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E. Cam

Publications and source records attributed to E. Cam.

14 recordsLinked to original sources

Hierarchical modeling of an invasive spread: The eurasian collared-dove streptopelia decaocto in the United States

Invasive species are regularly claimed as the second threat to biodiversity. To apply a relevant response to the potential consequences associated with invasions (e.g., emphasize management efforts to prevent new colonization or to eradicate the species in places where it has already settled), it is essential to understand invasion mechanisms and dynamics. Quantifying and understanding what influences rates of spatial spread is a key research area for invasion theory. In this paper, we develop a model to account for occupancy dynamics of an invasive species. Our model extends existing models to accommodate several elements of invasive processes; we chose the framework of hierarchical modeling to assess site occupancy status during an invasion. First, we explicitly accounted for spatial structure and how distance among sites and position relative to one another affect the invasion spread. In particular, we accounted for the possibility of directional propagation and provided a way of estimating the direction of this possible spread. Second, we considered the influence of local density on site occupancy. Third, we decided to split the colonization process into two subprocesses, initial colonization and recolonization, which may be ground-breaking because these subprocesses may exhibit different relationships with environmental variations (such as density variation) or colonization history (e.g., initial colonization might facilitate further colonization events). Finally, our model incorporates imperfection in detection, which might be a source of substantial bias in estimating population parameters. We focused on the case of the Eurasian Collared-Dove (Streptopelia decaocto) and its invasion of the United States since its introduction in the early 1980s, using data from the North American BBS (Breeding Bird Survey). The Eurasian Collared-Dove is one of the most successful invasive species, at least among terrestrial vertebrates. Our model provided estimation of the spread direction consistent with empirical observations. Site persistence probability exhibits a quadratic response to density. We also succeeded at detecting differences in the relationship between density and initial colonization vs. recolonization probabilities. We provide a map of sites that may be colonized in the future as an example of possible practical application of our work. ?? 2011 by the Ecological Society of America.

Ecological Applications

Occam's shadow: levels of analysis in evolutionary ecology - where to next?

Evolutionary ecology is the study of evolutionary processes, and the ecological conditions that influence them. A fundamental paradigm underlying the study of evolution is natural selection. Although there are a variety of operational definitions for natural selection in the literature, perhaps the most general one is that which characterizes selection as the process whereby heritable variation in fitness associated with variation in one or more phenotypic traits leads to intergenerational change in the frequency distribution of those traits. The past 20 years have witnessed a marked increase in the precision and reliability of our ability to estimate one or more components of fitness and characterize natural selection in wild populations, owing particularly to significant advances in methods for analysis of data from marked individuals. In this paper, we focus on several issues that we believe are important considerations for the application and development of these methods in the context of addressing questions in evolutionary ecology. First, our traditional approach to estimation often rests upon analysis of aggregates of individuals, which in the wild may reflect increasingly non-random (selected) samples with respect to the trait(s) of interest. In some cases, analysis at the aggregate level, rather than the individual level, may obscure important patterns. While there are a growing number of analytical tools available to estimate parameters at the individual level, and which can cope (to varying degrees) with progressive selection of the sample, the advent of new methods does not reduce the need to consider carefully the appropriate level of analysis in the first place. Estimation should be motivated a priori by strong theoretical analysis. Doing so provides clear guidance, in terms of both (i) assisting in the identification of realistic and meaningful models to include in the candidate model set, and (ii) providing the appropriate context under which the results are interpreted. Second, while it is true that selection (as defined) operates at the level of the individual, the selection gradient is often (if not generally) conditional on the abundance of the population. As such, it may be important to consider estimating transition rates conditional on both the parameter values of the other individuals in the population (or at least their distribution), and population abundance. This will undoubtedly pose a considerable challenge, for both single- and multi-strata applications. It will also require renewed consideration of the estimation of abundance, especially for open populations. Thirdly, selection typically operates on dynamic, individually varying traits. Such estimation may require characterizing fitness in terms of individual plasticity in one or more state variables, constituting analysis of the norms of reaction of individuals to variable environments. This can be quite complex, especially for traits that are under facultative control. Recent work has indicated that the pattern of selection on such traits is conditional on the relative rates of movement among and frequency of spatially heterogeneous habitats, suggesting analyses of evolution of life histories in open populations can be misleading in some cases.

