Nestedness in island faunas: novel insights into island biogeography through butterfly community profiles of colonization ability and migration capacity

Aim To relate variation in the migration capacity and colonization ability of island communities to island geography and species island occupancy. Location Islands off mainland Britain and Ireland. Methods Mean migration (transfer) capacity and colonization (establishment) ability (ecological indices), indexed from 12 ecological variables for 56 butterfly species living on 103 islands, were related to species nestedness, island and mainland source geography and indices using linear regression models, RLQ analysis and fourth‐corner analysis. Random creation of faunas from source species, rank correlation and rank regression were used to examine differences between island and source ecological indices, and relationships to island geography. Results Island butterfly faunas are highly nested. The two ecological indices related closely to island occupancy, nestedness rank of species, island richness and geography. The key variables related to migration capacity were island area and isolation; for colonization ability they were area, isolation and longitude. Compared with colonization ability, migration capacity was found to correlate more strongly with island species occupancy and species richness. For island faunas, the means for both ecological indices decreased, and variation increased, with increasing island species richness. Mean colonization ability and migration capacity values were significantly higher for island faunas than for mainland source faunas, but these differences decreased with island latitude. Main conclusions The nested pattern of butterfly species on islands off mainland Britain and Ireland relates strongly to colonization ability but especially to migration capacity. Differences in colonization ability among species are most obvious for large, topographically varied islands. Generalists with abundant multiple resources and greater migration capacity are found on all islands, whereas specialists are restricted to large islands with varied and long‐lived biotopes, and islands close to shore. The inference is that source–sink dynamics dominate butterfly distributions on British and Irish islands; species are capable of dispersing to new areas, but, with the exception of large and northern islands, facilities (resources) for permanent colonization are limited. The pattern of colonization ability and migration capacity is likely to be repeated for mainland areas, where such indices should provide useful independent measures for assessing the conservation status of faunas within spatial units.     

Composition and structure of Caribbean bat (Chiroptera) assemblages: effects of inter‐island distance,area, elevation and hurricane‐induced disturbance

Aim Although bats of the Caribbean have been studied extensively, previous work is largely restricted to zoogeography, phylogeography or the effects of island characteristics on species richness. Variation among islands in species composition that is related to geographical or environmental variation remains poorly understood for much of the Caribbean. Location Caribbean islands, including the Bahamas, Greater Antilles and Lesser Antilles. Methods Using presence–absence data, we assessed the extent to which island area, maximum island elevation, inter‐island distance and hurricane‐induced disturbance affected patterns of composition and nestedness for bats in the Bahamas, Greater Antilles and Lesser Antilles. Analyses were conducted for all species, as well as for two broadly defined guilds: carnivores and herbivores. Results For the Bahamas, only inter‐island distance accounted for variation in species composition between islands. For the Greater and Lesser Antilles, differences in island area and inter‐island distance accounted for differences in species composition between islands. Variation in species composition was not related significantly to differences in elevation or hurricane‐related disturbance. In general, results of analyses restricted to a particular broad guild (i.e. carnivores or herbivores) mirrored those for all bats. Bat species composition was nested significantly in each island group. Nestedness was stronger in the Greater Antilles and in the Lesser Antilles than in the Bahamas. Carnivore assemblages were nested significantly in the Greater and in the Lesser Antilles, but not in the Bahamas. In contrast, herbivore assemblages were nested significantly in each island group. Main conclusions Inter‐island distance had a greater effect on compositional similarity of Caribbean bat assemblages than did island area, elevation or disturbance related to hurricanes. Differential immigration and hierarchical habitat distributions associated with elevational relief are likely to be primary causes for nestedness of Caribbean bat assemblages.     

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.     

BIODIVERSITY RESEARCH: Nestedness for different reasons: the distributions of birds,lizards and small mammals on islands of an inundated lake

Aim We examined whether the community compositions of birds, lizards and small mammals were nested in a fragmented landscape in the Thousand Island Lake, China. We also assessed whether the mechanisms influencing nestedness differed among these taxonomic groups. Location Thousand Island Lake, China. Methods Presence/absence matrices were compiled for birds (42 islands) and lizards (42 islands) using line‐transect methods, and for small mammals (14 islands) using live‐trapping methods from 2006 to 2009. Nestedness was analysed using BINMATNEST, and statistical significance was assessed using the conservative null model 3. We used Spearman rank correlations and partial Spearman rank correlations to examine associations of nestedness and habitat variables (area, isolation, habitat diversity and plant richness) as well as life‐history traits (body size, habitat specificity, geographical range size and area requirement) related to species extinction and immigration tendencies. Results The community compositions of birds, lizards and small mammals were all significantly nested, but the causal factors underlying nestedness differed among taxonomic groups. For birds, island area, habitat specificity and area requirement were significantly correlated with nestedness after controlling for other independent variables. For lizards, habitat heterogeneity was the single best correlate of nestedness. For small mammals, island area, habitat heterogeneity and habitat specificity were significantly correlated with nestedness. The nested patterns of birds, lizards and small mammals were not attributable to passive sampling or selective colonization. Main conclusions The processes influencing nested patterns differed among taxonomic groups. Nestedness of bird assemblages was driven by selective extinction, and lizard assemblage was caused by habitat nestedness, while nestedness of small mammals resulted from both selective extinction and habitat nestedness. Therefore, we should take taxonomic differences into account when analysing nestedness to develop conservation guidelines and refrain from using single taxa as surrogates for others.     

