Null model analysis of communities on gradients

Aim I employed a novel null model and metric to uncover unusual species co‐occurrence patterns in a herpetofaunal assemblage of 49 species collected at discrete elevations along a gradient. Location Mount Kupe, Cameroon. Methods Using a construction algorithm that started from a matrix of 0s, a sample null space of 25,000 unique null matrices was generated by simultaneously conserving (1) the number of occurrences of each species, (2) site richness and (3) species range spans derived from the observed incidence matrix. I then compared the number of times each pair of confamilial species co‐occurred in the null space with the same number derived from the observed incidence matrix. Two cases dealing with embedded absences in species ranges were tested: (1) embedded absences were maintained, and (2) embedded absences were assumed to be sampling omissions and were replaced by presences. Results In the observed absence/presence assemblage there were 147 possible confamilial species pairs. Therefore, 5% or eight were expected by chance alone to have co‐occurrence patterns that differed from chance expectations by chance alone. Of these confamilial species pairs, 38 were congeneric and so 5% or two were expected to differ from chance expectations. For case (1) 16, and for case (2) 17 confamilial species pairs’ co‐occurrence patterns differed significantly from chance expectations. For case (1) nine congeneric species pairs, and for case (2) 10 congeneric pairs differed significantly from chance expectations. For case (1) four, and for case (2) five congeneric species pairs formed checkerboards (patterns of mutual exclusion). Results from case (1) were a proper subset of case (2) indicating that sampling omissions did not alter greatly the results. Main conclusions I have demonstrated that null models are valuable tools to analyse ecological communities provided that proper models are employed. The choice of the appropriate null space to analyse distributions is critical. The null model employed to analyse birds on islands of an archipelago can be adapted to analyse species along gradients provided an additional range constraint is added to the null model. Moreover, added precision to results can be obtained by analysing each species pair separately, particularly those in the same family or genus, as opposed to applying a community‐wide metric to the faunal assemblage. My results support some of the speculations of previous authors who were unable to demonstrate their suspicions analytically.     

Phylogeny and co‐occurrence of mammal species on Southeast Asian islands

Aim Islands have often been used as model systems in community ecology. The incorporation of information on phylogenetic relatedness of species in studies of island assemblage structure is still uncommon, but could provide valuable insights into the processes of island community assembly. We propose six models of island community assembly that make different predictions about the associations between co‐occurrences of species pairs on islands, phylogenetic relatedness and ecological similarity. We then test these models using data on mammals of Southeast Asian islands. Location Two hundred and forty islands of the Sundaland region of Southeast Asia. Methods We quantified the co‐occurrence of species pairs on islands, and identified pairs that co‐occur more frequently (positive co‐occurrence) or less frequently (negative co‐occurrence) than expected under null models. We then examined the distributions of these significantly deviating pairs with respect to phylogenetic relatedness and ecological differentiation, and compared these patterns with those predicted by the six community assembly models. We used permutation regression to test whether co‐occurrence patterns are predicted by relatedness, body size difference or difference in diet quality. Separate co‐occurrence matrices were analysed in this way for seven mammal families and four smaller subsets of the islands of Sundaland. Results In many matrices, average numbers of negative co‐occurrences were higher than expected under null models. This is consistent with assemblage structuring by competition, but may also result from low geographic overlap of species pairs, which contributes to negative co‐occurrences at the archipelago‐wide level. Distributions of species pairs within plots of phylogenetic distance × ecological differentiation were consistent with competition, habitat filtering or within‐island speciation models, depending on the taxon. Regressions indicated that co‐occurrence was more likely among closely related species pairs within the Viverridae and Sciuridae, but in most matrices phylogenetic distance was unrelated to co‐occurrence. Main conclusions Simple deterministic models linking co‐occurrence with phylogeny and ecology are a useful framework for interpreting distributions and assemblage structure of island species. However, island assemblages in Sundaland have probably been shaped by a complex idiosyncratic set of interacting ecological and evolutionary processes, limiting the predictive power of such models.     