Journal of Applied Statistics

Dispersal and individual quality in a long lived species

The idea of differences in individual quality has been put forward in numerous long-term studies in long-lived species to explain differences in lifetime production among individuals. Despite the important role of individual heterogeneity in vital rates in demography, population dynamics and life history theory, the idea of 'individual quality' is elusive. It is sometimes assumed to be a static or dynamic individual characteristic. When considered as a dynamic trait, it is sometimes assumed to vary deterministically or stochastically, or to be confounded with the characteristics of the habitat. We addressed heterogeneity in reproductive performance among individuals established in higher-quality habitat in a long-lived seabird species. We used approaches to statistical inference based on individual random effects permitting quantification of heterogeneity in populations and assessment of individual variation from the population mean. We found evidence of heterogeneity in breeding probability, not success probability. We assessed the influence of dispersal on individual reproductive potential. Dispersal is likely to be destabilizing in species with high site and mate fidelity. We detected heterogeneity after dispersal, not before. Individuals may perform well regardless of quality before destabilization, including those that recruited in higher-quality habitat by chance, but only higher-quality individuals may be able to overcome the consequences of dispersal. Importantly, results differed when accounting for individual heterogeneity (an increase in mean breeding probability when individuals dispersed), or not (a decrease in mean breeding probability). In the latter case, the decrease in mean breeding probability may result from a substantial decrease in breeding probability in a few individuals and a slight increase in others. In other words, the pattern observed at the population mean level may not reflect what happens in the majority of individuals.

Oikos

Long-term fitness consequences of early conditions in the kittiwake

1. The long - term fitness consequences of conditions during development are receiving growing attention: they are at the interface between ecological and evolutionary processes. We addressed the influence of the length of the rearing period and 'rank' on fitness components in a long -lived seabird species with deferred breeding: the kittiwake (Rissa tridactyla). Rank, which depends on hatching order, was used as a surrogate for dominance status in the brood. Rank could be viewed as a random factor affecting individuals regardless of their possible 'intrinsic quality' at birth. The length of the rearing period was used as a surrogate for parental effort. It reflects the interaction between numerous factors such as environmental conditions , parental quality, reproductive decisions and effort, and also offspring decisions and intrinsic quality at birth. 2. There was evidence of an influence of the length of the rearing period on local survival before recruitment. Individuals with shorter rearing periods had lower local survival during the first winter (e.g. the relationship was positive). They may incur higher mortality. In rank 1 prebreeders, this relationship was negative in older age-classes. Longer rearing periods and better condition at independence may be associated with stronger migrating ability, and prebreeders that have not yet made settlement decisions may emigrate permanently to distant locations. Such a complex pattern may reflect age-related changes in the relative contribution of mortality and permanent emigration to local survival. 3. The length of the rearing period had long - term consequences on reproductive performance. The relationship was positive but the rate of increase decreased slightly at higher values of the covariate. 4. There was an unambiguously negative influence of rank on survival before recruitment and recruitment probability, but not on demographic parameters specific to the reproductive stage. Juniors recruited later than elder siblings. The disadvantage of juniors may be expressed mainly in terms of higher mortality and disappearance from the population before recruitment.

Journal of Animal Ecology

Stratification based on reproductive state reveals contrasting patterns of age-related variation in demographic parameters in the kittiwake

Heterogeneity in individual quality can be a major obstacle when interpreting age-specific variation in life-history traits. Heterogeneity is likely to lead to within-generation selection, and patterns observed at the population level may result from the combination of hidden patterns specific to subpopulations. Population-level patterns are not relevant to hypotheses concerning the evolution of age-specific reproductive strategies if they differ from patterns at the individual level. We addressed the influence of age and a variable used as a surrogate of quality (yearly reproductive state) on survival and breeding probability in the kittiwake. We found evidence of an effect of age and quality on both demographic parameters. Patterns observed in breeders are consistent with the selection hypothesis, which predicts age-related increases in survival and traits positively correlated with survival. Our results also reveal unexpected age effects specific to subgroups: the influence of age on survival and future breeding probability is not the same in nonbreeders and breeders. These patterns are observed in higher-quality breeding habitats, where the influence of extrinsic factors on breeding state is the weakest. Moreover, there is slight evidence of an influence of sex on breeding probability (not on survival), but the same overall pattern is observed in both sexes. Our results support the hypothesis that age-related variation in demographic parameters observed at the population level is partly shaped by heterogeneity among individuals. They also suggest processes specific to subpopulations. Recent theoreticaI developments lay emphasis on integration of sources of heterogeneity in optimization models to account for apparently 'sub-optimal' empirical patterns. Incorporation of sources of heterogeneity is also the key to investigation of age-related reproductive strategies in heterogeneous populations. Thwarting 'heterogeneity's ruses' has become a major challenge: for detecting and understanding natural processes, and a constructive confrontation between empirical and theoretical studies.

