栖息地质量对两种网蛱蝶集合种群结构和分布的影响

在河北省赤城县研究了栖息地质量对大网蛱蝶Melitaea phoebe和金堇蛱蝶Euphydryas aurinia两种网蛱蝶集合种群结构和分布的影响。这两种网蛱蝶在约10 km_²的区域内共存,成虫期的蜜源植物几乎相同,大网蛱蝶的发生峰期比金堇蛱蝶晚约一个月,两者只有不到一周左右的时间重叠。大网蛱蝶和金堇蛱蝶幼虫的寄主植物分别是: 祁州漏芦（菊科）和华北蓝盆花（川续断科）。蜜源植物的丰度与两种网蛱蝶的局域种群大小呈正相关;祁州漏芦的密度对大网蛱蝶的局域种群大小影响很大,金堇蛱蝶的局域种群大小则与其寄主植物华北蓝盆花的高度正相关;斑块内平均植被高度与两种网蛱蝶的局域种群大小均呈正相关,植物多样性、植物均匀性和植被盖度均与金堇蛱蝶的局域种群大小负相关,与大网蛱蝶的关系不大。同时分析了其他因子如斑块的坡向、坡度等的影响。主要结论是：1）幼虫寄主植物的不同和成蝶飞行峰期的分离允许两种网蛱蝶在这样一个小的斑块区域内共存;2）蜜源是重要的限制因子,并且受气候随机性的影响很大,蜜源的波动可以很好地解释网蛱蝶集合种群在年度间的动态变化;3）大网蛱蝶和金堇蛱蝶的飞行、食物搜寻能力的不同以及各自寄主植物的生物学特性、空间分布的不同决定了它们具有不同的集合种群结构： 金堇蛱蝶是经典的集合种群,大网蛱蝶是源-汇集合种群;4）斑块质量和昆虫行为共同决定了两种网蛱蝶的集合种群结构和分布。     

Effect of cultivating croplands and grazing in arid grassland habitats on the conservation of melitaeine butterflies in a mountainous area in Northern China

In the study area (Yanjiaping Village, Hebei Province, China), grazing extensity varies at different loca-tions, small and discontinuous croplands are imbedded in some arid grassland, which are habitats for the melitaeine butterflies, Euphydryas aurinia and Melitaea phoebe. These two species of butterflies coexist in this area, in which grazing and cultivation are the main disturbances. Grazing and cultivation have a reciprocal effect on E. aurinia, rather than M. phoebe. We observed that E. aurinia preferred to occupy patches with moderate grazing and imbedded with small and discontinuous croplands, where E. aurinia also has high population density. The percentage of E. aurinia larval groups in the ribbings was significantly higher than that of M. phoebe, whereas larvae of both species tended to increase in recent years. Our data also showed that the population density and the patch occupancy rate of both E. aurinia and M. phoebe were the highest under moderate grazing. It indicates that cultivation of small and dis-continuous croplands within the patch has a significant effect on the population density of both spe-cies of melitaeine butterflies. Thus, to artificially create or maintain semi-natural habitats, comple-mented by moderate grazing, might be an ecological strategy to conserve melitaeine butterflies effec-tively. Considering the distinct impacts of cultivation and grazing on the population distribution and dynamics of the two different species, human disturbance in the mountainous area might be strategi-cally involved in proposing conservation plans for the target species in the future.     

Dynamic populations in a dynamic landscape: the metapopulation structure of the marsh fritillary butterfly

The marsh fritillary butterfly Euphydryas aurinia is an endangered species in most of northern Europe. We describe the metapopulation structure of E. aurinia in Finland, where the species has declined drastically in the past decades. We found two types of habitat patches suitable for the species: semi-permanent meadows and transient clearcuts in the forest. Patch area was the most significant variable predicting the occurrence of E. aurinia in a habitat patch. The species tended to be found in young rather than old clearcuts, apparently because the vegetation became too high in the latter. We used the incidence function model to simulate the metapopulation dynamics of E. aurinia in its dynamic landscape and discovered that the continued presence of the semi-permanent meadows is essential for the survival of the species in the study area in southeast Finland.     

