Viability analyses with habitat-based metapopulation models

Population viability analysis (PVA) models incorporate spatial dynamics in different ways. At one extreme are the occupancy models that are based on the number of occupied populations. The simplest occupancy models ignore the location of populations. At the other extreme are individual-based models, which describe the spatial structure with the location of each individual in the population, or the location of territories or home ranges. In between these are spatially structured metapopulation models that describe the dynamics of each population with structured demographic models and incorporate spatial dynamics by modeling dispersal and temporal correlation among populations. Both dispersal and correlation between each pair of populations depend on the location of the populations, making these models spatially structured. In this article, I describe a method that expands spatially structured metapopulation models by incorporating information about habitat relationships of the species and the characteristics of the landscape in which the metapopulation exists. This method uses a habitat suitability map to determine the spatial structure of the metapopulation, including the number, size, and location of habitat patches in which subpopulations of the metapopulation live. The habitat suitability map can be calculated in a number of different ways, including statistical analyses (such as logistic regression) that find the relationship between the occurrence (or, density) of the species and independent variables which describe its habitat requirements. The habitat suitability map is then used to calculate the spatial structure of the metapopulation, based on species-specific characteristics such as the home range size, dispersal distance, and minimum habitat suitability for reproduction. Received: April 1, 1999 / Accepted: October 29, 1999     

Metapopulation dynamics of the bog fritillary butterfly: modelling the effect of habitat fragmentation

Population viability analysis (PVA) and metapopulation theory are valuable tools to model the dynamics of spatially structured populations. In this article we used a spatially realistic population dynamic model to simulate the trajectory of a Proclossiana eunomia metapopulation in a network of habitat patches located in the Belgian Ardenne. Sensitivity analysis was used to evaluate the relative influence of the different parameters on the model output. We simulated habitat loss by removing a percentage of the original habitat, proportionally in each habitat patch. Additionally, we evaluated isolation and fragmentation effects by removing and dividing habitat patches from the network, respectively. The model predicted a slow decline of the metapopulation size and occupancy. Extinction risks predicted by the model were highly sensitive to environmental stochasticity and carrying capacity. For a determined level of habitat destruction, the expected lifetime of the metapopulation was highly dependent on the spatial configuration of the landscape. Moreover, when the proportion of removed habitat is above 40% of the original habitat, the loss of whole patches invariably leads to the strongest reduction in metapopulation viability.     

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.     

Matrix Matters:?Differences of Grand Skink Metapopulation Parameters in Native Tussock Grasslands and Exotic Pasture Grasslands

Modelling metapopulation dynamics is a potentially very powerful tool for conservation biologists. In recent years, scientists have broadened the range of variables incorporated into metapopulation modelling from using almost exclusively habitat patch size and isolation, to the inclusion of attributes of the matrix and habitat patch quality. We investigated the influence of habitat patch and matrix characteristics on the metapopulation parameters of a highly endangered lizard species, the New Zealand endemic grand skink (Oligosoma grande) taking into account incomplete detectability. The predictive ability of the developed zxmetapopulation model was assessed through cross-validation of the data and with an independent data-set. Grand skinks occur on scattered rock-outcrops surrounded by indigenous tussock (bunch) and pasture grasslands therefore implying a metapopulation structure. We found that the type of matrix surrounding the habitat patch was equally as important as the size of habitat patch for estimating occupancy, colonisation and extinction probabilities. Additionally, the type of matrix was more important than the physical distance between habitat patches for colonisation probabilities. Detection probability differed between habitat patches in the two matrix types and between habitat patches with different attributes such as habitat patch composition and abundance of vegetation on the outcrop. The developed metapopulation models can now be used for management decisions on area protection, monitoring, and the selection of translocation sites for the grand skink. Our study showed that it is important to incorporate not only habitat patch size and distance between habitat patches, but also those matrix type and habitat patch attributes which are vital in the ecology of the target species.     