Brittany

Inferences about nested subsets structure when not all species are detected

Comparisons of species composition among ecological communities of different size have often provided evidence that the species in communities with lower species richness form nested subsets of the species in larger communities. In the vast majority of studies, the question of nested subsets has been addressed using information on presence-absence, where a '0' is interpreted as the absence of a given species from a given location. Most of the methodological discussion in earlier studies investigating nestedness concerns the approach to generation of model-based matrices. However, it is most likely that in many situations investigators cannot detect all the species present in the location sampled. The possibility that zeros in incidence matrices reflect nondetection rather than absence of species has not been considered in studies addressing nested subsets, even though the position of zeros in these matrices forms the basis of earlier inference methods. These sampling artifacts are likely to lead to erroneous conclusions about both variation over space in species richness and the degree of similarity of the various locations. Here we propose an approach to investigation of nestedness, based on statistical inference methods explicitly incorporating species detection probability, that take into account the probabilistic nature of the sampling process. We use presence-absence data collected under Pollock?s robust capture-recapture design, and resort to an estimator of species richness originally developed for closed populations to assess the proportion of species shared by different locations. We develop testable predictions corresponding to the null hypothesis of a nonnested pattern, and an alternative hypothesis of perfect nestedness. We also present an index for assessing the degree of nestedness of a system of ecological communities. We illustrate our approach using avian data from the North American Breeding Bird Survey collected in Florida Keys.

Florida

On the estimation of species richness based on the accumulation of previously unrecorded species

Estimation of species richness of local communities has become an important topic in community ecology and monitoring. Investigators can seldom enumerate all the species present in the area of interest during sampling sessions. If the location of interest is sampled repeatedly within a short time period, the number of new species recorded is typically largest in the initial sample and decreases as sampling proceeds, but new species may be detected if sampling sessions are added. The question is how to estimate the total number of species. The data collected by sampling the area of interest repeatedly can be used to build species-accumulation curves: the cumulative number of species recorded as a function of the number of sampling sessions (which we refer to as ?species-accumulation data?). A classic approach used to compute total species richness is to fit curves to the data on species accumulation with sampling effort. This approach does not rest on direct estimation of the probability of detecting species during sampling sessions and has no underlying basis regarding the sampling process that gave rise to the data. Here we recommend a probabilistic, nonparametric estimator for species richness for use with species-accumulation data. We use estimators of population size that were developed for capture-recapture data, but that can be used to estimate the size of species assemblages using species-accumulation data. Models of detection probability account for the underlying sampling process. They permit variation in detection probability among species. We illustrate this approach using data from the North American Breeding Bird Survey (BBS). We describe other situations where species accumulation data are collected under different designs (e.g., over longer periods of time, or over spatial replicates) and that lend themselves to use of capture-recapture models for estimating the size of the community of interest. We discuss the assumptions and interpretations corresponding to each situation.

Alabama, Louisiana, Maine, Maryland, New Hampshire

Can non-breeding be a cost of breeding dispersal?

