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Beta diversity patterns are essential for understanding how biological communities are structured. Geographical and environmental factors, as well as species dispersal ability, are important drivers of beta diversity, but their relative importance may vary across spatial scales. In this study, we evaluate whether beta diversity changes across geographical scales and analyse how different drivers affect turnover patterns of native seed plants in an oceanic archipelago, the Azores (Portugal). Using a 500 × 500 m resolution grid, we selected cells that are covered by one of the following habitats: native forest, naturalized vegetation and seminatural pastures. We calculated species turnover at three spatial scales: 1) between islands, 2) between cells within each island, and finally 3) between cells of each of the habitats of interest in each island. We then calculated the contribution of dispersal syndromes (endozoochory, epizoochory, hydrochory and anemochory) to turnover at each of the scales. Lastly, we assessed the relationship between geographical and climatic distances and habitat type with turnover. Turnover was higher at the smallest spatial scale, particularly in seminatural pastures, and decreased with increasing spatial scales, a pattern potentially associated with the historical fragmentation and current patchy distribution of native forest and seminatural habitats in the Azores. Dispersal syndromes and habitat type had a negligible effect on turnover at all scales. Geographical distance had a positive effect on turnover at all scales, increasing its importance with scale. The relationship between turnover and climatic distance was only significant at the intermediate and small scales in specific islands and habitats. Therefore, scale plays an important role at determining the effect of the drivers of turnover, in particular geographical and climatic distance. These results highlight the need to carefully select the scale of analysis when studying turnover patterns, as well as identifying the potential drivers associated with each spatial scale. [ABSTRACT FROM AUTHOR]

Current understanding of ecological and evolutionary processes underlying island biodiversity is heavily shaped by empirical data from plants and birds, although arthropods comprise the overwhelming majority of known animal species, and as such can provide key insights into processes governing biodiversity. Novel high throughput sequencing (HTS) approaches are now emerging as powerful tools to overcome limitations in the availability of arthropod biodiversity data, and hence provide insights into these processes. Here, we explored how these tools might be most effectively exploited for comprehensive and comparable inventory and monitoring of insular arthropod biodiversity. We first reviewed the strengths, limitations and potential synergies among existing approaches of high throughput barcode sequencing. We considered how this could be complemented with deep learning approaches applied to image analysis to study arthropod biodiversity. We then explored how these approaches could be implemented within the framework of an island Genomic Observatories Network (iGON) for the advancement of fundamental and applied understanding of island biodiversity. To this end, we identified seven island biology themes at the interface of ecology, evolution and conservation biology, within which collective and harmonized efforts in HTS arthropod inventory could yield significant advances in island biodiversity research. [ABSTRACT FROM AUTHOR]

This document is a response to comments made by Carlos A. Góis‐Marques, José Madeira, and Miguel Menezes de Sequeira on a recent paper about plant-insect interactions in the Azores Islands. The authors address the criticisms and irregularities raised by the commenters and provide revised wording where necessary. They also clarify certain points and acknowledge the helpful contributions of the commenters. The authors express their preference for constructive criticism over lengthy public discussions and emphasize the importance of regional journals in the popularization of science. The research was supported by the J. E. Purkyně University Internal Grant Agency. [Extracted from the article]

Research on island species–area relationships (ISAR) has expanded to incorporate functional (IFDAR) and phylogenetic (IPDAR) diversity. However, relative to the ISAR, we know little about IFDARs and IPDARs, and lack synthetic global analyses of variation in form of these three categories of island diversity–area relationship (IDAR). Here, we undertake the first comparative evaluation of IDARs at the global scale using 51 avian archipelagic data sets representing true and habitat islands. Using null models, we explore how richness‐corrected functional and phylogenetic diversity scale with island area. We also provide the largest global assessment of the impacts of species introductions and extinctions on the IDAR. Results show that increasing richness with area is the primary driver of the (non‐richness corrected) IPDAR and IFDAR for many data sets. However, for several archipelagos, richness‐corrected functional and phylogenetic diversity changes linearly with island area, suggesting that the dominant community assembly processes shift along the island area gradient. We also find that archipelagos with the steepest ISARs exhibit the biggest differences in slope between IDARs, indicating increased functional and phylogenetic redundancy on larger islands in these archipelagos. In several cases introduced species seem to have 're‐calibrated' the IDARs such that they resemble the historic period prior to recent extinctions. [ABSTRACT FROM AUTHOR]

The fossil record of the Azores still lacks evidence of terrestrial invertebrates. Based on a study of the collections of Quaternary fossil plant imprints from the Azores, located in the depositories of public collections and also newly conducted field research, several samples of fossil leaves with traces of plant–animal interactions were found. These are evidence of marginal feeding (Phagophytichnus ekowskii) and anophionome mine (Cuniculonomus isp.). One specimen with traces of boring in fossil wood (Xylonichnus trypetus), previously only briefly mentioned in the literature, was also described in detail. In addition to a detailed description of the found traces, the present study also discusses the possible trace‐makers, using a comparison with the recent phytophagous fauna in relation to the original vegetation of the Azores (that includes laurisilva forest). It is thus possible to consider butterfly caterpillars (group Macrolepidoptera) or locust larvae (Orthoptera/Caelifera) as possible originators of marginal feeding. The ophionome mine could have been caused by the larvae of leaf‐miner flies (Agromyzidae) or moths from the group Microlepidoptera (Gracillariidae). Wood‐destroying beetles could be the cause of the boring in fossil wood. [ABSTRACT FROM AUTHOR]

Human activity and land management practices, in particular land use change, have resulted in the global loss of biodiversity. These types of disturbance affect the shape of macroecological patterns, and therefore analyzing these patterns can provide insights into how ecosystems are affected by land use change. We here use arthropod census data from 96 sites at Terceira Island in the Azores archipelago across four different land uses of increasing management intensity: native forest, exotic forest, semi‐natural pasture and intensive pasture, to examine the effects of land use type on three macroecological patterns: the species abundance distribution, the metabolic rate distribution of individuals and the species–area relationship. The maximum entropy theory of ecology (METE) has successfully predicted these patterns across habitats and taxa in undisturbed ecosystems, and thus provides a null expectation for their shapes. Across these patterns, we find that the forest habitats are the best fit by METE predictions, while the semi‐natural pasture is consistently the worst fit, and the intensive pasture is intermediately well fit. We show that the direction of failure of the METE predictions at the pasture sites is likely due to the hyper‐dominance of introduced spider species present there. We hypothesize that the particularly poor fit for the semi‐natural pasture is due to the mix of arthropod communities out of equilibrium, leading to greater heterogeneity in composition and complex dynamics that violate METE's assumption of static state variables. The comparative better fit for the intensive pasture plausibly results from more homogeneous arthropod communities that are well adapted to intensive management, and thus whose state variables are less in flux. Analyzing deviations from theoretical predictions across land use type provides useful information about how land use and disturbance affect ecosystems, and such comparisons could be useful across other habitats and taxa. [ABSTRACT FROM AUTHOR]

Biodiversity accumulates hierarchically by means of ecological and evolutionary processes and feedbacks. Within ecological communities drift, dispersal, speciation, and selection operate simultaneously to shape patterns of biodiversity. Reconciling the relative importance of these is hindered by current models and inference methods, which tend to focus on a subset of processes and their resulting predictions. Here we introduce massive ecoevolutionary synthesis simulations (MESS), a unified mechanistic model of community assembly, rooted in classic island biogeography theory, which makes temporally explicit joint predictions across three biodiversity data axes: (i) species richness and abundances, (ii) population genetic diversities, and (iii) trait variation in a phylogenetic context. Using simulations we demonstrate that each data axis captures information at different timescales, and that integrating these axes enables discriminating among previously unidentifiable community assembly models. MESS is unique in generating predictions of community‐scale genetic diversity, and in characterizing joint patterns of genetic diversity, abundance, and trait values. MESS unlocks the full potential for investigation of biodiversity processes using multidimensional community data including a genetic component, such as might be produced by contemporary eDNA or metabarcoding studies. We combine MESS with supervised machine learning to fit the parameters of the model to real data and infer processes underlying how biodiversity accumulates, using communities of tropical trees, arthropods, and gastropods as case studies that span a range of data availability scenarios, and spatial and taxonomic scales. [ABSTRACT FROM AUTHOR]