种群生存力分析:准确性和保护应用

目前已提出了五类估计濒危物种绝灭风险的种群生存力分析模型 ,即 :分析模型、单种群确定性模型、单种群随机模型、异质种群模型和显空间模型。模型的选择取决于物种的生活史特征和可用的数据。与用于保护实践的其他方法相比 ,种群生存力分析 (PVA)是相对准确的量化工具。然而 ,一些濒危物种种群统计学数据质量差和种群动态的有关假说模糊不清可能影响到模型预测的准确性 ,因此 ,要谨慎地使用PVA。在西方国家 ,PVA在濒危物种保护计划和管理中应用越来越广泛。它主要用于 :( 1)预测濒危物种未来的种群大小 ;( 2 )估计一定时间内物种的绝灭风险 ;( 3 )评估一套保护措施 ,确定哪个能使种群的存活时间最长 ;( 4)探索不同假说对小种群动态的影响 ;( 5 )指导濒危动物野外数据的搜集工作。我国的濒危物种很多 ,然而开展PVA研究的濒危物种却很少。应大力发展适合于模拟我国特有濒危物种及其保护问题的PVA模型     

Do invasive species undergo metapopulation dynamics? A case study of the invasive Caspian gull,Larus cachinnans,in Poland

Aim The mechanisms of initial dispersal and habitat occupancy by invasive alien species are fundamental ecological problems. Most tests of metapopulation theory are performed on local population systems that are stable or in decline. In the current study we were interested in the usefulness of metapopulation theory to study patch occupancy, local colonization, extinction and the abundance of the invasive Caspian gull (Larus cachinnans) in its initial invasion stages. Location Waterbodies in Poland. Methods Characteristics of the habitat patches (waterbodies, 35 in total) occupied by breeding pairs of Caspian gulls and an equal sample of randomly selected unoccupied patches were compared with t‐tests. Based on presence–absence data from 1989 to 2006 we analysed factors affecting the probability of local colonization, extinction and the size of local populations using generalized linear models. Results Occupied habitat patches were significantly larger and less isolated (from other habitat patches and other local populations) and were located closer to rivers than empty patches. The proximity of local food resources (fish ponds, refuse dumps) positively affected the occurrence of breeding pairs. The probability of colonization was positively affected by patch area, and negatively by distances to fish ponds, nearest habitat patch, nearest breeding colony and to a river, and by higher forest cover around the patch boundaries. The probability of extinction was lower in patches with a higher number of breeding pairs and with a greater area of islets. The extinction probability increased with distances to other local populations, other habitat patches, fish ponds and to refuse dumps and with a higher cover of forest around the patch boundaries. The size of the local population decreased with distances to the nearest habitat patch, local population, river, fish pond and refuse dump. Local abundance was also positively affected by the area of islets in the patch. Main conclusions During the initial stages of the invasion of Caspian gulls in Poland the species underwent metapopulation‐like dynamics with frequent extinctions from colonized habitat patches. The results prove that metapopulation theory may be a useful conceptual framework for predicting which habitats are more vulnerable to invasion.     

Ecologically Differentiated Rules of Thumb for Habitat Network Design – Lessons from a Formula

The present paper addresses the following typical question of metapopulation management: “What habitat distribution is optimal for metapopulation persistence if a habitat network with a given number and configuration of patches is considered?”. By utilizing a generic metapopulation model and performing a special model analysis, rules of thumb for optimum habitat distribution and a general ‘Principle of Optimality’ are derived. The whole study is based on the application of a formula for the mean lifetime of metapopulations derived in a previous study. Finally, some general conclusions are drawn concerning the potential of using PVA techniques for deriving tools for decision support for conservation management.     

