生境片段化对千岛湖地表甲虫物种多样性的影响

在千岛湖片段化景观中选取20个陆桥岛屿和8个大陆样点,从2012年7月到2014年4月,按季度(春、夏、秋季)6次采用巴氏陷阱法收集地表甲虫,分析其物种多度、组成、多样性和季节动态,以及不同岛屿上的地表甲虫的物种多样性与岛屿面积和隔离度等岛屿空间特征的关系.结果表明: 共收集记录到地表甲虫26科101种3370头.大陆和大岛地表甲虫的物种丰富度有显著差异,且小岛地表甲虫密度显著高于大陆;大陆地表甲虫的物种组成变化较大,而岛屿上分布的地表甲虫的物种组成则变化较小.地表甲虫的物种丰富度与岛屿面积呈显著正相关,密度与隔离度呈显著正相关.夏季岛屿上地表甲虫物种丰富度高于春秋两季,岛屿与大陆样地的Shannon指数、Simpson指数和Pielou均匀度指数均在夏季最高、秋季最低.     

Terrestrial bird community patterns on the coralline islands of the Dahlak Archipelago, Red Sea, Eritrea

Aim This study aims to explain the patterns of species richness and nestedness of a terrestrial bird community in a poorly studied region. Location Twenty‐six islands in the Dahlak Archipelago, Southern Red Sea, Eritrea. Methods The islands and five mainland areas were censused in summer 1999 and winter 2001. To study the importance of island size, isolation from the mainland and inter‐island distance, I used constrained null models for the nestedness temperature calculator and a cluster analysis. Results Species richness depended on island area and isolation from the mainland. Nestedness was detected, even when passive sampling was accounted for. The nested rank of islands was correlated with area and species richness, but not with isolation. Idiosyncrasies appeared among species‐poor and species‐rich islands, and among common and rare species. Cluster analysis showed differences among species‐rich islands, close similarity among species‐poor and idiosyncratic islands, and that the compositional similarity among islands decreased with increasing inter‐island distance. Thus, faunas of species‐poor, smaller islands were more likely to be subsets of faunas of species‐rich, larger islands if the distance between the islands was short. Main conclusions Species richness and nestedness were related to island area, and nestedness also to inter‐island distances but not to isolation from the mainland. Thus, nestedness and species richness are not affected in the same way by area and distance. Moreover, idiosyncrasies may have been the outcome of species distributions among islands being influenced also by non‐nested distributions of habitats, inter–specific interactions, and differences in species distributions across the mainland. Idiosyncrasies in nested patterns may be as important as the nested pattern itself for conservation – and conservation strategies based on nestedness and strong area effects (e.g. protection of only larger islands) may fail to preserve idiosyncratic species/habitats.     

Biogeographical determinants for total and endemic species richness in a continental archipelago

We examined the relationship between plant species richness and biogeographical variables (island area, island maximum elevation, distance from nearest inhabited island, distance from nearest mainland) using a data set comprising 201 islands of the Aegean archipelago. We found that endemic species richness was strongly correlated to total species richness. Single-island endemic species richness was most strongly correlated to island maximum elevation, and then to island area, with an apparent small island effect for islands smaller than 47 km². Total species richness was most strongly correlated to island area (with no apparent small island effect), and less strongly correlated to island maximum elevation. Distance from the mainland or other inhabited islands displayed limited predictive value in our data set. The slope of the relationship between species richness and geographical factors (island area, elevation, distance from island/mainland) was steeper for endemic species richness than for total richness. Finally, the different scales of endemicity (single-island endemics, island group endemics and Aegean regional endemics) displayed similar qualitative trends and only differed quantitatively. Thus, we conclude that different biogeographical factors act as drivers for total species richness than for endemic species richness.     