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Five decades ago, a landmark paper in Science titled The Cave Environment heralded caves as ideal natural experimental laboratories in which to develop and address general questions in geology, ecology, biogeography, and evolutionary biology. Although the 'caves as laboratory' paradigm has since been advocated by subterranean biologists, there are few examples of studies that successfully translated their results into general principles. The contemporary era of big data, modelling tools, and revolutionary advances in genetics and (meta)genomics provides an opportunity to revisit unresolved questions and challenges, as well as examine promising new avenues of research in subterranean biology. Accordingly, we have developed a roadmap to guide future research endeavours in subterranean biology by adapting a well‐established methodology of 'horizon scanning' to identify the highest priority research questions across six subject areas. Based on the expert opinion of 30 scientists from around the globe with complementary expertise and of different academic ages, we assembled an initial list of 258 fundamental questions concentrating on macroecology and microbial ecology, adaptation, evolution, and conservation. Subsequently, through online surveys, 130 subterranean biologists with various backgrounds assisted us in reducing our list to 50 top‐priority questions. These research questions are broad in scope and ready to be addressed in the next decade. We believe this exercise will stimulate research towards a deeper understanding of subterranean biology and foster hypothesis‐driven studies likely to resonate broadly from the traditional boundaries of this field. [ABSTRACT FROM AUTHOR]

Geographic range size predicts species' responses to land‐use change and intensification, but the reason why is not well established because many correlates of larger geographic ranges, such as realized niche breadth, may mediate species' responses to environmental change. Agricultural land uses (hereafter 'agroecosystems') have warm, dry and more variable microclimates than do cooler and wetter mature forests, so are predicted to filter for species that have warmer, drier and broader fundamental and realized niches. To test these predictions, we estimated species' realized niches, for temperature and precipitation, and geographic range sizes of 764 insect species by matching GBIF occurrence records to global climate layers, and modelled how species presence/absence in mature forest and nearby agroecosystems depend on species' realized niches or geographic ranges. The predicted species niche effects consistently matched the expected direction of microclimatic transition from mature forest to agroecosystems. We found a clear signal that species with preference for warmer and drier climates were more likely to be present in agroecosystems. In addition, the probability that species occurred in different land‐use types was predicted better by species' realized niche than their geographic range size. However, niche effects are often context‐dependent and varied amongst studies, taxonomic groups and regions used in this analysis: predicting which particular aspects of species' realized niche cause sensitivity to land‐use change, and the underpinning mechanisms, remains a major challenge for future research and multiple components of species' realized niches may be important to consider. Using realized niches derived from open‐source occurrence records can be a simple and widely applicable tool to help identify when biodiversity responds to the microclimate component of land‐use change. [ABSTRACT FROM AUTHOR]

A dramatic insect decline has been documented on the grasslands and forests of European or North American mainland. Yet, other parts of the world and other ecosystems remain much less studied with unknown patterns.Using a unique time‐series dataset, we investigate recent trends on abundance and richness of arthropods sampled in Azorean native forest over 6 years (2013–2018).We test the hypothesis that biodiversity erosion drivers are changing the diversity and relative species abundance structure (species abundance distribution, SAD) of endemics, native non‐endemics and exotic species over time. We also examine temporal trends in abundance for each individual species.In contrast with mainland studies, we observed no decline in overall arthropod diversity, but a clear increase in the diversity of exotic arthropods and some evidence of a tendency for decreasing abundance for some endemic species. We also document stronger species turnover for exotic species, but no specific changes in the SAD.We argue that many changes, particularly in unique systems such as islands, will be noticed not at the richness but mostly at compositional level. Special attention should be given to exotic species which are known to be one of the major drivers of biodiversity erosion on islands. [ABSTRACT FROM AUTHOR]

The study of biodiversity started as a single unified field that spanned both ecology and evolution and both macro and micro phenomena. But over the 20th century, major trends drove ecology and evolution apart and pushed an emphasis towards the micro perspective in both disciplines. Macroecology and macroevolution re‐emerged as self‐consciously distinct fields in the 1970s and 1980s, but they remain largely separated from each other. Here, we argue that despite the challenges, it is worth working to combine macroecology and macroevolution. We present 25 fundamental questions about biodiversity that are answerable only with a mixture of the views and tools of both macroecology and macroevolution. [ABSTRACT FROM AUTHOR]

Species abundance distributions (SADs) are one of the most widely used tools in macroecology, and it has become increasingly apparent that many empirical SADs can best be described as multimodal. However, only a few SAD models have been extended to incorporate multiple modes and no software packages are available to fit multimodal SAD models. In this study, we present an extension of the gambin SAD model to multimodal SADs.We derive the maximum likelihood equations for fitting the bimodal gambin distribution and generalize this approach to fit gambin models with any number of modes. We present these new functions, along with additional functions to aid in the analysis of multimodal SADs, within an updated r package ("gambin"; version 2.4.0) that enables the fitting, plotting and evaluating of gambin models with any number of modes.We use a mixture of simulations and empirical datasets to test our new models, including tests of the sensitivity of the model parameters to the number of individuals and the number of species in a sample. We show that the new multimodal gambin models perform well under a variety of circumstances, and that the application of these new models to empirical SAD and other macroecological (e.g., species range size distributions) datasets can provide interesting insights. The updated software package is simple to use and provides straightforward yet flexible statistical analyses of multimodality in SAD‐type datasets. [ABSTRACT FROM AUTHOR]

Aim: To quantify the influence of past archipelago configuration on present‐day insular biodiversity patterns, and to compare the role of long‐lasting archipelago configurations over the Pleistocene to configurations of short duration such as at the Last Glacial Maximum (LGM) and the present‐day. Location: 53 volcanic oceanic islands from 12 archipelagos worldwide—Azores, Canary Islands, Cook Islands, Galápagos, Gulf of Guinea, Hawaii, Madeira, Mascarenes, Pitcairn, Revillagigedo, Samoan Islands and Tristan da Cunha. Time period: The last 800 kyr, representing the nine most recent glacial–interglacial cycles. Major taxa studied: Land snails and angiosperms. Methods: Species richness data for land snails and angiosperms were compiled from existing literature and species checklists. We reconstructed archipelago configurations at the following sea levels: the present‐day high interglacial sea level, the intermediate sea levels that are representative of the Pleistocene and the low sea levels of the LGM. We fitted two alternative linear mixed models for each archipelago configuration using the number of single‐island endemic, multiple‐island endemic and (non‐endemic) native species as a response. Model performance was assessed based on the goodness‐of‐fit of the full model, the variance explained by archipelago configuration and model parsimony. Results: Single‐island endemic richness in both taxonomic groups was best explained by intermediate palaeo‐configuration (positively by area change, and negatively by palaeo‐connectedness), whereas non‐endemic native species richness was poorly explained by palaeo‐configuration. Single‐island endemic richness was better explained by intermediate archipelago configurations than by the archipelago configurations of the LGM or present‐day. Main conclusions: Archipelago configurations at intermediate sea levels—which are representative of the Pleistocene—have left a stronger imprint on single‐island endemic richness patterns on volcanic oceanic islands than extreme archipelago configurations that persisted for only a few thousand years (such as the LGM). In understanding ecological and evolutionary dynamics of insular biota it is essential to consider longer‐lasting environmental conditions, rather than extreme situations alone. [ABSTRACT FROM AUTHOR]