Metapopulation dynamics for spatially extended predator–prey interactions

Traditional metapopulation theory classifies a metapopulation as a spatially homogeneous population that persists on neighboring habitat patches. The fate of each population on a habitat patch is a function of a balance between births and deaths via establishment of new populations through migration to neighboring patches. In this study, we expand upon traditional metapopulation models by incorporating spatial heterogeneity into a previously studied two-patch nonlinear ordinary differential equation metapopulation model, in which the growth of a general prey species is logistic and growth of a general predator species displays a Holling type II functional response. The model described in this work assumes that migration by generalist predator and prey populations between habitat patches occurs via a migratory corridor. Thus, persistence of species is a function of local population dynamics and migration between spatially heterogeneous habitat patches. Numerical results generated by our model demonstrate that population densities exhibit periodic plane-wave phenomena, which appear to be functions of differences in migration rates between generalist predator and prey populations. We compare results generated from our model to results generated by similar, but less ecologically realistic work, and to observed population dynamics in natural metapopulations.     

Dynamics in a butterfly–plant–ant system: influence of habitat characteristics on turnover rates of the endangered lycaenid Maculinea alcon

Abstract.  1.  Maculinea alcon , a myrmecophilic, stenotopic lycaenid butterfly is restricted to wet heathlands, bogs, and nutrient-poor hay meadows. Due to intensification of agriculture and decrease of extensively grazed meadows, many suitable habitats have disappeared and the remaining ones are highly fragmented and deteriorated.
2. Historical distribution data and a comparison with the present occupation of patches show the decline of this critically endangered butterfly in north-west Germany. Most of the populations in north-west Germany are small and often geographically isolated.
3. In summer 2002, two-thirds of 77 investigated potential patches were unoccupied as a result of unsuitable habitat structure and habitat fragmentation.
4. Several habitat parameters were highly significantly correlated with the presence of M. alcon , in particular the distribution pattern of the host plant Gentiana pneumonanthe. Furthermore, butterflies were absent from many patches with an incidence probability below 50% with respect to patch size and isolation.
5. In the nature reserve Lüneburger Heide, part of the study area, M. alcon populations have been observed since 1995. Typical turnover of local populations could be detected during these years. Extinctions and re-colonisations have stabilised the presence of this species in a metapopulation in this nature reserve.
6. These data show the importance of different factors on different spatial levels influencing the presence of this endangered butterfly.     

Population viability analysis of the butterfly Lopinga achine in a changing landscape in Sweden

Metapopulation theory has generally focused only on the stochastic turn-over rate among populations and assumed that the number and location of suitable habitat patches will remain constant through time. This study combines in a PVA both the deterministic landscape dynamics and the stochastic colonisations and extinctions of populations for the butterfly Lopinga achine in Sweden. With data on occupancy pattern and the rate of habitat change, we built a simulation model and examined five different scenarios with different assumptions of landscape changes for L. achine . If no landscape changes would be expected, around 80 populations are predicted to persist during the next 100 yr. Adding the knowledge that many of the sites are unmanaged and that the host plant will slowly deteriorate as canopies close over, and adding environmental variation and synchrony, showed that the number of populations will decrease to around of 4.3 and 2.8 respectively, with an extinction risk of 34% – quite different from the first scenario based only on the metapopulation model. This study has shown the importance of incorporating both deterministic and stochastic events when making a reliable population viability analysis. Even though one can not expect that the long-term predictions of either occupied patches or extinction risks will be accurate quantitatively, the qualitative implications are correct. The extinction risk will be high if grazing is not applied to more patches than is the case today. The simulations indicate that an absolute minimum of 10–30 top-ranked patches needs to be managed for the persistence of the metapopulation of L. achine in the long term. The same problem of abandoned and overgrowing habitats affects many other threatened species in the European landscape and a similar approach could also be applied to them.     

Dispersal-induced desynchronization: from metapopulations to metacommunities

The conceptualization of fragmented populations in terms of metapopulation theory has become standard over the last three decades. It is well known that increases in between‐patch migration rates cause more synchronous population fluctuations and that this coherence increases the risk of global metapopulation extinction. Because species’ migration rates and the probability of individuals surviving migration events depend on the effective distance between patches, the benefit of improving conservation corridors or the matrix between habitat patches has been questioned. As populations occur in the context of larger communities, moving from a metapopulation to a metacommunity model framework is a natural extension to address the generality of these conclusions. We show how considering a metacommunity can modify the conclusion that decreasing the effective distance between habitat patches (via improving matrix quality or other measures) necessarily increases the degree of metapopulation synchrony. We show that decreases in effective between‐patch distance may deter population synchrony because of the simultaneous effect this change has on the migration patterns of other species. These results indicate that species interactions need to be considered when the effect of conservation measures on population synchrony, and ultimately persistence, is addressed.     

