Diversity of butterflies on British islands: ecological influences underlying the roles of area, isolation and the size of the faunal source

To distinguish between the influences of area and isolation on the butterfly faunas of British islands two approaches are adopted. First, species richness is related to island area, isolation and the size of the faunal source. Neither area nor isolation account for much variance in species richness, though area is more important than isolation. In contrast, species richness corresponds closely to the size of the faunal source on nearby islands and to that at proximate locations on adjacent mainlands. The second approach relates the incidence of species on islands to their ecological attributes. A very close relationship is found between species incidence on islands and those ecological variables that measure potential for migration and colonization and that resist extinction. The implications are that the majority of British islands in this survey are insufficiently isolated to prevent intermittent migrations of butterflies to them or so small as to generate frequent extinctions. Independent data indicate the capacity of many resident species to migrate distances in excess of the isolation of most of the islands. Some evidence also exists for the long-term survival of species on islands; important considerations in this respect are that most islands in the survey are large compared to habitat patches sustaining species on mainland Britain and that substantial portions of islands are retained in early seral stages or comprise long-lived stable habitats (e.g. peat mosses) that are particularly suitable for many British species.     

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.     

Species richness, rarity and endemicity on Italian offshore islands: complementary signals from island-focused and species-focused analyses

Aims To investigate the relative explanatory power of source faunas and geographical variables for butterfly incidence, frequency, richness, rarity, and endemicity on offshore islands. Location The western Italian offshore islands (Italy and Malta). Methods Thirty‐one islands were examined. Data were taken from our own field surveys and from the literature. Two approaches were undertaken, described as island‐focused and species‐focused, respectively. Offshore islands were allocated to their neighbouring source landmasses (Italian Peninsula, Sicily and Sardinia–Corsica) and compared with each other for faunal attributes, source and island geography. Generalized linear and stepwise multiple regression models were then used to determine the relationships of island species richness, rarity and endemicity with potential geographical predictors and source richness, rarity, and endemicity (island‐focused). Species frequency and incidence were assessed in relation to geographical and source predictors using stepwise linear and logistic regression, and inter‐island associations were examined using K‐Means clustering and non‐metric scaling (species‐focused). Results The analysis reveals firm evidence for the influence of the nearest large landmass sources on island species assemblages, richness, rarity and endemicity. A clear distinction in faunal affinities occurs between the Sardinian islands and islands lying offshore from the Italian mainland and Sicily. Islands neighbouring these three distinct sources differ significantly in richness, rarity and endemicity. Source richness, rarity, and endemicity have explanatory power for island richness, rarity, and endemicity, respectively, and together with island geography account for a substantial part of the variation in island faunas (richness 59%, rarity 60% and endemicity 64%). Source dominates the logistic regression parameters predicting the incidence of island species [13 (38%) of 34 species that could be analysed]; three ecological factors (source frequency, flight period and maximal altitude at which species live) explained 75% of the variation in the occurrence of species on the islands. Species found more frequently on islands occurred more frequently at sources, had longer flight periods, and occurred at lower altitudes at the sources. The incidence of most species on islands (84%) is correctly predicted by the same three variables. Main conclusions The Italian region of the Mediterranean Sea has a rich butterfly fauna comprising endemics and rare species as well as more cosmopolitan species. Analysis of island records benefited from the use of two distinct approaches, namely island‐focused and species‐focused, that sift distinct elements in island and source faunas. Clear contemporary signals appear in island–source relationships as well as historical signals. Differences among faunas relating to sources within the same region caution against assuming that contemporary (ecological) and historical (evolutionary) influences affect faunas of islands in different parts of the same region to the same extent. The implications of source–island relationships for the conservation of butterflies within the Italian region are considered, particularly for the long‐term persistence of species.     

Nestedness of butterfly assemblages in the Zhoushan Archipelago,China: area effects,life-history traits and conservation implications

The nested subset pattern (nestedness) of faunal assemblages has been a research focus in the fields of island biogeography and conservation biology in recent decades. However, relatively few studies have described nestedness in butterfly assemblages in oceanic archipelago systems. Moreover, previous studies often quantified nestedness using inappropriate nestedness metrics and random fill algorithms with high Type I errors. The aims of this study are to examine the existence of nestedness and underlying causal mechanisms of butterfly assemblages in the Zhoushan Archipelago, China. We used the line-transect method to determine butterfly occupancy and abundance on 42 study islands from July to August 2014. We obtained butterfly life-history traits (wingspan, body weight and minimum area requirement) by field work and island geographical features (area and isolation) from the literature. We used the recently developed metric WNODF to estimate nestedness. Partial Spearman rank correlation was used to evaluate the associations of nestedness and island geographical features as well as butterfly life-history traits related to species extinction risk and colonization ability. The butterfly assemblages were significantly nested. Island area and minimum area requirement of butterflies were significantly correlated with nestedness after controlling for other independent variables. In contrast, the nestedness of butterflies did not appear to result from passive sampling or selective colonization. However, multi-year studies are needed to confirm that target effects are not muddling these results. Our results indicate that selective extinction may be the main driver of nestedness of butterfly assemblages in our study system. From a conservation viewpoint, we should protect both large islands and species with large area requirement to maximize the number of species preserved.     