An important factor that hinders the management of non‐native species is a general lack of information regarding the biogeography of non‐natives, and, in particular, their rates of turnover. Here, we address this research gap by analysing differences in temporal beta‐diversity (using both pairwise and multiple‐time dissimilarity metrics) between native and non‐native species, using a novel time‐series dataset of arthropods sampled in native forest fragments in the Azores. We use a null model approach to determine whether temporal beta‐diversity was due to deterministic processes or stochastic colonisation and extinction events, and linear modelling selection to assess the factors driving variation in temporal beta‐diversity between plots. In accordance with our predictions, we found that the temporal beta‐diversity was much greater for non‐native species than for native species, and the null model analyses indicated that the turnover of non‐native species was due to stochastic events. No predictor variables were found to explain the turnover of native or non‐native species. We attribute the greater turnover of non‐native species to source‐sink processes and the close proximity of anthropogenic habitats to the fragmented native forest plots sampled in our study. Thus, our findings point to ways in which the study of turnover can be adapted for future applications in habitat island systems. The implications of this for biodiversity conservation and management are significant. The high rate of stochastic turnover of non‐native species indicates that attempts to simply reduce the populations of non‐native species in situ within native habitats may not be successful. A more efficient management strategy would be to interrupt source‐sink dynamics by improving the harsh boundaries between native and adjacent anthropogenic habitats. [ABSTRACT FROM AUTHOR]

Abstract: Changes in flagellate protist communities of subterranean termite Reticulitermes grassei across different locations were evaluated following four predictions: (i) Rural endemic (Portugal mainland) termite populations will exhibit high diversity of symbionts; (ii) invasive urban populations (Horta city, Faial island, Azores), on the contrary, will exhibit lower diversity of symbionts, showing high similarity of symbiont assemblages through environmental filtering; (iii) recent historical colonization of isolated regions—as the case of islands—will imply a loss of symbiont diversity; and (iv) island isolation will trigger a change in colony breeding structure toward a less aggressive behavior. Symbiont flagellate protist communities were morphologically identified, and species richness and relative abundances, as well as biodiversity indices, were used to compare symbiotic communities in colonies from urban and rural environments and between island invasive and mainland endemic populations. To evaluate prediction on the impact of isolation (iv), aggression tests were performed among termites comprising island invasive and mainland endemic populations. A core group of flagellates and secondary facultative symbionts was identified. Termites from rural environments showed, in the majority of observed colonies, more diverse and abundant protist communities, probably confirming prediction (i). Corroborating prediction (ii), the two least diverse communities belong to termites captured inside urban areas. The Azorean invasive termite colonies had more diverse protist communities than expected and prediction (iii) which was not verified within this study. Termites from mainland populations showed a high level of aggressiveness between neighboring colonies, in contrast to the invasive colonies from Horta city, which were not aggressive to neighbors according to prediction (iv). The symbiotic flagellate community of R. grassei showed the ability to change in a way that might be consistent with adaptation to available conditions, possibly contributing to optimization of the colonization of new habitats and spreading of its distribution area, highlighting R. grassei potential as an invasive species. [ABSTRACT FROM AUTHOR]

Abstract: Motivation: Past sea level fluctuations have shaped island area and archipelago configuration. The availability of global high‐resolution data on bathymetry and past sea levels allows reconstruction of island palaeo‐geography. Studies on the role of palaeo‐area often consider only the Last Glacial Maximum, which neglects the dynamics of island fusion and fission resulting from cyclic sea level fluctuations throughout the Quaternary. Currently, no global database is available to test the role of changing island geographies driven by Quaternary sea level change on evolutionary, ecological and archaeological patterns. However, data on island palaeo‐environments is crucial for understanding insular biodiversity distributions and human settlement patterns. Here, we present the Palaeo‐Islands and Archipelago Configuration (PIAC) database, containing sea level‐driven palaeo‐geography changes over the late Quaternary of 178 islands in 27 archipelagos, and discuss its relevance, limitations and uncertainties. The R functions developed to create the PIAC database are provided to allow calculations for other islands, time steps, sea‐level curves and higher spatio‐temporal resolutions. Main types of variables contained: Polygon shapefiles with archipelago configuration and tables with palaeo‐area per island. Spatial location and grain: The database has a global representation, with 27 archipelagos being covered: Aldabra, Azores, Balearic Islands, California Channel Islands, Canary Islands, Cape Verde, Comoros, Cook Islands, Crozet Islands, Dutch Caribbean, Galápagos, Gulf of Guinea, Hawaii, Inner Seychelles, Juan Fernández, Kuril Islands, Madeira, Marianas, Marquesas, Mascarenes, Phoenix Islands, Pitcairn Islands, Prince Edward Islands, Revillagigedo, Samoan Islands, Society Islands and Tristan da Cunha. All data are at 1 km2 spatial resolution. Time period and grain: The focus of this paper is on the last 35 kyr; data for the last 140 kyr are also provided. The grain is 1 kyr temporal resolution. Level of measurement: Data are per island, grouped per archipelago. Software format: The data were produced in the R programming language. [ABSTRACT FROM AUTHOR]

Aim Land-use change typically goes hand in hand with the introduction of exotic species, which mingle with indigenous species to form novel assemblages. Here, we compare the functional structure of indigenous and exotic elements of ground-dwelling arthropod assemblages across four land-uses of varying management intensity. Location Terceira Island (Azores, North Atlantic). Methods We used pitfall traps to sample arthropods in 36 sites across the four land-uses and collated traits related to dispersal ability, body size and resource use. For both indigenous and exotic species, we examined the impact of land-uses on trait diversity and tested for the existence of non-random assembly processes using null models. We analysed differences in trait composition among land-uses for both indigenous and exotic species with multivariate analyses. We used point-biserial correlations to identity traits significantly correlated with specific land-uses for each element. Results We recorded 86 indigenous and 116 exotic arthropod species. Under high-intensity land-use, both indigenous and exotic elements showed significant trait clustering. Trait composition strongly shifted across land-uses, with indigenous and exotic species being functionally dissimilar in all land-uses. Large-bodied herbivores dominated exotic elements in low-intensity land-uses, while small-bodied spiders dominated exotic elements in high-intensity land-uses. In contrast, with increasing land-use intensity, indigenous species changed from functionally diverse to being dominated by piercing and cutting herbivores. Main conclusions Our study revealed two main findings: first, in high-intensity land-uses, trait clustering characterized both indigenous and exotic elements; second, exotic species differed in their functional profile from indigenous species in all land-use types. Overall, our results provide new insights into the functional role of exotic species in a land-use context, suggesting that, in agricultural landscape, exotic species may contribute positively to the maintenance of some ecosystem functions. [ABSTRACT FROM AUTHOR]

