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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]

The ongoing decline of insect populations highlight the need for long-term ecological monitoring. As part of the "SLAM—Long Term Ecological Study of the Impacts of Climate Change on the Natural Forests of Azores" project, we investigated changes in arthropod diversity and community structure over a ten-year period (2012–2022) in the native forest of the island of Terceira (Azores). Focused on two arthropod assemblages (indigenous and non-indigenous species) monitored with SLAM traps, we asked if there was a distinguishable pattern in the diversity and structure of the studied arthropod subsets in a pristine island native forest. Species richness remained relatively constant. Endemic arthropods dominated and remained stable over time, indicating the forest's ecological stability. In contrast, the assemblage of native non-endemic arthropods underwent changes, including increased hyperdominance and decreased biomass. The introduced arthropod assemblage showed more erratic dynamics driven by species turnover. Results suggested that temporal variation in each subset may be due to different ecological processes and that niche filtering may limit the establishment and spread of introduced arthropods. This research contributes to our understanding of the temporal dynamics of arthropods in native island forests and highlights the need for ongoing conservation efforts to protect these fragile ecosystems. [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]

Climate change affects all ecosystems, but subterranean ecosystems are repeatedly neglected from political and public agendas. Cave habitats are home to unknown and endangered species, with low trait variability and intrinsic vulnerability to recover from human-induced disturbances. We studied the annual variability and cyclicity of temperatures in caves vis-à-vis surface in different climatic areas. We hypothesize that cave temperatures follow the average temperature pattern at the surface for each location with a slight delay in the signal, but we found three different thermal patterns occurring in caves: (1) high positive correlation and a similar thermal pattern to the surface, (2) low correlation and a slight thermal delay of the signal from the surface, and (3) high negative correlation with an extreme delay from the surface. We found daily thermal cycles in some caves, which may potentially control the circadian rhythms of cave organisms. Our results show that caves had lower thermal amplitude than the surface, and that thermal averages within caves approximately correspond to the to the annual average of surface temperature. Caves buffer external temperature and act as refugia for biota in extreme climatic events. Likewise, temperature increases at surface will lead to increment in caves, threatening subterranean biota and ecosystem services. [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]

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]

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]

Due to ongoing global changes, it is essential to establish a baseline record from which to determine future shifts in species distributions and community assembly patterns. In this context, we used digitised historical bryophyte distribution data along a 1021 m elevational gradient in Terceira Island (Azores) to determine how bryophyte species distribution varies with elevation and which spatial and climatic drivers contribute to this variation. We used ordinary least squares analysis to test for climatic and spatial data as explanatory variables for bryophyte richness and Mid-Domain Null simulations to assess the influence of spatial constraints on species distributions. Bryophyte richness follows a hump-shaped pattern, with mosses predominating in the first half of the gradient and liverworts in the second half. While moss richness did not correlate to any climatic variables and responded weakly to the presence of forest cover, liverwort presence was related to temperature, rainfall, humidity and the occurrence of native forest areas, suggesting that these plants are more sensitive to changes in their environment and can thus be used as better indicator species for climate change. Despite their inherent biases, our study shows that historically compiled data can be a valuable tool for preliminary assessment of macroecological patterns. [ABSTRACT FROM PUBLISHER]

Simple Summary: The vertical structure and the tree species in a forest have a strong influence on the communities of wood-inhabiting beetles. Little is known about how this influence varies between rare and common beetle species. We compared alpha and beta diversity patterns of common and rare species in the canopy of the Leipzig floodplain forest to assess their response to vertical structuring of the canopy and tree species. We assessed rarity firstly by using red list status and secondly based on the abundances of the beetle species. The understory displayed a significantly higher number of common species than the tree canopy. Conversely, the canopy harbored a higher number of rare species. The beetles' beta diversity patterns were predominantly shaped by differences in species composition, not by differences in species richness. Vertical structure had a higher influence on the beetle communities than tree species. Both factors had a higher influence on common than on rare beetle species. Our results indicate that studies carried out in the understory alone do not allow drawing conclusions regarding the biodiversity in the forest canopy, and thus regarding the overall community structure of xylobiont beetles in the canopy. Vertical stratification and host tree species are factors with a high influence on the structure of communities of xylobiont beetles. However, little is known about how this influence varies between common and rare species. Based on estimated species richness, we compared alpha and beta diversity patterns of common and rare species in the canopy of the Leipzig floodplain forest to assess their response to vertical stratification and tree species. We used two measures of rarity: threat level in red lists and abundance based on octaves. The understory displayed a significantly higher number of common species than the canopy strata. Conversely, the canopy strata harbored a higher number of rare species. Turnover was always dominant over richness differences in beta diversity partitions. Using Raup–Crick null models and non-metric multidimensional scaling, we found that the vertical strata accounted for 19% of the overall beta diversity of common species and for 15% of the overall beta diversity of rare species. The tree species accounted for 7% of the overall beta diversity of the common species and 3% of the beta diversity of the rare species. Our results indicate that studies carried out in the understory alone do not allow drawing conclusions regarding the biodiversity in the canopy strata, and thus regarding the overall community structure of xylobiont beetles in the canopy. [ABSTRACT FROM AUTHOR]

