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Aim Long‐term ecological data provide a stepped frame of island ecosystem transformation after successive waves of human colonization, essential to determine conservation and management baselines. However, the timing and ecological impact of initial human settlement on many islands is still poorly known. Here, we report analyses from a 4800‐year sedimentary sequence from Gran Canaria (Canary Islands), with the goal of disentangling forest responses to natural fire from early human pressure on the island. Location La Calderilla, a volcanic maar caldera at 1,770 m a.s.l. on Gran Canaria. Taxon Plants and fungi. Methods A core from the caldera infill was analysed for sediment properties, pollen, micro‐ and macrocharcoal, with radiocarbon and biochronology dating. Fossil data were statistically zoned and interpreted with the help of cross‐correlation and ordination analyses. Surface samples and a pollen–vegetation training set were used as modern analogues for vegetation reconstruction. Results Before human settlement (4,800–2,000 cal. yr bp), pine (Pinus canariensis) pollen dominated. Extensive dry pine forests characterized the highlands, although with temporary declining phases, followed by prompt (sub‐centennial scale) recovery. Towards 2,280 cal. yr bp there was a shift to open vegetation, marked by an increase in coprophilous spores. Coincidental with independent evidence of human settlement in the pine belt (2,000–470 cal. yr bp) there was a decline of pine and a peak in charcoal. Following historic settlement (470–0 cal. yr bp), pollen producers from anthropogenic habitats, secondary vegetation and coprophilous fungi increased in abundance, reflecting higher pressure of animal husbandry and farming. Modern moss polsters reflect extensive reforestation since 1950 ce (Common Era). Main conclusions From 4,800 cal. yr bp, the pristine vegetation covering the Gran Canaria highlands was a mosaic of dry pine forests and open vegetation. The pine forests sustained intense fires, which may well have promoted habitat diversity. Human interference was initiated around 2,280 cal. yr bp probably by recurrent cultural firing and animal husbandry, triggering a steady trend of forest withdrawal and expansion of grasses and scrubs, until the final disappearance of the pine forest locally in the 20th century. Grasslands were found to be of ancient cultural origin in the summit areas of Gran Canaria, although they underwent an expansion after the Castilian Conquest.

Plant colonization of islands may be limited by the availability of symbionts, particularly arbuscular mycorrhizal (AM) fungi, which have limited dispersal ability compared to ectomycorrhizal and ericoid (EEM) as well as orchid mycorrhizal (ORC) fungi. We tested for such differential island colonization within contemporary angiosperm floras worldwide. We found evidence that AM plants experience a stronger mycorrhizal filter than other mycorrhizal or non-mycorrhizal (NM) plant species, with decreased proportions of native AM plant species on islands relative to mainlands. This effect intensified with island isolation, particularly for non-endemic plant species. The proportion of endemic AM plant species increased with island isolation, consistent with diversification filling niches left open by the mycorrhizal filter. We further found evidence of humans overcoming the initial mycorrhizal filter. Naturalized floras showed higher proportions of AM plant species than native floras, a pattern that increased with increasing isolation and land-use intensity. This work provides evidence that mycorrhizal fungal symbionts shape plant colonization of islands and subsequent diversification., Delavaux et al. uses global plant databases to examine how the types of mycorrhizal fungi influence the colonization of plants on islands. Their results show that arbuscular mycorrhizal (AM) symbiosis limits plant species’ establishment on islands more than other mycorrhizal associations, and that this initial filter of AM plant species impacts diversification and plant invasion risks.

The destruction of forest for agricultural expansion has created a vast estate of human-modified land in tropical regions. One group of organisms that are particularly vulnerable to the loss of forest habitat are insectivorous birds. Despite this, few conservation strategies have been identified for this group in human-modified landscapes. We survey the use of 104 isolated trees by insectivorous birds in rural Assam, India. We used an information theoretic model comparison approach to determine the important variables driving insectivorous bird diversity within these isolated trees. Our work demonstrates that the conservation of large trees in human-modified landscapes may play an important role in maintaining bird diversity and ecological function beyond the forest edge. More specifically, we found that isolated Ficus trees hold assemblages with particularly high insectivore abundance, richness and functional diversity when compared to other isolated fruit and large trees. We argue that, where present, Ficus trees should be actively conserved in human-modified landscapes to maintain the composition of insectivore communities in a “Ficus first” strategy.

Tropical dry islands are currently facing major challenges derived from anthropogenic and climatic pressures. However, their trajectories of environmental change, which could provide relevant information applicable to biodiversity conservation, remain understudied. This is mainly due to poor micro-fossil preservation and irregular sediment deposition. Multi-proxy palaeoecological analyses spanning decades to 1000s of years can add perspective as to how vegetation, fungal communities, and the fauna responded to previous natural and anthropogenic disturbances. In Sao Nicolau, Cabo Verde, we used palaeoecological methods to analyse a highland soil profile (1000 m asl) dated to 5900 cal yr BP. We analysed how vegetation (abundances in pollen of native and introduced species, and leaf wax n-alkanes), ferns and fungal communities (abundance of non-pollen palynomorphs) varied over time in relation to fire (charcoal concentration) and erosion regimes (grain sizes and elemental composition). Between 5000 and 400 cal yr BP the highlands held native woody taxa such as Euphorbia tuckeyana, Dracaena draco subsp. caboverdeana, and Ficus, taxa that can be used for future reforestation programmes. From 400 cal yr BP to the present day, replacement of native taxa by introduced and cultivated taxa (Pinus, Eucalyptus, Asystasia, Opuntia) has occurred. Vegetation burning and grazing caused loss of vegetation and erosion, acting as conjoined drivers of scrubland degradation. This dataset helps to set historically contextualised restoration goals such as the re-introduction of native species, monitoring of recently introduced species and control of free grazing. This can serve as a model system for the conservation of tropical dry islands' biodiversity.

Cabo Verde was the first tropical archipelago colonized by Europeans. Historians have suggested that the first colonizers initiated archipelago-wide ecosystem degradation, loss of vegetation cover, and erosion. However, the human–environment interactions that led to the archipelago's current environmental status remain poorly understood. Here, we report the first palaeoecological study of past vegetation change and disturbance regimes for Cabo Verde. We present a 2130-yr old sediment sequence from a volcanic caldera (Cova de Paul) located at ~1200 m asl on Santo Antao Island, for which we analyzed fossil pollen, non-pollen palynomorphs (NPPs), charcoal, silica bodies, and grain size distribution. Our analyses do not show evidence of the presence of temperate, tropical or subtropical forests growing on the summits of Santo Antao in pre-human times. The pollen record shows that scrubland and grasslands dominated the highlands and underwent compositional changes ca. 1850 and 1300 cal yr BP. These shifts overlap with erosion phases and are linked to intensified seasonality. Steady rates of sedimentation marked the period 1230 to ca. 350 cal yr BP, but an increase in charcoal concentrations indicate a drying phase. Increases in regional and local fire, peaks of coprophilous fungi and the presence of New World crop pollen (Zea mays) are interpreted as the onset of Portuguese settlement of the highlands after 450 BP. Sustained erosion between ca. 350 and 100 cal yr BP indicate soil degradation, and the pollen record shows the increase in introduced herbaceous taxa (Rosaceae, Centaurea, Verbenaceae) and exotic tree taxa (e.g. Pinus), while shifting abundances of coprophilous fungi indicate changes in land-use. The record shows stabilization of soils in the last century due to recent afforestation of the highlands. Further palaeoecological studies have the potential to provide further detail of the long-term dynamics of Cabo Verde ecosystems and to inform conservation initiatives.

