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Islands in the tropical Pacific Ocean are renowned for high biodiversity and endemism despite having relatively small landmasses. However, our knowledge of how this biodiversity is formed remains limited. The taxon cycle, where well‐dispersed, earlier colonizers become displaced from coastal to inland habitats by new waves of colonizers, producing isolated, range‐restricted species, has been proposed to explain current biodiversity patterns. Here, we integrate the outcomes of phylogenetic studies in the region to investigate the sources, age, number of colonizations, and diversification of 16 archipelagos in the tropical and subtropical South Pacific. We then evaluate whether the results support the taxon cycle as a plausible mechanism for these observations. We find that most species in the Pacific arrived less than 5 Mya from geographically close sources, suggesting that colonization by new taxa is a frequent and ongoing process. Therefore, our findings are broadly consistent with the theory of the Taxon Cycle, which posits that ongoing colonization results in the gradual displacement of established lineages. Only the oldest archipelagos, New Caledonia and Fiji, do not conform to this trend, having proportionally less recent colonization events, suggesting that the taxon cycle may slow on older islands. This conclusion is further validated by New Caledonia having lower diversification rate estimates than younger islands. We found that diversification rates across archipelagos are negatively correlated with area and age. Therefore, a taxon cycle that slows with island age appears to be a suitable concept for understanding the dynamic nature and biodiversity patterns of the Pacific Islands. [ABSTRACT FROM AUTHOR]

Microsites are created by abiotic and biotic features of the landscape and may provide essential habitats for the persistence of biota. Forest canopies and understorey plants may moderate wind and solar radiation to create microclimatic conditions that differ considerably from regional climates. Skirt‐forming plants, where senescent leaves create hut‐like cavities around the stem, create microsites that are sheltered from ambient conditions and extreme weather events, constituting potential refuges for wildlife. We investigate day and night temperatures and humidity for four locations (grass tree cavities, soil, 20 cm above‐ground, 1 m above‐ground) in a South Australian forest with relatively open canopy of stringybark eucalypts (Eucalyptus baxteri, E. obliqua) and an understorey of skirt‐forming grass trees (Xanthorrhoea semiplana) at 5, 10, 20, and 40 m from the forest edge. We also measured the percentage of canopy and understorey covers. Generally, temperature and humidity differed significantly between more sheltered (grass tree cavities, soil) and open‐air microsites, with the former being cooler during the day and warmer and more humid during the night. Furthermore, our results suggest that canopy cover tends to decrease, and understorey cover tends to increase, the temperature of microsites. Distance to the edge was not significantly related to temperature for any microsite, suggesting that the edge effect did not extend beyond 10 m from the edge. Overall, grass trees influenced microclimatic conditions by forming a dense understorey and providing cavities that are relatively insulated. The capacity of grass tree cavities to buffer external conditions increased linearly with ambient temperatures, by 0.46°C per degree increase in maximum and 0.25°C per degree decrease in minimum temperatures, potentially offsetting climate warming and enabling persistence of fauna within their thermal limits. These climate moderation properties will make grass trees increasingly important refuges as extreme weather events become more common under anthropogenic climate change. [ABSTRACT FROM AUTHOR]

Aim: The island species–area relationship (ISAR) assumes that the area of islands is homogeneous, or scales with environmental heterogeneity across an archipelago, which is not always the case. We compare the performance of models that adjust or substitute for island area with measures of habitat diversity, island age and resource availability for two taxonomic groups. Location: Five hotspot archipelagos (Azores, Galapagos, Hawaii, Cape Verde, Canary Islands). Taxa: Vascular plants, birds. Methods: We used the mathematical framework of the power law to compare relevant models, treating the one containing only area as a null model against which others were compared. Data were collated from databases and the literature. Models were compared using linear regression within archipelagos and via mixed effect models with archipelago as a random effect. Results: Weighting of island area by habitat diversity and resource availability systematically improved statistical significance and model fits versus the area only power law. Models including island age did not show the same systematic improvement in model fits. For vascular plants, weighting islands by resource availability (energy and water) performed better than weighting by habitat diversity, although for birds these weightings performed equally well. Main Conclusions: Given that islands within archipelagos are fairly heterogeneous in climate, topography and geology, it is worth accounting for this in ISARs. Our results suggest that, for islands in volcanic hotspot archipelagos, this is best done by using direct measures of habitat diversity and resource availability rather than using island age as a proxy. We, therefore, recommend using direct measures, rather than proxies, when investigating the drivers of biodiversity patterns on islands. [ABSTRACT FROM AUTHOR]