One of the aims of island biogeography theory is to explain the number of species in an archipelago. Traditionally, the variables used to explain the species richness on an island are its area and distance to the mainland. However, increasing evidence suggests that accounting for other variables is essential for better estimates. In particular, the distance between islands should play a role in determining species richness. This work uses a Bayesian framework using Gaussian processes to assess whether distance to neighbouring islands (spatial autocorrelation) can better explain arthropod species richness patterns in the Azores Archipelago and in the Canary Islands. This method is flexible and allows the inclusion of other variables, such as maximum altitude above sea level (elevation). The results show that accounting for spatial autocorrelation provides the best results for both archipelagos, but overall, spatial autocorrelation seems to be more important in the Canary archipelago. Similarly, elevation plays a more important role in determining species richness in the Canary Islands. We recommend that spatial autocorrelation should always be considered when modelling an archipelago's species richness. [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]

The dispersal flights of West Indian drywood termite, Cryptotermes brevis (Walker) (Isoptera: Kalotermitidae) were surveyed in the major cities of Azores. The sampling device used to estimate termite density consisted of a yellow adhesive trap (size 45 by 24 cm), placed with an artificial or natural light source in a dark attic environment. In addition, data from two other projects were used to improve the knowledge about the geographical distribution of the species. The level of infestation in the two main Azorean towns differed, with high levels in the houses of Angra do Heroísmo, whereas in Ponta Delgada, there are fewer houses with high levels of infestation. The infestation in Ponta Delgada shows a pattern of spreading from the center outward to the city’s periphery, whereas in Angra do Heroísmo, there was a pattern of spreading outward from several foci. The heavy infestation observed in Angra do Heroísmo and the clear increase of infestation levels observed from 2010 to 2011 is a reason for concern and calls for an urgent application of an Integrated Pest Management (IPM) control strategy. [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]

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]

Azorean lava-tubes and volcanic pits adequately sampled for arthropod fauna were evaluated for species diversity and rarity. An iterative partial multiple regression analysis was performed to produce a multi-criteria index (Importance Value for Cave Conservation, IV-CC) incorporating arthropod species diversity indices but also including indices qualifying cave geological and management features (e.g., the diversity of geological structures, threats, accessibility). Additionally, we calculated complementarity solutions (irreplaceability and Fraction-of-Spare measures) for each cave with different targets per species, i.e., the minimum number of caves needed for each species to be represented either once or twice. Our results clearly show that to preserve all troglobiont arthropods endemic to the Azores, it is crucial to protect several caves per island. As many as 10 and 15 caves are needed to include one or two occurrences, respectively, per species. [ABSTRACT FROM AUTHOR]

An overview of the obligate hypogean fauna in Portugal (including Azores and Madeira archipelagos) is provided, with a list of obligated cave-dwelling species and subspecies, and a general perspective about its conservation. All the available literature on subterranean Biology of Portugal since the first written record in 1870 until today has been revised. A total of 43 troglobiont and 67 stygobiont species and subspecies from 12 orders have been described so far in these areas, included in the so-called Mediterranean hotspot of biodiversity. The subterranean fauna in Portugal has been considered moderately poor with some endemic relicts and it remains to be demonstrated if this fact is still true after investing in standard surveys in cave environments. The major problems related to the conservation of cave fauna are discussed, but it is clear that the protection of this specialized fauna implies an adequate management of surface habitats. [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]

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]