Aims The present-day biogeography of European amphibians has been hypothesized to have arisen from range expansion and recolonization of the northern part of the continent from southern late Pleistocene refugia, such that northern species generally possess large ranges while southerly species are mostly small-ranged. Here we test the hypothesis that these patterns are likely to be underpinned by biological traits associated with dispersal ability. We do this by analysing data for anurans and urodeles, the two main groups of European amphibians. Location Europe. Methods We built a database of biological traits (body size, fecundity, life span, habitat specialization) of European amphibians, excluding island endemics. We mapped the basic macroecological patterns of range size and position, and analysed the causal pathways for range size using structural equation models (SEMs). Results Amphibian species with a small range size are largely restricted to areas in southern Europe associated with putative Pleistocene refugia. Those present in northern Europe are exclusively large-ranged species whose distributions extend all the way from southern Europe. SEMs explained 54% of range size variation for anurans, with long life span and high fecundity being influential explanatory variables, and explained 61% of range size variation within urodeles, with measures of species fecundity being influential. Main conclusions Species that have successfully recolonized the north following deglaciation have the largest ranges for both groups of amphibians. These large-ranged species generally possess traits that indicate the potential for rapid range expansion, with differences apparent in the balance of traits between anurans and urodeles. The traits linked to northern distributions (and large range size) appear to be a mix of r and K traits, indicating that intermediate life-history strategies have proved to be optimal for range expansion into northern regions. These results integrate species biology with geographical history in explaining present-day patterns of species distribution, range size and diversity.

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.

Forest restoration is an increasingly important tool to offset and indeed reverse global deforestation rates. One low cost strategy to accelerate forest recovery is conserving scattered native trees that persist across disturbed landscapes and which may act as seedling recruitment foci. Ficus trees, which are considered to be critically important components of tropical ecosystems, may be particularly attractive to seed dispersers in that they produce large and nutritionally rewarding fruit crops. Here, we evaluate the effectiveness of remnant Ficus trees in inducing forest recovery compared to other common trees. We studied the sapling communities growing under 207 scattered trees, and collected data on seed rain for 55 trees in a modified landscape in Assam, India. We found that Ficus trees have more sapling species around them (species richness = 140.1 ± 9.9) than non-Ficus trees (79.5 ± 12.9), and significantly more saplings of shrub and large tree species. Sapling densities were twice as high under Ficus trees (median = 0.06/m2) compared to non-Ficus (0.03/m2), and seed rain densities of non-parent trees were significantly higher under Ficus trees (mean = 12.73 ± 3/m2/wk) than other fruit or non-zoochorous trees (2.19 ± 0.97/m2/wk). However, our regression model found that canopy area, used as a proxy for tree size, was the primary predictor of sapling density, followed by remnant tree type. These results suggest that large trees, and in particular large Ficus trees, may be more effective forest restoration agents than other remnant trees in disturbed landscapes, and therefore the conservation of these trees should be prioritized. Hindi Abstract वशविक वनोनमलन दर को अधोमख करन क लिए वन पनरदधार वासतविकता म एक तजी स बढता साधन ह। बिखर हए दशी पडो का सरकषण, जो कि विकषबध भदशय म बच हए ह और अकर निरमाण क कदर क रप म कारय कर सकत ह, वनो को तजी स सही हालत म लान की एक कम लागत की रणनीति ह। गलर परजाति क पड, विशष रप स बीज परसारको को आकरषित कर सकन वाल और उनम स व जो बड तथा पोषण की दषटि स लाभपरद फल उतपादित करत ह, उषणकटिबधीय पारिसथितिक ततर क अति महतवपरण घटक मान जात ह। यहा हमन वन पनरदधार उतपररण पर अनय साधारण और गलर परजाति क पडो क तलनातमक परभाव का मलयाकन किया ह। हमन भारत क असम राजय म एक रपातरित भदशय म २०७ बिखर हए पडो क नीच उगन वाल पौधो क समदायो का अधयन किया और ५५ पडो म बीज वषटि क तथय एकतर किय। हमन गर-गलर परजाति क पडो (परजाति परचरता=७९.५±१२.९) की तलना म गलर परजाति क पडो क नीच सारथक रप स अधिक (१४०.१±९.९) पौधो, झाडियो व बड पडो की परजातियो को उगत हए पाया। गर-गलर परजाति क पडो (मधयमान=०.०३/मी.२) की तलना म गलर परजाति क पडो क नीच उगन वाल पौधो का घनतव दो गना जयादा (०.०६/मी.२) था और गर-पतक पडो का बीज घनतव भी सारथक रप स फल या गर-जतपरकीरणित पडो (औसत=२.१९±०.९७/मी.२/सपताह) की तलना म गलर परजाति क पडो क नीच जयादा (१२.७३±३/मी.२/सपताह) पाया गया। हालाकि, हमार परतिगमन ततर न पाया कि बच हए पड परकार, पड आकार क एक परतिनिधि क रप म परयोग किय गए वितान कषतर क बाद, पौध घनतव का पराथमिक कारक थ। य परिणाम बतलात ह कि, बड पड और विशष रप स गलर परजाति क पड, समभवत: अधिक परभावी रप स विकषबध भदशय म वन पनरदधार अभिकरता हो सकत ह और इस कारण इन पडो का सरकषण पराथमिकता क आधार पर किया जाना चाहिय।

Aims: The general dynamic model (GDM) of oceanic island biogeography predicts how biogeographical rates, species richness and endemism vary with island age, area and isolation. Here, we used a simulation model to assess whether the isolation‐related predictions of the GDM may arise from low‐level process at the level of individuals and populations. Location: Hypothetical volcanic oceanic islands. Methods: Our model considers (a) an idealized island ontogeny, (b) metabolic constraints and (c) stochastic, spatially explicit and niche‐based processes at the level of individuals and populations (plant demography, dispersal, competition, mutation and speciation). Isolation scenarios involved varying the distance to mainland and the dispersal ability of the species pool. Results: For all isolation scenarios, we obtained humped temporal trends for species richness, endemic richness, proportion of endemic species derived from within‐island radiation, number of radiating lineages, number of species per radiating lineage and biogeographical rates. The proportion of endemics derived from mainland–island differentiation and of all endemics steadily increased over time. Extinction rates of endemic species peaked later than for non‐endemic species. Species richness and the number of endemics derived from mainland–island differentiation decreased with isolation as did rates of colonization, mainland–island differentiation and extinction. The proportion of all endemics and of radiated endemics, the number of radiated endemics, of radiating lineages, and of species per radiating lineage and the within‐island radiation rate all increased with isolation. Main conclusions: Our results lend strong support to most of the isolation‐related GDM predictions. New insights include an increasing proportion of endemics, particularly those arising from mainland–island differentiation, across isolation scenarios, as well as extinction trends of endemics differing from the overall extinction rates, with a much later peak. These results demonstrate how simulation models focusing on low ecological levels provide tools to assess biogeographical‐scale predictions and to develop more detailed predictions for further empirical tests.

Th e widespread destruction and fragmentation of natural habitats around the world creates a strong incentive to understand how species and communities respond to such pressures. Th e vast majority of research into habitat fragmentation has focused solely on species presence or absence. However, analyses using innovative functional methodologies off er the prospect of providing new insights into the key questions surrounding community structure in fragmented systems. A key topic in fragmentation research is nestedness (i.e. the ordered composition of species assemblages involving a signifi cant tendency for packing of the presence – absence matrix into a series of proper subsets). To date, nestedness analyses have been concerned solely with nestedness of species membership. Here, we capitalize on the publication of a recent nestedness index (traitNODF) in which the branch lengths of functional dendrograms are incorporated into the standard NODF nestedness index. Using bird community data from 18 forest-habitat-island studies, and measurements of eight continuous functional traits from over 1000 bird species, we conduct the fi rst synthetic analysis of nestedness from a functional perspective (i.e. a nestedness analysis which incorporates how similar species are in terms of their ecological traits). We use two null models to test the signifi cance of any observed functional nestedness, and investigate the role of habitat island area in driving functional nestedness. We also determine whether functional nestedness is driven primarily by species composition or by diff erences in species ’ traits. We found that the majority (94%) of datasets were functionally nested by island area when a permutation null model was used, although only 11 – 22% of datasets were signifi cantly functionally nested when a more conservative fi xed-fi xed null model was used. Species composition was always the most important driver of functional nestedness, but the eff ect of diff erences in species traits was occasionally quite large. Our results isolate the importance of island area in driving functional nestedness where it does occur and show that habitat loss results in the ordered loss of functional traits. Th is analysis demonstrates the potential insights that may derive from testing for ordered patterns of functional diversity.