Drivers of diversity in Macaronesian spiders and the role of species extinctions

Aim To identify the biogeographical factors underlying spider species richness in the Macaronesian region and assess the importance of species extinctions in shaping the current diversity. Location The European archipelagos of Macaronesia with an emphasis on the Azores and Canary Islands. Methods Seven variables were tested as predictors of single‐island endemics (SIE), archipelago endemics and indigenous spider species richness in the Azores, Canary Islands and Macaronesia as a whole: island area; geological age; maximum elevation; distance from mainland; distance from the closest island; distance from an older island; and natural forest area remaining per island – a measure of deforestation (the latter only in the Azores). Different mathematical formulations of the general dynamic model of oceanic island biogeography (GDM) were also tested. Results Island area and the proportion of remaining natural forest were the best predictors of species richness in the Azores. In the Canary Islands, area alone did not explain the richness of spiders. However, a hump‐shaped relationship between richness and time was apparent in these islands. The island richness in Macaronesia was correlated with island area, geological age, maximum elevation and distance to mainland. Main conclusions In Macaronesia as a whole, area, island age, the large distance that separates the Azores from the mainland, and the recent disappearance of native habitats with subsequent unrecorded extinctions seem to be the most probable explanations for the current observed richness. In the Canary Islands, the GDM model is strongly supported by many genera that radiated early, reached a peak at intermediate island ages, and have gone extinct on older, eroded islands. In the Azores, the unrecorded extinctions of many species in the oldest, most disturbed islands seem to be one of the main drivers of the current richness patterns. Spiders, the most important terrestrial predators on these islands, may be acting as early indicators for the future disappearance of other insular taxa.     

Cantharidin world on islands: Species diversity of canthariphilous arthropods in the Izu–Ogasawara Arc

Cantharidin is a defense chemical produced by only two beetle families, Meloidae and Oedemeridae. This compound also functions as an attractant for certain groups of arthropods (canthariphilous arthropods). Cantharidin‐producing beetles and canthariphilous arthropods build a cantharidin‐mediated community, called cantharidin world. In this study, to clarify the canthariphilous arthropod community formed on small islands, the arthropods were collected by cantharidin‐baited traps and compared among the mainland, peninsula and 11 small islands of the Izu–Ogasawara Arc, Japan. Control traps without cantharidin were also used to collect non‐canthariphilous arthropods. Cantharidin‐producing beetles were distributed across the study area, but the number of species was reduced on islands. We identified 10 species of five families as canthariphilous arthropods in this study area: two species of Anthicidae and Pyrochroidae (Coleoptera), four species of Ceratopogonidae (Diptera), three species of Formicidae (Hymenoptera) and one species of Podoctidae (Opiliones). The number of canthariphilous species was not correlated with island size (area) but was negatively correlated with the distance of island from the mainland. These tendencies were also seen in non‐canthariphilous fauna. The canthariphilous arthropods increased with the number of cantharidin‐producing beetle species, but the non‐canthariphilous fauna did not. Thus, the cantharidin world on islands seems to be affected not only by the geographic characteristics of islands, particularly the degree of isolation from the mainland, but also by the diversity of cantharidin‐producing members within it.     

Factors influencing vascular plant diversity on 22 islands off the coast of eastern North America

Aim The influence of physiographic and historical factors on species richness of native and non‐native vascular plants on 22 coastal islands was examined. Location Islands off the coast of north‐eastern USA and south‐eastern Canada between 41° and 45° N latitude were studied. Island size ranges from 3 to 26,668 ha. All islands were deglaciated between 15,000 and 11,000 yr bp ; all but the four New Brunswick islands were attached to the mainland until rising sea level isolated them between 14,000 and 3800 yr bp . Methods Island species richness was determined from floras compiled or revised since 1969. Simple and multiple regression and rank correlation analysis were employed to assess the relative influence of independent variables on species richness. Potential predictors included island area, latitude, elevation, distance from the mainland, distance from the nearest larger island, number of soil types, years since isolation, years since deglaciation, and human population density. Results Native vascular plant species richness for the 22 islands in this study is influenced most strongly by island area, latitude, and distance from the nearest larger island; richness increases with island area, but decreases with latitude and distance from the nearest larger island as hypothesized. That a similar model employing distance from the mainland does not meet the critical value of P confirms the importance of the stepping‐stone effect. Habitat diversity as measured by number of soil types is also an important predictor of native plant species richness, but at least half of its influence can be attributed to island area, with which it is correlated. Two historical factors, years since deglaciation and years since isolation, also appear to be highly correlated with native species richness, but their influence cannot be separated from that of latitude for the present sample size. Non‐native vascular plant species richness is influenced primarily by island area and present‐day human population density, although human population density may be a surrogate for the cumulative effect of several centuries of anthropogenic impacts related to agriculture, hunting, fishing, whaling, tourism, and residential development. Very high densities of ground‐nesting pelagic birds may account for the high percentage of non‐native species on several small northern islands. Main conclusions Many of the principles of island biogeography that have been applied to oceanic islands apply equally to the 22 islands in this study. Native vascular plant species richness for these islands is strongly influenced by physiographic factors. Influence of two historical factors, years since deglaciation and years since isolation, cannot be assessed with the present sample size. Non‐native vascular plant species richness is influenced by island area as well as by human population density; human population density may be a surrogate for other anthropogenic impacts.     