Binary matrices and checkerboard distributions of birds in the Bismarck Archipelago

Aim We examine a presence–absence matrix of the avifauna of the Bismarck Archipelago, for which the concept of competitively driven community assembly rules was formulated, to determine whether data support widespread competitive determination of geographical distributions. Location Bismarck Archipelago. Methods We obtained occurrences of 154 land and freshwater bird species on 31 islands. We calculated the observed number of checkerboards for all species pairs, for congeneric species pairs and for pairs of species within guilds, and employed randomization techniques to detect unusual co‐occurrence patterns. Results Compared with random expectations, there are excesses of checkerboard pairs within both genera and defined guilds, but a detailed examination shows that competition is a cogent possible explanation in few instances. For many checkerboard pairs, species are not widely interspersed but are regionally allopatric, which probably reflects historical biogeography and dispersal limitation. Most congeneric and intraguild checkerboards include a species classified as a supertramp; when supertramps are omitted, there are 11 congeneric checkerboards and four intraguild but heterogeneric checkerboards. Main conclusions In isolation, presence–absence matrices provide limited insight into the role of competition in structuring bird communities of the Bismarcks. A major problem is disentangling historical geography and colonization history of the archipelago from the present‐day ecology of the species. Examination of observed checkerboards from a geographically explicit perspective and with knowledge of colonization routes suggests that many checkerboards are likely to result, at least in part, from historical biogeography and supertramps. Although species may be forced into supertramp status by competition, other factors (e.g. habitat preference) may be causal, and biogeographical distributions alone cannot distinguish between causes.     

Geographical distribution pattern and interisland movements of Orii’s flying fox in Okinawa Islands, the Ryukyu Archipelago, Japan

The study of mobile animals such as flying foxes in insular habitats involves clarifying the population status on each island and determining the factors affecting movement patterns among the islands in their distributional range. We visited 25 of the Okinawa Islands and documented the number of Orii’s flying foxes Pteropus dasymallus inopinatus from August 2005 to May 2006. We also conducted a monthly road census on the main island (Okinawa-jima Island) and six adjacent islands from June 2006 to January 2007 and counted the number of fruit-bearing trees of the bats’ four main food plants. The results of classification and regression tree analysis suggested that distance from the main island was a primary factor in determining the distribution pattern and population size of this flying fox, whereas island area, number of plant species, and food availability did not directly affect population size. The number of flying foxes on each island tended to decrease with an increase in distance from the main island; no flying foxes existed on islands >30 km away from the main island. On the other hand, the results of the monthly census showed that the population size on each island fluctuated seasonally. Individuals may move between islands in response to seasonal changes in food availability. In conclusion, the distribution and abundance of Orii’s flying foxes in the Okinawa Islands may be determined by the rate of immigration/emigration, depending on each island’s distance from the main island. Seasonal changes in food availability may act as a trigger for interisland movement, but that movement may be restricted by island connectivity.     

An evaluation of randomization models for nested species subsets analysis

Randomization models, often termed “null” models, have been widely used since the 1970s in studies of species community and biogeographic patterns. More recently they have been used to test for nested species subset patterns (or nestedness) among assemblages of species occupying spatially subdivided habitats, such as island archipelagoes and terrestrial habitat patches. Nestedness occurs when the species occupying small or species-poor sites have a strong tendency to form proper subsets of richer species assemblages. In this paper, we examine the ability of several published simulation models to detect, in an unbiased way, nested subset patterns from a simple matrix of site-by-species presence-absence data. Each approach attempts to build in biological realism by following the assumption that the ecological processes that generated the patterns observed in nature would, if they could be repeated many times over using the same species and landscape configuration, produce islands with the same number of species and species present on the same number of islands as observed. In mathematical terms, the mean marginal totals (column and row sums) of many simulated matrices would match those of the observed matrix. Results of model simulations suggest that the true probability of a species occupying any given site cannot be estimated unambiguously. Nearly all of the models tested were shown to bias simulation matrices toward low levels of nestedness, increasing the probability of a Type I statistical error. Further, desired marginal totals could be obtained only through ad-hoc manipulation of the calculated probabilities. Paradoxically, when such results are achieved, the model is shown to have little statistical power to detect nestedness. This is because nestedness is determined largely by the marginal totals of the matrix themselves, as suggested earlier by Wright and Reeves. We conclude that at the present time, the best null model for nested subset patterns may be one based on equal probabilities of occurrence for all species. Examples of such models are readily available in the literature. Received: 3 February 1997 / Accepted: 21 September 1997     