Questions We addressed three questions: (1) what is the variation of α- and β-diversity of bryophyte communities across spatial scales in an elevational gradient; (2) is spatial variation characteristic of different phylogenetic (mosses/liverworts) and/or ecological groups (terrestrials/epiphytes); and (3) what is the contribution of species richness and species replacement ( true turnover) to β-diversity along the elevational gradient? Location La Palma, Canary Islands, Spain. Methods An elevational transect spanning 0-2200 m a.s.l was sampled at altitudinal intervals of ca. 200 m, two 10 m × 10 m plots were established at each elevation and inside each plot three 2 m × 2 m quadrats were randomly selected. Within each quadrat, bryophytes were sampled in three replicate microplots of 10 cm × 5 cm, for each substrate: soil, rocks, leaves, humus, decaying wood and tree trunks at three heights. We evaluated α- and β-diversity at plot and elevational level for all bryophytes and for phylogenetic and ecological groups. Values of β-diversity were further decomposed into replacement and richness differences. We tested the relationship between α- and β-diversity and elevation using linear models with and without a quadratic term (elevation2) in an AICc-based framework. Results A total of 121 species were recorded along the transect. Alpha-diversity showed a hump-shaped pattern with elevation for all bryophytes, liverworts and epiphytic species, however, terrestrial species and mosses displayed no clear pattern. Species replacement was the main driver of β-diversity for all bryophytes and also for mosses and terrestrial species, and at different scales (elevational belt and quadrat). The contribution of species replacement increased with elevation, except for epiphytes and terrestrials. Conclusions Our study contributes towards an understanding of the spatial organization of bryophyte diversity at different scales along an insular elevational gradient. We showed that bryophyte diversity was mostly influenced by species replacement at the largest scale. The observed differences in β-diversity decomposition between bryophyte phylogenetic and ecological groups could be dependent on climatic conditions and substrate availability. The strong differences observed along the elevational gradient related to true turnover are especially important in the current global change scenario. [ABSTRACT FROM AUTHOR]

Obtaining fundamental biodiversity metrics such as alpha, beta and gamma diversity for arthropods is often complicated by a lack of prior taxonomic information and/or taxonomic expertise, which can result in unreliable morphologically based estimates. We provide a set of standardized ecological and molecular sampling protocols that can be employed by researchers whose taxonomic skills may be limited, and where there may be a lack of robust a priori information regarding the regional pool of species. These protocols combine mass sampling of arthropods, classification of samples into parataxonomic units ( PUs) and selective sampling of individuals for mt DNA sequencing to infer biological species. We sampled ten lowland rainforest plots located on the volcanic oceanic island of Réunion (Mascarene archipelago) for spiders, a group with limited taxonomic and distributional data for this region. We classified adults and juveniles into PUs and then demonstrated the reconciliation of these units with presumed biological species using mt DNA sequence data, ecological data and distributional data. Because our species assignment protocol is not reliant upon prior taxonomic information, or taxonomic expertise, it minimizes the problem of the Linnean shortfall to yield diversity estimates that can be directly compared across independent studies. Field sampling can be extended to other arthropod groups and habitats by adapting our field sampling protocol accordingly. [ABSTRACT FROM AUTHOR]

Using a series of standardized sampling plots within forest ecosystems in remote oceanic islands, we reveal fundamental differences between the structuring of aboveground and belowground arthropod biodiversity that are likely due to large-scale species introductions by humans. Species of beetle and spider were sampled almost exclusively from single islands, while soil-dwelling Collembola exhibited more than tenfold higher species sharing among islands. Comparison of Collembola mitochondrial metagenomic data to a database of more than 80 000 Collembola barcode sequences revealed almost 30% of sampled island species are genetically identical, or near identical, to individuals sampled from often very distant geographic regions of the world. Patterns of mt DNA relatedness among Collembola implicate human-mediated species introductions, with minimum estimates for the proportion of introduced species on the sampled islands ranging from 45% to 88%. Our results call for more attention to soil mesofauna to understand the global extent and ecological consequences of species introductions. [ABSTRACT FROM AUTHOR]

ABSTRACT The general dynamic model of oceanic island biogeography ( GDM) has added a new dimension to theoretical island biogeography in recognizing that geological processes are key drivers of the evolutionary processes of diversification and extinction within remote islands. It provides a dynamic and essentially non-equilibrium framework generating novel predictions for emergent diversity properties of oceanic islands and archipelagos. Its publication in 2008 coincided with, and spurred on, renewed attention to the dynamics of remote islands. We review progress, both in testing the GDM's predictions and in developing and enhancing ecological-evolutionary understanding of oceanic island systems through the lens of the GDM. In particular, we focus on four main themes: ( i) macroecological tests using a space-for-time rationale; ( ii) extensions of theory to islands following different patterns of ontogeny; ( iii) the implications of GDM dynamics for lineage diversification and trait evolution; and ( iv) the potential for downscaling GDM dynamics to local-scale ecological patterns and processes within islands. We also consider the implications of the GDM for understanding patterns of non-native species diversity. We demonstrate the vitality of the field of island biogeography by identifying a range of potentially productive lines for future research. [ABSTRACT FROM AUTHOR]

Land-use change has profoundly impacted pollinator communities throughout the world. However, the processes through which it acts on pollinator diversity and composition are still poorly understood, especially in highly vulnerable island ecosystems., In this study, we investigated the distribution, abundance, richness and composition of flower-visiting insects to assess their response to land-use change in Terceira Island (Azores)., Flower-visiting insects were sampled over 2 years using a standardised protocol along 50 transects across five different habitats corresponding to a land-use gradient. Insect species were classified as indigenous or exotics. We assessed changes across habitats using multiple diversity indices, species abundance distribution models ( SAD) and species composition metrics (β-diversity), along with plant species composition., We observed that indigenous flower-visiting insects were dominant, both in abundance and species richness, across the entire land-use gradient. Species diversity varied only slightly across the gradient. SADs were lognormal in all habitats, with very few truly common and rare flower-visiting insects and a prevalence of species of intermediate abundance. Species replacement was significantly higher mainly between the two most contrasting habitats (i.e. natural forests and intensive pastures) but was significantly correlated with species replacement of host plant species across the gradient., Our results revealed that the Azorean flower-visiting insect communities were highly simplified across the entire gradient with little difference between habitats. In the absence of strong exotic competitors, indigenous flower-visiting insects expand their range and occupy new anthropogenic habitats, also facilitating the expansion of a large number of exotic plant species. [ABSTRACT FROM AUTHOR]

We investigated whether the equilibrium theory of island biogeography ( ETIB) can be applied to the meiofauna of groundwater-fed springs. We tested whether copepod species richness was related with spring area, discharge, and elevation. Additionally, five hypotheses are tested based on species distribution patterns, dispersal ability, and life-history characteristics of several guilds (stygobiotic, nonstygobiotic, cold stenotherm, and noncold stenotherm species). Thirty springs in the central Apennines (Italy) were considered. A multimodel selection procedure was applied to select best-fit models using both ordinary least-squares regressions and autoregressive models. Mantel tests were used to investigate the impact of spatial autocorrelation in determining interspring similarity (ßsor), pure turnover (ßsim), intersite nestedness (ßnest = ßsor − ßsim), and matrix nestedness (measured using NODF and other metrics). Explicit consideration of spatial correlations reduced the importance of predictors of overall species richness, noncold stenotherm species (both negatively affected by elevation), cold stenotherm species, and nonstygobiotic species, but increased the importance of area for the stygobiotic species. We detected nested patterns in all cases, except for the stygobites. Interspring distances were positively correlated with ßsor and ßnest (but not with ßsim) for the entire data set and for nonstygobiotic, cold stenotherm, and noncold stenotherm species. In the case of stygobites, interspring geographical distances were marginally correlated with ßsor and no correlation was found for ßsim and ßnest. We found support for ETIB predictions about species richness, which was positively influenced by area and negatively by elevation (which expresses the size of source of immigrants). Low turnover and high nestedness are consistent with an equilibrium scenario mainly regulated by immigration and extinction. Stygobites, which include many distributional and evolutionary relicts, have a low capability to disperse through the aquifers and tend to be mainly confined to the springs where they drifted out and were trapped by springbed sediments. [ABSTRACT FROM AUTHOR]