两种共存网蛱蝶幼期的生命表研究

在河北省赤城县大海坨国家自然保护区内,从2002年到2004年应用生命表方法研究了在同一网络斑块中共存的金堇蛱蝶Euphydryas aurinia和大网蛱蝶Melitaea phoebe两个世代的幼期各阶段的死亡情况,目的是了解影响种群动态的重要因子,为它们的长期保育提供信息。结果表明,金堇蛱蝶幼期总累积死亡率都较小,两个世代分别为59%和72%; 而大网蛱蝶总累积死亡率较高,两个世代分别为89%和80%。影响大网蛱蝶死亡的最重要因子是放牧,两个世代与放牧相关的k值分别为0.559和0.167;尤其是在越冬后大网蛱蝶幼虫组聚集取食阶段,导致两个世代分别有50%和25%的幼虫组死亡。寄生蜂在大网蛱蝶小种群下也分别使两个世代4%和9%的5～6龄期幼虫以及13%和24%的蛹死亡。金堇蛱蝶死亡主要发生在越冬期,与越冬死亡相关的k值两个世代分别为0.073和0.199, 而寄主植物的质量影响越冬期幼虫组死亡; 寄生蜂则对金堇蛱蝶种群的调控作用极小,只有在2003～2004世代有4.0% 的越冬后幼虫被寄生和7%的蛹被寄生。影响两种蛱蝶种群动态的关键因子不同,采取的保护措施应有所不同。在春季减少源斑块内的放牧,对于以源-汇集合种群形式存在的大网蛱蝶种群恢复和增长十分必要; 而对以经典集合种群形式存在的金堇蛱蝶, 通过适当植被管理提高繁殖区域内寄主植物质量,可以提高越冬期幼虫组存活率,有利于其长期持续生存。     

Genetic differentiation within metapopulations of Euphydryas aurinia and Melitaea phoebe in China

We analyzed genetic differentiation within metapopulations of two species of checkerspot butterfly, Euphydryas aurinia and Melitaea phoebe, in China. To generate genetic information, we used a new molecular technique, DALP – direct amplified length polymorphism. AMOVA results showed that most of the variation occurred among individuals within local populations of both E. aurinia and M. phoebe. However, while there was differentiation among local population in E. aurinia (P < 0.001), there was no subdivision in metapopulation of M. phoebe (P = 0.210). This is consistent with the behavior of M. phoebe adults being more dispersive than E. aurinia. Within the M. phoebe metapopulation, three neighboring patches were always occupied during the observation period (1998–2000). In addition, the number of individuals in these three populations accounted for the majority of M. phoebe larvae, and hence we conclude that the M. phoebe metapopulation might exist as a source-sink metapopulation. On the other hand, the E. aurinia metapopulation is an example of a classical metapopulation. Therefore, the conservation management of these two species should reflect these differences.     

两种共存网蛱蝶的卵块特征及其雌性成虫对产卵地的选择

对同一网络斑块中金堇蛱蝶(Euphydryas aurinia)和大网蛱蝶(Meiltaea phoebe)雌性成虫的产卵地进行了调查,以确定影响它们斑块质量的因素.结果表明,2种网蛱蝶雌性成虫均选择体积大的寄主植物、周围植被高度低、开放的区域产卵,它们的卵块主要分布在温暖向阳的坡面.金堇蛱蝶的卵块距离农田边缘为3.55±0.33 m (n=246),且集中分布在＜3 m区域内;大网蛱蝶卵块距离农田边缘为7.34±1.53 m (n=25),但在＜3 m区域内的数量少.研究表明,由于这2种网蛱蝶雌性成虫对产卵地要求的特异性,评价它们斑块质量以及对其制定保护管理措施时均应有所不同.     