The importance of terrestrial coastal ecosystems for maintaining healthy coral reef ecosystems remains understudied. Sea kraits are amphibious snakes that require healthy coral reefs for foraging, but little is known about their requirements of terrestrial habitats, where they slough their skin, digest prey, and breed. Using concurrent microclimate measurements and behavior surveys, we show that a small, topographically flat atoll in Fiji with coastal forest provides many microhabitats that relate to the behaviors of Yellow Lipped Sea Kraits, Laticauda colubrina. Microclimates were significantly related to canopy cover, leaf litter depth, and distance from the high‐water mark (HWM). Sea kraits were almost exclusively observed in coastal forest within 30 m of the HWM. Sloughing of skins only occurred within crevices of mature or dying trees. Resting L. colubrina were significantly more likely to occur at locations with higher mean diurnal temperatures, lower leaf litter depths, and shorter distances from the HWM. On Leleuvia, behavior of L. colubrina therefore relates to environmental heterogeneity created by old‐growth coastal forests, particularly canopy cover and crevices in mature and dead tree trunks. The importance of healthy coastal habitats, both terrestrial and marine, for L. colubrina suggests it could be a good flagship species for advocating integrated land‐sea management. Furthermore, our study highlights the importance of coastal forests and topographically flat atolls for biodiversity conservation. Effective conservation management of amphibious species that utilize land‐ and seascapes is therefore likely to require a holistic approach that incorporates connectivity among ecosystems and environmental heterogeneity at all relevant scales. [ABSTRACT FROM AUTHOR]

Conservation management to protect coastal ecosystems sometimes overlooks site‐specific fine‐scale heterogeneity. For example, while habitat loss is a known key driver of population declines in many shorebirds, these birds are also dependent on high‐quality habitats to maximize energy stores. Here we describe the microhabitats provided by beach‐cast wrack (washed up macroalgae and seagrasses), a resource threatened by harvesting and beach cleaning, and how shorebirds utilize these.We measured the temperature and absolute humidity at 10 cm above three substrates (fresh wrack, aged wrack and sand) and then related bird behaviour (roosting vs. foraging) to climatic and environmental data.Freshly beach‐cast wrack mostly provided cooler and less humid habitats, but warmer temperatures than aged wrack or sand in the early mornings. Microtopography created by shelter from prevailing winds and wrack depth modified these general trends.Generally, temperature predicted where shorebirds overall and the two most common species, the double‐banded plover Charadrius bicinctus and red‐necked stint Calidris ruficollis, were observed. During most of the day, foraging and roosting were more likely to occur on the warmer aged wrack. In the early morning, when fresh wrack provided the warmer temperatures, birds tended to roost and forage on fresh wrack.Synthesis and Applications. Beach‐cast wrack creates a complex mosaic of unique microclimates varying in space and time, which allows shorebirds to minimize energy expenditure by selecting the thermally most favourable habitats for roosting and foraging. Removal of beach‐cast wrack for commercial and aesthetic reasons thus reduces habitat quality and increases energy expenditure in shorebirds. Associated declines in energy stores may be contributing to declines in shorebird populations. Management of coastal ecosystems and shorebirds therefore needs to consider and maintain fine‐scale environmental heterogeneity at local scales. [ABSTRACT FROM AUTHOR]

