Effects of population characteristics and structure on estimates of effective population size in a house sparrow metapopulation

Effective population size (N_e) is a key parameter to understand evolutionary processes and the viability of endangered populations as it determines the rate of genetic drift and inbreeding. Low N_e can lead to inbreeding depression and reduced population adaptability. In this study, we estimated contemporary N_e using genetic estimators (LDNE, ONeSAMP, MLNE and CoNe) as well as a demographic estimator in a natural insular house sparrow metapopulation. We investigated whether population characteristics (population size, sex ratio, immigration rate, variance in population size and population growth rate) explained variation within and among populations in the ratio of effective to census population size (N_e/N_c). In general, N_e/N_c ratios increased with immigration rates. Genetic N_e was much larger than demographic N_e, probably due to a greater effect of immigration on genetic than demographic processes in local populations. Moreover, although estimates of genetic N_e seemed to track N_c quite well, the genetic N_e‐estimates were often larger than N_c within populations. Estimates of genetic N_e for the metapopulation were however within the expected range (<N_c). Our results suggest that in fragmented populations, even low levels of gene flow may have important consequences for the interpretation of genetic estimates of N_e. Consequently, further studies are needed to understand how N_e estimated in local populations or the total metapopulation relates to actual rates of genetic drift and inbreeding.     

From individual behavior to metapopulation dynamics: unifying the patchy population and classic metapopulation models

Spatially structured populations in patchy habitats show much variation in migration rate, from patchy populations in which individuals move repeatedly among habitat patches to classic metapopulations with infrequent migration among discrete populations. To establish a common framework for population dynamics in patchy habitats, we describe an individual-based model (IBM) involving a diffusion approximation of correlated random walk of individual movements. As an example, we apply the model to the Glanville fritillary butterfly (Melitaea cinxia) inhabiting a highly fragmented landscape. We derive stochastic patch occupancy model (SPOM) approximations for the IBMs assuming pure demographic stochasticity, uncorrelated environmental stochasticity, or completely correlated environmental stochasticity in local dynamics. Using realistic parameter values for the Glanville fritillary, we show that the SPOMs mimic the behavior of the IBMs well. The SPOMs derived from IBMs have parameters that relate directly to the life history and behavior of individuals, which is an advantage for model interpretation and parameter estimation. The modeling approach that we describe here provides a unified framework for patchy populations with much movements among habitat patches and classic metapopulations with infrequent movements.     

Effects of population subdivision and catastrophes on the persistence of a land snail metapopulation

We modeled the dynamics of a metapopulation of the land snail Arianta arbustorum in north-eastern Switzerland to investigate the effect of population subdivision on the persistence of a land snail metapopulation and to analyze the interaction between spatial factors, population subdivision, and catastrophes. We developed a spatially structured, stochastic, age-structured metapopulation model with field data from previous studies on the metapopulation in Switzerland and experimental and meteorological data. The model incorporated distance-dependent dispersal through stream banks, correlated environmental fluctuations, and catastrophes resulting from heavy rains. The results point to various complex interactions among factors involved in metapopulation dynamics and suggest that in some cases population subdivision may act to decrease threats from environmental fluctuations and catastrophes.     

The genetic effective size of a metapopulation

The structure of a population over time, space and categories of social and sexual role governs its ability to retain genetic variation in the face of drift. A metapopulation is an extreme form of spatial structure in which loosely coupled local populations 'turnover', that is, suffer extinction followed by recolonization from elsewhere within the metapopulation. These local populations turn over with a characteristic half-life. Based on a simulation model that incorporates both realistic features of population ecology and population genetics, the ability of such a metapopulation to retain genetic variation, which may be defined as proportional to its so-called effective population size, denoted N_e(^meta), can be one to two orders of magnitude lower than the maximum total number of individuals in the system. N_e(meta) depends on the persistence time associated with longevity of local populations (the turnover half-life), the average number of local populations extant in the metapopulation and the gene flow between local populations. Habitat fragmentation, which can create a metapopulation from a formerly continuously distributed species, may have unappreciated large genetic consequences for species impacted by human development.     