1. The positive abundance–occupancy and abundance–variance relationships are two of the most widely documented patterns in population and community ecology. 2. Recently, a general model has been proposed linking the mean abundance, the spatial variance in abundance, and the occupancy of species. A striking feature of this model is that it consists explicitly of the three variables abundance, variance and occupancy, and no extra parameters are involved. However, little is known about how well the model performs. 3. Here, we show that the abundance–variance–occupancy model fits extremely well to data on the abundance, variance and occupancy of a large number of arthropod species in natural forest patches in the Azores, at three spatial extents, and distinguishing between species of different colonization status. Indeed, virtually all variation about the bivariate abundance–occupancy and abundance–variance relationships is effectively explained by the third missing variable (variance in abundance in the case of the abundance–occupancy relationship, and occupancy in the case of the abundance–variance relationship). 4. Introduced species tend to exhibit lower densities, less spatial variance in these densities, and occupy fewer sites than native and endemic species. None the less, they all lie on the same bivariate abundance–occupancy and abundance–variance, and trivariate abundance–variance–occupancy, relationships. 5. Density, spatial variance in density, and occupancy appear to be all the things one needs to know to describe much of the spatial distribution of species. [ABSTRACT FROM AUTHOR]

1. Fifteen species richness estimators (three asymptotic based on species accumulation curves, 11 nonparametric, and one based in the species–area relationship) were compared by examining their performance in estimating the total species richness of epigean arthropods in the Azorean Laurisilva forests. Data obtained with standardized sampling of 78 transects in natural forest remnants of five islands were aggregated in seven different grains (i.e. ways of defining a single sample): islands, natural areas, transects, pairs of traps, traps, database records and individuals to assess the effect of using different sampling units on species richness estimations. 2. Estimated species richness scores depended both on the estimator considered and on the grain size used to aggregate data. However, several estimators (ACE, Chao1, Jackknife1 and 2 and Bootstrap) were precise in spite of grain variations. Weibull and several recent estimators [proposed by Rosenzweig et al. ( Conservation Biology, 2003, 17, 864–874), and Ugland et al. ( Journal of Animal Ecology, 2003, 72, 888–897)] performed poorly. 3. Estimations developed using the smaller grain sizes (pair of traps, traps, records and individuals) presented similar scores in a number of estimators (the above-mentioned plus ICE, Chao2, Michaelis–Menten, Negative Exponential and Clench). The estimations from those four sample sizes were also highly correlated. 4. Contrary to other studies, we conclude that most species richness estimators may be useful in biodiversity studies. Owing to their inherent formulas, several nonparametric and asymptotic estimators present insensitivity to differences in the way the samples are aggregated. Thus, they could be used to compare species richness scores obtained from different sampling strategies. Our results also point out that species richness estimations coming from small grain sizes can be directly compared and other estimators could give more precise results in those cases. We propose a decision framework based on our results and on the literature to assess which estimator should be used to compare species richness scores of different sites, depending on the grain size of the original data, and of the kind of data available (species occurrence or abundance data). [ABSTRACT FROM AUTHOR]

Aim This paper has two objectives. First, we examine how a variety of biotic, abiotic and anthropogenic factors influence the endemic and introduced arthropod richness on an oceanic island. Second, we look at the relationship between the endemic and introduced arthropod richness, to ask whether areas with high levels of endemic species richness deter invasions. Location The work was carried out on a young volcanic island, Terceira, in the Azores. Methods We used standard techniques to collect data on arthropod species richness. Environmental data were obtained from the CIELO climatic model and using GIS. The explanatory value of environmental variables on a small-scale gradient of endemic and exotic arthropod species richness was examined with generalized linear models (GLMs). In addition, the impact of both endemic and exotic species richness in the communities was assessed by entering them after the environmental variable(s) to see if they contributed significantly to the final model (the hierarchical method). Results Abiotic (climatic and geomorphological) variables gave a better explanation of the variation in endemic species richness, whereas anthropogenic variables explained most of the variation in introduced species richness. Furthermore, after accounting for all environmental variables, part of the unexplained variance in the endemic species richness is explained by the introduced species richness and vice-versa. That is, areas with high levels of endemic species richness had many introduced species. There is evidence of a somewhat inverse spatial distribution between a group of oceanic-type, forest-dwelling, endemic, relict arthropods and a group of more generalist endemic arthropods that are able to survive in disturbed marginal sites particularly rich in non-indigenous species. Main conclusions Richness of endemic species is mainly driven by abiotic factors such as a climatic axis (oceanic-type localities with lower temperatures and summer precipitations) and a binary variable CALD (location of sites in caldeiras or ravines), whereas richness of introduced species depends on disturbance related factors. However, after factoring out these major influences, there is a correlation between endemic and introduced richness, suggesting that – independent of the environmental and geographical factors that affect the distribution of endemic or introduced species – the richest endemic assemblages are more prone to invasion, due probably to a facilitation process. Inconclusive evidence suggests that non-indigenous species are limited to those sites under anthropogenic influence located mainly near forest edges, but the rate of expansion of those species to high-altitude, core pristine sites has still to be tested. [ABSTRACT FROM AUTHOR]