Wild and zoo animal interactive management and habitat conservation

This review considers interactive management of wild and zoo populations as a stratesy to support habitat preservation, help sustain key endangered species, and hasten the evolution of zoos and aquaria as proactive conservation organizations. Interactive management supports key species' subpopulations in an integrated fashion, using their study in nature as a way to understand wildlife habitats, ecological processes and conservation threats. In the face of human increase and habitat destruction, the survival of much wildlife will depend upon the utility of fragments of habitat and the survival of relatively small populations whose habitats are reduced or altered and whose numbers are capped. Under such conditions, interactive wild-captive metapopulation strategies may increase the security of key species.     

Metapopulation models for seasonally migratory animals

Metapopulation models are widely used to study species that occupy patchily distributed habitat, but are rarely applied to migratory species, because of the difficulty of identifying demographically independent subpopulations. Here, we extend metapopulation theory to describe the directed seasonal movement of migratory populations between two sets of habitat patches, breeding and non-breeding, with potentially different colonization and extinction rates between patch types. By extending the classic metapopulation model, we show that migratory metapopulations will persist if the product of the two colonization rates exceeds the product of extinction rates. Further, we develop a spatially realistic migratory metapopulation model and derive a landscape metric-the migratory metapopulation capacity-that determines persistence. This new extension to metapopulation theory introduces an important tool for the management and conservation of migratory species and may also be applicable to model the dynamics of two host-parasite systems.     

Long-term persistence of species and the SLOSS problem

The single large or several small (SLOSS) problem has been addressed in a large number of empirical and theoretical studies, but no coherent conclusion has yet been reached. Here I study the SLOSS problem in the context of metapopulation dynamics. I assume that there is a fixed total amount A(0) of habitat available, and I derive formulas for the optimal number n and area A of habitat patches, where n=A(0)/A. I consider optimality in two ways. First, I attempt to maximize the time to metapopulation extinction, which is a relevant measure for metapopulation viability for rare and threatened species. Second, I attempt to maximize the metapopulation capacity of the habitat patch network, which corresponds both with maximizing the distance to the deterministic extinction threshold and with maximizing the fraction of occupied patches. I show that in the typical case, a small number of large patches maximizes the metapopulation capacity, while an intermediate number of habitat patches maximizes the time to extinction. The main conclusion stemming from the analysis is that the optimal number of patches is largely affected by the relationship between habitat patch area and rates of immigration, emigration and local extinction. Here this relationship is summarized by a single factor zeta, termed the patch area scaling factor.     

Metapopulation viability analysis of the bog fritillary butterfly using RAMAS/GIS

Many species living in man-shaped landscapes are restricted to small natural habitat patches and form metapopulations; predicting their future is a central issue in applied ecology. We examined the viability of the bog fritillary butterfly Proclossiana eunomia Esper, a specialist glacial relict species, in a highly fragmented landscape (<1% of suitable habitat in 10 km²), by way of population viability analysis. We used comprehensive data from a long-term study in which a patchy population was monitored during ten consecutive years to parameterise a spatially structured metapopulation model using commercially available platform RAMAS/GIS 3.0. Population growth rate was density-dependent and modulated by various climatic variables acting on different developmental stages of the butterfly. Density dependence was probably related to larval parasitism by a specific parasitoid. Population size was negatively affected by an increase in the mean temperature. Dispersal was modelled as the observed proportion of movements between patches, taking into account the probability of emigration out of a given patch. Our model provided results close to the picture of the system drawn from the field data and was considered as useful in making predictions about the metapopulation. Demographic parameters proved to have a far higher impact on metapopulation persistence than dispersal or correlation of local dynamics. Scenarios simulating both global warming and management of habitat patches by rustic herbivore grazing indicated a decrease in the viability of the metapopulation. Our results prompted the regional nature conservation agency to modify the planned management regime. We urge conservation biologists to use structured population models including local population dynamics for viability analysis targeted to such threatened metapopulations in highly fragmented landscapes.     

