Spatial processes that maintain biodiversity in plant communities

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.     

Sequential colonization and diversification of Galapágos endemic land snail genus Bulimulus (Gastropoda, Stylommatophora)

Species richness on island or islandlike systems is a function of colonization, within-island speciation, and extinction. Here we evaluate the relative importance of the first two of these processes as a function of the biogeographical and ecological attributes of islands using the Galápagos endemic land snails of the genus Bulimulus, the most species-rich radiation of these islands. Species in this clade have colonized almost all major islands and are found in five of the six described vegetation zones. We use molecular phylogenetics (based on COI and ITS 1 sequence data) to infer the diversification patterns of extant species of Bulimulus, and multiple regression to investigate the causes of variation among islands in species richness. Maximum-likelihood, Bayesian, and maximum-parsimony analyses yield well-resolved trees with similar topologies. The phylogeny obtained supports the progression rule hypothesis, with species found on older emerged islands connecting at deeper nodes. For all but two island species assemblages we find support for only one or two colonization events, indicating that within-island speciation has an important role in the formation of species on these islands. Even though speciation through colonization is not common, island insularity (distance to nearest major island) is a significant predictor of species richness resulting from interisland colonization alone. However, island insularity has no effect on the overall bulimulid species richness per island. Habitat diversity (measured as plant species diversity), island elevation, and island area, all of which are indirect measures of niche space, are strong predictors of overall bulimulid land snail species richness. Island age is also an important independent predictor of overall species richness, with older islands harboring more species than younger islands. Taken together, our results demonstrate that the diversification of Galápagos bulimulid land snails has been driven by a combination of geographic factors (island age, size, and location), which affect colonization patterns, and ecological factors, such as plant species diversity, that foster within-island speciation.     

Genetic and phylogenetic consequences of island biogeography

Abstract.— Island biogeography theory predicts that the number of species on an island should increase with island size and decrease with island distance to the mainland. These predictions are generally well supported in comparative and experimental studies. These ecological, equilibrium predictions arise as a result of colonization and extinction processes. Because colonization and extinction are also important processes in evolution, we develop methods to test evolutionary predictions of island biogeography. We derive a population genetic model of island biogeography that incorporates island colonization, migration of individuals from the mainland, and extinction of island populations. The model provides a means of estimating the rates of migration and extinction from population genetic data. This model predicts that within an island population the distribution of genetic divergences with respect to the mainland source population should be bimodal, with much of the divergence dating to the colonization event. Across islands, this model predicts that populations on large islands should be on average more genetically divergent from mainland source populations than those on small islands. Likewise, populations on distant islands should be more divergent than those on close islands. Published observations of a larger proportion of endemic species on large and distant islands support these predictions.     

Local factors determine plant community structure on closely neighbored islands

Despite the recent popularity of the metacommunity concept, ecologists have not evaluated the applicability of different metacommunity frameworks to insular organisms. We surveyed 50 closely spaced islands in the Thousand-Island Lake of China to examine the role of local (environmental) and regional (dispersal) factors in structuring woody plant assemblages (tree and shrub species) on these islands. By partitioning the variation in plant community structure into local and regional causes, we showed that local environmental conditions, specifically island morphometric characteristics, accounted for the majority of the variation in plant community structure among the studied islands. Spatial variables, representing the potential importance of species dispersal, explained little variation. We conclude that one metacommunity framework-species sorting-best characterizes these plant communities. This result reinforces the idea that the traditional approach of emphasizing the local perspective when studying ecological communities continues to hold its value.     

Patch quality and context, but not patch number, drive multi-scale colonization dynamics in experimental aquatic landscapes

Colonization and extinction are primary drivers of local population dynamics, community structure, and spatial patterns of biological diversity. Existing paradigms of island biogeography, metapopulation biology, and metacommunity ecology, as well as habitat management and conservation biology based on those paradigms, emphasize patch size, number, and isolation as primary characteristics influencing colonization and extinction. Habitat selection theory suggests that patch quality could rival size, number, and isolation in determining rates of colonization and resulting community structure. We used naturally colonized experimental landscapes to address four issues: (a) how do colonizing aquatic beetles respond to variation in patch number, (b) how do they respond to variation in patch quality, (c) does patch context affect colonization dynamics, and (d) at what spatial scales do beetles respond to habitat variation? Increasing patch number had no effect on per patch colonization rates, while patch quality and context were critical in determining colonization rates and resulting patterns of abundance and species richness at multiple spatial scales. We graphically illustrate how variation in immigration rates driven by perceived predation risk (habitat quality) can further modify dynamics of the equilibrium theory of island biogeography beyond predator-driven effects on extinction rates. Our data support the importance of patch quality and context as primary determinants of colonization rate, occupancy, abundance, and resulting patterns of species richness, and reinforce the idea that management of metapopulations for species preservation, and metacommunities for local and regional diversity, should incorporate habitat quality into the predictive equation.     