Quantifying island isolation – insights from global patterns of insular plant species richness

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.     

Colonization of Sphagnum on land uplift islands in the Baltic Sea: time, area, distance and life history

Aim To test whether species richness of Sphagnum mosses on islands in a land uplift archipelago is related to island age, area or connectivity, and whether the frequency of different species can be predicted by their life history and autecology. Location The northern Stockholm archipelago in the Baltic Sea, east‐central Sweden, with a current land uplift rate of 4.4 mm year^?1. Methods We sampled 17 islands differing in area (0.55–55 ha), height (3.6–18 m, representing c. 800–4000 years of age) and distance from mainland (1.6–41 km). For each Sphagnum patch we measured area, height above sea level, horizontal distance from the shore and shading from vascular plants. Factors affecting island species richness, species frequency and habitats on the islands were tested by stepwise regressions. Species frequency was tested on nine life history and autecological variables, including estimated abundance and spore output on the mainland, habitat preference and distribution. Results We recorded 500 patches of 19 Sphagnum species, distributed in 83 rock pools on 14 islands. Island species richness correlated positively with island area and with degree of shelter by surrounding islands, while distance from the mainland, connectivity, height or age did not add to the model. Species frequency (number of colonized islands and rock pools) was mainly predicted by spore output on the mainland and by habitat preference (swamp forest species were more frequent than others), while spore size, for example, did not add to the model. Species differed in mean height above and horizontal distance from the shore, area of occupied rock pools and in the degree of shading of patches. The mean horizontal distance from the shore and the area of occupied rock pools correlated positively with the normal growth position above the water table among species. Spore capsules were found in only 2% of patches, mostly in the bisexual Sphagnum fimbriatum. Main conclusions The presence of Sphagnum in the Stockholm archipelago seems to be governed by regional spore production and habitat demands. Sphagnum does not appear to be dispersal limited at distances up to 40 km and time spans of centuries. Species with a high regional spore output have had a higher colonization rate, which, together with the rarity of spore capsules on the islands, indicate the mainland as a source for colonization rather than dispersal among islands. Swamp forest species seem more tolerant to the island conditions (summer droughts and some salt spray) than open mire species. The different distances from the sea occupied by the species indicate a slow, continuous succession and species replacement towards the island interior as islands are being uplifted and thus expand in area. This partly explains why larger islands harbour more species. Our results thus support some of the island biogeographical theories related to the species–area relationship.     

Evolutionary assembly of island faunas reverses the classic island–mainland richness difference in Anolis lizards

Aim Islands are widely considered to be species depauperate relative to mainlands but, somewhat paradoxically, are also host to many striking adaptive radiations. Here, focusing on Anolis lizards, we investigate if cladogenetic processes can reconcile these observations by determining if in situ speciation can reduce, or even reverse, the classical island–mainland richness discrepancy. Location Caribbean islands and the Neotropical mainland. Methods We constructed range maps for 203 mainland anoles from museum records and evaluated whether geographical area could account for differences in species richness between island and mainland anole faunas. We compared the island species–area relationship with total mainland anole diversity and with the richness of island‐sized mainland areas. We evaluated the role of climate in the observed differences by using Bayesian model averaging to predict island richness based on the mainland climate–richness relationship. Lastly, we used a published phylogeny and stochastic mapping of ancestral states to determine if speciation rate was greater on islands, after accounting for differences in geographical area. Results Islands dominated by in situ speciation had, on average, significantly more species than similarly sized mainland regions, but islands where in situ speciation has not occurred were species depauperate relative to mainland areas. Results were similar at the scale of the entire mainland, although marginally non‐significant. These findings held even after accounting for climate. Speciation has not been faster on islands; instead, when extinction was assumed to be low, speciation rate varied consistently with geographical area. When extinction was high, there was some evidence that mainland speciation was faster than expected based on area. Main conclusions Our results indicate that evolutionary assembly of island faunas can reverse the general pattern of reduced species richness on islands relative to mainlands.     