Toward ecologically scaled landscape indices

Nature conservation is increasingly based on a landscape approach rather than a species approach. Landscape planning that includes nature conservation goals requires integrated ecological tools. However, species differ widely in their response to landscape change. We propose a framework of ecologically scaled landscape indices that takes into account this variation. Our approach is based on a combination of field studies of spatially structured populations (metapopulations) and model simulations in artificial landscapes. From these, we seek generalities in the relationship among species features, landscape indices, and metapopulation viability. The concept of ecological species profiles is used to group species according to characteristics that are important in metapopulations' response to landscape change: individual area requirements as the dominant characteristic of extinction risk in landscape patches and dispersal distance as the main determinant of the ability to colonize patches. The ecological profiles and landscape indices are then integrated into two ecologically scaled landscape indices (ESLI): average patch carrying capacity and average patch connectivity. The field data show that the fraction of occupied habitat patches is correlated with the two ESLI. To put the ESLI into a perspective of metapopulation persistence, we determine the viability for six ecological profiles at different degrees of habitat fragmentation using a metapopulation model and computer-generated landscapes. The model results show that the fraction of occupied patches is a good indicator for metapopulation viability. We discuss how ecological profiles, ESLI, and the viability threshold can be applied for landscape planning and design in nature conservation.     

Colonization and extinction dynamics of an annual plant metapopulation in an urban environment

The metapopulation framework considers that the spatiotemporal distribution of organisms results from a balance between the colonization and extinction of populations in a suitable and discrete habitat network. Recent spatially realistic metapopulation models have allowed patch dynamics to be investigated in natural populations but such models have rarely been applied to plants. Using a simple urban fragmented population system in which favourable habitat can be easily mapped, we studied patch dynamics in the annual plant Crepis sancta (Asteraceae). Using stochastic patch occupancy models (SPOMs) and multi‐year occupancy data we dissected extinction and colonization patterns in our system. Overall, our data were consistent with two distinct metapopulation scenarios. A metapopulation (sensu stricto) dynamic in which colonization occurs over a short distance and extinction is lowered by nearby occupied patches (rescue effect) was found in a set of patches close to the city centre, while a propagule rain model in which colonization occurs from a large external population was most consistent with data from other networks. Overall, the study highlights the importance of external seed sources in urban patch dynamics. Our analysis emphasizes the fact that plant distributions are governed not only by habitat properties but also by the intrinsic properties of colonization and dispersal of species. The metapopulation approach provides a valuable tool for understanding how colonization and extinction shape occupancy patterns in highly fragmented plant populations. Finally, this study points to the potential utility of more complex plant metapopulation models than traditionally used for analysing ecological and evolutionary processes in natural metapopulations.     

Important population viability analysis parameters for giant pandas (Aliuropoda melanoleuca)

Population viability analysis(PVA) is a tool to evaluate the risk of extinction for endangered species and aid conservation decision-making.The quality of PVA output is dependent on parameters related to population dynamics and life-history;however,it has been difficult to collect this information for the giant panda(Aliuropoda melanoleuca),a rare and endangered mammal native to China,confined to some 30 fragmented habitat patches.Since giant pandas are long-lived,mature late,have lower reproductive rates,and show little sexual dimorphism,obtaining data to perform adequate PVA has been difficult.Here,we develop a parameter sensitivity index by modeling the dynamics of six giant panda populations in the Minshan Mountains,in order to determine the parameters most influential to giant panda populations.Our data shows that the giant panda populations are most sensitive to changes in four female parameters:initial breeding age,reproductive rate,mortality rate between age 0 and 1,and mortality rate of adults.The parameter sensitivity index strongly correlated with initial population size,as smaller populations were more sensitive to changes in these four variables.This model suggests that demographic parameters of females have more influence on the results of PVA,indicating that females may play a more important role in giant panda population dynamics than males.Consequently,reintroduction of female individuals to a small giant panda population should be a high priority for conservation efforts.Our findings form a technical basis for the coming program of giant panda reintroduction,and inform which parameters are crucial to successfully and feasibly monitoring wild giant panda populations.     