This article explores patterns of insect herbivore distribution in the canopy of the Laurisilva forests on seven islands in the Azores archipelago. To our knowledge, this is one of the first extensive study of this type in tree or shrub canopies of oceanic island ecosystems. One of the most frequently debated characteristics of such ecosystems is the likely prevalence of vague, ill-defined niches due to taxonomic disharmony, which may have implications for insect-plant interactions. For instance, an increase in ecological opportunities for generalist species is expected due to the lack of predator groups and reduced selection for chemical defence in host plants. The following two questions were addressed: 1) Are specialists species rare, and insect herbivore species randomly distributed among host plant species in the Azores? 2) Are the variances in insect herbivore species composition, frequency and richness explained by host plants or by regional island effects? We expect a proportional distribution of herbivore species between host plants, influenced by host frequency and distinct island effects; otherwise, deviation from expectation might suggest habitat preference for specific host tree crowns. Canopy beating tray samples were performed on seven islands, comprising 50 transects with 1 to 3 plant species each (10 replicates per species), giving 1320 samples from ten host species trees or shrubs in total. From a total of 129 insect herbivore species, a greater number of herbivore species was found onJuniperus brevifolia(s=65) andErica azorica(s=53). However, the number of herbivore species per individual tree crown was higher forE. azoricathan for any other host, on all islands, despite the fact that it was only the fourth more abundant plant. In addition, higher insect species richness and greater insect abundance were found on the trees of Santa Maria Island, the oldest in the archipelago. Insect species composition was strongly influenced by the presence ofE. azorica, which was the only host plant with a characteristic fauna across the archipelago, whereas the fauna of other plant crowns was grouped by islands. The great insect occurrence onE. azoricareflects strong habitat fidelity, but only four species were clearly specialists. Our findings indicate a broadly generalist fauna. The simplicity of Azorean Laurisilva contributed to the understanding of insect-plant mechanisms in canopy forest habitats. [ABSTRACT FROM AUTHOR]

We investigate the potential use of structural characteristics of vegetation as predictors of the densities of pasture arthropods in three Azorean islands. Two types of upland pastures, recent sown pastures(3-4 years old) and wet semi-natural old pastures (more than 35 years old), were studied in three Azorean islands (S. Maria, Terceira andPico). Three arthropod assemblages, of particular importance in pasture habitats, were assessed: insect forb-feeders, insect grass-feeders and web-building spiders. These are numerically abundant and represent a range of feeding strategies. Point quadrats for plants and suction (Vortis) for arthropods were used as sampling methods. Several vegetation indices were investigated, but only three (cover abundance of perennial forbs, cover abundance of perennial grasses and the totalvegetation alpha-diversity) were good predictors of invertebrate abundance. Vegetation structure, defined by a Height Index, was of minorpredictive value. In sown and semi-natural pastureland, diverse and structurally complex pasture sites support more individual insect forb-feeders, grass-feeders and web-building spiders per unit area than less complex ones. [ABSTRACT FROM AUTHOR]

Many stakeholders, from governments to civil society to businesses, lack the data they need to make informed decisions on biodiversity, jeopardising efforts to conserve, restore and sustainably manage nature. Here we review the importance of enhancing biodiversity monitoring, assess the challenges involved and identify potential solutions. Capacity for biodiversity monitoring needs to be enhanced urgently, especially in poorer, high-biodiversity countries where data gaps are disproportionately high. Modern tools and technologies, including remote sensing, bioacoustics and environmental DNA, should be used at larger scales to fill taxonomic and geographic data gaps, especially in the tropics, in marine and freshwater biomes, and for plants, fungi and invertebrates. Stakeholders need to follow best monitoring practices, adopting appropriate indicators and using counterfactual approaches to measure and attribute outcomes and impacts. Data should be made openly and freely available. Companies need to invest in collecting the data required to enhance sustainability in their operations and supply chains. With governments soon to commit to the post-2020 global biodiversity framework, the time is right to make a concerted push on monitoring. However, action at scale is needed now if we are to enhance results-based management adequately to conserve the biodiversity and ecosystem services we all depend on. [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]