Increase of island endemism with altitude – speciation processes on oceanic islands

Understanding speciation on oceanic islands is a major topic in current research on island biogeography. Within this context, it is not an easy task to differentiate between the influence of elevation as an indicator for habitat diversity and island age as an indicator for the time available for diversification. One reason for this is that erosion processes reduce the elevation of islands over time. In addition, the geographic distance to source ecosystems might differ among habitats, which could lead to habitat‐specific reduction of species immigration, niche occupation and diversification. We used the percentage of single island endemic species (pSIE) in five different zonal ecosystems (distributed in altitude) on the Canary Islands as an indicator for diversification. We tested whether diversification increases with altitude due to a greater ecological isolation of high elevation ecosystems on oceanic islands under the assumption of a low elevation source region on the mainland. In addition we tested whether the ‘hump‐shaped’ (unimodal) relationship between pSIE and island age as well as the linear relationship between species richness and pSIE is consistent across spatial scales. We also analyse a potential influence of island area and habitat area. We found that pSIE increases with elevation. The relations between species richness as well as age with pSIE are consistent across scales. We conclude that high elevation ecosystems are ecologically isolated. Surprisingly, the altitudinal belt with the strongest human influences has the highest values of pSIE. We successfully transfer the ‘general dynamic theory of island biogeography’ to the ecosystem scale, which provides multiple opportunities for future studies. With this approach we find that the effects of elevation on diversification can be separated from those of island age.     

Dung beetle assemblages on tropical land‐bridge islands: small island effect and vulnerable species

Aim Using dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) in a tropical land‐bridge island system, we test for the small island effect (SIE) in the species–area relationship and evaluate its effects on species richness and community composition. We also examine the determinants of species richness across island size and investigate the traits of dung beetle species in relation to their local extinction vulnerability following forest fragmentation. Location Lake Kenyir, a hydroelectric reservoir in north‐eastern Peninsular Malaysia. Methods We sampled dung beetles using human dung baited pitfall traps on 24 land‐bridge islands and three mainland sites. We used regression tree analyses to test for the SIE, as well as species traits related to local rarity, as an indication of extinction vulnerability. We employed generalized linear models (GLMs) to examine determinants for species richness at different scales and compared the results with those from conventional linear and breakpoint regressions. Community analyses included non‐metric multidimensional scaling, partial Mantel tests, nestedness analysis and abundance spectra. Results Regression tree analysis revealed an area threshold at 35.8 ha indicating an SIE. Tree basal area was the most important predictor of species richness on small islands (<35.8 ha). Results from GLMs supported these findings, with isolation and edge index also being important for small islands. The SIE also manifested in patterns of dung beetle community composition where communities on small islands (<35.8 ha) departed from those on the mainland and larger islands, and were highly variable with no significant nestedness, probably as a result of unexpected species occurrences on several small islands. The communities exhibited a low degree of spatial autocorrelation, suggesting that dispersal limitation plays a part in structuring dung beetle assemblages. Species with lower baseline density and an inability to forage on the forest edge were found to be rarer among sites and hence more prone to local extinction. Main conclusions We highlight the stochastic nature of dung beetle community composition on small islands and argue that this results in reduced ecosystem functionality. A better understanding of the minimum fragment size required for retaining functional ecological communities will be important for effective conservation management and the maintenance of tropical forest ecosystem stability.     