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.     

Effects of Fragmentation and Sea-Level Changes upon Frog Communities of Land-Bridge Islands off the Southeastern Coast of Brazil

We investigate the composition of anuran communities of land-bridge islands off the southeastern coast of Brazil. These islands provide natural long-term experiments on the effects of fragmentation in the Brazilian Atlantic Forest (BAF). We hypothesize that Pleistocene sea-level changes, in combination with other abiotic variables such as area and habitat diversity, has affected anuran species richness and community composition on these islands. Data from the literature and collections databases were used to produce species lists for eight land-bridge islands and for the mainland adjacent to the islands. We assess the effects of area, number of breeding habitats and distance to the mainland upon anuran species richness on land-bridge islands. Additionally we use nestedness analysis to quantify the extent to which the species on smaller and less habitat-diverse islands correspond to subsets of those on larger and more diverse ones. We found that area has both direct and indirect effects on anuran species richness on land-bridge islands, irrespective of distance to the mainland. However, on islands with comparable sizes, differences in species richness can be attributed to the number and quality of breeding habitats. Anuran communities on these islands display a nested pattern, possibly caused by selective extinction related to habitat loss. Common lowland pond-breeders were conspicuous by their absence. In the BAF, the conservation of fragments with a high diversity of breeding habitats could compensate for the generally negative effect of small area upon species richness. We suggest that sea-level changes have an important role in shaping composition of anuran species on coastal communities.     

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.     

Distributions of forest birds and butterflies in the Andaman islands, Bay of Bengal: nested patterns and processes

The distributional patterns of forest birds and butterflies in the Andaman islands, an oceanic chain located off SE Asia, were tested for nestedness. Both taxa were highly nested. Nestedness could be due to colonization or extinction processes, area or distance effects or nestedness of habitats. Nestedness in forest bird distributions were strongly influenced by area and habitat related factors. Habitats were significantly nested in all three island groups, however most strongly for the North Andamans. However forest bird distributions in the North Andamans, as indicated by row order in the packed matrix, was not correlated with habitat diversity, suggesting that habitat related factors alone cannot account for these patterns. Other causal influences could be passive sampling, where common and abundant species and habitats are more likely to have a widespread distribution than rare species and habitats. The nested subset pattern seen in two unrelated taxa suggests that the Andamans are extinction dominated and that the protection of forests on large islands is critical for the conservation of its biodiversity.     

Nested bird and micro-habitat assemblages in a peatland archipelago

Biotic assemblages of insular habitats are nested when poor assemblages are subsets of richer ones. Nestedness of species assemblages is frequent and may result from selective extinction or frequent colonization in insular habitats. It may also be created by a nested distribution of habitats among islands or by sampling bias. We sampled 67 isolated peatlands (7–843 ha) in southern Quebec, Canada, to measure nestedness of bird species assemblages among peatlands and assess the habitat nestedness hypothesis. Species and microhabitat assemblages were both strongly nested among peatlands. Whether sites were ranked by species richness, microhabitat richness or peatland area had no effect on nestedness. However, microhabitat nestedness was significantly reduced when sites were sorted by area rather than by microhabitat richness. As expected, if bird-microhabitat associations are responsible for the nested pattern of distribution, we found a positive correlation between the contributions of bird species and microhabitats to individual site nestedness. Nevertheless, microhabitat assemblages were significantly less nested than bird species assemblages, possibly because of frequent recolonization by birds or uneven sampling among sites. Received: 12 June 1998 / Accepted: 20 September 1998     

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.     

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.     

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.     

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.     