Inferring population and metapopulation dynamics of Liparis loeselii from single-census and inventory data

To conserve endangered species, information is needed on (meta)population responses to habitat quality and management. As possibilities for long-term studies are generally limited, it is important to obtain as much information as possible in a single field season.We obtained such single-census data for the orchid Liparis loeselii, a European Habitat Directive species. Stage structures of 15 Dutch dune and fen populations were related to vegetation structure, environmental indicators, and management. Botanical inventory records from 1930 to 2003 were used to infer population life spans.Cluster analysis did not reveal successional stage structure types. Dense populations with high recruitment mainly occurred in open, young-successional vegetation with high soil pH. High soil humidity and acidification negatively affected orchid densities. Early mowing was preferable over late mowing in dune slacks, because the latter reduced juvenile densities. The predominant population life span was three to eight years, and similar for dune slacks and fens. Longer life spans were occasionally observed at mown sites with influx of base-rich water.Our results suggest high metapopulation dynamics. Long-term metapopulation viability requires the formation of new habitat by dune slack formation in dunes and peat removal in fens. Population persistence can be prolonged to some extent by mowing, extensive grazing, or sod removal if natural habitat formation is impossible. Our study demonstrates that useful information on (meta)population ecology and viability can be obtained in a single field season.     

Climate, social factors and research disturbance influence population dynamics in a declining sociable weaver metapopulation

Population trends are determined by gains through reproduction and immigration, and losses through mortality and emigration. These demographic quantities and resulting population dynamics are affected by different external and internal drivers. We examined how these demographic quantities were affected by weather, research-induced disturbance, local density, colony site and year in a metapopulation of 17 sociable weaver (Philetairus socius) colonies over 17 years of study (4 years for reproduction). Most colonies declined, but at different rates. The four demographic quantities were related to different drivers. Survival strongly varied among years and colonies and was positively related to rainfall and negatively related to extreme temperature (together explaining 30 % of variation) and disturbance (measured as number of captures conducted at a colony; 7 %). There was a trend for a positive relationship between reproduction and rainfall (50 %). Movement was mainly related to local density: individuals were more likely to emigrate from small to large colonies and from colonies that were either well below or above their long-term mean. They were more likely to immigrate into colonies that were nearby, and below their mean size. We then quantified the effects of these relationships on metapopulation dynamics using a multi-site matrix projection model. Rainfall was potentially a strong driver of metapopulation dynamics. In addition, field-work disturbance might have contributed to the decline of this metapopulation but could not explain its full magnitude. Hence, through a combination of analytical methods we were able to obtain information on the main drivers affecting dynamics in a declining metapopulation.     

复合种群动态研究进展

复合种群动态研究是当今保护生物学和生态学中最热门的课题之一。在此较为详细地介绍了复合种群动态的有关概念和术语、经验研究和理论探索的最新进展以及相关的预测模型 ,对我国开展相关研究有参考价值。     

Spatial dynamics in a metapopulation network: recovery of a rare grasshopper Stauvoderus scalaris from population refuges

A characteristic feature of the spatial distribution of many species is patchiness. This spatial patchiness may be generated by very different processes, e.g. fragmentation, succession and extinction-colonisation dynamics. In this study, we apply a spatial realistic metapopulation model to analyse the occupancy pattern of a rare and endangered grasshopper, Stauroderus scalaris. found in an extensive network of 158 patches. When the study was initiated in 1985 the regional occupancy was 9.3% declining down to 7.1% in 1989. Then there was a spatial expansion of the population and in 1993 as many as 27.3% of the patches were occupied and 32.9% in 1995. During this expansion phase, the dynamics obeyed metapopulation principles: large patches and less isolated ones were more likely to be colonised. In the beginning, local extinction risks were negatively related to patch size and positively influenced by isolation. However, later on neither area nor isolation affected extinction probabilities. Altogether, 20 extinctions and 56 colonisations were observed. The shift in regional occupancy, with a growth of ca 20%, coincides with perturbations to the patch network and the warmest summer in 140 yr. Our results suggest that S. scalaris persists on a dynamic habitat mosaic, where refuges are crucial during adverse periods, and stochastic environmental factors (disturbances and climate), that are correlated over large areas, are generating population dynamic patterns that are hard to predict using current modelling techniques.     