Island biogeography extends to small-scale habitats: low competitor density and richness on islands may drive trait variation in nonnative plants

Previous island biogeography studies have quantified species richness on the scale of entire islands rather than smaller scales relevant to plant-to-plant competitive interactions. Further, they have not accounted for density compensation. Using mainland and island sites along the New England coast, we asked two questions. First, are both richness and density lower in small-scale habitats within islands than in similar mainland habitats? Second, do differences in competitor richness and density drive post-establishment trait variation in nonnative plant species? We used field surveys and individual-based rarefaction to estimate richness and density in 100-m² plots and demonstrated that island sites have significantly fewer species and individuals per unit area than mainland sites. We then conducted a field study in which we removed competing neighbors from nonnative plant individuals and found that when competitors were removed, individuals in low-competition environments demonstrated a lesser increase in vegetative growth but a greater increase in reproductive effort and herbivore tolerance relative to mainland individuals whose neighbors were also removed. We found that the central concept of island biogeography, i.e., that islands host fewer species than comparable mainland habitats, can be extended to smaller-scale habitats and that this difference in competitive pressure between mainland and island habitats can act as a driver of trait variation in nonnative plants.     

Effects of isolation,logging and dispersal on woody-species richness of islands

Isolation effects on species richness of woody plants were investigated in a system of islands that were created by the filling of the Clarks Hill Reservoir, Georgia. This reservoir was built between 1946–1954. Some islands were logged and cleared of woody plants prior to the filling of the reservoir and others were not logged. The presence of logged versus unlogged islands in the same system allowed us to test whether and how geographical isolation interacts with island history and species-specific dispersal properties in determining patterns of among-island variation in species number. Thirty-six years after the islands were created, logged islands had significantly fewer species of woody plants than unlogged ones. On logged islands, total number of woody species was negatively correlated with distance to the closest mainland (r=–0.95). On unlogged islands, variation in species number was very low (CV=4.9%) and was not correlated with distance to the mainland. These results indicate that the studied system as a whole has not yet reached equilibrium. However, the mean number of species on unlogged islands was very close to the intercept of the regression obtained for logged islands, suggesting that islands close to the mainland have already reached their equilibrium species richness. This conclusion is consistent with predictions of island biogeography theory. When species representing different dispersal properties were analyzed separately, statistically significant distance effects were obtained for bird-dispersed species (r=0.88) and for species with no adaptations to bird or wind dispersal (r=0.81). Wind-dispersed species did not show a decrease in species number with increasing isolation, but their relative frequency was positively and significantly correlated with distance to the mainland (r=0.94). Historical factors, as well as differences among species in dispersal properties, are important in explaining patterns of among-island variation in species number.     

Carabid beetle diversity and distribution in Boston Harbor Islands national park area (Coleoptera, Carabidae)

As part of an All Taxa Biodiversity Inventory in Boston Harbor Islands national park area, an inventory of carabid beetles on 13 islands was conducted. Intensive sampling on ten of the islands, using an assortment of passive traps and limited hand collecting, resulted in the capture of 6,194 specimens, comprising 128 species. Among these species were seven new state records for Massachusetts (Acupalpus nanellus,Amara aulica,Amara bifrons, Apenes lucidulus, Bradycellus tantillus, Harpalus rubripes and Laemostenus terricola terricola-the last also a new country record; in passing we report also new state records for Harpalus rubripes from New York and Pennsylvania, Amara ovata from Pennsylvania, and the first mainland New York records for Asaphidion curtum). For most islands, there was a clear relationship between species richness and island area. Two islands, however, Calf and Grape, had far more species than their relatively small size would predict. Freshwater marshes on these islands, along with a suite of hygrophilous species, suggested that habitat diversity plays an important role in island species richness. Introduced species (18) comprised 14.0% of the total observed species richness, compared to 5.5% (17 out of 306 species) documented for Rhode Island. We surmise that the higher proportion of introduced species on the islands is, in part, due to a higher proportion of disturbed and open habitats as well as high rates of human traffic. We predict that more active sampling in specialized habitats would bring the total carabid fauna of the Boston Harbor Islands closer to that of Rhode Island or eastern Massachusetts in richness and composition; however, isolation, human disturbance and traffic, and limited habitat diversity all contribute to reducing the species pool on the islands relative to that on the mainland.     