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.

Aim The concept of nestedness is important in determining the relative contribution to overall system diversity of different habitat patches within a fragmented system. Much of the previous work on nestedness has focused on islands within oceans (islands sensu stricto). The largest analysis of habitat island systems to date found significant nestedness to be a near universal feature, but the methods used have since been criticized as inappropriate. Thus, there is a need for an updated, critical examination of the prevalence, underlying drivers and implications of nestedness in multiple habitat island systems. Location Global. Methods Here, we collate 97 datasets from published habitat island studies, comprising multiple taxa. We use the NODF metric (nestedness metric based on overlap and decreasing fill) to estimate nestedness and determine significance using the four-step proportional–proportional algorithm to simulate presences/absence matrices. We investigate the role of habitat island area in driving observed nestedness. We use linear modelling to examine the impact of dataset characteristics on the degree of nestedness and assess the conservation biogeographic implications of nestedness in relation to strategic conservation planning. Results Significant nestedness occurred in only 9% of systems, whilst anti-nestedness (i.e. datasets less nested than expected by chance) occurred in 16% of systems. For the majority of datasets found to be significantly nested, we observed a relationship with fragment area, suggesting that structured extinctions may be important in determining the composition of certain habitat island communities. We found that the degree of nestedness in an archipelago is an important consideration for systematic conservation planning. Main conclusions Significant nestedness is considerably less common in habitat islands than previously reported. Strategic guidance for conservation planning should proceed on a case by case basis, and previous conservation recommendations based on the assumption of significant nestedness in most fragmented landscapes may need to be re-evaluated.

© The Author(s) 2017., Species abundance distributions (SAD) are central to the description of diversity and have played a major role in the development of theories of biodiversity and biogeography. However, most work on species abundance distributions has focused on one single spatial scale. Here we used data on arthropods to test predictions obtained with computer simulations on whether dispersal ability influences the rate of change of SADs as a function of sample size. To characterize the change of the shape of the SADs we use the moments of the distributions: the skewness and the raw moments. In agreement with computer simulations, low dispersal ability species generate a hump for intermediate abundance classes earlier than the distributions of high dispersal ability species. Importantly, when plotted as function of sample size, the raw moments of the SADs of arthropods have a power law pattern similar to that observed for the SAD of tropical tree species, thus we conjecture that this might be a general pattern in ecology. The existence of this pattern allows us to extrapolate the moments and thus reconstruct the SAD for larger sample sizes using a procedure borrowed from the field of image analysis based on scaled discrete Tchebichef moments and polynomials., This work was funded by the FCT project MOMENTOS (PTDC/ BIA-BIC/5558/2014). AMCS was supported by a Marie Curie Intra-European Fellowship (IEF 331623 ‘COMMSTRUCT’) and by a Juan de la Cierva Fellowship (IJCI-2014-19502) funded by the Spanish ‘Ministerio de Economía y Competitividad’. IRA was supported by a FCT Fellowship (SFRH/BPD/102804/2014)

Summary 1. The species abundance distribution (SAD) is an important concept in ecology, and much work has focused on the SAD in a theoretical context. However, less focus has been placed on the utility of SADs in applied ecology and biodiversity management, which therefore forms the focus of the present article. We illustrate that study of the SAD allows inferences beyond those that flow from many simple diversity indices, enabling workers to identify patterns in the commonness and rarity of species in a community. 2. First, we discuss how incorporating SAD analyses into the study of ecological communities can generate useful information for the management of biodiversity. In particular, we argue that deconstructing ecological assemblages into various subsets and analysing how each subset contributes to the overall SAD can reveal patterns of interest to managers. Secondly, we review the many applications of SADs in applied ecological fields, including disturbance ecology, conservation planning and conservation biological control. 3. Using examples we show that the SAD can be useful in applied ecology as it is visually intuitive, easy to implement in a broad variety of ecological contexts, and does not require substantial species-specific data. We provide a summary of the various methods available for plotting the SAD and illustrate how each method provides information of value for applied ecologists. 4. Using empirical and simulated data, we show that the SAD can provide early warning of the effects of disturbance on ecological communities and that a number of SAD models represent useful tools for comparing communities in a management context. 5. Synthesis and applications. Applied ecologists require tools that allow for relatively quick assessments of ecosystem health and/or the success of management prescriptions aimed at ameliorating the effects of disturbance. We demonstrate that that the methods reviewed herein provide such tools and that the species abundance distribution (SAD) has additional applied uses beyond traditional applications in disturbance ecology. We hope that this synthesis will provide a catalyst for advancing a more utilitarian SAD research agenda.

Summary 1. The integration of species distributions and evolutionary relationships is one of the most rapidly moving research fields today and has led to considerable advances in our understanding of the processes underlying biogeographical patterns. Here, we develop a set of metrics, the specific overrepresentation score (SOS) and the geographic node divergence (GND) score, which together combine ecological and evolutionary patterns into a single framework and avoids many of the problems that characterize community phylogenetic methods in current use. 2. This approach goes through each node in the phylogeny and compares the distributions of descendant clades to a null model. The method employs a balanced null model, is independent of phylogeny size, and allows an intuitive visualization of the results. 3. We demonstrate how this novel implementation can be used to generate hypotheses for biogeographical patterns with case studies on two groups with well-described biogeographical histories: a local-scale community data set of hummingbirds in the North Andes, and a large-scale data set of the distribution of all species of New World flycatchers. The node-based analysis of these two groups generates a set of intuitively interpretable patterns that are consistent with current biogeographical knowledge. 4. Importantly, the results are statistically tractable, opening many possibilities for their use in analyses of evolutionary, historical and spatial patterns of species diversity. The method is implemented as an upcoming R package nodiv, which makes it accessible and easy to use.

Aim The species–area relationship (SAR) is widely used in conservation science to predict the number of species likely to go extinct as a result of habitat loss. Often, studies employing the SAR use total species richness as the dependent variable. However, this overlooks the fact that habitat specialists and generalists differ in their susceptibility to habitat loss. We undertook a synthetic review of 23 habitat island datasets for birds to determine the impact of habitat generalists on the SAR. Location Global. Methods We sourced 19 habitat island datasets from the literature and combined these data with four of our own empirically gathered datasets. For each dataset, we classified all bird species as either forest habitat specialists or generalists. We then fitted the power SAR model (log–log and nonlinear forms) to the specialists, generalists and all species for each dataset and compared the resulting model parameters. We compared differences in the rate of change in richness with area between specialists and generalists using the first derivative of a multimodel SAR. Results We found that the slope of the power model was steeper for habitat specialists in the majority of datasets, and this difference was significant in 15 and 16 of the 23 datasets, for the nonlinear and log–log forms of the power model, respectively. Comparison of the multimodel SAR curve derivatives revealed further differences in the rate of change in species richness with area between subsets. Main conclusions The z values of both forms of the power model of the specialists' SARs were generally larger, often considerably so, than the values used in most SAR studies predicting extinctions from habitat loss. Thus, studies that have used z values derived from SAR studies using total richness may be underestimating the impact of habitat loss on specialist species, which are likely to be those of greatest conservation concern.

cited By 3; International audience; There is a broad consensus that habitat disturbance and introduction of non-indigenous species may dramatically modify community structure, particularly in insular ecosystems. However, it is less clear whether emergent macroecological patterns are similarly affected. The positive interspecific abundance-occupancy relationship (IAOR) is one of the most pervasive macroecological patterns, yet has rarely been analyzed for oceanic island assemblages. We use an extensive dataset for arthropods from six islands within the Azorean archipelago to test: (1) whether indigenous and non-indigenous species are distributed differently within the IAOR; and (2) to the extent that they are, can differences can be attributed to two indices of disturbance. We implemented modeling averaged methods using five of the most common IAOR models to derive an averaged IAOR fit for each island. After testing if species colonization status (indigenous versus non-indigenous) may explain the residuals of the IAOR, we identified true negative and positive outliers and tested the effect of colonization status on the likelihood of a species being a positive or negative outlier. We found that the indigenous and non-indigenous species are randomly distributed on both sides of the overall IAOR. Only for Flores Island, were non-indigenous species more aggregated than indigenous species. We were unable to detect a meaningful relationship between deviation from the IAOR and disturbance, despite the undoubted impact of both severe habitat loss and non-indigenous species on these oceanic islands. Our results show that the non-indigenous species have been integrated alongside indigenous species in the contemporary Azorean arthropod communities such that they are mostly undetectable by the study of the IAOR. © 2013 Elsevier Masson SAS.