Spatial ecology of host–parasitoid interactions: a threatened butterfly and its specialised parasitoid

Habitat conservation for threatened temperate insect species is often guided by one of two paradigms: a metapopulation approach focusing on patch area, isolation and number; or a habitat approach focusing on maintaining high quality habitat for the focal species. Recent research has identified the additive and interacting importance of both approaches for maintaining populations of threatened butterflies. For specialised host-parasitoid interactions, understanding the consequences of habitat characteristics for the interacting species is important, because (1) specialised parasitoids are particularly vulnerable to the consequences of fragmentation, and (2) altered interaction frequencies resulting from changes to habitat management or the spatial configuration of habitat are likely to have consequences for host dynamics. The spatial ecology of Cotesia bignellii, a specialist parasitoid of the threatened butterfly Euphydryas aurinia, was investigated at two spatial scales: within habitat patches (at the scale of individual aggregations of larvae, or ‘webs’) and among habitat patches (the scale of local populations). Parasitism rates were investigated in relation to larval web size, vegetation sward height and host density. Within patches, the probability of a larval webs being parasitized increased significantly with increasing number of larvae in the web, and parasitism rates increased significantly with increasing web isolation. The proportion of webs parasitized was significantly and negatively correlated with cluster density. Among habitat patches the proportion of parasitized webs decreased as cluster density increased. Clusters with a high proportion of larval webs parasitized tended to have lower parasitism rates per larval web. These results support the call for relatively large and continuous habitat patches to maintain stable parasitoid and host populations. Conservation efforts directed towards maintenance of high host plant density could allow E. aurinia to reduce parasitism risk, while providing C. bignellii with sufficient larval webs to allow population persistence.     

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.     

Three ways of assessing metapopulation structure in the butterfly Plebejus argus

1. Three independent methods were used to investigate population structure in the butterfly Plebejus argus . First, migration and dispersal ability were measured by mark–release–recapture in seven adjacent habitat patches, and by release of butterflies in unoccupied habitat. Secondly, colonization of newly created habitat was observed over 7 years. Finally, genetic differentiation of local populations within a metapopulation was investigated. Sampled local populations included parts of the mark–release–recapture study area.
2. Plebejus argus is relatively sedentary: the maximum movement detected was 395 m, and only 2% of individuals moved further than 100 m between recaptures on different days. None the less, adjacent local populations in the mark–release–recapture study area were linked by occasional migration, with ≈ 1.4% of individuals moving between patches separated by 13–200 m.
3. Despite low mobility, observed colonizations occurred rapidly over distances of 1 km. Because P. argus occurs at high population densities, 1.4% migration can generate enough migrants to colonize newly suitable habitat quickly at this spatial scale.
4. Mark–release–recapture data were used to predict that there would be limited genetic differentiation through drift between local populations at this spatial scale. The prediction was supported by allele frequency data for the same local populations.
5. Genetic differentiation often indicates higher levels of migration than are revealed by the movements of marked individuals. This study shows that when experimental releases and extensive marking are undertaken in areas that are large relative to most movements, indirect measures of gene flow and direct measures of dispersal can concur.
6. Evidence from the three different approaches was complementary, indicating that P. argus occurs as metapopulations within the study area.     

What is a patch in a dynamic metapopulation? Mobility of an endangered woodland butterfly,Euphydryas maturna

In metapopulations of butterflies inhabiting transient forest openings, the dynamic character of biotopes prevents unequivocal delimitation of habitat patches, complicating analyses of inter-patch dispersal. We analyse mobility of one such metapopulation, using mark-recapture data on an endangered butterfly, the scarce fritillary Euphydryas maturna , at its last site in the Czech Republic. The butterfly inhabits woodland clearings, its population numbered 190 individuals in 2002. We modelled movements in eight habitat patchworks delimited according to different criteria and superimposed over the forest, using the Virtual Migration model. Resulting mobility parameters were generally robust against patchwork definitions, the main distinction was between patchworks based on presence of adults and consisting of small patches and patchworks based on distributions of larval nests and consisting of large patches. Emigration constant decreased towards the latter, whereas males suffered higher migration mortality than females in the former. Patchworks delimited according to presence of adults performed better than patchworks based on distribution of larval nests. A comparison of our parameters with those found for the species in Finland showed that the Czech butterflies were more prone to leave individual patches and suffered higher migration mortality, which combined into considerably shorter survival. We conclude that transferring dispersal parameters among different regions and populations is more risky than using less suitable definitions of habitat patches within single region.     

