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1.
Aim To use molecular data to test for dispersal structuring in the immigration history of an amphidromous community on an island. Location The Caribbean island of Puerto Rico. Methods Mitochondrial DNA sequences were obtained from 11 amphidromous species, including shrimps, fish and a gastropod, sampled from throughout the island. The timing of population expansion (TE) in each species was calculated using nucleotide variation and molecular clock dating methods. The order of species accumulation was then reconstructed (oldest to most recent estimate for TE), and groups of species with non‐overlapping estimates for TE were identified. The temporal span and average immigration rate for each group were calculated and compared with expectations of two previously published models of island immigration [the ‘dispersal‐structured model of island recolonization’ ( Whittaker & Jones, Oikos, 1994 , 69 , 524–529), which predicts short phases of rapid immigration followed by extended phases with relatively slow immigration rates; and the ‘colonization window hypothesis’ ( Carine, Taxon, 2005 , 54 , 895–903), which suggests that opportunities for island colonization are temporally constrained to discrete waves of colonization]. Results The molecular data indicated the immigration history of Puerto Rican amphidromous fauna from the late Pleistocene through the Holocene and identified two groups of species with non‐overlapping estimates for TE and one group that overlapped with the other two groups. The temporal span, average immigration rate and lack of discreteness between all three groups indicated a continuum of immigration rather than distinct phases of species arrivals. Main conclusions This study did not support the expectations of the immigration models and suggested that amphidromous species from Puerto Rico comprise a single class of marine‐based dispersers. The immigration sequence we report probably reflects a recolonization chronology in this community, in keeping with the notion of species turnover through time. Four areas of future research into the immigration history of amphidromous species on islands are identified, and indicated the possibility that equilibrium processes govern long‐term community change in amphidromous biota on islands.  相似文献   

2.
Abiotic environmental change, local species extinctions and colonization of new species often co‐occur. Whether species colonization is driven by changes in abiotic conditions or reduced biotic resistance will affect community functional composition and ecosystem management. We use a grassland experiment to disentangle effects of climate warming and community diversity on plant species colonization. Community diversity had dramatic impacts on the biomass, richness and traits of plant colonists. Three times as many species colonized the monocultures than the high diversity 17 species communities (~30 vs. 10 species), and colonists collectively produced 10 times as much biomass in the monocultures than the high diversity communities (~30 vs. 3 g/m2). Colonists with resource‐acquisitive strategies (high specific leaf area, light seeds, short heights) accrued more biomass in low diversity communities, whereas species with conservative strategies accrued most biomass in high diversity communities. Communities with higher biomass of resident C4 grasses were more resistant to colonization by legume, nonlegume forb and C3 grass colonists, but not by C4 grass colonists. Compared with effects of diversity, 6 years of 3°C‐above‐ambient temperatures had little impact on plant colonization. Warmed subplots had ~3 fewer colonist species than ambient subplots and selected for heavier seeded colonists. They also showed diversity‐dependent changes in biomass of C3 grass colonists, which decreased under low diversity and increased under high diversity. Our findings suggest that species colonization is more strongly affected by biotic resistance from residents than 3°C of climate warming. If these results were extended to invasive species management, preserving community diversity should help limit plant invasion, even under climate warming.  相似文献   

3.
Milesi  Silvia V.  Melo  Adriano S.  Dolédec  Sylvain 《Hydrobiologia》2019,838(1):183-192

Initial colonists of empty habitats tend to differ from those arriving later in terms of species identity and traits. We evaluated the dynamics of the functional attributes in aquatic insect communities during a colonization experiment under natural conditions. We tested whether the late stages of colonization show higher functional richness, diversity and specialization than early successional stages. We used 60 artificial slate samplers that were removed after 1, 3, 5, 10, 15 and 30 days of colonization. We considered five traits (with a total of 17 trait categories): feeding habits, dispersal medium, body size, body shape and locomotion. With these traits, we computed a global specialization index at the community level. Large shredders with a cylindrical body shape and fly dispersal while flying were prominent in late colonization. In contrast, early colonists tended to have flattened body and to disperse through water. Functional diversity and functional richness significantly increased in late colonization, resulting in a more specialized community. Our results suggest that any interference during the processes involved in stream insect colonization can be reflected in the community through the decrease or even lack of functional attributes.