Mediterranean island biogeography: Analysis of fern species distribution in the system of islets around Sicily

Abstract

The aim of this article was to test the way in which geographical factors influence island floras in the Mediterranean basin, using ferns as target organisms, and the islands surrounding Sicily as location. A matrix with presence/absence data concerning fern taxa in the 16 islands studied was compiled. Cluster analysis, principal co-ordinates analysis (PCoA), principal components analysis (PCA) and a Bayesian analysis were performed. For each island, the total number of fern taxa was regressed against three factors: island area, island elevation and isolation. All the analyses pointed to affinities between islands according to their different geological composition, independently from their geographic position. A clear positive island species/area relationship (ISAR) was shown only for the volcanic islands. The island species/(area×elevation) relationship (ISAER), on the contrary, was unsatisfactory. The main features of interest are the following: (1) the clear division of the islands into two groups, volcanic vs. sedimentary; (2) the floristic richness of the volcanic compared to sedimentary islands and (3) the uniqueness of the pteridophyte flora of Pantelleria. This seems to demonstrate that the lower number of taxa in the islands farthest away from the “mainland” (Sicily, Tunisia) is not due to isolation, but due to another factor, probably habitat availability.     

Not as the crow flies: assessing effective isolation for island biogeographical analysis

Aim To evaluate the role of island isolation in explaining the distribution of vascular plant species in a dense freshwater archipelago, specifically comparing conventional measures of island isolation with landscape measures of island isolation. Location Data were collected from 35 islands within Massasauga Provincial Park on the eastern shores of the Georgian Bay, Ontario, Canada. Methods Sampled islands were located using stratified random selection based on location and size variation. The number of species was recorded along stratified random transects. Island isolation variables included distance to the mainland, distance to the nearest island, largest gap in a stepping‐stone sequence, distance to the closest upwind point of land, and a landscape measure of island isolation. The landscape measure of isolation was quantified as the percentage of the land area within 100, 250, 500, 1000, 1500 and 2000 m of each island’s perimeter. The isolation variables were calculated within a geographical information system (GIS). Dependent variables in the regression analyses included species richness, the logarithm of species richness and residuals of the species–area relationship. Independent variables included island isolation variables and their logarithmic transformations. Results Isolation plays a role, albeit small, in explaining species richness in the study area. In the regression analyses, the landscape measure of isolation provided a better fit than conventional measures of island isolation. Islands with less land than water within a 250‐m buffer were more effectively isolated and had fewer species present than islands surrounded by a greater proportion of water. Main conclusions Consistent with the species–isolation relationship, fewer species were present on more isolated islands within the Massasauga study area, as elucidated using a series of island buffers in a GIS. Applying a landscape measure of isolation to similar dense, freshwater archipelagos may elucidate species–isolation patterns not evident through conventional, straight‐line distance measurements of island isolation. The low value of the regression coefficients as well as the isolation history and high density of the Massasauga islands suggests caution in extending the results, especially to dissimilar archipelagos.     

Insular black files (Diptera: Simuliidae) of North America: tests of colonization hypotheses

Aim To understand factors that facilitate insular colonization by black flies, we tested six hypotheses related to life‐history traits, phylogeny, symbiotes, island area, and distance from source areas. Location Four northern islands, all within 150 km of the North American mainland, were included in the study: Isle Royale, Magdalen Islands, Prince Edward Island, and Queen Charlotte Islands. Methods Immature black flies and their symbiotes were surveyed in streams on the Magdalen Islands, and the results combined with data from similar surveys on Isle Royale, Prince Edward Island, and the Queen Charlotte Islands. Black flies were analysed chromosomally to ensure that all sibling species were revealed. Tests of independence were used to examine the frequency of life‐history traits and generic representation of black flies on islands vs. source areas. Results A total of 13–20 species was found on each of the islands, but no species was unique to any of the islands. The simuliid faunas of the islands reflected the composition of their source areas in aspects of voltinism (univoltine vs. multivoltine), blood feeding (ornithophily vs. mammalophily), and phylogeny (genus Simulium vs. other genera). Five symbiotic species were found on the most distant island group, the Magdalen Islands, supporting the hypothesis that obligate symbiotes are effectively transported to near‐mainland islands. An inverse relationship existed between the number of species per island and distance from the source. The Queen Charlotte Islands did not conform to the species–area relationship. Main conclusions The lack of precinctive insular species and an absence of life‐history and phylogenetic characteristics related to the presence of black flies on these islands argue for gene flow and dispersal capabilities of black flies over open waters, possibly aided by winds. However, the high frequency of precinctive species on islands 500 km or more from the nearest mainland indicates that at some distance beyond 100 km, open water provides a significant barrier to colonization and gene exchange. An inverse relationship between number of species and distance from the source suggests that as long as suitable habitat is present, distance plays an important role in colonization. Failure of the Queen Charlotte Islands to conform to an area–richness trend suggests that not all resident species have been found.     

