity and sensitivity of the stochastic growth rate have examined the response to perturbations in the vital rates in a complete set of possible environments. However, it is often of interest to examine the response to perturbations occurring in only a subset of the possible environments. This has been done for periodic time-varying models elsewhere, but here we describe a recently published method for calculating the environment-specific sensitivity and elasticity of the stochastic growth rate and apply this method to data. These environment-specific perturbation analyses provide a logical way of dividing the sensitivity and elasticity among the environments. They give important insight into the selection regime in different environments and can provide valuable information for making management decisions and management evaluations in stochastic environments.     

The importance of space, time, and stochasticity to the demography and management of Alliaria petiolata

As population modeling is increasingly called upon to guide policy and management, it is important that we understand not only the central tendencies of our study systems, but the consequences of their variation in space and time as well. The invasive plant Alliaria petiolata (garlic mustard) is actively managed in the United States and is the focus of a developing biological control program. Two weevils (Coleoptera: Curculionidae: Ceutorhynchus) that reduce fecundity (C. alliariae) and rosette survival plus fecundity (C. scrobicollis) are under consideration for release pending host specificity testing. We used a demographic modeling approach to (1) quantify variability in A. petiolata growth and vital rates and (2) assess the potential for single- or multiple-agent biocontrol to suppress growth of 12 A. petiolata populations in Illinois and Michigan studied over three plant generations. We used perturbation analyses and simulation models with stochastic environments to estimate stochastic growth rates (lambda(S)) and predict the probability of successful management using either a single biocontrol agent or two agent species together. Not all populations exhibited invasive dynamics. Estimates of lambda(S) ranged from 0.78 to 2.21 across sites, while annual, deterministic growth (lambda) varied up to sevenfold within individual sites. Given our knowledge of the biocontrol agents, this analysis suggests that C. scrobicollis alone may control A. petiolata at up to 63% of our study sites where lambda >1, with the combination of both agents predicted to succeed at 88% of sites. Across sites and years, the elasticity rankings were dependent on lambda. Reductions of rosette survival, fecundity, or germination of new seeds are predicted to cause the greatest reduction of lambda in growing populations. In declining populations, transitions affecting seed bank survival have the greatest effect on lambda. This contrasts with past analyses that varied parameters individually in an otherwise constant matrix, which may yield unrealistic predictions by decoupling natural parameter covariances. Overall, comparisons of stochastic and deterministic growth rates illustrate how analyses of individual populations or years could misguide management or fail to characterize complex traits such as invasiveness that emerge as attributes of populations rather than species.     

Combined effects of temperature and salinity on the survival and growth of the larvae of Pandalus jordani (Decapoda: Pandalidae)

Survival and growth over an environmental range of temperature and salinities were examined in order to help assess the importance of these environmental factors in affecting the distribution, abundance and survival of larvae and provide greater understanding of factors affecting fluctuations in adult Pandalus jordani Rathbun population sizes. Larvae were shown to have a wide tolerance to salinity, especially in the early stages, but a relatively narrow tolerance to temperature. The optimal temperatures for survival, 8° to 11°C, were also optimal for growth as reflected by maximal growth increments and body size. It is therefore felt that fluctuations in temperature as seen within and between successive larval seasons would have profound effects on larval survival, growth rates and size at metamorphosis to the benthic juvenile phase.