Bat metacommunity structure on Caribbean islands and the role of endemics

Aim We evaluate characteristics of species ranges (i.e. coherence, species turnover and range boundary clumping) to determine the structure of bat metacommunities and metaensembles from Caribbean islands. We evaluate the effects of endemic species on that structure, and quantify associations between island characteristics and latent environmental gradients that structure these metacommunities and metaensembles. Location Sixty‐five Caribbean islands throughout the Bahamas, Greater Antilles and Lesser Antilles. Methods Metacommunity structure is an emergent property of a set of ecological communities at different sites defined by species distributions across geographic or environmental gradients. We analysed elements of metacommunity structure (coherence, range turnover and range boundary clumping) to determine the best‐fit pattern for metacommunities from all Caribbean islands, as well as from the Bahamas, the Greater Antilles and the Lesser Antilles separately. For each island group, analyses were conducted for all bats and for each of two broadly defined guilds (i.e. carnivores and herbivores). In addition, analyses were conducted for all species and for a subset in which endemic species were removed from the fauna. Spearman rank correlations identified island characteristics (area, elevation, latitude, longitude) that were associated significantly with island scores for ordination axes based on reciprocal averaging. Results Metacommunity structure for all bats and for carnivores was similar for each island group, with Clementsian distributions (i.e. discrete communities with groups of species replacing other groups of species along the gradient) for all islands, the Bahamas and the Lesser Antilles, but with nested distributions for the Greater Antilles. Herbivore distributions were random for the Bahamas, but were Clementsian for all other island groups. Removal of endemic species affected the best‐fit model of metacommunity structure in only 3 of 12 cases. In general, ordination scores for islands were correlated with longitude or latitude, but not with island area or elevation. Main conclusions Characteristics of bat species ranges and associated metacommunity structure were primarily dependent on the number and geographic arrangement of primary sources of colonization, and not on interspecific interactions, species‐specific levels of environmental tolerance, or the physical characteristics of islands. Endemic species did not greatly affect metacommunity structure in Caribbean bats.     

A neglected facet of island biogeography: The role of internal spatial dynamics in area effects

The classical theory of island biogeography has as its basic variable the presence or absence of species on entire islands, and as its basic processes colonization and extinction rates on entire islands as functions of island area, distance, and so forth. Yet for many organisms with limited dispersal abilities, it may be more reasonable to consider larger islands as comprised of an ensemble of local populations coupled by within-island dispersal. Conceptual arguments and a simple patch occupancy model are used to examine the potential relevance of such internal spatial dynamics in explaining area effects, expressed via the probability that a species is present per unit area as a function of total island area. The model suggests that strong area effects depend on a rather fine balance between local colonization and extinction rates. A fruitful direction of future research should be the application of patch dynamic theory to classic island biogeographic questions and systems.     