Breeding habitat selection and dispersal are crucial processes that affect many components of fitness. Breeding dispersal entails costs, one of which has been neglected: dispersing animals may miss breeding opportunities because breeding dispersal requires finding a new nesting site and mate, two time- and energy-consuming activities. Dispersers are expected to be prone to non-breeding. We used the kittiwake (Rissa tridactyla) to test whether breeding dispersal influences breeding probability. Breeding probability was associated with dispersal, in that both were negatively influenced by private information (previous individual reproductive success) and public information (average reproductive success of conspecifics) about patch quality. Furthermore, the probability of skipping breeding was 1.7 times higher in birds that settled in a new patch relative to those that remained on the same patch. Finally, non-breeders that resumed breeding were 4.4 times more likely to disperse than birds that bred in successive years. Although private information may influence breeding probability directly, the link between breeding probability and public information may be indirect, through the influence of public information on breeding dispersal, non-breeding thus being a cost of dispersal. These results support the hypothesis that dispersal may result in not being able to breed. More generally, non-breeding (which can be interpreted as an extreme form of breeding failure) may reveal costs of various previous activities. Because monitoring the non-breeding portion of a population is difficult, non-breeders have been neglected in many studies of reproduction trade-offs.

Behavioral Ecology and Sociobiology

Individual covariation in life-history traits: Seeing the trees despite the forest

We investigated the influence of age on survival and breeding rates in a long-lived species Rissa tridactyla using models with individual random effects permitting variation and covariation in fitness components among individuals. Differences in survival or breeding probabilities among individuals are substantial, and there was positive covariation between survival and breeding probability; birds that were more likely to survive were also more likely to breed, given that they survived. The pattern of age-related variation in these rates detected at the individual level differed from that observed at the population level. Our results provided confirmation of what has been suggested by other investigators: within-cohort phenotypic selection can mask senescence. Although this phenomenon has been extensively studied in humans and captive animals, conclusive evidence of the discrepancy between population-level and individual-level patterns of age-related variation in life-history traits is extremely rare in wild animal populations. Evolutionary studies of the influence of age on life-history traits should use approaches differentiating population level from the genuine influence of age: only the latter is relevant to theories of life-history evolution. The development of models permitting access to individual variation in fitness is a promising advance for the study of senescence and evolutionary processes.

Brittany

Disentangling sampling and ecological explanations underlying species-area relationships

We used a probabilistic approach to address the influence of sampling artifacts on the form of species-area relationships (SARs). We developed a model in which the increase in observed species richness is a function of sampling effort exclusively. We assumed that effort depends on area sampled, and we generated species-area curves under that model. These curves can be realistic looking. We then generated SARs from avian data, comparing SARs based on counts with those based on richness estimates. We used an approach to estimation of species richness that accounts for species detection probability and, hence, for variation in sampling effort. The slopes of SARs based on counts are steeper than those of curves based on estimates of richness, indicating that the former partly reflect failure to account for species detection probability. SARs based on estimates reflect ecological processes exclusively, not sampling processes. This approach permits investigation of ecologically relevant hypotheses. The slope of SARs is not influenced by the slope of the relationship between habitat diversity and area. In situations in which not all of the species are detected during sampling sessions, approaches to estimation of species richness integrating species detection probability should be used to investigate the rate of increase in species richness with area.

Maryland

Test for age-specificity in survival of the common tern

Much effort in life-history theory has been addressed to the dependence of life-history traits on age, especially the phenomenon of senescence and its evolution. Although senescent declines in survival are well documented in humans and in domestic and laboratory animals, evidence for their occurrence and importance in wild animal species remains limited and equivocal. Several recent papers have suggested that methodological issues may contribute to this problem, and have encouraged investigators to improve sampling designs and to analyse their data using recently developed approaches to modelling of capture-mark-recapture data. Here we report on a three-year, two-site, mark-recapture study of known-aged common terns (Sterna hirundo) in the north-eastern USA. The study was nested within a long-term ecological study in which large numbers of chicks had been banded in each year for > 25 years. We used a range of models to test the hypothesis of an influence of age on survival probability. We also tested for a possible influence of sex on survival. The cross-sectional design of the study (one year's parameter estimates) avoided the possible confounding of effects of age and time. The study was conducted at a time when one of the study sites was being colonized and numbers were increasing rapidly. We detected two-way movements between the sites and estimated movement probabilities in the year for which they could be modelled. We also obtained limited data on emigration from our study area to more distant sites. We found no evidence that survival depended on either sex or age, except that survival was lower among the youngest birds (ages 2-3 years). Despite the large number of birds included in the study (1599 known-aged birds, 2367 total), confidence limits on estimates of survival probability were wide, especially for the oldest age-classes, so that a slight decline in survival late in life could not have been detected. In addition, the cross-sectional design of this study meant that a decline in survival probability within individuals (actuarial senescence) could have been masked by heterogeneity in survival probability among individuals (mortality selection). This emphasizes the need for the development of modelling tools permitting separation of these two phenomena, valid under field conditions in which the recapture probabilities are less than one.