It has become increasingly recognized that multiple processes can generate similar shapes of species abundance distributions (SADs), with the result that the fit of a given SAD model cannot unambiguously provide evidence in support of a given theory or model. An alternative approach to comparing the fit of different SAD models to data from a single site is to collect abundance data from a variety of sites, and then build models to analyse how different SAD properties (e.g. form, skewness) vary with different predictor variables. Such a biogeographical approach to SAD research is potentially very revealing, yet there has been a general lack of interest in SADs in the biogeographical literature. In this Perspective, we address this issue by highlighting findings of recent analyses of SADs that we consider to be of intrinsic biogeographical interest. We use arthropod data drawn from the Azorean archipelago to further highlight how analyses of SAD form and function may be biogeographically informative. We hope that, by reviewing a number of novel approaches, our article may prove to be a catalyst for a greater interest in analysing SADs in biogeography. The Azorean data are from project BALA, funded by Direcção Regional dos Recursos Florestais (Project: 17.01-080203, 1999–2004) and PO Açores 2016–2018 PROAAcXXIs. P.A.V.B. was partly supported by project FCT-PTDC/BIA-BIC/5558/2014–MOMENTOS. info:eu-repo/semantics/publishedVersion

Aims: The 50th anniversary of the publication of the seminal book, The Theory of Island Biogeography, by Robert H. MacArthur and Edward O. Wilson, is a timely moment to review and identify key research foci that could advance island biology. Here, we take a collaborative horizon-scanning approach to identify 50 fundamental questions for the continued development of the field. Location: Worldwide. Methods: We adapted a well-established methodology of horizon scanning to identify priority research questions in island biology, and initiated it during the Island Biology 2016 conference held in the Azores. A multidisciplinary working group prepared an initial pool of 187 questions. A series of online surveys was then used to refine a list of the 50 top priority questions. The final shortlist was restricted to questions with a broad conceptual scope, and which should be answerable through achievable research approaches. Results: Questions were structured around four broad and partially overlapping island topics, including: (Macro)Ecology and Biogeography, (Macro)Evolution, Community Ecology, and Conservation and Management. These topics were then subdivided according to the following subject areas: global diversity patterns (five questions in total); island ontogeny and past climate change (4); island rules and syndromes (3); island biogeography theory (4); immigration–speciation–extinction dynamics (5); speciation and diversification (4); dispersal and colonization (3); community assembly (6); biotic interactions (2); global change (5); conservation and management policies (5); and invasive alien species (4). Main conclusions: Collectively, this cross-disciplinary set of topics covering the 50 fundamental questions has the potential to stimulate and guide future research in island biology. By covering fields ranging from biogeography, community ecology and evolution to global change, this horizon scan may help to foster the formation of interdisciplinary research networks, enhancing joint efforts to better understand the past, present and future of island biotas., B.C.E. was supported by Spanish grant CGL2013-42589-P awarded to B.C.E. by the MINECO and co-financed by FEDER, and Spanish grant S20141203_002597 awarded to B.C.E. by the Organismo Autónomo Parques Nacionales of the MAGRAMA.

Next-Generation Biogeography In 1876, Alfred Russel Wallace mapped the zoogeographical regions of the world, based on the distributions and taxonomic relationships of broadly defined mammalian families. Wallace's classification of zoogeographical regions became a cornerstone of modern biogeography and a reference for a wide variety of biological disciplines, including global biodiversity and conservation sciences. Holt et al. (p. 74 , published online 20 December) present a next-generation map of wallacean zoogeographic regions, incorporating phylogenetic data on >20,000 vertebrate species to discern and characterize their natural biogeographic patterns.

The marked gradient in species diversity between the equator (where it is highest) and the poles (where it is lowest) suggests a simple environmental explanation for this phenomenon. However, as Willis and Whittaker point out in their Perspective, the environmental variables driving species diversity are different depending on the size and time scale over which species diversity is measured.

Aim We collate and analyse data for land snail diversity and endemism, as a means of testing the explanatory power of the general dynamic model of oceanic island biogeography (GDM): a theoretical model linking trends in species immigration, speciation and extinction to a generalized island ontogeny. Location Eight oceanic archipelagos: Azores, Canaries, Hawaii, Galapagos, Madeira, Samoa, Society, Tristan da Cunha. Methods Using data obtained from literature sources we examined the power of the GDM through its derivative ATT2 model (i.e. diversity metric = b1 + b2Area + b3Time + b4Time2), in comparison with all the possible simpler models, e.g. including only area or time. The diversity metrics considered were the number of (1) native species, (2) archipelagic endemic species, and (3) single-island endemic species. Models were evaluated using both log-transformed and untransformed diversity data by means of linear mixed effect models. For Hawaii and the Canaries, responses of different major taxonomic groups were also analysed separately. Results The ATT2 model was always included within the group of best models and, in many cases, was the single-best model and was particularly successful in fitting the log-transformed diversity metrics. In four archipelagos, a hump-shaped relationship with time (island age) is apparent, while the other four archipelagos show a general increase of species richness with island age. In Hawaii and the Canaries outcomes vary between different taxonomic groups. Main conclusions The GDM is an intentionally simplified representation of environmental and diversity dynamics on oceanic islands, which predicts a simple positive relationship between diversity and island area combined with a humped response to time. We find broad support for the applicability of this model, especially when a full range of island developmental stages is present. However, our results also show that the varied mechanisms of island origins and the differing responses of major taxa should be taken into consideration when interpreting diversity metrics in terms of the GDM. This heterogeneity is reflected in the fact that no single model outperforms all the other models for all datasets analysed.

We study how endemic, native and introduced arthropod species richness, abundance, diversity and community composition vary between four different habitat types (native forest, exotic forest of Cryptomeria japonica, semi-natural pasture and intensive pasture) and how arthropod richness and abundance change with increasing distance from the native forest in adjacent habitat types in Santa Maria Island, the Azores. Arthropods were sampled in four 150 m long transects in each habitat type. Arthropods were identified to species level and classified as Azorean endemic, single-island endemic (SIE), native, or introduced. The native forest had the highest values for species richness of Azorean endemics, SIEs and natives; and also had highest values of Azorean endemic diversity (Fisher’s alpha). In contrast, the intensive pasture had the lowest values for endemic and native species richness and diversity, but the highest values of total arthropod abundance and introduced species richness and diversity. Arthropod community composition was significantly different between the four habitat types. In the semi-natural pasture, the number of SIE species decreased with increasing distance from the native forest, and in the exotic forest the abundance of both Azorean endemics and SIEs decreased with increasing distance from the native forest. There is a gradient of decreasing arthropod richness and abundance from the native forest to the intensive pasture. Although this study demonstrates the important role of the native forest in arthropod conservation in the Azores, it also shows that unmanaged exotic forests have provided alternative habitat suitable for some native species of forest specialist arthropods, particularly saproxylic beetles.