Temporal dynamics and nestedness of an oceanic island bird fauna

Aim To examine temporal variation in nestedness and whether nestedness patterns predict colonization, extinction and turnover across islands and species. Location Dahlak Archipelago, Red Sea. Method The distributions of land birds on 17 islands were recorded in two periods 30 years apart. Species and islands were reordered in the Nestedness Temperature Calculator, software for assessing degrees of nestedness in communities. The occupancy probability of each cell, i.e. species–island combinations, was calculated in the nested matrix and an extinction curve (boundary line) was specified. We tested whether historical and current nested ranks of species and islands were correlated, whether there was a relationship between occupancy probability (based on the historical data) and number of extinctions or colonizations (regression analyses) and whether the boundary line could predict extinctions and colonizations (chi‐square analyses). Results Historical and current nested ranks of islands and species were correlated but changes in occupancy patterns were common, particularly among bird species with intermediate incidence. Extinction and turnover of species were higher for small than large islands, and colonization was negatively related to isolation. As expected, colonizations were more frequent above than below the boundary line. Probability of extinction was highest at intermediate occupancy probability, giving a quadratic relationship between extinction and occupancy probability. Species turnover was related to the historical nested ranks of islands. Colonization was related negatively while extinction and occupancy turnover were related quadratically to historical nested ranks of species. Main conclusions Some patterns of the temporal dynamics agreed with expectations from nested patterns. However, the accuracy of the predictions may be confounded by regional dynamics and distributions of idiosyncratic, resource‐limited species. It is therefore necessary to combine nestedness analysis with adequate knowledge of the causal factors and ecology of targeted species to gain insight into the temporal dynamics of assemblages and for nestedness analyses to be helpful in conservation planning.     

Turnover of passerine birds on islands in the Aegean Sea (Greece)

Aim We wish to determine the effect of migratory status on turnover rates in island birds. Because turnover is influenced by factors other than migratory status, we also considered the influence of body size and physical characteristics of the islands inhabited on the probabilities of extinction and immigration. Location The Mediterranean islands of Delos, Astypalea, Paros, Naxos and Lesvos in the Aegean Sea, Greece. Methods The passerine birds of these islands were surveyed between 1954 and 1961 by G.E. Watson, and were resurveyed between 1988 and 1992. The effects of migratory status and body size on the probabilities of extinction and immigration were examined by G‐tests of linear trend in proportion, and analysis of variance, respectively. A combined analysis of migratory status, body size and physical characteristics of the islands was carried out using logistic regressions of the probabilities of extinction and immigration on these factors. Results Species number on each island changed little between surveys, with no island's species number changing by more than one species. Twelve population extinctions and 11 immigrations were recorded. The smallest island, Delos (6 km²), had the highest annualized relative turnover rate (1.08), while the four larger islands (96–1614 km²) had lower and mutually similar rates (0.21–0.27). Populations on higher elevation islands were less likely to go extinct. There is no evidence for an effect of body size on the probabilities of extinction or immigration. Migratory status affected extinction and immigration probabilities differently: migratory species were more likely to immigrate, but less likely to go extinct. Main conclusions The position of the Aegean islands along a major north–south flyway may account for the observed effects of migratory status. The annual passage of large numbers of migrants may, via the rescue effect, decrease the chances of extinction, while at the same time increasing the chances of colonization of unoccupied islands. The likelihood of both extinction and immigration involves a complex interaction between life‐history traits and island characteristics. The effects of migratory status will depend not only on consideration of vagility, vulnerability and stochasticity identified by previous authors, but also upon the location of the islands in relationship to migratory pathways.     

Metacommunity,mainland‐island system or island communities? Assessing the regional dynamics of plant communities in a fragmented landscape

Understanding the regional dynamics of plant communities is crucial for predicting the response of plant diversity to habitat fragmentation. However, for fragmented landscapes the importance of regional processes, such as seed dispersal among isolated habitat patches, has been controversially debated. Due to the stochasticity and rarity of among‐patch dispersal and colonization events, we still lack a quantitative understanding of the consequences of these processes at the landscape‐scale. In this study, we used extensive field data from a fragmented, semi‐arid landscape in Israel to parameterize a multi‐species incidence‐function model. This model simulates species occupancy pattern based on patch areas and habitat configuration and explicitly considers the locations and the shapes of habitat patches for the derivation of patch connectivity. We implemented an approximate Bayesian computation approach for parameter inference and uncertainty assessment. We tested which of the three types of regional dynamics – the metacommunity, the mainland‐island, or the island communities type – best represents the community dynamics in the study area and applied the simulation model to estimate the extinction debt in the investigated landscape. We found that the regional dynamics in the patch‐matrix study landscape is best represented as a system of highly isolated ‘island’ communities with low rates of propagule exchange among habitat patches and consequently low colonization rates in local communities. Accordingly, the extinction rates in the local communities are the main drivers of community dynamics. Our findings indicate that the landscape carries a significant extinction debt and in model projections 33–60% of all species went extinct within 1000 yr. Our study demonstrates that the combination of dynamic simulation models with field data provides a promising approach for understanding regional community dynamics and for projecting community responses to habitat fragmentation. The approach bears the potential for efficient tests of conservation activities aimed at mitigating future losses of biodiversity.     