Source pools and disharmony of the world's island floras

Island disharmony refers to the biased representation of higher taxa on islands compared to their mainland source regions and represents a central concept in island biology. Here, we develop a generalizable framework for approximating these source regions and conduct the first global assessment of island disharmony and its underlying drivers. We compiled vascular plant species lists for 178 oceanic islands and 735 mainland regions. Using mainland data only, we modelled species turnover as a function of environmental and geographic distance and predicted the proportion of shared species between each island and mainland region. We then quantified the over‐ or under‐representation of families on individual islands (representational disharmony) by contrasting the observed number of species against a null model of random colonization from the mainland source pool, and analysed the effects of six family‐level functional traits on the resulting measure. Furthermore, we aggregated the values of representational disharmony per island to characterize overall taxonomic bias of a given flora (compositional disharmony), and analysed this second measure as a function of four island biogeographical variables. Our results indicate considerable variation in representational disharmony both within and among plant families. Examples of generally over‐represented families include Urticaceae, Convolvulaceae and almost all pteridophyte families. Other families such as Asteraceae and Orchidaceae were generally under‐represented, with local peaks of over‐representation in known radiation hotspots. Abiotic pollination and a lack of dispersal specialization were most strongly associated with an insular over‐representation of families, whereas other family‐level traits showed minor effects. With respect to compositional disharmony, large, high‐elevation islands tended to have the most disharmonic floras. Our results provide important insights into the taxon‐ and island‐specific drivers of disharmony. The proposed framework allows overcoming the limitations of previous approaches and provides a quantitative basis for incorporating functional and phylogenetic approaches into future studies of island disharmony.     

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.     

Response to Collins et al. (2011)

We note serious problems in Collins et al. (Journal of Biogeography, 2011, doi: 10.1111/j.1365‐2699.2011.02506.x ): failure to use over 80% of the available data; failure to use one of the two available archipelagoes; mistaken inclusion of four species; and reliance on a grossly inadequate number of null matrices. Curing the paper of these problems would have strengthened the evidence for checkerboards and the role of competition.     

The form of the curves: a direct evaluation of MacArthur & Wilson's classic theory

1. We calculate the yearly numbers of bird species immigrating to – and becoming extinct on – 13 small islands of the British Isles, using a long and relatively complete data record.
2. We estimate the size of the colonist pool for each island using four methods.
3. We assume that immigrations and extinctions are distributed binomially, and use a maximum likelihood method to fit concave immigration and extinction functions to the data, utilizing all four species pool estimates.
4. Extinction rates increase significantly and consistently with increasing numbers of breeding species on each island. For nine of the 13 islands the extinction functions are significantly concave.
5. Immigration rates decrease consistently with increasing numbers of breeding species on each island. Seven islands have significantly concave immigration functions.
6. Immigration rates and extinction rates decline consistently, but not significantly, with island distance and island size, respectively. The number of breeding species does not always reflect the number of species likely to have reached an island. Moreover, some species may choose not to breed when their chance of extinction is high. These factors, plus the modest range of island areas and distances in our database, reduce our chances of finding the theoretically predicted effects of area and distance on extinction and immigration rates.     

Determinants of the diversity of plants, birds and mammals of coastal islands of the Humboldt current systems: implications for conservation

Sound conservation plans for islands require understanding the processes underlying to the patterns of species richness and composition. Larger islands are often the targets of conservation assuming that the island area mainly determines species richness, and that species composition is nested across islands. However, in small-island these patterns could be altered because of stochastic processes, and species assemblages could be disharmonious. In addition, human impact could further modify the distribution pattern and diversity. Here we use the case of seven islands from the coastal system of Coquimbo as a model to address the role of environmental variables and human impacts on species richness and assembly rules of plants, birds, and mammals. We hypothesize (a) the existence of a small-island effect, and the prevalence of habitat diversity and anthropogenic impacts as main drivers of species richness, and (b) the existence of disharmonious assemblages, characterized by a low degree of nestedness and random patterns of species co-occurrence. Our results showed that (a) species richness is mainly correlated with habitat diversity, and only weakly related to island area supporting the ‘small-island effect’ and (b) species composition is highly structured, but that such structure may be the result of anthropogenic activities. Nestedness was observed in plants and landbirds, while co-occurrence patterns were only detected in plants. Assemblages in small-islands departed from the nestedness pattern and maintain rare species. Currently, only three of the seven islands are protected by national regulations, excluding the smaller ones that are subjected to human disturbance and invasive mammals. Our study suggests that it necessary to include all the islands in a major protected area to preserve both richness and species composition of a number of representative islands of the Humboldt current systems. We showed that conservation plans solely based on island area might not be robust.     

