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 speciesÁarea 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 speciesÁarea 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 B600 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.

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. Nineteen areas in seven of the nine Azorean islands were evaluated for species diversity and rarity based on soil epigean arthropods. Fifteen out of the 19 study areas are managed as Natural Forest Reserves and the remaining four were included due to their importance as indigenous forest cover. Four of the 19 areas are not included in the European Conservation network, NATURA 2000. Two sampling replicates were run per study area, and a total of 191 species were collected; 43 of those species (23%) are endemic to the archipelago and 12 have yet to be described. To produce an unbiased multiple-criteria index (importance value for conservation, IV-C) incorporating diversity and rarity based indices, an iterative partial multiple regression analysis was performed. In addition, an irreplaceability index and the complementarity method (using both optimisation and heuristic methods) were used for priority-reserves analyses. It was concluded that at least one well-managed reserve per island is absolutely necessary to have a good fraction of the endemic arthropods preserved. We found that for presence/absence data the suboptimal complementarity algorithm provides solutions as good as the optimal algorithm. For abundance data, optimal solutions indicate that most reserves are needed if we want that at least 50% of endemic

Islands harbour evolutionary and ecologically unique biota, which are currently disproportionately threatened by a multitude of anthropogenic factors, including habitat loss, 2568 Biodivers Conserv (2018) 27:2567-2586 1 3 invasive species and climate change. Native forests on oceanic islands are important refugia for endemic species, many of which are rare and highly threatened. Long-term monitoring schemes for those biota and ecosystems are urgently needed: (i) to provide quantitative baselines for detecting changes within island ecosystems, (ii) to evaluate the effectiveness of conservation and management actions, and (iii) to identify general ecological patterns and processes using multiple island systems as repeated 'natural experiments'. In this contribution, we call for a Global Island Monitoring Scheme (GIMS) for monitoring the remaining native island forests, using bryophytes, vascular plants, selected groups of arthropods and vertebrates as model taxa. As a basis for the GIMS, we also present new, optimized monitoring protocols for bryophytes and arthropods that were developed based on former standardized inventory protocols. Effective inventorying and monitoring of native island forests will require: (i) permanent plots covering diverse ecological gradients (e.g. elevation, age of terrain, anthropogenic disturbance); (ii) a multiple-taxa approach that is based on standardized and replicable protocols; (iii) a common set of indicator taxa and community properties that are indicative of native island forests' welfare, building on, and harmonized with existing sampling and monitoring efforts; (iv) capacity building and training of local researchers, collaboration and continuous dialogue with local stakeholders; and (v) long-term commitment by funding agencies to maintain a global network of native island forest monitoring plots.

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.

Worldwide, lava caves host colorful microbial mats. However, little is known about the diversity of these microorganisms, or what role they may play in the subsurface ecosystem. White and yellow microbial mats were collected from four lava caves each on the Azorean island of Terceira and the Big Island of Hawai’i, to compare the bacterial diversity found in lava caves from two widely separated archipelagos in two different oceans at different latitudes. Scanning electron microscopy of mat samples showed striking similarities between Terceira and Hawai’ian microbial morphologies. 16S rRNA gene clone libraries were constructed to determine the diversity within these lava caves. Fifteen bacterial phyla were found across the samples, with more Actinobacteria clones in Hawai’ian communities and greater numbers of Acidobacteria clones in Terceira communities. Bacterial diversity in the subsurface was correlated with a set of factors. Geographical location was the major contributor to differences in community composition (at the OTU level), together with differences in the amounts of organic carbon, nitrogen and copper available in the lava rock that forms the cave. These results reveal, for the first time, the similarity among the extensive bacterial diversity found in lava caves in two geographically separate locations and contribute to the current debate on the nature of microbial biogeography.

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 landuses 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 highintensity 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.

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