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4.
The composition of communities of sessile organisms, and the change in species diversity with time, is a spatially explicit phenomenon. Three spatial factors clearly affect diversity: (1) the structure and heterogeneity of the landscape that limits species immigration and ultimate community size; (2) neighborhood interactions that determine colonization and extinction rates and influence residence times of local populations; and (3) disturbances that open spatially contiguous areas for recolonization by less abundant species. The importance of these three factors was first reviewed and then examined with a spatially explicit, multi-species model of plant dispersal, competition and establishment, with an assumption of neutrality (all species had equivalent life histories) that reduced the initial dimensionality of the problem. The simulations assumed that the probability of immigration was a linear function of mainland abundance and distance to islands, similar to the equilibrium theory of island biogeography and the unified neutral theory of biodiversity. The rate of increase in species richness was not constant across island sizes, declining as island area became very large. This pattern was explained by the spatial dynamics of colonization and establishment, a non-random process that cannot be explained by passive sampling alone. Simulations showed that population establishment depended critically on rare long-distance dispersal events while population persistence was achieved by the formation of aggregated species distributions that developed through restricted dispersal and local competitive interactions. Nevertheless, species richness always declined to a single species in the absence of disturbances, while up to 40 species could persist to 10,000 years when spatially dependent mortality was added. Further explorations with spatially explicit models will be required to fully appreciate the consequence of land use change and altered disturbance regimes on patterns of species distribution and the maintenance of diversity.  相似文献   

5.
An important goal in ecology is to discern under what habitat conditions community structure is primarily regulated by local ecological interactions and under what conditions community structure is more regulated by the pool of available colonists. I conducted a seed addition experiment in successional grassland to evaluate the relative significance of neighbourhood biotic interactions and propagule availability in regulating plant colonization and species richness along a natural gradient of grassland productivity. In undisturbed field plots, seed additions of 34 species led to an increase in species richness in locations of low productivity, an effect that declined in magnitude as productivity increased. In disturbed plots, seed additions led to a relatively constant increase in species richness at all levels of productivity. The results support the hypothesis that the role of propagule availability in regulating colonization dynamics and species richness declines in significance relative to local-scale competitive interactions as habitat productivity increases.  相似文献   

6.
Island biodiversity has long fascinated biologists as it typically presents tractable systems for unpicking the eco‐evolutionary processes driving community assembly. In general, two recurring themes are of central theoretical interest. First, immigration, diversification, and extinction typically depend on island geographical properties (e.g., area, isolation, and age). Second, predictable ecological and evolutionary trajectories readily occur after colonization, such as the evolution of adaptive trait syndromes, trends toward specialization, adaptive radiation, and eventual ecological decline. Hypotheses such as the taxon cycle draw on several of these themes to posit particular constraints on colonization and subsequent eco‐evolutionary dynamics. However, it has been challenging to examine these integrated dynamics with traditional methods. Here, we combine phylogenomics, population genomics and phenomics, to unravel community assembly dynamics among Pheidole (Hymenoptera, Formicidae) ants in the isolated Fijian archipelago. We uphold basic island biogeographic predictions that isolated islands accumulate diversity primarily through in situ evolution rather than dispersal, and population genomic support for taxon cycle predictions that endemic species have decreased dispersal ability and demography relative to regionally widespread taxa. However, rather than trending toward island syndromes, ecomorphological diversification in Fiji was intense, filling much of the genus‐level global morphospace. Furthermore, while most endemic species exhibit demographic decline and reduced dispersal, we show that the archipelago is not an evolutionary dead‐end. Rather, several endemic species show signatures of population and range expansion, including a successful colonization to the Cook islands. These results shed light on the processes shaping island biotas and refine our understanding of island biogeographic theory.  相似文献   

7.
In the recently broadened context of insular biogeographic theory, artificial substrates might well be considered as initially barren islands with the surrounding natural community acting as a source pool of potential colonists. In an attempt to relate protozoan communities forming on polyurethane foam (PF) substrates to their environments, a series of field experiments were performed in several aquatic and semi-aquatic (wetlands) systems in Michigan, Virginia, and Colorado. Results were considered in the context of island colonization theory.Given relatively stable environmental conditions and sufficient time to colonize, protozoa formed highly replicable communities on PF substrates; MacArthur & Wilson's equation for noninteractive island colonization, S = Seq ·(l-eGt ), accuratelt described the process of species accrual in most cases. Sites were compared using non-linear regression estimates of the parameter, G, which reflects the rate at which substrates attained equilibrium species numbers. In general, the rate of equilibrium acquisition reflected differences in productivity and structure between the protozoan communities from the various habitats and systems studied.  相似文献   