Founder events as determinants of within-island and among-island genetic structure of Daphnia metapopulations

The genetic structure of metapopulations offers insights into the genetic consequences of local extinction and recolonization. We studied allozyme variation in rock pool metapopulations of two species of waterfleas (Daphnia) with the aim to understand how these dynamics influence genetic differentiation. We screened 138 populations of D. magna and 65 populations of D. longispina from an area in the archipelago of southern Finland. The pools from which they were sampled are separated by distances between 1.5 and 4710 m and located on a total of 38 islands. The genetic population structure of the two species was strikingly similar, consistent with their similar metapopulation ecology. The mean F(PT) value (differentiation among pools with respect to the total metapopulation) was 0.55 and a hierarchical analysis showed that genetic differentiation was strong (>0.25) among pools within islands as well as among whole islands. Within islands, pairwise genetic differentiation increased with geographic distance, indicating isolation by distance due to spatially limited dispersal. Previous studies have shown strong founder events occurring during colonization in our metapopulation. We suggest that the genetic population structure in the studied metapopulations is largely explained by three consequences of these founder events: (i) strong drift during colonization, (ii) local inbreeding, which results in hybrid vigour and increased effective migration rates after subsequent immigration, and (iii) effects of selection through hitchhiking of neutral genes with linked loci under selection.     

Why do some small islands lack vegetation? Evidence from long‐term introduction experiments

Classic island biogeography theory predicts that very small islands, near the extreme lower end of the species–area relationship, should support very few species. At times no species may be present, however, due to randomness in the immigration–extinction dynamics. Alternatively, a lack of vegetation on very small islands may indicate that such islands do not contain the appropriate habitat for the establishment or long‐term survival of plants, or that disturbances are too frequent or intense. These potential mechanisms were evaluated in the central Exumas, Bahamas, where surveys of 117 small islands revealed that over a third of the islands supported no terrestrial plant life. Area and exposure were significant predictors of whether a small island was vegetated or not in multiple logistic regressions. No islands naturally devoid of vegetation were colonized over a 17‐yr period, and only two naturally vegetated islands lost all vegetation. Experimental introductions of two species –Sesuvium portulacastrum and Borrichia arborescens– revealed that a number of islands naturally lacking vegetation were able to sustain introduced populations over the long term (up to 15 yr). Drought and hurricanes appeared to have reduced the establishment success and possibly long‐term survival of the introductions, although some populations survived four major hurricanes. Turnover rates of both introduced species were often an order of magnitude higher on the experimental introduction islands than on other islands in the archipelago. It appears many of the islands in this system that naturally lack vegetation may be physically capable of supporting terrestrial plant life, yet have no plants primarily due to barriers to colonization.     

Spatial patterns of coexistence of competing species in patchy habitat

The single-species spatially realistic patch occupancy metapopulation model is, in this study, extended to a metacommunity of many competing species. Competition is assumed to reduce the local carrying capacity (effective patch area), which in turn increases local extinction rates and reduces colonization rates because of smaller population sizes. Each species is described by three parameters: pre-competitive abundance (equilibrium incidence of patch occupancy, which reflects the rate of colonization in relation to extinction rate), the spatial range of migration, and competitive ability. The model ignores spatio–temporal correlations caused by interspecific interactions, because in metacommunities of unequal competitors inhabiting heterogeneous landscapes, correlations in the occurrence of species are driven more by patch heterogeneity than by competition. The model allows the calculation of multispecies equilibria in patchy habitats without simulations. In general, the number of coexisting species in the metacommunity increases with decreasing strength of competition, increasing rate of colonization, and decreasing range of migration. Habitat heterogeneity in the form of spatial variation in patch areas tends to facilitate coexistence. Poor competitors may coexist with superior competitors in the patch network if the former have higher colonization rates (competition–colonization trade-off). When migration distances are short, competition leads to spatial pattern formation: Species tend to have restricted spatial distributions in the network, but contrary to intuitive expectations, often the distributions of many species are nested. Having more dispersive species enhances both local and global diversity, whereas more local migration decreases local but increases global diversity.     