Aggregate statistical measures and metapopulation dynamics

There are two main types of metapopulation models. Spatially implicit models are analytically tractable but neglect spatial heterogeneities. Spatially explicit models are more realistic but too complex. In this paper, I build a bridge between both approximations. I derive a new metapopulation model using a well-known technique in population genetics. Spatial heterogeneities are captured by an aggregate statistical measure of spatial correlation. When this correlation is zero, i.e., space is homogeneous, the model becomes the well-known Levins' model. As spatial correlation increases, equilibrium patch occupancy decreases from what would be expected under the spatially homogeneous assumption. I proceed by testing how well spatial complexities from a spatially explicit simulation can be encapsulated by such an aggregate statistical measure.     

Structured models of metapopulation dynamics

I develop models of metapopulation dynamics that describe changes in the numbers of individuals within patches. These models are analogous to structured population models, with patches playing the role of individuals. Single species models which do not include the effect of immigration on local population dynamics of occupied patches typically lead to a unique equilibrium. The models can be used to study the distributions of numbers of individuals among patches, showing that both metapopulations with local outbreaks and metapopulations without outbreaks can occur in systems with no underlying environmental variability. Distributions of local population sizes (in occupied patches) can vary independently of the total population size, so both patterns of distributions of local population sizes are compatible with either rare or common species. Models which include the effect of immigration on local population dynamics can lead to two positive equilibria, one stable and one unstable, the latter representing a threshold between regional extinction and persistence.     

Effects of demographic parameters on metapopulation size and persistence: an analytical stochastic model

An analytical stochastic metapopulation model is developed. It describes how individuals will be distributed among patches as a function of density-dependent birth, death and emigration rates, and the probability of successful dispersal. The model includes demographic stochasticity, but not catastrophes, environmental stochasticity or variation in patch size and suitability. All patches are equally likely to be colonized by migrants. The model predicts: (a) mean and variance of the number of individuals per patch; (b) probability distribution of individuals per patch; (c) mean number of individuals in transit; and (d) turn-over rate and expected persistence time of a single patch. The model shows that (a) dispersal rates must be intermediate in order to ensure metapopulation persistence; (b) the mean number of individuals per patch is often well below the carrying capacity; (c) long transit times and/or high mortality during dispersal reduce the mean number of individuals per patch; (d) density-dependent emigration responses will usually increase metapopulation size and persistence compared with density-independent dispersal; (e) an increase in the per capita net growth rate can both increase and decrease metapopulation size and persistence depending on whether dispersal rates are high or low; (f) density-independent birth, death, and emigration rates lead to a spatial pattern described by the negative binomial distribution.     

Optimal patch use and metapopulation dynamics

Summary We compared the metapopulation dynamics of predator—prey systems with (1) adaptive global dispersal, (2) adaptive local dispersal, (3) fixed global dispersal and (4) fixed local dispersal by predators. Adaptive dispersal was modelled using the marginal value theorem, such that predators departed patches when the instantaneous rate of prey capture was less than the long-term rate of prey capture averaged over all patches, scaled to the movement time between patches. Adaptive dispersal tended to stabilize metapopulation dynamics in a similar manner to conventional fixed dispersal models, but the temporal dynamics of adaptive dispersal models were more unpredictable than the smooth oscillations of fixed dispersal models. Moreover, fixed and adaptive dispersal models responded differently to spatial variation in patch productivity and the degree of compartmentalization of the system. For both adaptive dispersal and fixed dispersal models, localized (stepping-stone) dispersal was more strongly stabilizing than global (island) dispersal. Variation among predators in the probability of dispersal in relation to local prey density had a strong stabilizing influence on both within-patch and metapopulation dynamics. These results suggest that adaptive space use strategies by predators could have important implications for the dynamics of spatially heterogeneous trophic systems.     

