Metapopulation dynamics: Does it help to have more of the same?

With the interest in conservation biology shifting towards processes from patterns, and to populations from communities, the theory of metapopulation dynamics is replacing the equilibrium theory of island biogeography as the population ecology paradigm in conservation biology. The simplest models of metapopulation dynamics make predictions about the effects of habitat fragmentation - size and isolation of habitat patches - on metapopulation persistence. The simple models may be enriched by considerations of the effects of demographic and environmental stochasticity on the size and extinction probability of local populations. Environmental stochasticity affects populations at two levels: it makes local extinctions more probable, and it also decreases metapopulation persistence time by increasing the correlation of extinction events across populations. Some controversy has arisen over the significance of correlated extinctions, and how they may affect the optimal subdivision of metapopulations to maximize their persistence time.     

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.     

Local and Regional Determinants of Colonisation-Extinction Dynamics of a Riparian Mainland-Island Root Vole Metapopulation

The role of local habitat geometry (habitat area and isolation) in predicting species distribution has become an increasingly more important issue, because habitat loss and fragmentation cause species range contraction and extinction. However, it has also become clear that other factors, in particular regional factors (environmental stochasticity and regional population dynamics), should be taken into account when predicting colonisation and extinction. In a live trapping study of a mainland-island metapopulation of the root vole (Microtus oeconomus) we found extensive occupancy dynamics across 15 riparian islands, but yet an overall balance between colonisation and extinction over 4 years. The 54 live trapping surveys conducted over 13 seasons revealed imperfect detection and proxies of population density had to be included in robust design, multi-season occupancy models to achieve unbiased rate estimates. Island colonisation probability was parsimoniously predicted by the multi-annual density fluctuations of the regional mainland population and local island habitat quality, while extinction probability was predicted by island population density and the level of the recent flooding events (the latter being the main regionalized disturbance regime in the study system). Island size and isolation had no additional predictive power and thus such local geometric habitat characteristics may be overrated as predictors of vole habitat occupancy relative to measures of local habitat quality. Our results suggest also that dynamic features of the larger region and/or the metapopulation as a whole, owing to spatially correlated environmental stochasticity and/or biotic interactions, may rule the colonisation – extinction dynamics of boreal vole metapopulations. Due to high capacities for dispersal and habitat tracking voles originating from large source populations can rapidly colonise remote and small high quality habitat patches and re-establish populations that have gone extinct due to demographic (small population size) and environmental stochasticity (e.g. extreme climate events).     

Extinction dynamics and the regional persistence of a tree frog metapopulation

The concept of a metapopulation acknowledges local extinctions as a natural part of the dynamics of a patchily distributed population. However, if extinctions are not balanced by recolonizations or if there is a high degree of spatial synchrony of local extinctions, this poses a threat to and will reduce the metapopulation persistence time. Here we show that, in a metapopulation network of 378 pond patches used by the tree frog (Hyla arborea), even though extinctions are frequent (mean extinction probability p(e) = 0.24) they pose no threat to the metapopulation as they are balanced by recolonizations (p(c) = 0.33). In any one year there was a pattern of large populations tending to persist while small populations became extinct. The total number of individuals belonging to populations that went extinct was small (< 5%) compared with those populations that persisted. A spatial autocorrelation analysis indicated no clustering of local extinctions. The tree frog metapopulation studied consisted of a set of larger, persistent populations mixed with smaller populations characterized by high turnover dynamics.     

Extinction risks in metapopulations of a beetle inhabiting hollow trees predicted from time series

Ancient and dead trees are declining habitats harbouring many threatened species. These habitats are naturally patchy, and inhabiting species might exhibit metapopulation dynamics at a small spatial scale. In this study, the demography and metapopulation dynamics was analysed for Osmoderma eremita , which is an endangered beetle species associated with tree hollows in Europe. Extinction risks of O. eremita populations were predicted using Monte Carlo simulations based on time series of population assessments. Predicted occurrence patterns were consistent with field observations from an area with many small stands in which the populations are believed to have been more or less isolated from each other during the last 150–200 yr. Population growth was found to be density dependent. Carrying capacity was proportional to the volume of wood mould (i.e. loose material of dead wood in the tree hollows), which varied widely between hollow trees. This generates large differences in local extinction risks between hollow trees. The predicted metapopulation extinction risk was much higher if the habitat dynamics (formation, gradual increase and deterioration of tree hollows) were taken into consideration than in predictions yielded by models in which the amount of wood mould was assumed to be constant over time. Thus, this system has features from both mainland-island metapopulations and habitat-tracking metapopulations, and is rather far from a classic metapopulation. For the long-term persistence of the species in hollow trees, the habitat dynamics seem to be more important than demographic processes. Since the formation and deterioration of suitable tree are partly stochastic processes, there is a considerable extinction risk for many O. eremita populations, because they mainly rely on only one or a few trees with large amounts of wood mould.     