Aim To assess the hypotheses that compound leaves of trees in the Amazon forest are an adaptation to drought and/or rapid growth. Location Amazon rain forest, South America. Methods Genera from 137 permanent forest plots spread across Amazonia were classified into those with compound leaves and those with simple leaves. Metrics of compound leaf prevalence were then calculated for each plot and regression models that accounted for spatial autocorrelation were used to identify associations between climate variables and compound leaf structure. We also tested for associations between compound leaf structure and a variety of ecological variables related to life history and growth strategies, including wood density, annual increase in diameter and maximum height. Results One plant family, Fabaceae, accounts for 53% of compound-leaved individuals in the dataset, and has a geographical distribution strongly centred on north-east Amazonia. On exclusion of Fabaceae from the analysis we found no significant support for the seasonal drought hypothesis. However, we found evidence supporting the rapid growth hypothesis, with possession of compound leaves being associated with faster diameter growth rates and lower wood densities. Main conclusion This study provides evidence that possession of compound leaves constitutes one of a suite of traits and life-history strategies that promote rapid growth in rain forest trees. Our findings highlight the importance of carefully considering the geographical distribution of dominant taxa and spatial clustering of data points when inferring ecological causation from environment–trait associations.

Aim We assess whether and how datasets collected by the general public, so-called citizen science programmes, can improve biogeographical studies and contribute to large-scale conservation target-setting. Location Worldwide. Methods We first set a general framework highlighting the prerequisites of a relevant dataset for conservation biogeography. We then illustrate how many different citizen science programmes currently running in different countries can be placed within this framework. Results We highlight that citizen science is particularly useful to address issues spanning large temporal and spatial extents. We then show how datasets based on citizen science can be used to investigate major aspects of global change impacts on biodiversity. We further highlight why these programmes are also particularly valuable in developing the preventative and educational component of conservation biogeography. Main conclusions Conservation biogeography requires considerable amounts of data collected over large spatial and/or temporal extents. Beyond increasing technical advances to collect and analyse these data, citizen science seems to be a highly valuable tool in many aspects. However, while citizen science programmes are now popular and increasingly used in several countries, they are lacking in many others. We argue that the development of citizen science programmes should be encouraged as they can both be highly valuable for conservation biogeography and promote the reconnection between people and nature and more generally between people and science.

Conservation biogeography involves the application of biogeographical principles, theories, and analyses to problems regarding biodiversity conservation. The field was formally defined in 2005, and considerable research has been conducted in the ensuing 5 years. This editorial sets the context for 16 contributions in a special issue of Diversity and Distributions on developments and challenges in conservation biogeography. Papers are grouped into the following main themes: species distribution modelling; data requirements; approaches for assigning conservation priorities; approaches for integrating information from numerous disparate sources; special challenges involving invasive species; and the crucial issue of determining how elements of biodiversity are likely to respond to rapid climate change. One paper provides a synthesis of requirements for a robust conservation biogeography for freshwater ecosystems. Conservation biogeography is well poised to make a significant contribution to the process of providing policy makers with objectively formulated scenarios and options for the effective management of biodiversity. The editorial, and the papers in the special issue, deliberate on many of the exciting developments in play in the field, and the many complex challenges that lie ahead.

Leaves in tropical forests come in an enormous variety of sizes and shapes, each of which can be ultimately viewed as an adaptation to the complex problem of optimising the capture of light for photosynthesis. However, the fact that many different shape "strategies" coexist within a habitat demonstrate that there are many other intrinsic and extrinsic factors involved, such as the differential investment in support tissues required for different leaf lamina shapes. Here, we take a macrogeographic approach to understanding the function of different lamina shape categories. Specifically, we use 106 permanent plots spread across the Amazon rainforest basin to: 1) describe the geographic distribution of some simple metrics of lamina shape in plots from across Amazonia, and; 2) identify and quantify relationships between key environmental parameters and lamina shape in tropical forests. Because the plots are not randomly distributed across the study area, achieving this latter objective requires the use of statistics that can account for spatial auto-correlation. We found that between 60–70% of the 2791 species and 83 908 individual trees in the dataset could be classified as having elliptic leaves (= the widest part of the leaf is on an axis in the middle fifth of the long axis of the leaf). Furthermore, the average Amazonian tree leaf is 2.5 times longer than it is wide and has an entire margin. Contrary to theoretical expectations we found little support for the hypothesis that narrow leaves are an adaptation to dry conditions. However, we did find strong regional patterns in leaf lamina length-width ratios and several significant correlations with precipitation variables suggesting that water availability may be exerting an as yet unrecognised selective pressure on leaf shape of rainforest trees. Some support was found for the hypothesis that narrow leaves are an adaptation to low nutrient soils. Furthermore, we found a strong correlation between the proportion of trees with non-entire laminas (dissected, toothed, etc.) and mean annual temperature once again supporting the well documented association that provides a basis for reconstructing past temperature regimes.

The species–area relationship (SAR) between different biological provinces is one of the most interesting, but least explored aspects of the well-known species–area pattern. Following the usage that a biological province is a region whose species have for the most part evolved within it, rather than immigrating from somewhere else, we propose that islands can be considered equivalent to biological provinces for single island endemic species (SIEs). Hence, analyses of the relationships between numbers of SIEs and island area can be used as model systems to explore the form of inter-provincial SARs. We analyzed 13 different data sets derived from 11 sources, using the power (log–log) model. Eleven of the SIE–area relationships were statistically significant, explaining high proportions of the variance in SIE numbers (R2 0.57–0.95). The z-values (slopes) of the statistically significant SIE–area relationships ranged from 0.47 to 1.13, with a mean value of 0.80 (SD±0.24). All the island systems in which SIE represent >50% of species exhibited z-values for the SARs of native species higher than those deemed typical of archipelagic SARs. The SIE–area slopes are quite similar to those previously calculated for inter-provincial SARs, indicating that, when speciation becomes the dominant process adding to the species richness of assemblages, high z-values should be anticipated, regardless of the biogeographical scale of the study system.

Aim MacArthur and Wilson’s dynamic equilibrium model of island biogeography provides a powerful framework for understanding the ecological processes acting on insular populations. However, their model is known to be less successful when applied to systems and processes operating on evolutionary and geological timescales. Here, we present a general dynamic model (GDM) of oceanic island biogeography that aims to provide a general explanation of biodiversity patterns through describing the relationships between fundamental biogeographical processes – speciation, immigration, extinction – through time and in relation to island ontogeny. Location Analyses are presented for the Azores, Canaries, Galapagos, Marquesas and Hawaii. Methods We develop a theoretical argument from first principles using a series of graphical models to convey key properties and mechanisms involved in the GDM. Based on the premises (1) that emergent properties of island biotas are a function of rates of immigration, speciation and extinction, (2) that evolutionary dynamics predominate in large, remote islands, and (3) that oceanic islands are relatively short-lived landmasses showing a characteristic humped trend in carrying capacity (via island area, topographic variation, etc.) over their life span, we derive a series of predictions concerning biotic properties of oceanic islands. We test a subset of these predictions using regression analyses based largely on data sets for native species and single-island endemics (SIEs) for particular taxa from each archipelago, and using maximum island age estimates from the literature. The empirical analyses test the power of a simple model of diversity derived from the GDM: the log(Area) + Time + Time2 model (ATT2), relative to other simpler time and area models, using several diversity metrics. Results The ATT2 model provides a more satisfactory explanation than the alternative models evaluated (for example the standard diversity–area models) in that it fits a higher proportion of the data sets tested, although it is not always the most parsimonious solution. Main conclusions The theoretical model developed herein is based on the key dynamic biological processes (migration, speciation, extinction) combined with a simple but general representation of the life cycle of oceanic islands, providing a framework for explaining patterns of biodiversity, endemism and diversification on a range of oceanic archipelagos. The properties and predictions derived from the model are shown to be broadly supported (1) by the empirical analyses presented, and (2) with reference to previous phylogenetic, ecological and geological studies.