Metapopulation Dynamics of the Mistletoe and Its Host in Savanna Areas with Different Fire Occurrence

Mistletoes are aerial hemiparasitic plants which occupy patches of favorable habitat (host trees) surrounded by unfavorable habitat and may be possibly modeled as a metapopulation. A metapopulation is defined as a subdivided population that persists due to the balance between colonization and extinction in discrete habitat patches. Our aim was to evaluate the dynamics of the mistletoe Psittacanthus robustus and its host Vochysia thyrsoidea in three Brazilian savanna areas using a metapopulation approach. We also evaluated how the differences in terms of fire occurrence affected the dynamic of those populations (two areas burned during the study and one was fire protected). We monitored the populations at six-month intervals. P. robustus population structure and dynamics met the expected criteria for a metapopulation: i) the suitable habitats for the mistletoe occur in discrete patches; (ii) local populations went extinct during the study and (iii) colonization of previously non-occupied patches occurred. The ratio of occupied patches decreased in all areas with time. Local mistletoe populations went extinct due to two different causes: patch extinction in area with no fire and fire killing in the burned areas. In a burned area, the largest decrease of occupied patch ratios occurred due to a fire event that killed the parasites without, however, killing the host trees. The greatest mortality of V. thyrsoidea occurred in the area without fire. In this area, all the dead trees supported mistletoe individuals and no mortality was observed for parasite-free trees. Because P. robustus is a fire sensitive species and V. thyrsoidea is fire tolerant, P. robustus seems to increase host mortality, but its effect is lessened by periodic burning that reduces the parasite loads.     

Dispersal and the metapopulation paradigm in amphibian ecology and conservation: are all amphibian populations metapopulations?

Amphibians are frequently characterized as having limited dispersal abilities, strong site fidelity and spatially disjunct breeding habitat. As such, pond‐breeding species are often alleged to form metapopulations. Amphibian species worldwide appear to be suffering population level declines caused, at least in part, by the degradation and fragmentation of habitat and the intervening areas between habitat patches. If the simplification of amphibians occupying metapopulations is accurate, then a regionally based conservation strategy, informed by metapopulation theory, is a powerful tool to estimate the isolation and extinction risk of ponds or populations. However, to date no attempt to assess the class‐wide generalization of amphibian populations as metapopulations has been made. We reviewed the literature on amphibians as metapopulations (53 journal articles or theses) and amphibian dispersal (166 journal articles or theses for 53 anuran species and 37 salamander species) to evaluate whether the conditions for metapopulation structure had been tested, whether pond isolation was based only on the assumption of limited dispersal, and whether amphibian dispersal was uniformly limited. We found that in the majority of cases (74%) the assumptions of the metapopulation paradigm were not tested. Breeding patch isolation via limited dispersal and/or strong site fidelity was the most frequently implicated or tested metapopulation condition, however we found strong evidence that amphibian dispersal is not as uniformly limited as is often thought. The frequency distribution of maximum movements for anurans and salamanders was well described by an inverse power law. This relationship predicts that distances beneath 11–13 and 8–9 km, respectively, are in a range that they may receive one emigrating individual. Populations isolated by distances approaching this range are perhaps more likely to exhibit metapopulation structure than less isolated populations. Those studies that covered larger areas also tended to report longer maximum movement distances – a pattern with implications for the design of mark‐recapture studies. Caution should be exercised in the application of the metapopulation approach to amphibian population conservation. Some amphibian populations are structured as metapopulations – but not all.