Both population size and patch quality affect local extinctions and colonizations

Currently, the habitat of many species is fragmented, resulting in small local populations with individuals occasionally dispersing between the remaining habitat patches. In a solitary bee metapopulation, extinction probability was related to both local bee population sizes and pollen resources measured as host plant population size. Patch size, on the other hand, had no additional predictive power. The turnover rate of local bee populations in 63 habitat patches over 4 years was high, with 72 extinction events and 31 colonization events, but the pollen plant population was stable with no extinctions or colonizations. Both pollen resources and bee populations had strong and independent effects on extinction probability, but connectivity was not of importance. Colonizations occurred more frequently within larger host plant populations. For metapopulation survival of the bee, large pollen plant populations are essential, independent of current bee population size.     

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.     

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.     

生境破碎化对动物种群存活的影响

生境破碎是生物多样性下降的主要原因之一。通常以岛屿生物地理学、异质种群生物学和景观生态学的理论来解释不同空间尺度中生境破碎化的生态学效应。生境破碎化引起面积效应、隔离效应和边缘效应。这些效应通过影响动物种群的绝灭阈值、分布和多度、种间关系以及生态系统过程,最终影响动物种群的存活。野外研究表明,破碎化对动物的影响,因物种、生境类型和地理区域不同而有所变化,因此,预测物种在破碎生境中的存活比较困难。研究热点集中于：确定生境面积损失和生境斑块的空间格局对破碎景观中物种绝灭的相对影响,破碎景观中物种的适宜生境比例和绝灭阈值,异质种群动态以及生态系统的生态过程。随着3S技术的发展,生境破碎化模型趋于复杂,而发展有效的模型和验证模型将成为一项富有挑战性的任务。     

The applicability of metapopulation theory to large mammals

Metapopulation theory has become a common framework in conservation biology and it is sometimes suggested that a metapopulation approach should be used for management of large mammals. However, it has also been suggested that metapopulation theory would not be applicable to species with long generations compared to those with short ones. In this paper, we review how and on what empirical ground metapopulation terminology has been applied to insects, small mammals and large mammals. The review showed that the metapopulation term sometimes was used for population networks which only fulfilled the broadest possible definition of a metapopulation, i.e. they were subpopulations connected by migrating individuals. We argue that the metapopulation concept should be reserved for networks that also show some kind of metapopulation dynamics. Otherwise it applies to almost all populations and loses its substance. We found much empirical support for metapopulation dynamics in both insects and small mammals, but not in large mammals. A possible reason is the methods used to confirm the existence of metapopulation dynamics. For insects and small mammals, the common approach is to study population turnover through patch occupancy data. Such data is difficult to obtain for large mammals, since longer temporal scales need to be covered to record extinctions and colonizations. Still, many populations of large mammals are exposed to habitat fragmentation and the resulting subpopulations sometimes have high risks of extinction. If there is migration between the subpopulations, the metapopulation framework could provide valuable information on their population dynamics. We suggest that a metapopulation approach can be interesting for populations of large mammals, when there are discrete breeding subpopulations and when these subpopulations have different growth rates and demographic fates. Thus, a comparison of the subpopulations’ demographic fates, rather than subpopulation turnover, can be a feasible alternative for studies of metapopulation dynamics in large mammals.     

How much does an individual habitat fragment contribute to metapopulation dynamics and persistence?

We derive measures for assessing the value of an individual habitat fragment for the dynamics and persistence of a metapopulation living in a network of many fragments. We demonstrate that the most appropriate measure of fragment value depends on the question asked. Specifically, we analyse four alternative measures: the contribution of a fragment to the metapopulation capacity of the network, to the equilibrium metapopulation size, to the expected time to metapopulation extinction and the long-term contribution of a fragment to colonization events in the network. The latter measure is comparable to density-dependent measures in general matrix population theory, though some differences are introduced by the fact that "density dependence" is spatially localized in the metapopulation context. We show that the value of a fragment depends not only on the properties of the landscape but also on the properties of the species. Most importantly, variation in fragment values between the habitat fragments is greatest in the case of rare species that occur close to the extinction threshold, as these species are likely to be restricted to the most favorable parts of the landscape. We expect that the measures of habitat fragment value described and analysed here have applications in landscape ecology and in conservation biology.     

Is the population turnover of patchy‐distributed annuals determined by dormancy dynamics or dispersal processes?