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.     

Conservation biogeography of large Mediterranean islands. Butterfly impoverishment, conservation priorities and inferences for an ecologica "island paradigm"

Conservation biogeography is considered the Cinderella of biological conservation. Nevertheless biogeography provides the basis for establishing species distributions over space and time, therefore conservation priorities among areas and individual species. We demonstrate that there is no need to simplify analyses by using subsets of species (rare species, endemics) as surrogates. In doing so, we apply strict biogeographical techniques to determine butterfly impoverishment on three of the west Mediterranean's largest islands (Sardinia, Corsica and Sicily). The analyses performed on species, both collectively and individually, reveal that regional species richness in the Mediterranean zone can be largely predicted by latitude, altitude and latitudinal range (maximum minus minimum latitude), but that Sardinia and Corsica have clearly impoverished faunas. Logistic regression at individual species level demonstrates that several species, predicted to be present in these islands on the basis of their continental distributions, are actually absent. When compared with species that are present in these islands, such missing species are disclosed as having ecological traits which reduce their colonization capability. Probabilities of occurrence are calculated for each species on each island; they reflect the potential for each butterfly species to migrate to and colonise each island, and can be considered as a measure of conservation value. As such, species present on islands but having low immigration probabilities are predicted to represent isolated populations from the mainland that are unlikely to re-colonize the islands in the case of extinction. Island endemic species and races are shown to have lower occurrence probabilities compared to widespread species occurring on islands and illustrate the usefulness of occurrence probabilities for identifying isolated populations in need of conservation attention.     

舟山群岛蝶类群落嵌套分布格局及其影响因素

片断化生境中群落的物种组成常呈现嵌套分布格局。2013年7-8月, 我们在浙江舟山群岛采用截线法对28个岛屿上的蝴蝶群落进行了野外调查, 探讨了岛屿物种嵌套分布格局及其影响因素。通过测量采集标本获得蝶类的生活史特征(最小需求面积、翅展和体重), 查阅文献资料获得蝶类的栖息地特征(岛屿面积、距最近大陆距离和距最近大岛距离), 分析了影响蝶类群落嵌套结构的因素。研究结果显示: (1)舟山群岛蝶类群落符合嵌套分布格局; (2)岛屿面积和物种最小需求面积对嵌套格局的形成有显著影响; (3)舟山群岛蝶类群落嵌套格局的形成支持选择性灭绝假说; (4)随机检验零模型结果显示该嵌套分布格局并非采样偏差造成的。因此, 在制定舟山群岛区域蝶类保护措施时, 应优先考虑那些分布在面积较大岛屿的和最小需求面积较大的物种。     

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.     

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.     

Contemporary geography dominates butterfly diversity gradients within the Aegean archipelago (Lepidoptera: Papilionoidea, Hesperioidea)

Aim We compare the influence of contemporary geography and historical influences on butterfly diversity for islands in the Aegean archipelago. Location The Aegean archipelago (Greece) and two islands (Cyprus and Megisti) in the Levantine Sea. Methods Thirty‐one islands were examined. Data are taken from own surveys (Coutsis and Olivier) and from the literature. Stepwise multiple regression is used to determine relationships between species richness, frequency, rarity and endemicity against potential geographical predictors. Stepwise logit regression is used to determine geographical predictors of species incidence on islands. Inter‐island and inter‐species associations have been examined using multivariate ordination and clustering techniques. Results The Aegean butterfly fauna is characterized by decreasing diversity and rarity, and increasing homogeneity, from the periphery to the present geographical centre of the archipelago (Cyclades). Diversity and rarity are shown to relate closely to species richness, and species richness, in turn, is largely explained by contemporary geography, particularly the degree of isolation from the nearest mainland sources of Greece or Turkey, and island dimensions. Islands towards the centre of the archipelago are characterized by a group of mobile species (n ≥ 20 species) with extensive ranges across Europe; species that would have recolonized Santorini (Thira) following the VI6 eruption there c. 1630 bc . Endemic components, indicative of autochthonous evolutionary events, are few (5% of species are endemic) compared to known sedentary organisms (molluscs and isopods), but exceed those for more mobile animals (i.e. birds); their distribution is mainly confined to large isolated islands along the Aegean arc (i.e. Kriti) and in the Dodecanese group. Main conclusions Contemporary geography, i.e. processes currently operating in ecological time, dominates butterfly diversity gradients (species richness, frequency, rarity and incidence) in the archipelago. Two reasons are suggested to account for the lack of endemism and the pattern of decreasing diversity into the Cyclades. First, relict butterfly elements may have become extinct on all but a few larger islands, particularly from environmental changes since the Neolithic (fire and overgrazing). Second, colonization from the continental landmasses is ongoing with more mobile species transferring even to the most isolated islands.     