Patterns and processes in insular floras affected by hurricanes

Aim To investigate species compositions, rates of species turnover, species–area and species–distance relationships and patterns of nestedness in the floras of small Bahamian islands, by comparing two groups of islands that had been differentially affected by two hurricanes. Location Small islands occurring on either side of Great Exuma near Georgetown, Bahamas. Methods We surveyed the plant species of 44 small islands over a 5‐year period from 1998 to 2002. Hurricanes Lili and Michelle occurred in 1996 and 2001, respectively; both storms affected small islands on the more exposed south‐west side of Great Exuma to a greater degree than small islands on the more protected north‐east side. A set of 27 islands was surveyed in 1998 and 2002 to evaluate species turnover. Stepwise multiple linear regression analyses and an information‐theoretic approach (the Akaike information criterion) were used to elucidate the importance of area and distance as predictors of plant species number. We compared a piecewise linear regression model with a simple linear regression of species number against area to determine whether a small island effect existed. Nestedness patterns were evaluated by Wilcoxon two‐sample tests to analyse occurrence sequences. Results Species turnover was low in an absolute sense (overall = 0.74% year^?1), yet was over three times higher than that documented in a nearby archipelago in the absence of hurricanes. Both vegetated area and distance were important predictor variables for exposed islands but not for protected islands. Some support was found for a small island effect for the exposed islands based on a piecewise linear regression model. Both island groups revealed significant nestedness at the level of the assemblage (both P < 0.001). On exposed islands, 65–79% (depending upon the method of calculation) of all species were significantly nested, but only 47% of all species were significantly nested on protected islands. Main conclusions Overall, these insular floras seem highly resistant to hurricane‐force disturbances. Species turnover was low (< 1% year^?1) in an absolute sense, particularly in comparison with rates for other taxa. Higher degrees of nestedness and significant species–area and species–distance relationships for exposed islands indicated stronger patterns of community assembly. It is likely that disturbance is a major structuring force for the exposed islands, although the type of disturbances that mediate these patterns may not be primarily hurricane‐force storms.     

Plant species persistence and turnover on small Bahamian islands

I conducted surveys of the plant species occupying 136 small islands in the Exuma Cays and 58 small islands near Andros, Bahamas. Most species occurred on relatively few islands, and most islands contained relatively few species. Identities of the most common species differed between the two archipelagos. Comparisons with earlier surveys revealed species extinctions and immigrations. Turnover was relatively low on both a per island and a per species basis on both archipelagos, although significant spatial variation in turnover rates between archipelagos was found. Most islands experienced no turnover; islands on which turnover did occur were larger and had higher species richness. Likewise, most species did not turnover, although much variation existed in turnover rates among those that did. Experimental introductions of two species to very small islands naturally devoid of vegetation revealed that these islands could support plant life. One species survived on eight of ten islands for >9 years, including the effects of a moderate (class 2) hurricane. This hurricane caused substantial damage and loss of plant biomass, but resulted in no species extinctions on 30 small islands. Data for the small islands in this region, now spanning almost a decade, reveal that most populations are persistent over periods of years to decades, rarely going extinct or immigrating. Even moderate hurricanes seem to have little impact on species compositions.     

Sheltered from the storm? Population viability analysis of a rare endemic under periodic catastrophe regimes

Rare species are important targets for biodiversity conservation efforts because rarity often equates to small populations and increased endangerment. Rare species are prone to stochastic extinction events and may be particularly susceptible to catastrophes. Therefore, understanding how rare species respond to disturbances is critical for evaluating extinction risk and guiding conservation managers. Population viability analyses (PVAs) are essential for assessing rare species' status yet they seldom consider catastrophic events. Accordingly, we present a PVA of a rare tropical epiphyte, Lepanthes caritensis (Orchidaceae), under simulated disturbance regimes to evaluate its demographics and extinction risk. We aimed to test how demographic models incorporating catastrophes affect population viability estimates. Our goal was to better guide management of these orchids and other rare plants. Results revealed L. caritensis numbers have declined recently, but projected growth rates indicated that most subpopulations should increase in size if undisturbed. Still, projection models show that moderate catastrophes reduce growth rates, increase stochasticity in subpopulation sizes, and elevate extinction risk. Severe catastrophes had a more pronounced effect in simulations; growth rates fell below replacement level, there was greater variation in projected population sizes, and extinction risk was significantly higher. PVAs incorporating periodic catastrophes indicate that rare species may have greater extinction probabilities than standard models suggest. Thus, precautionary conservation measures should be taken in disturbance prone settings and we encourage careful monitoring after environmental catastrophes. Future rare plant PVAs should incorporate catastrophes and aim to determine if rescue and reintroduction efforts are necessary after disturbances to insure long-term population viability.     