Aim The general dynamic model (GDM) of oceanic island biogeography integrates rates of immigration, speciation and extinction in relation to a humped trajectory of island area, species carrying capacity and topographic complexity through time, based on a simplified island ontogeny. In practice, many islands have more complex ontogenies, featuring surfaces of varying age. Here, we extend the GDM to apply at a local scale within islands, and test the predictions analytically within individual islands. Location El Hierro, La Palma and Tenerife (Canary Islands). Methods Following the GDM logic, we derive predictions for the distributions and richness of single island endemics (SIEs) across island landscapes of different age. We test these predictions by means of generalized linear models and binominal tests using gridded species occurrence data for vascular plant SIE species and a set of climatic, topographic and terrain age variables. We also examined phylogenetic divergence times for a subset of endemic lineages. Results Geological age, in interaction with slope, and topographic variables, best explained SIE richness at the landscape scale. About 70% of SIEs had ranges strongly biased to, or largely restricted to old terrain. Available phylogenetic divergence times of SIEs of radiated plant lineages suggested an origin on the older parts of the islands. Metrics of anthropogenic disturbance and habitat availability were unrelated to the observed SIE pattern. Main conclusions Our findings support the hypothesis that SIEs have evolved and accumulated on older and topographically complex terrain, while colonization processes predominate on the youngest parts. These results imply that evolutionary processes shape species distributions at the landscape scale within islands. This opens the perspective of extending the GDM framework to understand processes at a local scale within individual islands.

[Background] In this contribution we present detailed distribution and abundance data for arthropod species identified during the BALA ¿ Biodiversity of Arthropods from the Laurisilva of the Azores (1999-2004) and BALA2 projects (2010-2011) from 18 native forest fragments in seven of the nine Azorean islands (all excluding Graciosa and Corvo islands, which have no native forest left)., [New information] Of the total 286 species identified, 81% were captured between 1999 and 2000, a period during which only 39% of all the samples were collected. On average, arthropod richness for each island increased by 10% during the time frame of these projects. The classes Arachnida, Chilopoda and Diplopoda represent the most remarkable cases of new island records, with more than 30% of the records being novelties. This study stresses the need to expand the approaches applied in these projects to other habitats in the Azores, and more importantly to other less surveyed taxonomic groups (e.g. Diptera and Hymenoptera). These steps are fundamental for getting a more accurate assessment of biodiversity in the archipelago., AMCS was supported by a Marie Curie Intra-European Fellowship (IEF 331623 ‘COMMSTRUCT’) and by a Juan de la Cierva Fellowship (IJCI-2014-19502) funded by the Spanish ‘Ministerio de Economía y Competitividad’.

We provide the first fossil pollen and charcoal analysis from the island of Gran Canaria (Canary Islands). The pollen record obtained from Laguna de Valleseco (870 m a.s.l.) spans the late Holocene (c. 4500–1500 cal. years BP) and thereby captures the impact of human colonization. During the earliest period, pollen composition resembled contemporary thermophilous communities, with palms (Phoenix canariensis) and junipers (Juniperus cf. turbinata) being the dominant trees, suggesting that these elements were more widespread in the past. Vegetation in Valleseco began to change at around 2300 cal. years BP, 400 years before the earliest archaeological evidence of human presence in the island (c. 1900 cal. years BP). Our data show an increased frequency of fires at that time, coinciding with the decline of palms and the increase of grasses, indicating that humans were present and were transforming vegetation, thus showing that the demise of Gran Canaria’s forest began at an early point in the prehistoric occupation of the island. In the following centuries there were no signs of forest recovery. Pollen from cultivated cereals became significant, implying the introduction of agriculture in the site, by 1800 cal. years BP. The next shift in vegetation (c. 1600 cal. years BP) involved the decrease of grasses in favour of shrubs and trees like Morella faya, suggesting that agriculture was abandoned at the site.

Oceanic islands host a disproportionately high fraction of endangered or recently extinct endemic species. We report on species extinctions among endemic Azorean beetles following 97% habitat loss since AD 1440. We infer extinctions from historical and contemporary records and examine the influence of three predictors: geographical range, habitat specialization and body size. Of 55 endemic beetle species investigated (out of 63), seven can be considered extinct. Single-island endemics (SIEs) were more prone to extinction than multi-island endemics. Within SIEs restricted to native habitat, larger species were more extinction-prone. We thus show a hierarchical path to extinction in Azorean beetles: species with small geographical range face extinction first, with the larger bodied ones being the most threatened. Our study provides a clear warning of the impact of habitat loss on island endemic biotas. This work was funded by Direcção Regional dos Recursos Florestais (PROJ.17.01–080203), Fundação para a Cie̊ncia e a Tecnologia FCT-PTDC/BIA-BEC/100182/2008, FCT-PTDC/BIA-BIC/119255/2010. info:eu-repo/semantics/publishedVersion

The conversion of forest to agriculture across the world's tropics, and the limited space for protected areas, has increased the need to identify effective conservation strategies in human-modified landscapes. Isolated trees are believed to conserve elements of ecological structure, providing micro-sites for conservation in matrix landscapes, and facilitating seed dispersal and forest restoration. Here we investigate the role of isolated Ficus trees, which are of critical importance to tropical forest ecosystems, in conserving frugivore composition and function in a human-modified landscape in Assam, India. We surveyed the frugivorous birds feeding at 122 isolated Ficus trees, 33 fruit trees, and 31 other large trees across a range of 32 km from the nearest intact forest. We found that Ficus trees attracted richer and more abundant assemblages of frugivores than the other tree categories. However, incidence function estimates revealed that forest specialist species decreased dramatically within the first kilometre of the forest edge. Despite this, species richness and functional diversity remained consistent across the human-modified landscape, as habitat generalists replaced forest-dependent frugivores, and accounted for most of the ecological function found in Ficus trees near the forest edge. We recommend that isolated Ficus trees are awarded greater conservation status, and suggest that their conservation can support ecologically functional networks of frugivorous bird communities.

Many of the world's rural populations are dependent on the local provision of economically and medicinally important plant resources. However, increasing land-use intensity is depleting these resources, reducing human welfare, and thereby constraining development. Here we investigate a low cost strategy to manage the availability of valuable plant resources, facilitated by the use of isolated Ficus trees as restoration nuclei. We surveyed the plants growing under 207 isolated trees in Assam, India, and categorized them according to their local human-uses. We found that Ficus trees were associated with double the density of important high-grade timber, firewood, human food, livestock fodder, and medicinal plants compared to non-Ficus trees. Management practices were also important in determining the density of valuable plants, with grazing pressure and land-use intensity significantly affecting densities in most categories. Community management practices that conserve isolated Ficus trees, and restrict livestock grazing and high-intensity land-use in their vicinity, can promote plant growth and the provision of important local resources.

Diversity indices have been widely used in ecological research, but they remain problematic in that different indices may rank communities inconsistently. This problem can be solved by using diversity ordering methods, the output of which is a diversity profile in graphical form for each community being compared. In this paper, we demonstrate that existing diversity ordering methods can be classified into four groups and that within-group methods are essentially equivalent, while among-group methods are not. We find that the intrinsic diversity-related methods—i.e., the group containing the right tail-sum method, the logarithmic dominance plot, the majorization method, and the k-dominance plot—provide the most stringent test of diversity ordering, and we recommend the right tail-sum method as the method of preference for practical purposes.