In species with fragmented distribution, regional turnover dynamics is given by the processes of local population extinction and patch (re)colonization by migrants spreading from neighboring occupied patches. In plants with dormant stages (e.g. seeds) and limited dispersal capacity, regional dynamics based on dispersal processes can be overridden by pseudo-turnover determined by signals inducing or breaking dormancy (e.g. due to changes in habitat quality) resulting in a low importance of habitat configuration and size.
In this study, I investigated the turnover dynamics of 5 annual plant species growing on ant mounds of Lasius flavus over three years. I analyzed whether the grassland-scale dynamics of these annuals is influenced by dispersal processes, or alternatively, by pseudo-turnover of soil seed populations. For that purpose I 1) searched for populations formed from soil seeds only, 2) compared the relative contribution of the soil seed bank and seed rain for population restoration after disappearance from the vegetation and 3) investigated whether colonization and extinction events are affected by patch isolation. I assumed if population turnover was rather a result of the soil seed bank dynamics then spatial effects would be hard to detect.
In spite of the presence of populations formed from soil seed and the relatively more important soil seed bank for potential population reestablishment, turnover dynamics followed the predictions of metapopulation theory. Population appearance was more probable in larger and less isolated patches. Probability of disappearance increased with decrease of population size that was negatively influenced by the patch size and its isolation. These findings indicate dispersal processes to be important in the turnover dynamics and only limited contribution of soil seed populations. Their small effectiveness is probably related to the low chance of recurrent disturbance on the mound surface.     

Demographic consequences of population subdivision on the long-furred woolly mouse opossum (Micoureus paraguayanus) from the Atlantic Forest

Habitat destruction and fragmentation severely affected the Atlantic Forest. Formerly contiguous populations may become subdivided into a larger number of smaller populations, threatening their long-term persistence. The computer package VORTEX was used to simulate the consequences of habitat fragmentation and population subdivision on Micoureus paraguayanus, an endemic arboreal marsupial of the Atlantic Forest. Scenarios simulated hypothetical populations of 100 and 2000 animals being partitioned into 1–10 populations, linked by varying rates of inter-patch dispersal, and also evaluated male-biased dispersal. Results demonstrated that a single population was more stable than an ensemble of populations of equal size, irrespective of dispersal rate. Small populations (10–20 individuals) exhibited high instability due to demographic stochasticity, and were characterized by high rates of extinction, smaller values for metapopulation growth and larger fluctuations in population size and growth rate. Dispersal effects on metapopulation persistence were related to the size of the populations and to the sexes that were capable of dispersing. Male-biased dispersal had no noticeable effects on metapopulation extinction dynamics, whereas scenarios modelling dispersal by both sexes positively affected metapopulation dynamics through higher growth rates, smaller fluctuations in growth rate, larger final metapopulation sizes and lower probabilities of extinction. The present study highlights the complex relationships between metapopulation size, population subdivision, habitat fragmentation, rate of inter-patch dispersal and sex-biased dispersal and indicates the importance of gaining a better understanding of dispersal and its interactions with correlations between disturbance events.     

Metapopulation dynamics of a flightless alpine insect Hemideina maori in a naturally fragmented habitat

Abstract 1. Despite widespread acceptance of metapopulation theory, the effects that inter-patch dispersal and variability in patch size have on metapopulation dynamics in insects are two issues that require further study. In addition, previous studies of metapopulations have tended to focus on organisms with high dispersal capabilities such as some species of butterfly and bird.
2. Mountain stone weta Hemideina maori are a long-lived, flightless orthopteran that live on island rock outcrops or tors in the alpine region of southern New Zealand. A total of 480 adults and 789 juveniles was marked over three seasons on four large and 14 small tors to assess the effects of habitat fragmentation on the population dynamics of H. maori .
3. Only 12 adults (2.5% of marked adults and 4.0% of recaptured adults) and two juveniles (0.3% of marked juveniles and 0.7% of recaptured juveniles) dispersed between tors. The mean dispersal distance was 361 m (range = 36–672 m). Larger tors supported larger populations and had a higher number of emigrants and immigrants while smaller tors had proportionally higher emigration and immigration rates. Although adults on large and small tors had similar mean lifespans, five extinction events and three recolonisation events occurred during the study period, all on small tors.
4. Hemideina maori conform to many of the predictions of metapopulation theory even though they are flightless, show relatively low dispersal rates, and occur at low densities. Extinction and colonisation events are more common on small tors but may be relatively unimportant for the long-term survival of the metapopulation because they occur on the smallest habitat patches, which support the smallest proportion of the overall population.     