Modelling population persistence on islands: mammal introductions in the New Zealand archipelago

Islands are likely to differ in their susceptibility to colonization or invasion due to variation in factors that affect population persistence, including island area, climatic severity and habitat modification. We tested the importance of these factors in explaining the persistence of 164 introductions of six mammal species to 85 islands in the New Zealand archipelago using survival analysis and model selection techniques. As predicted by the theory of stochastic population growth, extinction risk was the greatest in the period immediately following introduction, declining rapidly to low probability by ca 25 years. This suggests that initially small populations were at greatest risk of extinction and that populations which survived for 25 years were likely to persist subsequently for much longer. Islands in the New Zealand archipelago become colder and windier with increasing latitude, and the probability of mammal populations persisting on islands declined steeply with increasing latitude. Hence, our results suggest that climatic suitability was an important determinant of the outcome of these invasions. The form of the relationship between latitude and persistence probability differed among species, emphasizing that the outcome of colonization attempts is species-environment specific.     

The small‐island effect: fact or artefact?

Positive relationships between species richness and sampling area are perhaps the most pervasive patterns in nature. However, the shape of species–area relationships is often highly variable, for reasons that are poorly understood. One such source of variability is the "small-island effect", which refers to a decrease in the capacity of sampling area to predict species richness on small islands. Small-island effects have been attributed to a variety of processes, including spatial subsidies, habitat characteristics and ocean-born disturbances. Here, we show that small-island effects can be generated by logarithmic data transformations, which are commonly applied to both axes of species–area relationships. To overcome this problem, we derive several null models to test for non-random variability in the capacity of island area to predict species richness and apply them to data sets on island plant communities in Canada and New Zealand. Both archipelagos showed evidence for small-island effects using traditional breakpoint regression techniques on log-log axes. However, null model analyses revealed different results. The capacity of sampling area to predict species richness in the Canadian archipelago was actually lowest at intermediate island size classes. In the New Zealand archipelago, island area was similarly capable of predicting species richness across the full range of island sizes, indicating the small-island effect detected by breakpoint regression is an artifact of logarithm data transformation. Overall results show that commonly used regression techniques can generate spurious small-island effects and that alternative analytic procedures are needed to detect non-random patterns in species richness on small islands.     

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.     

THE RELEVANCE OF GENE FLOW IN METAPOPULATION DYNAMICS OF AN OCEANIC ISLAND ENDEMIC,OLEA EUROPAEA SUBSP. GUANCHICA

Theoretical and empirical studies suggest that geographical isolation and extinction–recolonization dynamics are two factors causing strong genetic structure in metapopulations, but their consequences in species with high dispersal abilities have not been tested at large scales. Here, we investigated the effect of population age structure and isolation by distance in the patterns of genetic diversity in a wind‐pollinated, zoochorous tree (Olea europaea subsp. guanchica) sporadically affected by volcanic events across the Canarian archipelago. Genetic variation was assessed at six nuclear microsatellites (nDNA) and six chloroplast fragments (cpDNA) in nine subpopulations sampled on four oceanic islands. Subpopulations occurring on more recent substrates were more differentiated than those on older substrates, but within‐subpopulation genetic diversity was not significantly different between age groups for any type of marker. Isolation‐by‐distance differentiation was observed for nDNA but not for cpDNA, in agreement with other metapopulation studies. Contrary to the general trend for island systems, between‐island differentiation was extremely low, and lower than differentiation between subpopulations on the same island. The pollen‐to‐seed ratio was close to one, two orders of magnitude lower than the average estimated for other wind‐pollinated, animal‐dispersed plants. Our results showed that population turnover and geographical isolation increased genetic differentiation relative to an island model at equilibrium, but overall genetic structure was unexpectedly weak for a species distributed among islands. This empirical study shows that extensive gene flow, particularly mediated by seeds, can ameliorate population subdivision resulting from extinction–recolonization dynamics and isolation by distance.