8.
The native land-snail fauna of the Hawaiian islands was investigated from a combined perspective of ecological and historical, vicariant, and dispersalist biogeography. There were more than 750 described, valid species; almost all were endemic to the archipelago, many to single islands. Path analysis showed that island area, per se, had the strongest influence on numbers of species. Island altitude and number of plant communities, both strongly related to area and both dimensions of habitat diversity, also had major influences. The influence of island age was complex. A direct effect, older islands having more species, was more than counterbalanced by the strong indirect effects of age on area and altitude: older islands are smaller and lower, and smaller, lower islands had fewer species. Distance of an island from a source of colonization was of minor importance. Species richness thus appears to be related almost exclusively to evolutionary radiation in situ and not to an equilibrium between immigration and extinction. Islands need not be extremely isolated for evolutionary radiation to be more important than immigration/extinction dynamics in determining species richness, but isolation is a relative term dependent on the dispersal abilities of the organisms in question. Numbers of recorded species were also strongly correlated with collecting effort on each island, a result that stands as a warning to others involved in such studies. Numbers of species in different families were not evenly distributed across islands. Notably, Kauai had more amastrids and helicinids and fewer achatinellids than predicted; Oahu had more amastrids but fewer pupillids and succineids than predicted; Hawaii exhibited the opposite pattern from Oahu. These patterns may partly reflect the vagaries of collecting/describing effort, but some may be due to the combined effects of historical factors and competitive exclusion. The distribution of shell height/diameter was bimodal with a distinct absence of more or less equidimensional species, a general pattern seen in other faunas. Among the pulmonates, tall species predominated, suggesting a relative lack of opportunity for globular/flat species. Notably, amastrids occurred in both modes, evidence that, at least in part, ecological not taxonomic factors underlie the bimodality. The proportions of tall and globular/flat species did not vary among islands. Prosobranchs were mostly low-spired but generally less flat than the pulmonates in the low-spired mode. The islands were probably colonized originally by small taxa. Large, tall shells are found only on Kauai and Niihau, the oldest of the main islands, suggesting that opportunities for such species are probably available on other islands.  相似文献   

9.
Question: Do regional species pools, landscape isolation or on‐site constraints cause plants from different guilds to vary in their ability to colonize restored wetlands? Location: Iowa, Minnesota, and South Dakota, USA. Methods: Floristic surveys of 41 restored wetlands were made three and 12 years after reflooding to determine changes in local species pools for eight plant guilds. The effect of landscape isolation on colonization efficiency was evaluated for each guild by plotting local species pools against distance to nearby natural wetlands, and the relative importance of dispersal vs. on‐site constraints in limiting colonization was explored by comparing the local species pools of restored and natural wetlands within the region. Results: Of the 517 wetland plant taxa occurring in the region, 50% have established within 12 years. The proportion of the regional species pool represented in local species pools differed among guilds, with sedge‐meadow perennials, emergent perennials and floating/submersed aquatics least represented (33‐36%) and annual guilds most represented (74‐94%). Colonization‐to‐extinction ratios suggest that floating/submersed aquatics have already reached a species equilibrium while sedge‐meadow and emergent perennials are still accumulating species. Increasing distance to nearest wetlands decreased the proportion of the regional species pool present in local pools for all guilds except native annuals and woody plants. The maximum proportion predicted, assuming no distance constraint, was comparable to the lowest‐diversity natural wetlands for most perennial guilds, and also lower than what was achieved in a planted, weeded restoration. Conclusions: A biotic constraints seem to limit the colonization of floating/submersed aquatics into natural or restored wetlands, whereas all other guilds are potentially constrained by dispersal or biotic factors (i.e. competition from invasive species). Using species pools to evaluate restoration progress revealed that immigration potential varies considerably among guilds, that local species richness does not necessarily correspond to immigration limitations, and that some guilds (e.g. sedge‐meadow perennials) will likely benefit more than others from being planted at restoration sites.  相似文献   