Butterflies of European islands: the implications of the geography and ecology of rarity and endemicity for conservation

Depending on their faunal content islands can function as important ‘vehicles’ for conservation. In this study, we examine data on 440 butterfly species over 564 European islands in 10 island groups. To determine the status of the butterfly fauna, we have adopted two approaches, island-focused and species-focused, examined using principal components analysis and regression modelling. In the former, we relate species richness, rarity and endemicity to island geography (area, elevation, isolation and location in latitude and longitude); in the latter, species occurrence on islands is examined in relation to distribution, range, range boundaries, and altitudinal limits on the continent as well as species’ ecology (number of host plants) and morphology (wing expanse). Species on islands are also assessed for their status on the continental mainland, their distributional dynamics (extinctions, distribution changes) and conservation status (Red Data Book, European Habitat Directive, Species of European Conservation Concern and Bern Convention listing. Unexpectedly, we find that a large fraction of the European butterfly species is found on the islands (63.4%; 59% on small islands) comprising some 6.2% of the land area of Europe. Although species occurring on the islands tend, on the whole, to have lower conservation status and are not declining over Europe, 45 species are endemics restricted to the islands. Species richness shows only a weak locational pattern and is related as expected to isolation from the continental source and island area; but, both rarity and endemicity have distinctive geographical bias to southern Europe, on islands now under increasing pressure from climate change and increasingly intensive human exploitation. The vulnerability of species on islands is emphasised in the relationship of island occurrence (% occurrence and presence/absence of species on any island) with continental distributions. A large proportion of the variation (84%) is accounted by continental distribution, the southern range limit and lower altitudinal limit. Most species (69%) occur on very few islands (<5%). In view of ongoing species dynamics on islands, migrations and extinctions of species, island repositories of species depend in large part on conservation of butterflies at continental sources. The unique faunas and rare species on islands also depend on appropriate concern being given to the island faunas. Conservation of European islands is thus a two-way process, sustaining sources and conserving island refuges. Residuals from the regressions (islands with more or fewer species, rare and endemic species; species occurring more or less frequently than expected on islands) provide warning signals of regions and islands deserving immediate attention.     

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.     

Influence of island geography,age and landscape on species composition in different animal groups

Aim To study the importance of ecological and geographical factors in explaining arthropod species composition on islands. Location The Aeolian Islands, a volcanic archipelago in the central Mediterranean, near Sicily. Methods The influence of island area, age, distance to the mainland, distance to the nearest island and land cover categories on species composition of arthropod groups was analysed using canonical correspondence analysis (CCA). The use of multiple animal groups in the same archipelago allowed the development of two complementary approaches based on CCA – a ‘taxon‐focused’ approach and an ‘island‐focused’ approach – to elucidate, respectively, how different taxa respond to the same environmental factors, and which factors are mainly responsible for the composition of the faunas in different locations. Results Island area was an important factor in explaining species composition in Chilopoda, Orthoptera and Tenebrionidae. Distance to the mainland was important mainly for Carabidae. Distance to the closest island was important for many groups. By contrast, island age exerted a significant influence only on the species composition of Orthoptera. Various groups were influenced by a combination of broad‐leaved forest and natural grassland. Main conclusions The example of the arthropods of the Aeolian Islands indicates that the influence of a given island characteristic on species composition varies among groups, although measures of inter‐island isolation were typically more important for taxa than isolation from the mainland source. This suggests that colonization of islands may occur mostly by stepping‐stone dispersal.     