Migration and Allee effects in the six-spot burnet moth Zygaena filipendulae

Abstract 1. Migration into local populations may increase the likelihood of persistence but emigration may decrease the persistence of small and isolated populations. The dispersal behaviour of a day-flying moth Zygaena filipendulae was examined to determine whether emigration is correlated positively or negatively with population size and host plant density.
2. A mark–release–recapture study showed that most moths moved small distances (< 40 m on average) and only 6% of movements were > 100 m.
3. Twenty-five individuals moved between populations, a measured exchange rate of 8%. Moths were more likely to move between patches that were close together and they moved to relatively large patches.
4. The fraction of residents increased with increasing population size in the patch and increasing host plant cover. Relatively high proportions of individuals left small patches with small moth populations.
5. Moths released in grassland lacking Lotus corniculatus (the host plant) tended to leave the area and biased their movement towards host plant areas, whereas those released within an area containing L. corniculatus tended to stay in that area.
6. Biased movement away from small populations and areas of low host plant density (normally with low population density) was found. This migration-mediated Allee effect is likely to decrease patch occupancy in metapopulations, the opposite of the rescue effect. The effects on metapopulation persistence are not known.     

Area-dependent migration by ringlet butterflies generates a mixture of patchy population and metapopulation attributes

Interpretation of spatially structured population systems is critically dependent on levels of migration between habitat patches. If there is considerable movement, with each individual visiting several patches, there is one ”patchy population”; if there is intermediate movement, with most individuals staying within their natal patch, there is a metapopulation; and if (virtually) no movement occurs, then the populations are separate (Harrison 1991, 1994). These population types actually represent points along a continuum of much to no mobility in relation to patch structure. Therefore, interpretation of the effects of spatial structure on the dynamics of a population system must be accompanied by information on mobility. We use empirical data on movements by ringlet butterflies, Aphantopus hyperantus, to investigate two key issues that need to be resolved in spatially-structured population systems. First, do local habitat patches contain largely independent local populations (the unit of a metapopulation), or merely aggregations of adult butterflies (as in patchy populations)? Second, what are the effects of patch area on migration in and out of the patches, since patch area varies considerably within most real population systems, and because human landscape modification usually results in changes in habitat patch sizes? Mark-release-recapture (MRR) data from two spatially structured study systems showed that 63% and 79% of recaptures remained in the same patch, and thus it seems reasonable to call both systems metapopulations, with some capacity for separate local dynamics to take place in different local patches. Per capita immigration and emigration rates declined with increasing patch area, while the resident fraction increased. Actual numbers of emigrants either stayed the same or increased with area. The effect of patch area on movement of individuals in the system are exactly what we would have expected if A. hyperantus were responding to habitat geometry. Large patches acted as local populations (metapopulation units) and small patches simply as locations with aggregations (units of patchy populations), all within 0.5 km². Perhaps not unusually, our study system appears to contain a mixture of metapopulation and patchy-population attributes.     

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.     

Genetic diversity and migration patterns of the aquatic macrophyte Potamogeton malaianus in a potamo-lacustrine system