10.
The colonization of ball-clay ponds by macroinvertebrates and macrophytes   总被引:4,自引:0,他引:4  
SUMMARY. 1. The rate and nature of colonization of ball-clay ponds by aquatic macrophytes and macroinvertebrates were studied by comparing communities present in a scries of ten ponds of similar dimensions, but of different ages, ranging from 6 months to 15 years.
2. Multivariate analyses of the biotic data distinguished between'neutral'and'acid'ponds; further analyses concentrated on the former type.
3. Initial colonization of neutral ponds was rapid with a predictable sequence of species arrival, probably reflecting short dispersal distances. Variations in macroinvertebrate invasion times were related to dispersal strategy and ability. As the pond aged, successional changes, linked with macrophyte colonization, produced a shift in dominance from algivores and predators towards epiphyton grazers and detritivores. Some evidence of succession of species within invertebrate orders was found.
4. Low pH affected colonization principally by preventing the establishment of acid-intolerant immigrant species, but also by retarding succession.
5. It is concluded that non-interactive models of insular colonization, with constant immigration and extinction rates, are of limited applicability to pond colonization in temperate areas.  相似文献   

11.
We examined the species-area relationship for three historically distinct subsets of Lesser Antillean birds identified by molecular phylogenetic analysis of island and continental populations. The groups comprised recent colonists from continental or Greater Antillean source populations, old taxa having recently expanded distributions within the Lesser Antilles, and old endemic taxa lacking evidence of recent dispersal between islands. The number of young taxa was primarily related to distance from the source of colonists in South America. In a multiple regression, the logarithmic slope of the species-area relationship for this group was shallow (0.066+/-0.016). Old endemic taxa were restricted to islands with high elevation, and within this subset, species richness was related primarily to island area, with a steep slope (0.719+/-0.110). The number of recently spread endemic taxa was related primarily to island elevation, apparently reflecting the persistence of such populations on islands with large areas of forested and montane habitats. Historical analysis of the Lesser Antillean avifauna supports the dynamic concept of island biogeography of MacArthur and Wilson, rather than the more static view of David Lack, in that colonists exhibit dispersal limitation and extinction plays a role in shaping patterns of diversity. However, the avifauna of the Lesser Antilles is probably not in equilibrium at present, and the overall species-area relationship might reflect changing proportions of historically distinguishable subsets of species.  相似文献   

12.
According to the equilibrium theory of island biogeography, high colonization ability of species is associated with low exponents (z) of the species–area relationship (SAR) and weak spatial patterns in species number and dissimilarity. However, the relationship between z and the strength of these spatial patterns has not been investigated systematically. We used a multispecies metapopulation model to investigate these relationships in an archipelago of islands. We conclude that this relationship can only be predicted if either the dispersal ability or the power of establishment of species is known. With species richness limited by establishment, we generated high z‐values associated with weak spatial patterns in species number and dissimilarity. If species richness was constrained by the dispersal ability of species, we observed low to medium z‐values but strong spatial patterns. If the dispersal ability and the abilities of species to establish were both high, z‐values and spatial pattern tend to be low and species numbers were limited by the size of the regional species pool.  相似文献   

13.
Binckley CA  Resetarits WJ 《Oecologia》2007,153(4):951-958
The specific dispersal/colonization strategies used by species to locate and colonize habitat patches can strongly influence both community and metacommunity structure. Habitat selection theory predicts nonrandom dispersal to and colonization of habitat patches based on their quality. We tested whether habitat selection was capable of generating patterns of diversity and abundance across a transition of canopy coverage (open and closed canopy) and nutrient addition by investigating oviposition site choice in two treefrog species (Hyla) and an aquatic beetle (Tropisternus lateralis), and the colonization dynamics of a diverse assemblage of aquatic insects (primarily beetles). Canopy cover produced dramatic patterns of presence/absence, abundance, and species richness, as open canopy ponds received 99.5% of propagules and 94.6% of adult insect colonists. Nutrient addition affected only Tropisternus oviposition, as females oviposited more egg cases at higher nutrient levels, but only in open canopy ponds. The behavioral partitioning of aquatic landscapes into suitable and unsuitable habitats via habitat selection behavior fundamentally alters how communities within larger ecological landscapes (metacommunities) are linked by dispersal and colonization.  相似文献   