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 size is not the only consideration. Ranking priorities for the conservation of butterflies on Italian offshore islands

Ecological and historical factors virtually create a unique faunal assemblage on each island. From this perspective every island deserves protection. However, economic limitations usually restrict conservation efforts to particularly important areas. As part of the SLOSS issue (the relative importance of single large or several small areas), there is the long debated question of whether it is better to protect few large areas (islands) or several small areas (islands). Here, we assess the butterfly faunas of the Italian offshore islands, using several biodiversity measures, in order to highlight priorities for conserving butterfly richness, rarity and endemicity. First, the nested pattern of butterfly fauna was investigated to determine the relative importance of large and small islands. Then, residuals were assessed for the species-area relationship and for multiple regressions of richness, rarity and endemicity against geographic variables. Subsequently, two other indices were calculated: Biodiversity Conservation Concern and an index scoring islands in the order that maximizes the cumulative percentage of total, endemic, and rare species. The results clearly indicate that although greatest concern is for the island having the largest butterfly fauna in the sample (Elba), the importance of several small islands should not be ignored. This is primarily due to the substantial impact of source areas and consequently the occurrence of several rare and endemic species occurring on small islands as well as on large islands.     

How island size and isolation affect bee and wasp ensembles on small tropical islands: a case study from Kepulauan Seribu,Indonesia

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.     

Nestedness of desert bat assemblages: species composition patterns in insular and terrestrial landscapes

Nested patterns of community composition exist when species at depauperate sites are subsets of those occurring at sites with more species. Nested subset analysis provides a framework for analyzing species occurrences to determine non-random patterns in community composition and potentially identify mechanisms that may shape faunal assemblages. We examined nested subset structure of desert bat assemblages on 20 islands in the southern Gulf of California and at 27 sites along the Baja California peninsula coast, the presumable source pool for the insular faunas. Nested structure was analyzed using a conservative null model that accounts for expected variation in species richness and species incidence across sites (fixed row and column totals). Associations of nestedness and island traits, such as size and isolation, as well as species traits related to mobility, were assessed to determine the potential role of differential extinction and immigration abilities as mechanisms of nestedness. Bat faunas were significantly nested in both the insular and terrestrial landscape and island size was significantly correlated with nested structure, such that species on smaller islands tended to be subsets of species on larger islands, suggesting that differential extinction vulnerabilities may be important in shaping insular bat faunas. The role of species mobility and immigration abilities is less clearly associated with nestedness in this system. Nestedness in the terrestrial landscape is likely due to stochastic processes related to random placement of individuals and this may also influence nested patterns on islands, but additional data on abundances will be necessary to distinguish among these potential mechanisms.     

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.     

Mammalian biogeography of the Alexander Archipelago of Alaska: a north temperate nested fauna

Aim A large number of studies have analysed the distribution of mammals within archipelagos, yet few have focused on islands that were heavily glaciated and subsequently colonized following deglaciation. Location We explored the relative effects of island area and isolation on faunal composition based on twenty-three mammalian taxa of twenty-four islands of the Alexander Archipelago, Southeast Alaska. Methods We used regression of log-transformed variables and several indices of nestedness. Results These faunas showed significant nested structure using tests of nestedness and regression models. Unlike most landbridge and mainland archipelagos studied previously, isolation appears to be the primary factor determining patterns of species richness. Main conclusions Colonization ability of particular taxa, rather than extinction, has determined this nested structure. We suggest that other higher latitude archipelagos may show similar historical patterns.     

Determinants of bird species richness,endemism, and island network roles in Wallacea and the West Indies: is geography sufficient or does current and historical climate matter?

Island biogeography has greatly contributed to our understanding of the processes determining species' distributions. Previous research has focused on the effects of island geography (i.e., island area, elevation, and isolation) and current climate as drivers of island species richness and endemism. Here, we evaluate the potential additional effects of historical climate on breeding land bird richness and endemism in Wallacea and the West Indies. Furthermore, on the basis of species distributions, we identify island biogeographical network roles and examine their association with geography, current and historical climate, and bird richness/endemism. We found that island geography, especially island area but also isolation and elevation, largely explained the variation in island species richness and endemism. Current and historical climate only added marginally to our understanding of the distribution of species on islands, and this was idiosyncratic to each archipelago. In the West Indies, endemic richness was slightly reduced on islands with historically unstable climates; weak support for the opposite was found in Wallacea. In both archipelagos, large islands with many endemics and situated far from other large islands had high importance for the linkage within modules, indicating that these islands potentially act as speciation pumps and source islands for surrounding smaller islands within the module and, thus, define the biogeographical modules. Large islands situated far from the mainland and/or with a high number of nonendemics acted as links between modules. Additionally, in Wallacea, but not in the West Indies, climatically unstable islands tended to interlink biogeographical modules. The weak and idiosyncratic effect of historical climate on island richness, endemism, and network roles indicates that historical climate had little effects on extinction‐immigration dynamics. This is in contrast to the strong effect of historical climate observed on the mainland, possibly because surrounding oceans buffer against strong climate oscillations and because geography is a strong determinant of island richness, endemism and network roles.