Aim Higher-elevation areas on islands and continental mountains tend to be separated by longer distances, predicting higher endemism at higher elevations; our study is the first to test the generality of the predicted pattern. We also compare it empirically with contrasting expectations from hypotheses invoking higher speciation with area, temperature and species richness. Location Thirty-two insular and 18 continental elevational gradients from around the world. Methods We compiled entire floras with elevation-specific occurrence information, and calculated the proportion of native species that are endemic ('percent endemism') in 100-m bands, for each of the 50 elevational gradients. Using generalized linear models, we tested the relationships between percent endemism and elevation, isolation, temperature, area and species richness. Results Percent endemism consistently increased monotonically with elevation, globally. This was independent of richness-elevation relationships, which had varying shapes but decreased with elevation at high elevations. The endemism-elevation relationships were consistent with isolation-related predictions, but inconsistent with hypotheses related to area, richness and temperature. Main conclusions Higher per-species speciation rates caused by increasing isolation with elevation are the most plausible and parsimonious explanation for the globally consistent pattern of higher endemism at higher elevations that we identify. We suggest that topography-driven isolation increases speciation rates in mountainous areas, across all elevations and increasingly towards the equator. If so, it represents a mechanism that may contribute to generating latitudinal diversity gradients in a way that is consistent with both present-day and palaeontological evidence. [ABSTRACT FROM AUTHOR]

Aim To infer colonization and speciation history for a closely related complex of nine species within the enigmatic Canary Island Laparocerus weevil radiation of 128 species. Using molecular dating and the spatial and temporal context that islands provide, we evaluate the possible explanations of incomplete lineage sorting and gene flow for the origin of shared genetic variation among species from different islands. Location Canary Islands (Gran Canaria, Tenerife, La Palma and El Hierro). Methods We collected a total of 173 specimens from 37 sites distributed across the four islands. Phylogenetic analyses of mitochondrial (COII) and nuclear (ITS2) sequence data and molecular dating techniques were used to infer the origin of the group in the archipelago and their history of colonization and differentiation. Results Gran Canaria appeared to be the geographical origin of the complex. An unexpected result was that mtDNA revealed each of the single species on La Palma and El Hierro to be the product of more than one colonization event from more than one source island. In both cases nuclear ITS2 data revealed these multiple colonizations to have been followed by admixture. Main conclusions The two gene trees present very different topologies, with a rather simple colonization history required to explain the pattern of nuclear gene relationships, while the mtDNA gene tree implicates a much more complex history of colonization. Explanations of incomplete lineage sorting are ruled out and a history of colonization and speciation for the L. tessellatus complex involving genetic admixture is inferred. [ABSTRACT FROM AUTHOR]

1. Colour polymorphisms are common across animals and are often the result of complex selection regimes. Philaenus spumarius ( Linnaeus) ( Hemiptera, Aphrophoridae) shows a widely studied dorsal colour polymorphism with several described phenotypes whose variation in their occurrence and frequency, as well as their maintenance across time, have been reported. Several selective influences have been suggested to play a role, but the mechanisms underlying the maintenance of this polymorphism are still poorly understood. 2. To explore the adaptive significance of the colour polymorphism of P. spumarius, an experiment was conducted in captivity under semi-natural conditions to measure survival, reproductive success, and duration of egg maturation. 3. It was found that there was higher longevity, a higher number of oviposition events, and a higher number of eggs laid by trilineatus phenotype females than by typicus and marginellus, supporting previous reports of an increase in trilineatus frequency during the season. The duration of egg maturation did not differ among phenotypes. 4. The higher longevity and fertility of the trilineatus phenotype may compensate, for example, the higher rate of attack by parasitoids and/or higher solar radiation reflectance in this phenotype, which have already been reported in previous studies, constituting a possible mechanism for the maintenance of this polymorphism. [ABSTRACT FROM AUTHOR]

A key challenge in island biogeography is to quantity the role of dispersal in shaping biodiversity patterns among the islands of a given archipelago. Here, we propose such a framework. Dispersal within oceanic archipelagos may be conceptualized as a spatio-temporal process dependent on: (1) the spatial distribution of islands, because the probability of successful dispersal is inversely related to the spatial distance between islands and (2) the chronological sequence of island formation that determines the directional asymmetry of dispersal (hypothesized to be predominantly from older to younger islands). From these premises, directional network models may be constructed, representing putative connections among islands. These models may be translated to eigenfunctions in order to be incorporated into statistical analysis. The framework was tested with 12 datasets from the Hawaii, Azores, and Canaries. The explanatory power of directional network models for explaining species composition patterns, assessed by the Jaccard dissimilarity index, was compared with simpler time-isolation models. The amount of variation explained by the network models ranged from 5.5% (for Coleoptera in Hawaii) to 60.2% (for Pteridophytes in Canary Islands). In relation to the four studied taxa, the variation explained by network models was higher for Pteridophytes in the three archipelagos. By the contrary, small fractions of explained variation were observed for Coleoptera (5.5%) and Araneae (8.6%) in Hawaii. Time-isolation models were, in general, not statistical significant and explained less variation than the equivalent directional network models for all the datasets. Directional network models provide a way for evaluating the spatio-temporal signature of species dispersal. The method allows building scenarios against which hypotheses about dispersal within archipelagos may be tested. The new framework may help to uncover the pathways via which species have colonized the islands of a given archipelago and to understand the origins of insular biodiversity. [ABSTRACT FROM AUTHOR]

The species abundance distribution (SAD) has been a central focus of community ecology for over fifty years, and is currently the subject of widespread renewed interest. The gambin model has recently been proposed as a model that provides a superior fit to commonly preferred SAD models. It has also been argued that the model's single parameter (α) presents a potentially informative ecological diversity metric, because it summarises the shape of the SAD in a single number. Despite this potential, few empirical tests of the model have been undertaken, perhaps because the necessary methods and software for fitting the model have not existed. Here, we derive a maximum likelihood method to fit the model, and use it to undertake a comprehensive comparative analysis of the fit of the gambin model. The functions and computational code to fit the model are incorporated in a newly developed free-to-download R package (gambin). We test the gambin model using a variety of datasets and compare the fit of the gambin model to fits obtained using the Poisson lognormal, logseries and zero-sum multinomial distributions. We found that gambin almost universally provided a better fit to the data and that the fit was consistent for a variety of sample grain sizes. We demonstrate how α can be used to differentiate intelligibly between community structures of Azorean arthropods sampled in different land use types. We conclude that gambin presents a flexible model capable of fitting a wide variety of observed SAD data, while providing a useful index of SAD form in its single fitted parameter. As such, gambin has wide potential applicability in the study of SADs, and ecology more generally. [ABSTRACT FROM AUTHOR]

It is becoming increasingly apparent that the species abundance distribution of many ecological communities contains multiple modes, a phenomenon that has been largely overlooked. Here, we test for multiple modes in the species abundance distribution using a combination of one, two and three mode Poisson lognormal distributions and an extensive arthropod dataset from the Azores. We consider the abundance distribution of twelve native laurisilva forest fragments and the combination of fragments within five islands, allowing us to detect whether patterns are consistent across scales. An information theoretic approach is employed to determine the best model in each case. To explore the processes driving multimodal abundance distributions we tested various potential mechanisms. We classified species as core if they are present in over half the fragments in our study, and as satellite if they are sampled in fewer than half the fragments. Furthermore, species are classified based on body size, whether they are indigenous (i.e. endemic or native non-endemic) or introduced to the Azores, abundance in land uses other than native forest, and dispersal ability. We find that models incorporating multiple modes perform best for most fragments and islands. A large number of communities are bimodal, comprising a mode of very rare species and a mode of relatively common species. Deconstructing the full assemblages into their constituent subsets reveals that the combination of ecologically different groups of species into a single sample underpins the multimodal pattern. Specifically, the rarer mode prevailingly contains a higher proportion of satellite taxa, introduced species and species that are more adapted to anthropogenic land uses that surround the native forest. [ABSTRACT FROM AUTHOR]