14.
The extent that biotic interactions and dispersal influence species ranges and diversity patterns across scales remains an open question. Answering this question requires framing an analysis on the frontier between species distribution modelling (SDM), which ignores biotic interactions and dispersal limitation, and community ecology, which provides specific predictions on community and meta‐community structure and resulting diversity patterns such as species richness and functional diversity. Using both empirical and simulated datasets, we tested whether predicted occurrences from fine‐resolution SDMs provide good estimates of community structure and diversity patterns at resolutions ranging from a resolution typical of studies within reserves (250 m) to that typical of a regional biodiversity study (5 km). For both datasets, we show that the imprint of biotic interactions and dispersal limitation quickly vanishes when spatial resolution is reduced, which demonstrates the value of SDMs for tracking the imprint of community assembly processes across scales.  相似文献   

15.
岛屿生物地理学理论的核心过程是岛屿物种的周转, 包括迁入与灭绝。本研究旨在探讨扩散能力差异对岛屿繁殖鸟类群落动态的影响。2007年4月至2013年6月, 采用样线法调查了千岛湖36个陆桥岛屿的繁殖鸟类, 依据扩散能力强弱将其划分为两类, 结合陆桥岛屿参数, 并运用逻辑斯蒂回归模型和最大似然法, 来研究鸟类扩散能力的不同对其周转率的影响。结果表明, 千岛湖繁殖鸟类扩散能力强的物种具有较高周转率且受岛屿参数约束较小, 而扩散能力弱的物种周转率较低且对岛屿参数变化更敏感。因此, 千岛湖陆桥岛屿繁殖鸟类的扩散能力显著影响其群落动态。  相似文献   

16.
Isolation is a driving factor of species richness and other island community attributes. Most empirical studies have investigated the effect of isolation measured as distance to the nearest continent. Here we expanded this perspective by comparing the explanatory power of seventeen isolation metrics in sixty‐eight variations for vascular plant species richness on 453 islands worldwide. Our objectives were to identify ecologically meaningful metrics and to quantify their relative importance for species richness in a globally representative data set. We considered the distances to the nearest mainland and to other islands, stepping stone distances, the area of surrounding landmasses, prevailing wind and ocean currents and climatic similarity between source and target areas. These factors are closely linked to colonization and maintenance of plant species richness on islands. We tested the metrics in spatial multi‐predictor models accounting for area, climate, topography and island geology. Besides area, isolation was the second most important factor determining species richness on the studied islands. A model including the proportion of surrounding land area as the isolation metric had the highest predictive power, explaining 86.1% of the variation. Distances to large islands, stepping stone distances and distances to climatically similar landmasses performed slightly better than distance to the nearest mainland. The effect of isolation was weaker for large islands suggesting that speciation counteracts the negative effect of isolation on immigration on large islands. Continental islands were less affected by isolation than oceanic islands. Our results suggest that a variety of immigration mechanisms influence plant species richness on islands and we show that this can be detected at macro‐scales. Although the distance to the nearest mainland is an adequate and easy‐to‐calculate measure of isolation, accounting for stepping stones, large islands as source landmasses, climatic similarity and the area of surrounding landmasses increases the explanatory power of isolation for species richness.  相似文献   

17.
For the mammalian faunas of 24 landbridge islands in the Gulf of Maine (0.003–279 km2 in size), area accounts for 86% of variance in species richness. The slope, z , of the species-area curve is 0.247. For the seven largest islands (>10km2), the non-equilibrium hypothesis of relaxation following saturation in the post-Pleistocene is suggested by (1) elevated slope of the species-area curve (0.353), (2) correlation of species richness with island age ( r =- 0.81) and water depth to mainland ( r = -0.70), (3) highly non-random nested subsets of species ranked by island area, and (4) discontinuity with the extremely depauperate faunas of oceanic islands of the eastern North Atlantic. The alternative hypothesis of a dynamic equilibrium determined by recurrent immigration and extinction is supported by (1) documented turnover in 16 species, (2) correlation of species-area residuals with distance ( r = - 0.90), (3) distribution dependent upon vagility with reduction or absence of hibernators and other poor dispersers, (4) low levels of endemism, and (5) congruence of community structure with that of mainland fauna for both trophic level and body size.
I conclude that while some insular populations may be relictual, the faunal composition of most of these islands is dependent on recurrent colonization, much of which takes place over ice bridges. However, true equilibrium is perturbed by climatic shifts, range expansions, and human disturbance.  相似文献   