The empirical Bayes approach as a tool to identify non-random species associations

A statistical challenge in community ecology is to identify segregated and aggregated pairs of species from a binary presence–absence matrix, which often contains hundreds or thousands of such potential pairs. A similar challenge is found in genomics and proteomics, where the expression of thousands of genes in microarrays must be statistically analyzed. Here we adapt the empirical Bayes method to identify statistically significant species pairs in a binary presence–absence matrix. We evaluated the performance of a simple confidence interval, a sequential Bonferroni test, and two tests based on the mean and the confidence interval of an empirical Bayes method. Observed patterns were compared to patterns generated from null model randomizations that preserved matrix row and column totals. We evaluated these four methods with random matrices and also with random matrices that had been seeded with an additional segregated or aggregated species pair. The Bayes methods and Bonferroni corrections reduced the frequency of false-positive tests (type I error) in random matrices, but did not always correctly identify the non-random pair in a seeded matrix (type II error). All of the methods were vulnerable to identifying spurious secondary associations in the seeded matrices. When applied to a set of 272 published presence–absence matrices, even the most conservative tests indicated a fourfold increase in the frequency of perfectly segregated “checkerboard” species pairs compared to the null expectation, and a greater predominance of segregated versus aggregated species pairs. The tests did not reveal a large number of significant species pairs in the Vanuatu bird matrix, but in the much smaller Galapagos bird matrix they correctly identified a concentration of segregated species pairs in the genus Geospiza. The Bayesian methods provide for increased selectivity in identifying non-random species pairs, but the analyses will be most powerful if investigators can use a priori biological criteria to identify potential sets of interacting species.     

Island size, isolation, or interspecific competition? The breeding distribution of the Parus guild in the Danish archipelago

The Parus guild (Parus spp., Sitta, Certhia, and Regulus) is distributed as a complex mosaic within the Danish archipelago, with from one to eight species on different islands. We assessed the roles of island isolation, island size, and interspecific competition in determining the breeding species compositions of this guild on 53 Danish islands. Small, isolated islands supported fewer species than larger, nearshore islands. These effects, however, were largely restricted to a few sedentary species (P. cristatus, P. palustris, S. europaea) that are known to be poor dispersers/colonizers. In some cases, these three species were also absent from large, nearshore islands with suitable habitat, suggesting that habitat availability was not always responsible for the absence of a species. Monte Carlo simulations suggested that the pattern of species presence/absence was not a result of interspecific interactions. Thus, although a number of previous studies have documented interspecific competition among members of the Parus guild, our results suggest that such competition is not responsible for the unusual pattern of species distribution within the Danish archipelago. Received: 28 October 1996 / Accepted: 7 February 1997     

Sampling design may obscure species–area relationships in landscape-scale field studies

We investigated 1) the role of area per se in explaining anuran species richness on reservoir forest islands, after controlling for several confounding factors. We also assessed 2) how sampling design affects the inferential power of island species–area relationships (ISARs) aiming to 3) provide guidelines to yield reliable estimates of area-induced species losses in patchy systems. We surveyed anurans with autonomous recording units at 151 plots located on 74 islands and four continuous forest sites at the Balbina Hydroelectric Reservoir landscape, central Brazilian Amazonia. We applied semi-log ISAR models to assess the effect of sampling design on the fit and slope of species–area curves. To do so, we subsampled our surveyed islands following both a 1) stratified and 2) non-stratified random selection of 5, 10, 15, 20 and 25 islands covering 1) the full range in island size (0.45–1699 ha) and 2) only islands smaller than 100 ha, respectively. We also compiled 25 datasets from the literature to assess the generality of our findings. Island size explained ca half of the variation in species richness. The fit and slope of species–area curves were affected mainly by the range in island size considered, and to a very small extent by the number of islands surveyed. In our literature review, all datasets covering a range of patch sizes larger than 300 ha yielded a positive ISAR, whereas the number of patches alone did not affect the detection of ISARs. We conclude that 1) area per se plays a major role in explaining anuran species richness on forest islands within an Amazonian anthropogenic archipelago; 2) the inferential power of island species–area relationships is severely degraded by sub-optimal sampling designs; 3) at least 10 habitat patches spanning three orders of magnitude in size should be surveyed to yield reliable species–area estimates in patchy systems.     