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]

The Azorean archipelago, recognized as one of the world's biodiversity hotspots, is home to a diverse and unique community of arthropod species, highlighting a notable degree of endemism. However, the native forests that support these species are facing significant degradation due to habitat loss and fragmentation. In this study, we aimed to determine the ideal season for measuring the biological integrity of forest sites using a biological integrity index (IBI) based on arthropod communities captured with Sea, Land, and Air Malaise (SLAM) traps. Drawing on more than thirty years of research experience in the Azorean forests, we selected twelve reference sites, six representing preserved native forest and six representing disturbed native forest, and compared how IBI values vary between seasons. IBI values exhibited consistent variations between seasons in disturbed sites, indicating that measuring the biological integrity in these areas can be conducted at any time of the year without a specific seasonal preference. In contrast, significant differences were observed in pristine forest sites, with the winter season and the combination of winter and spring data (cold semester) showing notably higher values compared to other seasons and semesters. This finding suggests that measuring the biological integrity of preserved sites is best optimized in the cold seasons, while the detection of exotic species impact is most effective in summer and autumn. Consequently, if resources are limited, monitoring efforts should be concentrated in the winter and summer seasons to obtain the maximum and minimum values of IBI, respectively. Additionally, our study suggests that the summer season is the optimal time to detect potentially invasive exotic species. [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]

The aims of this study were to predict the potential distribution of two introduced Mustelidae, Mustela nivalis and M. putorius in the Azores archipelago (Portugal), and evaluate the relative contribution of environmental factors from native and introduced ranges to predict species distribution ranges in oceanic islands. We developed two sets of Species Distribution Models using MaxEnt and distribution data from the native and introduced ranges of the species to project their potential distribution in the archipelago. We found differences in the predicted distributions for the models based on introduced and on native occurrences for both species, with different most important variables being selected. Climatic variables were most important for the introduced range models, while other groups of variables (i.e., human-disturbance) were included in the native-based models. Most of the islands of the Azorean archipelago were predicted to have suitable habitat for both species, even when not yet occupied. Our results showed that predicting the invaded range based on introduced range environmental conditions predicted a narrower range. These results highlight the difficulty to transfer models from native to introduced ranges across taxonomically related species, making it difficult to predict future invasions and range expansion. [ABSTRACT FROM AUTHOR]

Climate change is causing shifts in species distributions worldwide. Understanding how species distributions will change with future climate change is thus critical for conservation planning. Impacts on oceanic islands are potentially major given the disproportionate number of endemic species and the consequent risk that local extinctions might become global ones. In this study, we use species climate envelope models to evaluate the current and future potential distributions of Azorean endemic species of bryophytes, vascular plants, and arthropods on the Islands of Terceira and São Miguel in the Azores archipelago (Macaronesia). We examined projections of climate change effects on the future distributions of species with particular focus on the current protected areas. We then used spatial planning optimization software (PRION) to evaluate the effectiveness of protected areas at preserving species both in the present and future. We found that contractions of species distributions in protected areas are more likely in the largest and most populated island of São Miguel, moving from the coastal areas towards inland where the current protected areas are insufficient and inadequate to tackle species distribution shifts. There will be the need for a revision of the current protected areas in São Miguel to allow the sustainable conservation of most species, while in Terceira Island the current protected areas appear to be sufficient. Our study demonstrates the importance of these tools for informing long-term climate change adaptation planning for small islands. [ABSTRACT FROM AUTHOR]