1. Previously, the Yangtze River connected thousands of shallow lakes which together formed a potamo-lacustrine system capable of sustaining a rich variety of submerged macrophytes.
2.  Potamogeton malaianus is one of the dominant submerged macrophytes in many lakes of this area. Genetic variation and population structure of P. malaianus populations from ten lakes in the potamo-lacustrine system were assessed using inter-simple sequence repeat markers.
3. Twelve primer combinations produced a total of 166 unambiguous bands of which 117 (70.5%) were polymorphic. Potamogeton malaianus exhibited a moderate level of population genetic diversity ( P _P = 70.5%, H _E = 0.163 and I =  0.255), as compared with that of plants in the same habitat and range. The main factors responsible for this moderate value were the plant's mixed breeding system (both sexual and asexual) and the hydrological connectivity among habitats.
4.  F statistics, calculated using different approaches, consistently revealed a moderate genetic differentiation among populations, contributing about 20% of total genetic diversity. An estimate of gene flow (using F _ST) suggested that gene flow played a more important role than genetic drift in the current population genetic structure of P. malaianus ( Nm  = 1.131).
5. The genetic diversity of P. malaianus did not increase downstream. A high level of linkage–disequilibrium at the whole population level suggested that metapopulation processes may affect genetic structure. The migration pattern of P. malaianus was best explained by a two-dimensional stepping stone model, indicating that bird-mediated dispersal could greatly influence gene movements among lakes.     

Metapopulation structure and movements in five species of checkerspot butterflies

We studied the patterns and rates of migration among habitat patches for five species of checkerspot butterflies (Lepidoptera: Melitaeini) in Finland: Euphydryas aurinia, E. maturna, Melitaea cinxia, M. diamina and M. athalia. We applied the virtual migration (VM) model to mark-release-recapture data collected from multiple populations. The model includes parameters describing migration and survival rates and how they depend on the areas and connectivities of habitat patches. The number of individuals captured varied from 73 to 1,123, depending on species and sex, and the daily recapture probabilities varied between 0.09-0.52. The VM model fitted the data quite well. The results show that the five species are broadly similar in their movement rates and patterns, though, e.g. E. maturna tends to move shorter distances than the other species. There is no indication of any phylogenetic component in the parameter values. The parameter values estimated for each species suggest that a large percentage (80-90%) of migration events were successful in the landscapes that were studied. The area of the habitat patch had a substantial effect on emigration and immigration rates, such that butterflies were more likely to leave small than large patches and large patches were more likely than small patches to receive immigrants.     

Is the matrix a sea? Habitat specificity in a naturally fragmented landscape

Abstract.  1. Metapopulation and island biogeography theory assume that landscapes consist of habitat patches set in a matrix of non-habitat. If only a small proportion of species conform to the patch–matrix assumptions then metapopulation theory may only describe special cases rather than being of more general ecological importance.
2. As an initial step towards understanding the prevalence of metapopulation dynamics in a naturally fragmented landscape, the distribution of beetle species in three replicates of three habitat types was examined, including rainforest and eucalypt forest (the habitat patches), and buttongrass sedgeland (the matrix), in south-west Tasmania, Australia.
3. Ordination methods indicated that the buttongrass fauna was extremely divergent from the fauna of forested habitats. Permutation tests showed that the abundance of 13 of 17 commonly captured species varied significantly among habitats, with eight species confined to eucalypts or rainforest, and three species found only in buttongrass. Approximately 60% of species were confined to forested habitat implying that metapopulation theory has the potential to be very important in the forest–buttongrass landscape.
4. Although floristically the rainforest and eucalypts were extremely distinct, the beetle faunas from eucalypts and rainforests overlapped substantially. Therefore rainforest patches connected by eucalypt forest represent continuous habitat for most species.
5. Other studies report a wide range of values for the proportion of patch-specific species in fragmented landscapes. Understanding the environmental or historical conditions under which a high proportion of species become patch specialists would help to identify where spatial dynamic theory may be especially applicable, and where habitat loss and fragmentation poses the greatest threat to biodiversity.     

Mass aerial migration in the carabid beetle Notiophilus biguttatus

Abstract.  1. Catches in traps at 12 and 200 m above ground, and observations with entomological radar, were used to document the high-altitude windborne movements of carabid beetles over agricultural land in south-east U.K. during July 1999, 2000, 2002, and 2004.
2.  Notiophilus biguttatus (Fabricius) was found to be the most abundant species at altitude, with flights by reproductively immature adults resulting in millions of beetles passing through a 1 km² window during the brief migration period in July 2002.
3. This result was unexpected in view of the conclusions of an earlier, classic Dutch study by den Boer and colleagues that classified N. biguttatus as a poorly dispersing species that tended to become isolated in remnants of natural habitat.
4. Reasons for the discrepancy between the two studies are discussed. It is hypothesised that the high mobility seen in the N. biguttatus populations results from a recent adaptation of the beetle's migration syndrome allowing it to exploit the mosaic of temporary habitat patches making up the increasingly intensive agroecosystems of northern Europe.     