Evidence for rapid faunal changes on islands in a man-made lake

Summary Few studies of island biogeography have been made on islands in which the time of insularization is precisely known. We tested the effects of island formation on ant species diversity in a man-made lake in South Africa, to determine whether island effects are detectable after only 16 years of insularization. The number of ant species observed at trap-line censuses on islands was significantly correlated with island size (r=0.608; P<0.05) and ant species diversity was generally low compared with similar mainland habitats. Mean species number for all islands, including landbridge islands, was 5.5±3.3 species, and on mainland sites was 7.9±2.85 species. Island effects were more marked on islands <20 ha, which had a mean of 3.3±2.5 species per island. Species number on islands was inversely related to densities of the aggressive Anoplolepis custodiens and A. steingroeveri. These two species were only patchily distributed on mainlands, but these ants were nearly ubiquitous on small islands. Several lines of evidence suggest that this single species domination may be responsible for island effects. Island sites also differed in the number of ant species in different trophic groupings, tending to have fewer granivorous species than the mainland sites, but species in other diet groups were similar in both island and mainland habitats. We conclude that there have been marked changes in the ant faunas on islands smaller than 20 ha apparently due to changes in abundance of the dominant ant species. However, the causes of these changes are unknown.     

Predicting community structure in snakes on Eastern Nearctic islands using ecological neutral theory and phylogenetic methods

Predicting species presence and richness on islands is important for understanding the origins of communities and how likely it is that species will disperse and resist extinction. The equilibrium theory of island biogeography (ETIB) and, as a simple model of sampling abundances, the unified neutral theory of biodiversity (UNTB), predict that in situations where mainland to island migration is high, species-abundance relationships explain the presence of taxa on islands. Thus, more abundant mainland species should have a higher probability of occurring on adjacent islands. In contrast to UNTB, if certain groups have traits that permit them to disperse to islands better than other taxa, then phylogeny may be more predictive of which taxa will occur on islands. Taking surveys of 54 island snake communities in the Eastern Nearctic along with mainland communities that have abundance data for each species, we use phylogenetic assembly methods and UNTB estimates to predict island communities. Species richness is predicted by island area, whereas turnover from the mainland to island communities is random with respect to phylogeny. Community structure appears to be ecologically neutral and abundance on the mainland is the best predictor of presence on islands. With regard to young and proximate islands, where allopatric or cladogenetic speciation is not a factor, we find that simple neutral models following UNTB and ETIB predict the structure of island communities.     

A global comparison of plant invasions on oceanic islands

Oceanic islands have long been considered to be particularly vulnerable to biotic invasions, and much research has focused on invasive plants on oceanic islands. However, findings from individual islands have rarely been compared between islands within or between biogeographic regions. We present in this study the most comprehensive, standardized dataset to date on the global distribution of invasive plant species in natural areas of oceanic islands. We compiled lists of moderate (5–25% cover) and dominant (>25% cover) invasive plant species for 30 island groups from four oceanic regions (Atlantic, Caribbean, Pacific, and Western Indian Ocean). To assess consistency of plant behaviour across island groups, we also recorded present but not invasive species in each island group.We tested the importance of different factors discussed in the literature in predicting the number of invasive plant species per island group, including island area and isolation, habitat diversity, native species diversity, and human development. Further we investigated whether particular invasive species are consistently and predictably invasive across island archipelagos or whether island-specific factors are more important than species traits in explaining the invasion success of particular species.We found in total 383 non-native spermatophyte plants that were invasive in natural areas on at least one of the 30 studied island groups, with between 3 and 74 invaders per island group. Of these invaders about 50% (181 species) were dominants or co-dominants of a habitat in at least one island group. An extrapolation from species accumulation curves across the 30 island groups indicates that the total current flora of invasive plants on oceanic islands at latitudes between c. 35°N and 35°S may eventually consist of 500–800 spermatophyte species, with 250–350 of these being dominant invaders in at least one island group. The number of invaders per island group was well predicted by a combination of human development (measured by the gross domestic product (GDP) per capita), habitat diversity (number of habitat types), island age, and oceanic region (87% of variation explained). Island area, latitude, isolation from continents, number of present, non-native species with a known invasion history, and native species richness were not retained as significant factors in the multivariate models.Among 259 invaders present in at least five island groups, only 9 species were dominant invaders in at least 50% of island groups where they were present. Most species were invasive only in one to a few island groups although they were typically present in many more island groups. Consequently, similarity between island groups was low for invader floras but considerably higher for introduced (but not necessarily invasive) species – especially in pairs of island groups that are spatially close or similar in latitude. Hence, for invasive plants of natural areas, biotic homogenization among oceanic islands may be driven by the recurrent deliberate human introduction of the same species to different islands, while post-introduction processes during establishment and spread in natural areas tend to reduce similarity in invader composition between oceanic islands. We discuss a number of possible mechanisms, including time lags, propagule pressure, local biotic and abiotic factors, invader community assembly history, and genotypic differences that may explain the inconsistent performance of particular invasive species in different island groups.     