Simple Summary: Ecosystem services (ESs) and disservices (EDs) are routinely estimated from changes in service provider densities without measuring their actual levels. By using the sentinel approach (i.e., exposing a plant, seeds, and prey models in a standardized way), we tested how coriander (Coriandrum sativum) strips planted in mixed orchards on Terceira Island (Azores, Portugal) affected ESs/EDs via herbivory on lettuce plants, seed predation on wheat and weed seeds, and predation on artificial caterpillars. Vertebrates had more influence than invertebrates on ESs and EDs. Herbivory (ED) after 2 weeks was similar in the coriander and the control plots, while seed predation was higher in the control than in the coriander plots on both wheat grain (an ED: 30.8% vs. 15.3%) and weed seeds (an ES: 2.5% vs. 0.4%). Vertebrate predation (ES) rates after 48 h were significantly higher in the control (9%) than in the coriander plots (3%), while no difference was observed for invertebrate predation. Coriander strips did not support increased ES/reduced ED levels in this habitat. The sentinel approach is effective to quantitatively compare multiple ESs/EDs under different farming management strategies. The effect of flower strips on ecosystem services (ESs) and disservices (EDs) is routinely assessed following changes in service provider densities without measuring the associated levels of ES/EDs. By using the sentinel approach (i.e., exposing a plant, seeds, and prey models in a standardized way), we tested how coriander (Coriandrum sativum) strips planted in mixed orchards on Terceira Island (Azores, Portugal) affected herbivory on lettuce plants, seed predation on wheat and weed seeds, and predation on artificial caterpillars. Vertebrates had more influence than invertebrates on ESs/EDs. Herbivory (ED) after 2 weeks was similar in the coriander and the control plots (mean ± SD; 2.3% ± 3.3% vs. 2.2% ± 2.9%, n = 32 for both). Seed predation was higher in the control than in the coriander plots for both grain (ED; 30.8% ± 38.9% vs. 15.3% ± 10.8%, n = 18 for both) and weed seeds (ES; 2.5% ± 4.1% vs. 0.4% ± 0.5%, n = 18 for both). Vertebrate predation (ES) rates after 48 h were significantly higher in the control (estimate 9%, 95% CI: 4–20%) than in the coriander plots (3%, 1–8%), while no difference was observed for invertebrate predation. Coriander strips did not support increased ES/reduced ED levels in this setting. The tools used can be effective to quantitatively compare multiple ESs/EDs under different farming management strategies. [ABSTRACT FROM AUTHOR]

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]

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]

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]

Background: Functional trait-based approaches link species diversity patterns to ecosystem functioning. In the context of global change, understanding these links is vital for developing holistic biodiversity management strategies. Bryophytes, important ecosystem components owing to their biogeochemical functions, have not been the focus of many functional studies. Aims: This is the first assessment of bryophyte functional diversity in the Azores archipelago, aiming to uncover multivariate trait richness and composition patterns along the elevational gradient on Terceira Island. Methods: Based on five water acquisition and retention traits of leafy liverworts and mosses, we calculated functional diversity metrics within and among six bryophyte communities sampled along a 1021-m elevational transect. Results: Trait composition differed significantly between coastal and mountain communities. Mosses presented inrolled leaves and ornamented leaf cells at low elevation but not at high elevation. These patterns were associated with an uphill shift from drier and warmer conditions to a moister and cooler environment. Conclusions: Future climatic changes might affect bryophyte functional diversity patterns in Terceira Island, particularly for mosses. These results can be directly compared with those obtained for other archipelagos where the same protocol has been applied, allowing a joint assessment of insular vegetation functional diversity patterns. [ABSTRACT FROM PUBLISHER]

We report 18 species of wild bees plus the domesticated honeybee from the Azores, which adds nine species to earlier lists. One species, Hylaeus azorae, seems to be a single island endemic, and three species are possibly native (Colletes eous, Halictus villosulus, and Hylaeus pictipes). All the remaining bee species are most likely accidental introductions that arrived after human colonization of the archipelago in the 15th century. Bee diversity in the Azores is similar to bee diversity of Madeira and Cape Verde but nearly ten times lower than it is in the Canary Islands. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Species richness on oceanic islands has been related to a series of ecological factors including island size and isolation (i.e. the Equilibrium Model of Island Biogeography, EMIB), habitat diversity, climate (i.e., temperature and precipitation) and more recently island ontogeny (i.e. the General Dynamic Model of oceanic island biogeography, GDM). Here we evaluate the relationship of these factors with the diversity of bryophytes in the Macaronesian region (Azores, Madeira, Canary Islands and Cape Verde). The predictive power of EMIB, habitat diversity, climate and the GDM on total bryophyte richness, as well as moss and liverwort richness (the two dominant bryophyte groups), was evaluated through ordinary least squares regressions. After choosing the best subset of variables using inference statistics, we used partial regression analyses to identify the independent and shared effects of each model. The variables included within each model were similar for mosses and liverworts, with orographic mist layer being one of the most important predictors of richness. Models combining climate with either the GDM or habitat diversity explained most of richness variation (up to 91%). There was a high portion of shared variance between all pairwise combinations of factors in mosses, while in liverworts around half of the variability in species richness was accounted for exclusively by climate. Our results suggest that the effects of climate and habitat are strong and prevalent in this region, while geographical factors have limited influence on Macaronesian bryophyte diversity. Although climate is of great importance for liverwort richness, in mosses its effect is similar to or, at least, indiscernible from the effect of habitat diversity and, strikingly, the effect of island ontogeny. These results indicate that for highly vagile taxa on oceanic islands, the dispersal process may be less important for successful colonization than the availability of suitable ecological conditions during the establishment phase. [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]