Book chapter

Influence of behavioural tactics on recruitment and reproductive trajectory in the kittiwake

Many studies have provided evidence that, in birds, inexperienced breeders have a lower probability of breeding successfully. This is often explained by lack of skills and knowledge, and sometimes late laying dates in the first breeding attempt. There is growing evidence that in many species with deferred reproduction, some prebreeders attend breeding places, acquire territories and form pairs. Several behavioural tactics assumed to be associated with territory acquisition have been described in different species. These tactics may influence the probability of recruiting in the breeding segment of the population, age of first breeding, and reproductive success in the first breeding attempt. Here we addressed the influence of behaviour ('squatting') during the prebreeding period on demographic parameters (survival and recruitment probability) in a long-lived colonial seabird species: the kittiwake. We also investigated the influence of behaviour on reproductive trajectory. Squatters have a higher survival and recruitment probability, and a higher probability of breeding successfully in the first breeding attempt in all age-classes where this category is represented. The influence of behaviour is mainly expressed in the first reproduction. However, there is a relationship between breeding success in the first occasion and subsequent occasions. The influence of breeding success in the first breeding attempt on the rest of the trajectory may indirectly reflect the influence of behaviour on breeding success in the first occasion. The shape of the reproductive trajectory is influenced by behaviour and age of first breeding. There is substantial individual variation from the mean reproductive trajectory, which is accounted for by heterogeneity in performance among individuals in the first attempt, but there is no evidence of individual heterogeneity in the rate of change over time in performance in subsequent breeding occasions

Book chapter

Model-based estimation of individual fitness

Fitness is the currency of natural selection, a measure of the propagation rate of genotypes into future generations. Its various definitions have the common feature that they are functions of survival and fertility rates. At the individual level, the operative level for natural selection, these rates must be understood as latent features, genetically determined propensities existing at birth. This conception of rates requires that individual fitness be defined and estimated by consideration of the individual in a modelled relation to a group of similar individuals; the only alternative is to consider a sample of size one, unless a clone of identical individuals is available. We present hierarchical models describing individual heterogeneity in survival and fertility rates and allowing for associations between these rates at the individual level. We apply these models to an analysis of life histories of Kittiwakes (Rissa tridactyla ) observed at several colonies on the Brittany coast of France. We compare Bayesian estimation of the population distribution of individual fitness with estimation based on treating individual life histories in isolation, as samples of size one (e.g. McGraw & Caswell, 1996).

Book chapter

Relative species richness and community completeness: avian communities and urbanization in the mid-Atlantic states

The idea that local factors govern local richness has been dominant for years, but recent theoretical and empirical studies have stressed the influence of regional factors on local richness. Fewer species at a site could reflect not only the influence of local factors, but also a smaller regional pool. The possible dependency of local richness on the regional pool should be taken into account when addressing the influence of local factors on local richness. It is possible to account for this potential dependency by comparing relative species richness among sites, rather than species richness per se. We consider estimation of a metric permitting assessment of relative species richness in a typical situation in which not all species are detected during sampling sessions. In this situation, estimates of absolute or relative species richness need to account for variation in species detection probability if they are to be unbiased. We present a method to estimate relative species richness based on capture-recapture models. This approach involves definition of a species list from regional data, and estimation of the number of species in that list that are present at a site-year of interest. We use this approach to address the influence of urbanization on relative richness of avian communities in the Mid-Atlantic region of the United States. There is a negative relationship between relative richness and landscape variables describing the level of urban development. We believe that this metric should prove very useful for conservation and management purposes because it is based on an estimator of species richness that both accounts for potential variation in species detection probability and allows flexibility in the specification of a 'reference community.' This metric can be used to assess ecological integrity, the richness of the community of interest relative to that of the 'original' community, or to assess change since some previous time in a community.

Delaware, Maryland, New Jersey, New York, Pennsylv