Testing island biogeography theory with visitation rates of birds to British islands

Aim We consider three hypotheses – MacArthur and Wilson’s island biogeography theory (IBT), Lack’s habitat diversity idea and the ‘target effect’– that explain the pattern of decreased species richness on small and distant islands. Location We evaluate these hypotheses using a detailed dataset on the occurrence and abundance of terrestrial birds on nine islands off the coast of Britain and the Republic of Ireland. Methods  Unlike previous studies, we compile data on species that visit the islands, rather than just those that breed on them. We divided the species into five mutually exclusive categories based upon their migratory status and where they regularly breed: British residents, summer visitors to Britain, winter visitors to Britain, and vagrants from Europe or beyond Europe. For each species group on each island we calculated the average number of species visiting each year. We then regressed the average number of species against island area and distance to the mainland (all variables were log‐transformed). We also compared the average number of species visiting each island with the average number of species breeding on each island. Results  Average number of visiting British residents decreased significantly with increasing island distance, but showed no relationship with island area. There was no significant relationship between island area or island distance and average number of summer or winter visitors. European and non‐European vagrants likewise showed no relationship between numbers of species visiting and island distance. However, the relationship between island area and number of visiting species was significant for both these categories; as island area increases so too does the number of visiting species. Main conclusions  As predicted by IBT, there were fewer visiting species on more distant islands. There were substantially more visitors to each island than breeding species, supporting Lack’s argument that lower bird richness is not a result of varying immigration rates (as predicted by IBT) but rather a result of some other island property, e.g. fewer resources. Birds make a decision to either leave an island or stay and breed. The target effect was also clearly demonstrated by the increase in European and non‐European breeders with increasing island size.     

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.     

Assemblage-level responses of phyllostomid bats to tropical forest fragmentation: land-bridge islands as a model system

Aim Working within a system of high structural contrast between fragments and the surrounding matrix, we assessed patterns of species loss and changes in species composition of phyllostomid bats on artificial land‐bridge islands relative to mainland assemblages, and evaluated the responses of bats to forest edges. We further examined the relative influence of local‐scale characteristics (e.g. vegetation structure, island area) versus landscape attributes (e.g. forest cover, patch density) and the importance of spatial scale in determining phyllostomid species richness and composition on islands. Location Islands in Gatún Lake and adjacent mainland peninsulas in the Barro Colorado Nature Monument, Panama. Methods Bats were sampled over a 2‐year period on 11 islands as well as at forest‐edge and interior sites on adjacent mainland, resulting in > 8400 captures. Results The islands harboured a less diverse and structurally simplified phyllostomid bat fauna. Islands far from the mainland were especially species‐poor. This decline in species richness was associated with compositional shifts towards assemblages strongly dominated by frugivores with good dispersal abilities. Members of other ensembles, most importantly gleaning animalivores, were much less common or absent. Although overall species composition was not significantly altered, species richness at continuous forest‐edge sites was significantly lower compared with that at interior sites. Distance from the mainland and amount of forest cover in the landscape were the best predictors of species richness and assemblage composition. Responses were scale‐dependent. At the local scale, species richness was independent of island area but was correlated positively with distance from the mainland. In contrast, area effects became more important at larger spatial scales, suggesting that many species use multiple fragments. Main conclusions Our results underline the conservation value of small habitat remnants, which, even when embedded in a hostile matrix, can support a relatively diverse bat fauna, provided that there is a low degree of patch isolation and spatial proximity to larger tracts of continuous forest. Although the results at the assemblage level were inconclusive, we demonstrate that certain bat species and ensembles, particularly gleaning animalivores, exhibit high edge‐sensitivity. Our results point to habitat loss rather than changes in landscape configuration as the main process after isolation underlying phyllostomid bat responses, suggesting that conservation efforts should focus on habitat preservation instead of trying to minimize fragmentation per se at the expense of habitat amount.