Summary: Complete sampling of all dimensions of biodiversity is a formidable task, even for small areas. Undersampling is the norm, and the underquantification of diversity is a common outcome. Estimators of taxon diversity (TD) are widely used to correct for undersampling. Yet, no similar strategy has been developed for phylogenetic (PD) or functional (FD) diversity. We propose three ways of estimating PD and FD, building on estimators originally developed for TD: (i) correcting PD and FD values based on the completeness of TD; (ii) fitting asymptotic functions to accumulation curves of PD and FD; and (iii) adapting nonparametric estimators to PD and FD data. Using trees as a common framework for the estimation of PD and FD, we tested the approach with European mammal and Azores Islands arthropod data. We demonstrated that different methods were able to considerably reduce the undersampling bias and often correctly estimated true diversity using a fraction of the samples necessary to reach complete sampling. Besides the utility of knowing the true diversity of an assemblage from incomplete samples, the use of estimators may present further advantages. For instance, comparisons between sites or time periods are possible only if either sampling is complete or sampling effort is equivalent and sufficient to allow sensible comparisons. Also, as PD and FD asymptote faster than TD, comparisons between these different dimensions may require unbiased values. The framework now proposed combines taxon, phylogenetic and functional diversity into a single framework, offering a tool for future developments involving these different facets of biological diversity. [ABSTRACT FROM AUTHOR]

Aim To propose a unified framework for quantifying taxon ( Tβ), phylogenetic ( Pβ) and functional ( Fβ) beta diversity via pairwise comparisons of communities, which allows these types of beta diversity to be partitioned into ecologically meaningful additive components. Location Global, with case studies in Europe and the Azores archipelago. Methods Using trees as a common representation for taxon, phylogenetic and functional diversity, we partition total beta diversity (βtotal) into its replacement (turnover, βrepl) and richness difference (βrich) components according to which part of a global tree was shared by or unique to communities that were being compared. We demonstrate the application of this framework using artificial and empirical examples (mammals in Europe and epigean arthropods in the Azores). Results Our empirical examples show that comparing Pβ and Fβ with the most commonly used Tβ revealed previously hidden patterns of beta diversity. More importantly, we demonstrate that partitioning Pβtotal and Fβtotal into their respective βrepl and βrich components facilitates the detection of more complex patterns than using the overall coefficients alone, further elucidating the different forces operating in community assembly. Main conclusions The methods presented here allow the integration and full comparison of Tβ, Pβ and Fβ. They provide a tool for effectively disentangling the replacement (turnover) and richness difference components of the different biodiversity facets within the same methodological framework. [ABSTRACT FROM AUTHOR]

Beta diversity and nestedness are central concepts of ecology and biogeography and evaluation of their relationships is in the focus of contemporary ecological and conservation research. Beta diversity patterns are originated from two distinct processes: the replacement (or turnover) of species and the loss (or gain) of species leading to richness differences. Nested distributional patterns are generally thought to have a component deriving from beta diversity which is independent of replacement processes. Quantification of these phenomena is often made by calculating a measure of beta diversity, and the resulting value being subsequently partitioned into a contribution by species replacement plus a fraction shared by beta diversity and nestedness. Three methods have been recently proposed for such partitioning, all of them based on pairwise comparisons of sites. In this paper, the performance of these methods was evaluated on theoretical grounds and tested by a simulation study in which different gradients of dissimilarity, with known degrees of species replacement and species loss, were created. Performance was also tested using empirical data addressing land-use induced changes in endemic arthropod communities of the Terceira Island in the Azores. We found that the partitioning of βcc (dissimilarity in terms of the Jaccard index) into two additive fractions, β-3 (dissimilarity due to species replacement) plus βrich (dissimilarity due to richness differences) reflects the species replacement and species loss processes across the simulated gradients in an ecologically and mathematically meaningful way, whilst the other two methods lack mathematical consistency and prove conceptually self-contradictory. Moreover, the first method identified a selective local extinction process for endemic arthropods, triggered by land-use changes, while the latter two methods overweighted the replacement component and led to false conclusions. Their basic flaw derives from the fact that the proposed replacement and nestedness components (deemed to account for species loss) are not scaled in the same way as the measure that accounts for the total dissimilarity (Sørensen and Jaccard indices). We therefore recommend the use of βcc=β-3+βrich, since its components are scaled in the same units and their responses are proportional to the replacement and the gain/loss of species. [ABSTRACT FROM AUTHOR]

Aim We explore the island species-area relationships ( ISARs) of several plant groups differing in dispersal ability. We examine whether: (i) the ISAR slope is higher for less dispersive groups (diminishing from not-wind-dispersed seed plants to wind-dispersed seed plants, pteridophytes and bryophytes); and (ii) the regional richness is higher than that predicted by the ISAR of its constituent islands for groups with lower dispersal ability. Additionally, we relate both patterns to the compositional dissimilarity (beta diversity) between islands. Location The Macaronesian archipelagos of the Azores, Madeira and Canary Islands. Methods ISARs were estimated using the conventional power model. Differences in slopes among taxa were analysed through ANCOVA tests. We assessed the deviation of the total richness of the Macaronesian flora from that predicted by the ISAR and calculated several measures of beta diversity (turnover, nestedness-resultant dissimilarity and nestedness). Analyses were repeated after excluding the two Canarian islands of Lanzarote and Fuerteventura, which have unique climatic and geological conditions in the region. Results All ISAR models were significant except for pteridophytes (for which it was significant when excluding Lanzarote and Fuerteventura). ISAR slopes did not differ among taxa. Regional richness followed ISAR predictions for bryophytes and pteridophytes when Lanzarote and Fuerteventura were excluded, while the total number of Macaronesian seed plants (particularly not-wind-dispersed species) always fell above the ISAR. Turnover was higher in seed plants than in bryophytes and pteridophytes, and the opposite occurred with nestedness. Main conclusions Differences in dispersal ability influence the compositional dissimilarity between these islands, but not the rate of species increase with area. This may be because ISAR slopes are mostly determined by within-island processes of species accumulation, while differences in dispersal generate different between-island patterns. The lack of relationship between the ISAR slope and species replacement or nestedness prevents its use as a proxy for beta diversity. [ABSTRACT FROM AUTHOR]

. 1. The aim of this paper was to investigate the arthropod colonisation of a recently erupted volcano in the framework of a general model of colonisation kinetics. 2. We analysed the diversity of arthropod communities at three locations on Faial Island (Azores) using a well-defined disturbance gradient: (i) a site that is new land added by the eruption of Capelinhos Volcano of 1957; (ii) a site moderately affected by this eruption; and (iii) a pristine site not affected. We calculated the recolonisation times at the disturbed sites using species richness at the undisturbed site as an equilibrium value (last erupted 900-1000 years ago). 3. Species with different distributional ranges (endemic, native non-endemic and introduced) have different colonisation kinetics. Introduced exotic species were particularly rapid in colonising the erupted volcano, reaching a number of species greater than that observed in the undisturbed area. By contrast, native non-endemic species had more difficulty in recolonising the erupted area, and no endemic has reached it. The volcano community is dominated by a few species with high abundance and shows low richness and strong dominance in comparison with the undisturbed community. The moderately disturbed site supports a rich and well-balanced arthropod community. 4. Although the erupted volcano has a species richness even slightly higher than the undisturbed site, this is a consequence of the high colonisation ability of introduced species, and its arthropod community is strongly disharmonic. [ABSTRACT FROM AUTHOR]