物种保护的理论基础——从岛屿生物地理学理论到集合种群理论

全球面临着生境破碎化的危机,物种保护已成为人类面临的重大课题,并不是所有的人对岛屿生物地理学理论的产生及其关注的海洋岛屿都很熟悉,但是越来越多生物赖以生存的自然栖息地的丧失和破碎化都是有目共睹的,岛屿生物地理学和集合种群理论是目前物种保护的两个基本理论,物种迁入率和绝灭率的动态变化决策岛屿上的物种丰富度是岛屿生物地理学理论的核心内容,而集合种群理论关注的是局部种群之间个体迁移的动态以及物种的续存条件,在概述两个理论形成、发展及其核心内容的基础上,着重比较它们的异同点以及在生态学理论和实践中的应用,并论述物种保护理论范式从岛屿生物地理学向集合种群理论转变的基本背景和原因。     

Transient dynamics in metapopulation response to perturbation

Transient time in population dynamics refers to the time it takes for a population to return to population-dynamic equilibrium (or close to it) following a perturbation in the environment or in population size. Depending on the direction of the perturbation, transient time may either denote the time until extinction (or until the population has decreased to a lower equilibrium level), or the recovery time needed to reach a higher equilibrium level. In the metapopulation context, the length of the transient time is set by the interplay between population dynamics and landscape structure. Assuming a spatially realistic metapopulation model, we show that transient time is a product of four factors: the strength of the perturbation, the ratio between the metapopulation capacity of the landscape and a threshold value determined by the properties of the species, and the characteristic turnover rate of the species, adjusted by a factor depending on the structure of the habitat patch network. Transient time is longest following a large perturbation, for a species which is close to the threshold for persistence, for a species with slow turnover, and in a habitat patch network consisting of only a few dynamically important patches. We demonstrate that the essential behaviour of the n-dimensional spatially realistic Levins model is captured by the one-dimensional Levins model with appropriate parameter transformations.     

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.     

Metapopulation theory for fragmented landscapes

We review recent developments in spatially realistic metapopulation theory, which leads to quantitative models of the dynamics of species inhabiting highly fragmented landscapes. Our emphasis is in stochastic patch occupancy models, which describe the presence or absence of the focal species in habitat patches. We discuss a number of ecologically important quantities that can be derived from the full stochastic models and their deterministic approximations, with a particular aim of characterizing the respective roles of the structure of the landscape and the properties of the species. These quantities include the threshold condition for persistence, the contributions that individual habitat patches make to metapopulation dynamics and persistence, the time to metapopulation extinction, and the effective size of a metapopulation living in a heterogeneous patch network.     

Metapopulation processes and persistence in remnant water vole populations

We examined the spatial distribution of water vole populations in four consecutive years and investigated whether the regional population processes of extinction, recolonisation and migration influence distribution and persistence. We examined how such regional processes are influenced by spatial variation in habitat quality. In addition, we assessed the relevance of metapopulation concepts for understanding the dynamics of species that deviate from classical metapopulation assumptions and developing conservation measures for them. Populations were patchy and discrete, and the patchy distribution was not static between years. Population turnover occurred even in the absence of predatory mink, which only influenced the network of populations at the end of the study. Most populations were clustered close together in the upper tributaries. Local population persistence was predominantly influenced by population size: large populations were more persistent. Recolonisation rates were influenced by isolation and habitat quality. The isolation estimates which best explained the distribution of water vole populations incorporated straight‐line distances, suggesting water voles disperse overland. The distribution of recolonised sites indicated that dispersing voles actively selected habitat on the basis of its quality. Water voles depart from some of the assumptions made by frequently used metapopulation models. In particular there is no clear binary distinction between suitable and non‐suitable habitat. Accounting for variation in habitat quality before investigating temporal changes in population distribution allowed us to demonstrate that the key metapopulation processes were important. The significance of regional population processes relative to local population processes may have increased in declining, fragmented populations compared to pristine regional populations. We hypothesise that although mink predation is likely to eventually cause regional extinction in many areas, metapopulation processes have delayed this decline. Consequently, conservation measures should take into account mink predation rates and regional population processes, before considering aspects of habitat quality.     

Extinction, colonization, and species occupancy in tidepool fishes

Despite the increasing sophistication of ecological models with respect to the size and spatial arrangement of habitat, there is relatively little empirical documentation of how species dynamics change as a function of habitat size and the fraction of habitat occupied. In an assemblage of tidepool fishes, I used maximum-likelihood estimation to test whether models which included habitat size provided a better fit to empirical data on extinction and colonization probabilities than models that assumed constant probabilities over all habitats. I found species differences in how extinction and colonization probabilities scaled with habitat size (and hence local population size). However, there was little evidence for a relationship between extinction and colonization probabilities and the fraction of occupied tidepools, as assumed in simple metapopulation models. Instead, colonization and extinction were independent of the fraction of occupied tidepools, favoring a MacArthur-Wilson island-mainland model. When I incorporated declines in extinction probability with tidepool volume in a simple simulation model, I found that predicted occupancy could change greatly, especially when colonization was low. However, the predicted fraction of occupied patches in the simulation model changed little when I incorporated the range of values reported here for extinction and colonization and the rate at which they scale with habitat size. Quantifying extinction and colonization patterns of natural populations is fundamental to understanding how species are distributed spatially and whether metapopulation models of species occupancy provide explanatory power for field populations. Received: 14 March 1997 / Accepted: 21 September 1997