Measles metapopulation dynamics: a gravity model for epidemiological coupling and dynamics

Infectious diseases provide a particularly clear illustration of the spatiotemporal underpinnings of consumer-resource dynamics. The paradigm is provided by extremely contagious, acute, immunizing childhood infections. Partially synchronized, unstable oscillations are punctuated by local extinctions. This, in turn, can result in spatial differentiation in the timing of epidemics and, depending on the nature of spatial contagion, may result in traveling waves. Measles epidemics are one of a few systems documented well enough to reveal all of these properties and how they are affected by spatiotemporal variations in population structure and demography. On the basis of a gravity coupling model and a time series susceptible-infected-recovered (TSIR) model for local dynamics, we propose a metapopulation model for regional measles dynamics. The model can capture all the major spatiotemporal properties in prevaccination epidemics of measles in England and Wales.     

Egg size evolution and energetic constraints on population dynamics.

We use population models that are based on dynamic energy budget models for individuals in order to study the evolution of offspring size and its relationship to the evolution of population dynamics. We show the existence of alternative evolutionarily stable strategies for offspring investment strategy resulting from a trade off between offspring number and time-to-maturity. The model predicts egg energy in Daphnia magna well, and suggests that the observed egg energy in D. magna is the result of selection for minimal egg investment constrained by minimum viable egg energy, combined with selection for a juvenile energy reserve. The selection for minimal egg size pushes populations toward chaotic dynamics. However, the minimum viable egg size combined with low efficiency of conversion of energy to new biomass is sufficient to keep population dynamics out of chaos.     

Potential metapopulation structure and the effects of habitat quality on population size of the endangered False Ringlet butterfly

The False Ringlet (Coenonympha oedippus) is a European butterfly species, endangered due to the severe loss and fragmentation of its habitat. In Hungary, two remaining populations of the butterfly occur in lowland Purple Moorgrass meadows. We studied a metapopulation occupying twelve habitat patches in Central Hungary. Our aim was to reveal what measures of habitat quality affect population size and density of this metapopulation, estimate dispersal parameters and describe phenology of subpopulations. Local population sizes and dispersal parameters were estimated from an extensive mark–release–recapture dataset, while habitat quality was characterized by groundwater level, cover of grass tussocks, bush cover, height of vegetation and grass litter at each habitat patch. The estimated size of the metapopulation was more than 3,000 individuals. We estimated a low dispersal capacity, especially for females, indicating a very low probability of (re)colonization. Butterfly abundance and density in local populations increased with higher grass litter, lower groundwater level and larger area covered by tussocks. We suppose that these environmental factors affect butterfly abundance by determining the microclimatic conditions for both larvae and adult butterflies. Our results suggest that the long-term preservation of the studied metapopulation needs the maintenance of high quality habitat patches by appropriate mowing regime and water regulation. Management also should facilitate dispersal to strengthen metapopulation structure with creating stepping-stones or gradually increase habitat quality in present matrix.     

Long-term dynamics in a metapopulation of the American pika

A 20-yr study of a metapopulation of the American pika revealed a regional decline in occupancy in one part of a large network of habitat patches. We analyze the possible causes of this decline using a spatially realistic metapopulation model, the incidence function model. The pika metapopulation is the best-known mammalian example of a classical metapopulation with significant population turnover, and it satisfies closely the assumptions of the incidence function model, which was parameterized with data on patch occupancy. The model-predicted incidences of patch occupancy are consistent with observed incidences, and the model predicts well the observed turnover rate between four metapopulation censuses. According to model predictions, the part of the metapopulation where the decline has been observed is relatively unstable and prone to large oscillations in patch occupancy, whereas the other part of the metapopulation is predicted to be persistent. These results demonstrate how extinction-colonization dynamics may produce spatially correlated patterns of patch occupancy without any spatially correlated processes in local dynamics or extinction rate. The unstable part of the metapopulation gives an empirical example of multiple quasi equilibria in metapopulation dynamics. Phenomena similar to those observed here may cause fluctuations in species' range limits.     

The effect of conspecific attraction on metapopulation dynamics

Random dispersal direction is assumed in all current metapopulation models. This assumption is called into question by recent experiments demonstrating that some species disperse preferentially to sites occupied by conspecifies. We incorporate conspecific attraction into two metapopulation models which differ in type of dispersal, the Levins model and a two-dimensional stepping-stone model. In both models, conspecific attraction lowers the proportion of occupied habitat patches within a metapopulation at equilibrium.