Aim We used a phylogenetic framework to examine island colonization and predictions pertaining to differentiation within Macaronesian Tarphius (Insecta, Coleoptera, Zopheridae), and explain the paucity of endemics in the Azores compared with other Macaronesian archipelagos. Specifically, we test whether low diversity in the Azores could be due to recent colonization (phylogenetic lineage youth), cryptic speciation (distinct phylogenetic entities within species) or the young geological age of the archipelago. Location Macaronesian archipelagos (Azores, Madeira and the Canary Islands), northern Portugal and Morocco. Methods Phylogenetic analyses of mitochondrial and nuclear genes of Tarphius beetles of the Azores, other Macaronesian islands and neighbouring continental areas were used to investigate the origin of island biodiversity and to compare patterns of colonization and differentiation. A comparative nucleotide substitution rate test was used to select the appropriate substitution rate to infer clade divergence times. Results Madeiran and Canarian Tarphius species were found to be more closely related to each other, while Azorean taxa grouped separately. Azorean taxa showed concordance between species and phylogenetic clades, except for species that occur on multiple islands, which segregated by island of origin. Divergence time estimates revealed that Azorean Tarphius are an old group and that the most recent intra-island speciation event on Santa Maria, the oldest island, occurred between 3.7 and 6.1 Ma. Main conclusions Our phylogenetic approach provides new evidence to understand the impoverishment of Azorean endemics: (1) Tarphius have had a long evolutionary history within the Azores, which does not support the hypothesis of fewer radiation events due to recent colonization; (2) the current taxonomy of Azorean Tarphius does not reflect common ancestry and cryptic speciation is responsible for the underestimation of endemics; (3) intra-island differentiation in the Azores was found only in the oldest island, supporting the idea that young geological age of the archipelago limits the number of endemics; and (4) the lack of evidence for recent intra-island diversification in Santa Maria could also explain the paucity of Azorean endemics. Phylogenetic reconstructions of other species-rich taxa that occur on multiple Macaronesian archipelagos will reveal whether our conclusions are taxon specific, or of a more general nature. [ABSTRACT FROM AUTHOR]

Species abundance distributions are an essential tool in describing the biodiversity of ecological communities. We now know that their shape changes as a function of the size of area sampled. Here we analyze the scaling properties of species abundance distributions by using the moments of the logarithmically transformed number of individuals. We find that the moments as a function of area size are well fitted by power laws and we use this pattern to estimate the species abundance distribution for areas larger than those sampled. To reconstruct the species abundance distribution from its moments, we use discrete Tchebichef polynomials. We exemplify the method with data on tree and shrub species from a 50 ha plot of tropical rain forest on Barro Colorado Island, Panama. We test the method within the 50 ha plot, and then we extrapolate the species abundance distribution for areas up to 5 km2. Our results project that for areas above 50 ha the species abundance distributions have a bimodal shape with a local maximum occurring for the singleton classes and that this maximum increases with sampled area size. [ABSTRACT FROM AUTHOR]

Carine & Schaefer ( Journal of Biogeography, 2010, 37, 77-89) suggest that the lack of past climate oscillations in the Azores may have contributed to the low plant endemism in this archipelago compared to that of the Canary Islands, a pattern they term the Azorean diversity enigma. Here we challenge their hypothesis, and discuss how the particular characteristics of the Azores may have driven current diversification patterns in this archipelago. We argue that the restricted number of Azorean endemic species and their wide distribution is explicable by the geological, geographical and ecological attributes of the archipelago. That is, the Azores are too young, too small, and too environmentally homogeneous to have hosted many in situ diversification events, so they do not host as many endemic species as other Macaronesian archipelagos, such as Madeira and especially the Canary Islands. [ABSTRACT FROM AUTHOR]

Aim To examine whether island parasitoid faunas are biased towards generalists when compared with the mainland and their species pool, and to evaluate the effects of climate, island characteristics and regional factors on the relative proportions of idiobionts (i.e. generalists) and koinobionts (i.e. specialists) of two parasitic wasp families, Braconidae and Ichneumonidae. Location Seventy-three archipelagos distributed world-wide. Methods We used data on the distribution and biology obtained from a digital catalogue and several literature sources. We related level of generalism, measured as the ratio between the number of idiobiont and koinobiont species, to climatic, physiographic and regional factors using generalized linear models. We compared models by means of Akaike weighting, and evaluated the spatial structure of their residuals. We used partial regressions to determine whether the final models account for all latitudinal structure in the level of generalism. Results Islands host comparatively more idiobionts than continental areas. Although there is a latitudinal gradient in the level of generalism of island faunas correlating with both environmental factors and island characteristics, the most important determinant of island community structure is their source pool. This effect is stronger for ichneumonids, where generalism is higher in the Indomalayan region, arguably due to the higher diversity of endophytic hosts in its large rain forests. Main conclusions The level of generalism of island parasitoid faunas is largely constrained by regional factors, namely by the structure of the species pool, which emphasizes the importance of including regional processes in our understanding of the functioning of ecological communities. The fact that generalist species are more predominant in islands with a large cover of rain forests pinpoints the importance of the indirect effects of ecological requirements on community structure, highlighting the complex nature of geographical gradients of diversity. [ABSTRACT FROM AUTHOR]

Island communities are exposed to several evolutionary and ecological processes that lead to changes in their diversity and structure compared to mainland biotas. These phenomena have been observed for various taxa but not for parasitoids, a key group in terms of community diversity and functioning. Here we use the parasitoid communities associated with the moth Acroclita subsequana (Lepidoptera: Tortricidae) in the Macaronesian region, to test whether species richness differs between islands and mainland, and whether island parasitoid faunas are biased towards generalist species. Host larvae were collected on several islands and adjacent mainland, carefully searched for ectoparasitoid larvae and dissected to recover any endoparasitoids. Parasitoids were classified as idiobionts, which usually have a wide host range (i.e. generalists), or koinobionts that are considered specialists. Mainland species richness was lower than expected by chance, with most of the species being koinobionts. On the other hand, island communities showed a greater proportion of idiobiont species. Overall parasitism rates were similar between islands and mainland, but islands had higher rates of parasitism by idiobionts than expected by chance, and mainland areas showed the highest koinobiont parasitism rates. Th ese results suggest that island parasitoid communities are dominated by generalists, in comparison to mainland communities. Several hypotheses may explain this pattern: (1) generalist parasitoids might have better dispersal abilities; (2) they may be less constrained by ' sequential dependencies ' ; and (3) island parasitoids probably have fewer competitors and/or predators, thus favouring the establishment of generalists. New studies including multiple hosts, other habitats, and/or more islands are necessary to identify which of these processes shape island parasitoid communities. [ABSTRACT FROM AUTHOR]

Aim To establish the extent to which archipelagos follow the same species–area relationship as their constituent islands and to explore the factors that may explain departures from the relationship. Location Thirty-eight archipelagos distributed worldwide. Methods We used ninety-seven published datasets to create island species–area relationships (ISARs) using the Arrhenius logarithmic form of the power model. Observed and predicted species richness of an archipelago and of each of its islands were used to calculate two indices that determined whether the archipelago followed the ISAR. Archipelagic residuals ( ArcRes) were calculated as the residual of the prediction provided by the ISAR using the total area of the archipelago, standardized by the total richness observed in the archipelago. We also tested whether any characteristic of the archipelago (geological origin and isolation) and/or taxon accounts for whether an archipelago fits into the ISAR or not. Finally, we explored the relationship between ArcRes and two metrics of nestedness. Results The archipelago was close to the ISAR of its constituent islands in most of the cases analysed. Exceptions arose for archipelagos where (i) the slopes of the ISAR are low, (ii) observed species richness is higher than expected by the ISAR and/or (iii) distance to the mainland is small. The archipelago's geological origin was also important; a higher percentage of oceanic archipelagos fit into their ISAR than continental ones. ArcRes indicated that the ISAR underpredicts archipelagic richness in the least isolated archipelagos. Different types of taxon showed no differences in ArcRes. Nestedness and ArcRes appear to be related, although the form of the relationship varies between metrics. Main conclusions Archipelagos, as a rule, follow the same ISAR as their constituent islands. Therefore, they can be used as distinct units themselves in large-scale biogeographical and macroecological studies. Departure from the ISAR can be used as a crude indicator of richness-ordered nestedness, responsive to factors such as isolation, environmental heterogeneity, number and age of islands. [ABSTRACT FROM AUTHOR]