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     

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.     

Small off-shore islands can serve as important refuges for endemic beetle conservation

Although large islands generally support a richer insect fauna than small islands, many large islands, which are more often inhabited, have lost numerous species because of human activities and introduced organisms. To clarify the consequences of endemic insect conservation on small islands near inhabited islands, we compared the species richness, abundance, and composition of two beetle groups (Coleoptera: Cerambycidae and Mordellidae) captured using Malaise traps among three islands (Chichijima, 24.0 km²; Anijima, 7.85 km²; Nishijima, 0.49 km²) in the oceanic Ogasawara (Bonin) Island group in the northwestern Pacific during June–July 2006 and 2007. Chichijima, the largest island, is inhabited, while Anijima and Nishijima are not. The numbers of cerambycid and mordellid species previously recorded were positively correlated with island area. However, the total numbers of cerambycid and mordellid species we captured in Malaise traps were not correlated with island area because we were unable to collect many species previously documented on Chichijima. The numbers of cerambycid and mordellid species per trap did not differ significantly among islands and years, although the deviance was well explained by the island variable. We captured greater numbers of cerambycid and mordellid individuals on Chichijima than on Anijima and Nishijima, and the numbers of cerambycid and mordellid individuals per trap significantly differed among islands and between years. Redundancy analysis (RDA) showed that the species composition of cerambycids and mordellids differed among the three islands. Whereas endangered species were rarely captured on Chichijima, alien or non-endemic species were frequently collected. Cerambycid and mordellid beetles on Chichijima may have been deleteriously affected by recent forest disturbance and introduced organisms. Therefore, conserving insect fauna on uninhabited island “refugia” is important for preserving the insect diversity of the Ogasawara Islands.     

Responses of Primates to Landscape Change in Amazonian Land‐bridge islands—a Multi‐scale Analysis

Large hydroelectric dams are one of the current drivers of habitat loss across Amazonian forests. We investigated how the primate community at a hydroelectric dam in Brazilian Amazonia responded to changes in the landscape and local habitat structure of land‐bridge islands after 21 yr of post‐isolation history. The Balbina Dam, constructed in 1986, inundated 3129 km² of primary forests and created more than 3500 variable‐sized islands. We conducted primate and habitat structure surveys on 20 islands from 5 to 1815 ha, and extracted forest patch and landscape metrics for each island. The number of primate species per island varied between 0 and 7 species. Primate composition varied substantially according to both island area and forest cover remaining within the landscape, whereas island area alone was the most significant predictor of richness. Locally, tree density and vertical stratification were the most significant explanatory variables of primate composition and richness. A model containing area effects had the most explanatory power regarding site occupancy for most species. Individually, each species responded differently, with howler and brown capuchin monkeys showing greater tolerance to cope with habitat changes. Body size was also an important predictor of primate occupancy. We recommend protecting large fragments and enhancing the suitability of surrounding habitats to ensure primate conservation in most Neotropical fragmented landscapes. Given the flat topography of hydroelectric reservoirs, which mainly favors the formation of small islands, and the escalating hydropower development plans in Amazonia, our findings provide evidence for pervasive detrimental impacts of dams on primate communities.