Dispersal, distribution, patch network and metapopulation dynamics of the dingy skipper butterfly (Erynnis tages)

Two general approaches have usually been taken towards understanding the distributions and dynamics of localised species in heterogeneous landscapes, namely habitat characterisation and metapopulation dynamics. We show how habitat and metapopulation dynamics interact to generate a highly localised distribution of a butterfly, despite the extremely widespread nature of the butterfly’s host plant. Egg placement, macro-habitat requirements and dispersal were studied for the butterfly Erynnis tages, in North Wales, where it shows a restricted distribution relative to that of its host plant, Lotus corniculatus. Females laid eggs disproportionately on large plants growing in hollows, with intermediate cover of bare ground and high cover of L. corniculatus. Ideal macro-habitat, studied at 100-m grid resolution, consisted of areas with high host plant densities, sheltered from wind, with light or no grazing or cutting. These specialised conditions are represented as localised patches in the landscape, and define the potential habitat network, within which metapopulation dynamics take place. Although there was a moderate (22%) level of exchange of individual E. tages among local populations, the total number of potential colonists in the whole system was low because source population sizes were small (≤200 individuals at peak in any site in 1997 and 1998). Four unoccupied but apparently suitable 500-m grid squares were colonised between 1997 and 1998, and isolated habitat was less likely to be occupied. Overall, our study suggests that long-term regional persistence of E. tages is very likely to depend on metapopulation processes within the restricted patch network, rather than on the long-term survival of local populations. Received: 25 May 1999 / Accepted: 9 August 1999     

Single-species metapopulation dynamics: concepts, models and observations

This paper outlines a conceptual and theoretical framework for single-species metapopulation dynamics based on the Levins model and its variants. The significance of the following factors to metapopulation dynamics are explored: evolutionary changes in colonization ability; habitat patch size and isolation; compensatory effects between colonization and extinction rates; the effect of immigration on local dynamics (the rescue effect); and heterogeneity among habitat patches. The rescue effect may lead to alternative stable equilibria in metapopulation dynamics. Heterogeneity among habitat patches may give rise to a bimodal equilibrium distribution of the fraction of patches occupied in an assemblage of species (the core-satellite distribution). A new model of incidence functions is described, which allows one to estimate species' colonization and extinction rates on islands colonized from mainland. Four distinct kinds of stochasticity affecting metapopulation dynamics are discussed with examples. The concluding section describes four possible scenarios of metapopulation extinction.     

Asynchronization of local population dynamics and persistence of a metapopulation: a lesson from an endangered composite plant, Aster kantoensis

Fragmentation of a large habitat makes local populations less linked to others, and a whole population structure changes to a metapopulation. The smaller a local population is, the more strengthened extinction factors become. Then, frequent extinctions of local populations threaten persistence of the metapopulation unless recolonizations occur rapidly enough after local extinctions. Spatially structured models have been more widely used for predicting future population dynamics and for assessing the extinction risk of a metapopulation. In this article, we first review such spatially structured models that have been applied to conservation biology, focusing on effects of asynchronization among local population dynamics on persistence of the whole metapopulation. Second, we introduce our ongoing project on extinction risk assessment of an endangered composite biennial plant, Aster kantoensis, in the riverside habitat, based on a lattice model for describing its spatiotemporal population dynamics. The model predicted that the extinction risk of A. kantoensis depends on both the frequency of flood occurrence and the time to coverage of a local habitat by other competitively stronger perennials. Finally, we present a measure (Hassell and Pacala's CV ²) for quantifying the effect of asynchronization among local population dynamics on the persistence of a whole metapopulation in conservation ecology. Received: January 12, 2000 / Accepted: February 8, 2000