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]

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]

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]

Simple Summary: We studied the diversity of arthropods in native forests along a 1000 m elevation gradient on Terceira Island, Azores (Portugal). These forests form an isolated and threatened habitat with unique endemic species. We analysed the change in alpha and beta diversity of arthropod species with elevation and if the diversity of endemic, native non-endemic and introduced species responds differently to elevation. Resident arthropods were sampled using SLAM (Sea, Land and Air Malaise) traps between 2014 and 2018. Spiders (Araneae), beetles (Coleoptera), true bugs (Hemiptera) and barklice (Psocoptera), as well as endemic, native and introduced species, were analysed separately. Total species richness decreases with elevation for all species, Coleoptera and Psocoptera, and particularly so for introduced species, but peaks at mid-high elevation for Araneae and endemic species. These patterns are probably driven by unfavourable climatic conditions at higher elevations while being influenced by human disturbance at lower elevations. Total species diversity along the whole elevation gradient is shaped by this decreasing richness as well as the replacement of species at different elevations. We present an analysis of arthropod diversity patterns in native forest communities along the small elevation gradient (0–1021 m a.s.l.) of Terceira island, Azores (Portugal). We analysed (1) how the alpha diversity of Azorean arthropods responds to increasing elevation and (2) differs between endemic, native non-endemic and introduced (alien) species, and (3) the contributions of species replacement and richness difference to beta diversity. Arthropods were sampled using SLAM traps between 2014 and 2018. We analysed species richness indicators, the Hill series and beta diversity partitioning (species replacement and species richness differences). Selected orders (Araneae, Coleoptera, Hemiptera and Psocoptera) and endemic, native non-endemic and introduced species were analysed separately. Total species richness shows a monotonic decrease with elevation for all species and Coleoptera and Psocoptera, but peaks at mid-high elevation for Araneae and endemic species. Introduced species richness decreases strongly with elevation especially. These patterns are most likely driven by climatic factors but also influenced by human disturbance. Beta diversity is, for most groups, the main component of total (gamma) diversity along the gradient but shows no relation with elevation. It results from a combined effect of richness decrease with elevation and species replacement in groups with many narrow-ranged species. [ABSTRACT FROM AUTHOR]

Measuring progress toward international biodiversity targets requires robust information on the conservation status of species, which the International Union for Conservation of Nature (IUCN) Red List of Threatened Species provides. However, data and capacity are lacking for most hyperdiverse groups, such as invertebrates, plants, and fungi, particularly in megadiverse or high‐endemism regions. Conservation policies and biodiversity strategies aimed at halting biodiversity loss by 2020 need to be adapted to tackle these information shortfalls after 2020. We devised an 8‐point strategy to close existing data gaps by reviving explorative field research on the distribution, abundance, and ecology of species; linking taxonomic research more closely with conservation; improving global biodiversity databases by making the submission of spatially explicit data mandatory for scientific publications; developing a global spatial database on threats to biodiversity to facilitate IUCN Red List assessments; automating preassessments by integrating distribution data and spatial threat data; building capacity in taxonomy, ecology, and biodiversity monitoring in countries with high species richness or endemism; creating species monitoring programs for lesser‐known taxa; and developing sufficient funding mechanisms to reduce reliance on voluntary efforts. Implementing these strategies in the post‐2020 biodiversity framework will help to overcome the lack of capacity and data regarding the conservation status of biodiversity. This will require a collaborative effort among scientists, policy makers, and conservation practitioners. Article Impact Statement: Global conservation policies require a strategy to obtain robust data for measuring progress against biodiversity targets. [ABSTRACT FROM AUTHOR]

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]