Where high species richness and high human population density coincide, potential exists for conflict between the imperatives of species conservation and human development. We examine the coincidence of at-risk bird species richness and human population in the countries of the tropical Andes. We then compare the performance of the expert-driven Important Bird Areas (IBA) scheme against a hypothetical protected-areas network identified with a systematic reserve selection algorithm seeking to maximize at-risk bird species representation. Our aim is to assess the degree to which: IBAs contain a higher richness of at-risk species than would be expected by chance, IBAs contain more people than would be expected by chance, and IBAs are congruent with complementary areas that maximize species representation with an equivalent number of sites. While the correlation of richness and population was low for the region as a whole, representation of all at-risk bird species required many sites to be located in areas of high human population density. IBA sites contained higher human population densities than expected by chance ( P < 0.05) and were markedly less efficient in representing at-risk bird species of the region than sites selected using the reserve selection algorithm. Moreover, overlap between IBAs and these latter sites was very limited. Expert-driven selection procedures may better reflect existing sociopolitical forces, including land ownership and management regimes, but are limited in their ability to develop an efficient, integrated network of sites to represent priority species. Reserve selection algorithms may serve this end by optimizing complementarity in species representation among selected sites, whether these sites are adopted independently or as a supplement to the existing reserve network. As tools of site selection, they may be particularly useful in areas such as the tropical Andes where complex patterns of species disjunction and co-occurrence make the development of representative reserve networks particularly difficult. Furthermore, they facilitate making spatially explicit choices about how reserve sites are located in relation to human populations. We advocate their use not in replacement of approaches such as the IBA initiative but as an additional, complementary tool in ensuring that such reserve networks are developed as efficiently as practically possible.

This paper identifies the historical precedents and recent advances in descriptive and analytical aspects of invasive plant ecology. The paper takes a global perspective that focuses primarily on natural and semi-natural systems. The dynamics of plant invasions depend on the unique combination of species and recipient environments in light of short-lived, but highly influential, stochastic events. Spreading from the original point of establishment can be virtually instantaneous or follow a prolonged timelag. Range extension proceeds according to a variety of patterns dependent on the interplay between dispersal modes and landscape characteristics. The impacts of plant invasions are all-encompassing: biodiversity loss, economic impacts and aesthetic impacts occasioned by the loss of traditional cultural or natural landscapes. From the conservation perspective, costs are incalculable, but undoubtedly high. The impacts of invasive plants on natural ecosystems occur across all levels of biotic organization and, in the worst case, result in global extinctions and modification of fundamental ecosystem properties that make restoration practically impossible. Plant invasions occur across all habitat types and have spawned complementary theories, which are briefly presented within particular contexts.

Aim Global patterns in primary productivity in natural ecosystems are important for interpreting ecological processes and patterns of biodiversity. Net primary productivity (NPP) on land has long been thought to be greatest in tropical forests and to decrease towards the poles. However, it has recently been claimed that the NPP of mid-latitude forests is as great as, or even greater than, that of tropical forests and that ecologically relevant productivity peaks at mid-latitudes. Here we evaluate these hypotheses by testing for relationships between latitude and productivity using a range of forest productivity datasets. Location Global. Methods We apply ordinary least squares regression and t-test analyses to published latitude–productivity data for forests, specifically updated to include an expanded dataset for the previously data-poor tropics, and we evaluate the relationship between the primary productivity of forests and modelled vascular plant species richness. Results Contrary to the recent claims, we found strong support for a negative relationship between latitude and annual NPP of forests with all datasets, and NPP was significantly greater in tropical forests than in temperate forests. Vascular plant richness was positively correlated with NPP. Main conclusions NPP of forests increases towards the equator. Given that species richness also increases towards the equator, and that vascular plant richness correlates with NPP, these results are consistent with recent meta-analyses showing that the relationships between productivity and species richness of both plants and animals in natural ecosystems are predominantly positive. These results are congruent with ecological theories that predict a positive relationship between species richness and productivity, and they indicate that there is no need to explain peaked richness–productivity relationships over broad spatial extents, since they do not appear to exist.

Scattered fig (Ficus spp. Moraceae) trees are critically important for biodiversity conservation in tropical rural landscapes. By providing large fruit crops, they help maintain seed dispersal networks and facilitate forest restoration. The conservation of fig trees scattered across rural landscapes is therefore vital for the preservation of ecosystem services and biodiversity beyond the borders of protected areas. Given the threats to scattered fig trees, it is increasingly important to identify potentially effective local conservation strategies that accommodate existing perceptions of their value. We used ethnographic techniques to assess the attitudes of villagers towards fig trees in the village of Komargoan and its surroundings in Assam, India. As reported for other parts of South Asia, we found fig trees have significant sacred status, which included taboos against cutting them down. However, we discovered mixed and sometimes contradictory understandings of the religious attributes of fig trees, which were sometimes believed to be inhabited by gods or ancestral spirits. The benefits most commonly associated with fig trees by interviewees were their aesthetic beauty, large size, and shade during the daytime heat. When the presence of these trees incurred economic costs, their religious, aesthetic, and practical benefits were not sufficient reasons to prevent people from cutting them down, although often saplings would be planted in another place as compensation. Unexpectedly, figs were only planted by respected members of the community, usually older men, who had sufficient social status. Any conservation strategy aiming to sustain the abundance of figs in rural Assam is more likely to be successful if these cultural views are taken into account.

Vegetation history in the Canary Islands, one of the most biodiverse regions within Europe, has recently and for the first time, been the subject of palaeoecological studies. The interpretation of fossil records may be limited by several uncertainties regarding howwell the different vegetation types are represented in the pollen rain. In this study we address this key knowledge gap within Canarian vegetation science, taking the island of Tenerife as a model. Based on quantitative and qualitative data we analysed pollen–vegetation relationships to test whether different vegetation types can be discriminated by means of their pollen rain composition, to identify underand over-represented taxa, and to test the match between modern pollen rain and fossil pollen assemblages. We found clear differences in pollen composition and abundance among major Canarian vegetation types. We also identified good indicator taxa of the local occurrence of their source plants by their high fidelity and low dispersibility index scores. Extra-regional types (taxa without potential source plants in the Canary Islands) were not detected in our traps. However, several important floristic elements are either over-represented (Pinus, Morella and Ericaceae trees, Chenopodiaceae, Poaceae, Rumex and Urticaceae herbs and shrubs) or under-represented (Lauraceae trees, Fabaceae and Euphorbia shrubs). Results indicate that pollen composition is a good reflection of vegetation in Canarian ecosystems and can be used effectively to reconstruct past environments.

© 2014 John Wiley & Sons Ltd/CNRS. The study of islands as model systems has played an important role in the development of evolutionary and ecological theory. The 50th anniversary of MacArthur and Wilson's (December 1963) article, 'An equilibrium theory of insular zoogeography', was a recent milestone for this theme. Since 1963, island systems have provided new insights into the formation of ecological communities. Here, building on such developments, we highlight prospects for research on islands to improve our understanding of the ecology and evolution of communities in general. Throughout, we emphasise how attributes of islands combine to provide unusual research opportunities, the implications of which stretch far beyond islands. Molecular tools and increasing data acquisition now permit re-assessment of some fundamental issues that interested MacArthur and Wilson. These include the formation of ecological networks, species abundance distributions, and the contribution of evolution to community assembly. We also extend our prospects to other fields of ecology and evolution - understanding ecosystem functioning, speciation and diversification - frequently employing assets of oceanic islands in inferring the geographic area within which evolution has occurred, and potential barriers to gene flow. Although island-based theory is continually being enriched, incorporating non-equilibrium dynamics is identified as a major challenge for the future., This work was supported by the FRB (Fondation pour la Recherche sur la Biodiversite), through its Centre for Synthesis and Analysis of Biodiversity (CESAB), and the ‘Laboratoire d’Excellence’ TULIP (ANR-10-LABX- 41; ANR-11-IDEX-0002-02). BHW acknowledges funding from the EU Seventh Framework Programme under grant 263958 (RUN-Emerge) and 267243 (Plant Fellows).