Temporal variability of nestedness and idiosyncratic species in stream insect assemblages

Aims The nested subset pattern has been widely studied in the last 20 years, and recent syntheses have challenged the prevalence of this pattern in nature. We examined the degree of nestedness, its temporal variability and its environmental correlates in stream insects of a boreal drainage system. We also examined differences between nested and idiosyncratic species in site occupancy, niche position and niche breadth. Location Koutajoki drainage basin in northern Finland. Methods We used (i) nestedness analyses with three null models for testing the significance of nestedness; (ii) Spearman rank correlation to examine the correlates of nestedness; (iii) outlying mean index analysis to analyse the niche characteristics of species; (iv) and t‐test to examine differences in niche breadth, niche position and site occupancy of idiosyncratic and other nested species. Results Stream insect assemblages were significantly nested in each of the three study years. The maximally packed matrices were significantly nested according to the nestedness calculator based on null models I (species frequencies and site richness equiprobable) and II (species frequencies fixed and site richness equiprobable), but non‐significant based on a conservative null model III (species frequencies and site richness fixed to those of the observed matrix). The most important correlate of nestedness was stream size, whereas isolation, productivity (total phosphorus) and habitat heterogeneity exhibited non‐significant relationship with nestedness. Idiosyncratic species occurred, on average, at more sites than nested species, mirroring the restricted distributions of several nested species that were inclined towards species‐rich sites. Idiosyncratic and nested species also differed in niche position and niche breadth, with idiosyncratic species having, on average, less marginal niche positions and wider niches than nested species. Main conclusions Stream size correlated with nestedness, possibly because small streams were inhabited only by species able to persist under, or colonize shortly after, disturbances, while most species could occur at larger sites where disturbances are less severe. From the conservation perspective, our findings suggest that stream size really matters, given that sites with high species richness and many rare species are more likely to occur in larger streams. However, also the requirements of idiosyncratic species should be accommodated in conservation planning.     

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.     

Fine‐tuning the nested structure of pollination networks by adaptive interaction switching,biogeography and sampling effect in the Galápagos Islands

The structure of pollination networks, particularly its nestedness, contain important information on network assemblages. However, there is still limited understanding of the mechanisms underlying nested pollination network structures. Here, we investigate the role of adaptive interaction switching (AIS), island area, isolation, age and sampling effort in explaining the nestedness of pollination networks across ten Galápagos Islands. The AIS algorithm is inspired by Wallace's elimination of the unfit, where a species constantly replaces its least profitable mutualistic partner with a new partner selected at random. To explain network structures, we first use a dynamic model that includes functional response of pollination and AIS, with only species richness and binary connectance as input (hereafter the AIS model). Thereafter, other explanatory variables (isolation, area, age and sampling effort) were added to the model. In four out of ten islands, the pollination network was significantly nested, and predictions from the AIS model correlated with observed structures, explaining 69% variation in nestedness. Overall, in terms of independent contribution from hierarchical partitioning of variation in observed nestedness, the AIS model predictions contributed the most (37%), followed by sampling effort (28%) and island area (22%), with only trivial contributions from island isolation and age. Therefore, adaptive switching of biotic interactions seems to be key to ensure network function, with island biogeographic factors being only secondary. Although large islands could harbour more diverse assemblages and thus foster more nested structures, sufficient sampling proves to be essential for detecting non‐random network structures.     

Nested distribution of amphibians in the Zhoushan archipelago, China: can selective extinction cause nested subsets of species?

Because of their poor dispersal ability, amphibians are well suited for testing the selective extinction theory on islands. Amphibian fauna in the Zhoushan archipelago, China, exhibit a high level of nestedness (C = 0.893), and the species number is lower on islands than on similar sized areas on the mainland. No correlation was found between island-specific species richness and the nearest distance from a larger island, distance from the mainland or density of human population. These results suggest that no amphibian colonisation has occurred in the archipelago since island isolation 7000–9000 years ago. Furthermore, the results imply that selective extinction contributes to the nestedness of amphibians in the Zhoushan archipelago. The incidence of a species on the islands is significantly correlated with log area of the smallest island occupied by the species and the number of provinces on the Chinese mainland in which the species occur. However, there is no correlation with average body length of adults and island occurrence. It is concluded that (1) the area of the smallest island occupied by a species is a good estimate of the minimum area for a viable population of the species and a good predictor of species incidence on islands, (2) species with a restricted distribution range are more vulnerable to extinction from islands than those with a wide distribution range and (3) the effect of body size on occurrence on the islands is uncertain, and may be specific to the archipelago and taxa studied. The observed nestedness of amphibian assemblages has two implications for conservation: (1) not only can all the species found in several small reserves be found on a large reserve of the same total size, but additional species can be found on the single large reserve; (2) for a reserve to maintain viable populations of all species in a region it should be at least as large as the smallest island occupied by the most vulnerable species. Received: 16 December 1996 / Accepted: 22 September 1997     

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.     

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.