18.
An aim of community ecology is to understand the patterns of competing species assembly along environmental gradients. All species interact with their environments. However, theories of community assembly have seldom taken into account the effects of species that are able to engineer the environment. In this modeling study, we integrate the species' engineering trait together with processes of immigration and local dispersal into a theory of community assembly. We quantify the species' engineering trait as the degree to which it can move the local environment away from its baseline state towards the optimum state of the species (species‐environment feedback). We find that, in the presence of immigration from a regional pool, strong feedback can increase local species richness; however, in the absence of continual immigration, species richness is a declining function of the strength of species‐environment feedback. This shift from a negative effect of engineering strength on species richness to a positive effect, as immigration rate increases, is clearer when there is spatial heterogeneity in the form of a gradient in environmental conditions than when the environment is homogeneous or it is randomly heterogeneous. Increasing the scale over which local dispersal occurs can facilitate species richness when there is no species‐environment feedback or when the feedback is weak. However, increases in the spatial scale of dispersal can reduce species richness when the species‐environment feedback is strong. These results expand the theoretical basis for understanding the effects of the strength of species‐environment feedback on community assembly.  相似文献   

19.
Dispersal plays a prominent role in most conceptual models of community assembly. However, direct measurement of dispersal across a whole community is difficult at ecologically relevant spatial scales. For cryptic organisms, such as fungi and bacteria, the scale and importance of dispersal limitation has become a major point of debate. We use an experimental island biogeographic approach to measure the effects of dispersal limitation on the ecological dynamics of an important group of plant symbionts, ectomycorrhizal fungi. We manipulated the isolation of uncolonized host seedlings across a natural landscape and used a range of molecular techniques to measure the dispersal rates of ectomycorrhizal propagules and host colonization. Some species were prolific dispersers, producing annual spore loads on the order of trillions of spores per km(2). However, fungal propagules reaching host seedlings decreased rapidly with increasing distance from potential spore sources, causing a concomitant reduction in ectomycorrhizal species richness, host colonization and host biomass. There were also strong differences in dispersal ability across species, which correlated well with the predictable composition of ectomycorrhizal communities associated with establishing pine forest. The use of molecular tools to measure whole community dispersal provides a direct confirmation for a key mechanism underlying island biogeography theory and has the potential to make microbial systems a model for understanding the role of dispersal in ecological theory.  相似文献   

20.
Aim To study the effects of isolation and size of small tropical islands on species assemblages of bees (superfamily Apoidea) and wasps (superfamily Vespoidea). Location Twenty islands in the Kepulauan Seribu Archipelago off the coast of west Java, Indonesia. The size of surveyed islands ranges between 0.75 and 41.32 ha; their distance from the coast of Java varies between 3 and 62 km. Methods Field work was conducted from February to May 2005. Bees and wasps were caught with a sweep net during sampling units of 15 min, continuing until four consecutive samples revealed no new species. Total species richness was quantified by the estimators Chao 2, first‐order jackknife and Michaelis–Menten. The software binmatnest was used to test for nestedness of species assemblages. Similarities of species composition between islands were quantified by Sørensen’s similarity index. Results Eighty‐two species were recorded on the 20 surveyed islands. Species richness declined with increasing isolation of islands from the source area, Java. Although the size of the largest island exceeded that of the smallest island by a factor of almost 60, island size only very weakly affected species richness of bees; no effect of island size was found for wasps. Mean body size of species decreased with increasing island isolation. Nestedness of island faunas was only weakly developed. Species composition of both superfamilies was affected by island isolation, but not by island size. Main conclusions While the species–isolation relationship on the very small islands of Kepulauan Seribu followed the prediction of MacArthur and Wilson’s equilibrium theory, the absence of a species–area relationship indicated a weak ‘small‐island effect’, at least in wasps. The combination of an only weakly developed pattern of nested species subsets, the shift in species compositions and the decline of mean body size with increasing island isolation from the source area indicates that biotic interactions and different species traits contribute to the shaping of communities of bees and wasps within the archipelago. The potential of biotic interactions for generating distribution patterns of species within the archipelago is also emphasized by the observed restriction of some species with apparently high dispersal abilities to outer islands.  相似文献   

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