Aim Current weaknesses of diversity theory include: a failure to distinguish different biogeographical response variables under the general heading of diversity; and a general failure of ecological theory to deal adequately with geographical scale. Our aim is to articulate the case for a top-down approach to theory building, in which scale is addressed explicitly and in which different response variables are clearly distinguished. Location The article draws upon both theoretical contributions and empirical analyses from all latitudes, focusing on terrestrial ecosystems and with some bias towards (woody) plants. Methods We review current diversity theory and terminology in relation to scale of applicability. As a starting point in developing a general theory, we take the issue of geographical gradients in species richness as a main theme and evaluate the extent to which commonly cited theories are likely to operate at scales from the macro down to the local. Results A degree of confusion surrounds the use of the terms alpha, beta and gamma diversity, and the terms local, landscape and macro-scale are preferred here as a more intuitive framework. The distinction between inventory and differentiation diversity is highlighted as important as, in terms of scale of analysis, are the concepts of focus and extent. The importance of holding area constant in analysis is stressed, as is the notion that different environmental factors exhibit measurable heterogeneity at different scales. Evaluation of several of the most common diversity theories put forward for the grand clines in species richness, indicates that they can be collapsed to dynamic hypotheses based on climate or historical explanations. The importance of the many ecological/ biological mechanisms that have been proposed is evident mainly at local scales of analysis, whilst at the macro-scale they are dependent largely upon climatic controls for their operation. Local communities have often been found not to be saturated, i.e. to be non-equilibrial. This is argued, perhaps counter-intuitively, to be entirely compatible with the persistence through time of macro-scale patterns of richness that are climatically determined. The review also incorporates recent developments in macroecology, Rapoport’s rule, trade-offs, and the importance of isolation, landscape impedance and geometric constraints on richness (the mid-domain effect) in generating richness patterns; highlighting those phenomena that are contributory to the first-order climatic pattern, and those, such as the geometric constraints, that may confound or obscure these patterns. Main conclusions A general theory of diversity must necessarily cover many disparate phenomena, at various scales of analysis, and cannot therefore be expressed in a simple formula, but individual elements of this general theory may be. In particular, it appears possible to capture in a dynamic climate-based model and ‘capacity rule’, the form of the

Aim Few data sets exist that allow measurement of long-term extinction and turnover rates for islands of the size of the three main islands of the Krakatau group. We test the reliability of previous estimates of plant species extinction and examine structure within the extinction data. Location The data analysed are for the three older Krakatau islands: Rakata, Sertung and Panjang in the Sunda Strait, Indonesia. Methods Our analysis is based on a comprehensive database incorporating all species records for each island since recolonization began after the 1883 sterilization, plus attributes such as distribution, phylogeny, population status and dispersal mechanism for each species. We employ a combination of univariate and multi-term analyses in analysing structure, and derive Minimal Adequate Models using binary logistic analyses of variance and covariance. We compare the 1883–1934 data set with the contemporary flora as represented by (1) 1979–83 records (as used in previous analyses) and (2) 1979–94 data (original). Results The improved data for the contemporary flora reduces the number of missing species by one-third. We show that a variety of estimates of extinction rate can be produced depending on what assumptions are made concerning the status of particular species groups. Structural features in the extinction data persist despite the reduction in overall numbers of losses. Losses relate to: (1) the number of islands on which a species originally occurred, (2) the primary dispersal mode, and (3) the original abundance of a species (e.g. whether it was known to have established a successful resident population, and whether it was in decline or increasing in c. 1930). The ‘best’ descriptive model employs the variables denoted under (3). A high proportion of losses comprised species introduced by people and rare or ephemeral species. Losses of ‘residents’ that had colonized naturally could largely be accounted for by reference to (1) successional loss of habitat and, to a lesser degree, (2) other habitat disturbance or loss. Main conclusions Previous analyses, based on a more limited data set, have significantly over-estimated extinction from the Krakatau flora. Few naturally colonizing and established species have become extinct. The findings indicate that caution is necessary in interpreting ‘headline’ island ecological rates, and in analysing and modelling such data. Examination of structural features of the data appear to be valuable both in providing ecological insights in their own right, and in enabling refinements to estimates of extinction and thus turnover.

1 This paper offers a commentary on the development of island ecological theory since the publication of MacArthur & Wilson’s equilibrium theory in the 1960s. I distinguish the simple model at the core of their Equilibrium Theory of Island Biogeography (ETIB) and the broader body of their theory, which embraces evolutionary as well as ecological patterns — all, however, within the overarching framework or assumption of equilibrium. 2 The basic problems with the ETIB have long been known, and its status as a ruling paradigm has been the subject of concern for more than two decades. With the development of nonequilibrium ideas in ecology, island biogeographers arguably now have viable theoretical frameworks to set alongside or around the ETIB. Four conditions are highlighted as extremes: i) dynamic equilibrium; ii) dynamic nonequilibrium; iii) ‘static’ equilibrium; and iv) ‘static’ nonequilibrium: together providing a conceptual framework for island ecological analyses. 3 The importance of scale is stressed and attention is drawn to Haila’s spatial-temporal continuum as an organizational device. It is argued that the processes represented within the ETIB (and by extension, other island theories) may be prominent within only a limited portion of this continuum, while elsewhere they are generally subsumed by other dominant processes. 4 Colonization and ecosystem development of near-shore islands constitute just a special case of ecological succession, and thus the development of theories of island assembly may benefit accordingly from efforts to incorporate ideas from the ecological succession literature. 5 The desirability of specifying answerable questions is stressed, as is the need to build a greater degree of complexity into the development of island ecological models. Notwithstanding which, it is also recognized that key advances are often brought about by simple, but bold models, of the form exemplified elsewhere in this issue.

cited By 27; International audience; Analyses of species-diversity patterns of remote islands have been crucial to the development of biogeographic theory, yet little is known about corresponding patterns in functional traits on islands and how, for example, they may be affected by the introduction of exotic species. We collated trait data for spiders and beetles and used a functional diversity index (FRic) to test for nonrandomness in the contribution of endemic, other native (also combined as indigenous), and exotic species to functional-trait space across the nine islands of the Azores. In general, for both taxa and for each distributional category, functional diversity increases with species richness, which, in turn scales with island area. Null simulations support the hypothesis that each distributional group contributes to functional diversity in proportion to their species richness. Exotic spiders have added novel trait space to a greater degree than have exotic beetles, likely indicating greater impact of the reduction of immigration filters and/or differential historical losses of indigenous species. Analyses of species occurring in native-forest remnants provide limited indications of the operation of habitat filtering of exotics for three islands, but only for beetles. Although the general linear (not saturating) pattern of trait-space increase with richness of exotics suggests an ongoing process of functional enrichment and accommodation, further work is urgently needed to determine how estimates of extinction debt of indigenous species should be adjusted in the light of these findings. © 2014 PNAS.

Published in 2001, The Unified Neutral Theory of Biodiversity and Biogeography (UNTB) emphasizes the importance of stochastic processes in ecological community structure, and has challenged the traditional niche-based view of ecology. While neutral models have since been applied to a broad range of ecological and macroecological phenomena, the majority of research relating to neutral theory has focused exclusively on the species abundance distribution (SAD). Here, we synthesize the large body of work on neutral theory in the context of the species abundance distribution, with a particular focus on integrating ideas from neutral theory with traditional niche theory. First, we summarize the basic tenets of neutral theory; both in general and in the context of SADs. Second, we explore the issues associated with neutral theory and the SAD, such as complications with fitting and model comparison, the underlying assumptions of neutral models, and the difficultly of linking pattern to process. Third, we highlight the advances in understanding of SADs that have resulted from neutral theory and models. Finally, we focus consideration on recent developments aimed at unifying neutral- and niche-based approaches to ecology, with a particular emphasis on what this means for SAD theory, embracing, for instance, ideas of emergent neutrality and stochastic niche theory. We put forward the argument that the prospect of the unification of niche and neutral perspectives represents one of the most promising future avenues of neutral theory research.