Habitat destruction is the leading cause of species extinctions. However, there is typically a time-lag between the reduction in habitat area and the eventual disappearance of the remnant populations. These “surviving but ultimately doomed” species represent an extinction debt. Calculating the magnitude of such future extinction events has been hampered by potentially inaccurate assumptions about the slope of species–area relationships, which are habitat- and taxon-specific. We overcome this challenge by applying a method that uses the historical sequence of deforestation in the Azorean Islands, to calculate realistic and ecologically-adjusted species–area relationships. The results reveal dramatic and hitherto unrecognized levels of extinction debt, as a result of the extensive destruction of the native forest:>95%, in<600 yr. Our estimations suggest that more than half of the extant forest arthropod species, which have evolved in and are dependent on the native forest, might eventually be driven to extinction. Data on species abundances from Graciosa Island, where only a very small patch of secondary native vegetation still exists, as well as the number of species that have not been found in the last 45 yr, despite the extensive sampling effort, offer support to the predictions made. We argue that immediate action to restore and expand native forest habitat is required to avert the loss of numerous endemic species in the near future. [ABSTRACT FROM AUTHOR]

Aim Researchers measuring beta diversity have rarely concerned themselves with the problems of how complete the species lists of studied communities are, and of how the varying degrees of completeness can actually change estimates of beta diversity. No comprehensive assessment has been made regarding the behaviour of most beta diversity indices when applied to incomplete samples, a situation which is more common than usually recognized. Our objective was to assess the behaviour and robustness of a number of beta diversity measures for incidence data from undersampled communities. Location Mainland Portugal and the Azorean archipelago (North Atlantic). Methods Data from intensive sampling of spiders in mainland Portugal and arthropods in Azores were collected. We examined the properties of 15 beta diversity measures developed for incidence data. We simulated varying degrees of completeness, whereas computing beta diversity for selected pairs of samples. The robustness of these beta diversity accumulation curves was assessed for the purpose of finding the best measures for undersampled communities. Results The Harrison et al.β-2 and the Williams β-3 are particularly robust to undersampling. These measures are also insensitive to differences of alpha diversity (species richness) between communities, and therefore to nestedness. Colwell & Coddington βcc and the related Jaccard βj and Gaston et al.βg performed best of the measures sensitive to alpha diversity. They performed poorly, however, when compared communities exhibited very low values of beta diversity. In such cases, the Routledge βr performed the best. Main conclusions No index was found to perform without bias in all circumstances. Overall, β-2, β-3 and βcc (or related measures βj and βg) are recommended as they seem to be the most robust to undersampling. [ABSTRACT FROM AUTHOR]

A major focus of geographical ecology and macroecology is to understand the causes of spatially structured ecological patterns. However, achieving this understanding can be complicated when using multiple regression, because the relative importance of explanatory variables, as measured by regression coefficients, can shift depending on whether spatially explicit or non-spatial modeling is used. However, the extent to which coefficients may shift and why shifts occur are unclear. Here, we analyze the relationship between environmental predictors and the geographical distribution of species richness, body size, range size and abundance in 97 multi-factorial data sets. Our goal was to compare standardized partial regression coefficients of non-spatial ordinary least squares regressions (i.e. models fitted using ordinary least squares without taking autocorrelation into account; “OLS models” hereafter) and eight spatial methods to evaluate the frequency of coefficient shifts and identify characteristics of data that might predict when shifts are likely. We generated three metrics of coefficient shifts and eight characteristics of the data sets as predictors of shifts. Typical of ecological data, spatial autocorrelation in the residuals of OLS models was found in most data sets. The spatial models varied in the extent to which they minimized residual spatial autocorrelation. Patterns of coefficient shifts also varied among methods and datasets, although the magnitudes of shifts tended to be small in all cases. We were unable to identify strong predictors of shifts, including the levels of autocorrelation in either explanatory variables or model residuals. Thus, changes in coefficients between spatial and non-spatial methods depend on the method used and are largely idiosyncratic, making it difficult to predict when or why shifts occur. We conclude that the ecological importance of regression coefficients cannot be evaluated with confidence irrespective of whether spatially explicit modelling is used or not. Researchers may have little choice but to be more explicit about the uncertainty of models and more cautious in their interpretation. [ABSTRACT FROM AUTHOR]

Aim R. J. Whittaker et al. recently proposed a ‘general dynamic model of oceanic island biogeography’ (GDM), providing a general explanation of island biodiversity patterns by relating fundamental biogeographical processes – speciation, immigration, extinction – to area (A) and time (T; maximum island geological age). We adapt their model, which predicts a positive relationship with area combined with a humped relationship to time (designated the ATT2 model), to study the factors promoting diversification on the Azores for several arthropod groups. Location The Azorean archipelago (North Atlantic; 37–40° N, 25–31° W). Methods We use the number of single-island endemics (SIEs) as a measure of diversification, to evaluate four different predictions for the variation in SIEs between different islands, derived from the GDM theory and our knowledge of the fauna and history of the Azores. We calculated the number of SIEs for seven out of the nine Azorean islands and six groups of species (all arthropods, beetles, cavernicolous and non-cavernicolous species, and taxa with high and low dispersal abilities). Several variables accounting for island characteristics (area, geological age, habitat diversity and isolation) and generalized linear models were used to evaluate the reliability of each prediction. Results A linear and positive relationship between SIEs and an AT (area + time) model was the most parsimonious explanation for overall arthropod diversification. However, cavernicolous species showed the opposite pattern (more SIEs inhabiting the youngest islands). Also, isolation was an important predictor of diversification for all groups except for the species with high dispersal ability; while the former were negatively related to the distance from the main source of colonizing lineages (Santa Maria island in most cases), the latter were related to area. Dispersal ability was also a key factor affecting the diversification of most groups of species. Main conclusions In general, the diversification of Azorean arthropods is affected by age, area and isolation. However, different groups are affected by these factors in different ways, showing radically different patterns. Although the ATT2 model fails to predict the diversification pattern of several groups, it provides a framework for integrating these deviations into a general theory. Further improvements of the GDM theory need to take into account the particular traits of each group and the role of isolation in shaping island diversity. [ABSTRACT FROM AUTHOR]

The species–area relationship is one of the strongest empirical generalizations in geographical ecology, yet controversy persists about some important questions concerning its causality and application. Here, using more accurate measures of island surface size for five different island systems, we show that increasing the accuracy of the estimation of area has negligible impact on the fit and form of the species–area relationship, even though our analyses included some of the most topographically diverse island groups in the world. In addition, we show that the inclusion of general measurements of environmental heterogeneity (in the form of the so-called choros model), can substantially improve the descriptive power of models of island species number. We suggest that quantification of other variables, apart from area, that are also critical for the establishment of biodiversity and at the same time have high explanatory power (such as island age, distance, productivity, energy, and environmental heterogeneity), is necessary if we are to build up a more predictive science of species richness variation across island systems. [ABSTRACT FROM AUTHOR]