The extraordinary diversity of orchids has captivated scientists for more than a century, yet their complex spatial patterns at large scales remain poorly resolved. On islands, orchid diversity patterns are especially puzzling. While some islands are centres of orchid diversity, orchids are underrepresented on most islands. To disentangle such complex patterns, key functional differences among orchids must be considered – a distinction seldom made in biogeographical analyses. Using a global dataset of 454 islands, we tested prominent hypotheses in island biogeography, while simultaneously making the distinction between epiphytes and two terrestrial life forms (geophytes and non‐geophytes). Orchid diversity was unevenly distributed across islands and life forms. Epiphytic orchid diversity strongly increased with temperature, illustrating the near confinement of epiphytes to the tropics. Geophytes became proportionally more important with increasing seasonality, highlighting their ability to withstand harsh climatic conditions. Epiphytes and non‐geophytes both displayed responses (e.g. negative relationship with seasonality) related to their dependence on consistently favourable conditions, possibly because of the absence of subterranean storage organs. This highlights that the factors explaining orchid diversity differ strongly with, and are related to, life form. We suggest that key functional differences within and across plant families be considered in future studies to better understand drivers of complex diversity patterns. [ABSTRACT FROM AUTHOR]

Aims: Trees dominate the biomass in many ecosystems and are essential for ecosystem functioning and human well-being. They are also one of the best-studied functional groups of plants, with vast amounts of biodiversity data available in scattered sources. We here aim to illustrate that an efficient integration of these data could produce a more holistic understanding of vegetation. Methods: To assess the extent of potential data integration, we use key databases of plant biodiversity to: (a) obtain a list of tree species and their distributions; (b) identify coverage of and gaps in different aspects of tree biodiversity data; and (c) discuss large-scale patterns of tree biodiversity in relation to vegetation. Results: Our global list of trees included 58,044 species. Taxonomic coverage varies in three key databases, with data on the distribution, functional traits, and molecular sequences for about 84%, 45% and 44% of all tree species, which is >10% greater than for plants overall. For 28% of all tree species, data are available in all three databases. However, less data are digitally accessible about the demography, ecological interactions, and socio-economic role of tree species. Integrating and imputing existing tree biodiversity data, mobilization of non-digitized resources and targeted data collection, especially in tropical countries, could help closing some of the remaining data gaps. Conclusions: Due to their key ecosystem roles and having large amounts of accessible data, trees are a good model group for understanding vegetation patterns. Indeed, tree biodiversity data are already beginning to elucidate the community dynamics, functional diversity, evolutionary history and ecological interactions of vegetation, with great potential for future applications. An interoperable and openly accessible framework linking various databases would greatly benefit future macroecological studies and should be linked to a platform that makes information readily accessible to end users in biodiversity conservation and management. [ABSTRACT FROM AUTHOR]

Species abundance distributions (SADs) characterise the distribution of individuals among species. SADs have rarely been explored on islands and the ecological processes shaping SADs are still not fully understood. Notably, the relative importance of disturbance regime in shaping plant SADs remains poorly known. We investigate the relative importance of disturbance regime and island geography on the shape of SADs. We computed SADs for local tree communities in 1‐ha forest plots on 20 tropical islands in the Indo‐Pacific region. We used generalized linear models to analyse how the shape parameter of the gambin SAD model was related to the number of trees and the number of species. Regression analyses were also used to investigate how the shape of SADs, the number of trees and the number of species were related to cyclone disturbance (power dissipation index) and geography (island area and isolation), with direct and indirect (i.e. through the number of trees and species) effects assessed using variance partitioning. Cyclone disturbance was the best predictor of the shape of SADs, with higher power dissipation index producing more lognormal‐like distributions. This effect was mostly due to cyclones increasing the number of trees and decreasing the number of species. Island area affected the shape of SADs through its effect on the number of species, and larger islands were associated with higher species richness and more logseries‐like distributions. The effect of cyclones was stronger on smaller islands. Our results illustrate that disturbances can affect SADs in complex ways; directly and indirectly by impacting the number of species and individuals in communities, and these effects may be moderated by island‐specific characteristics, such as island area or isolation. Our results therefore suggest that multiple, interacting processes shape SADs and that studying SADs has the potential to contribute important new insights to the field of island biogeography. [ABSTRACT FROM AUTHOR]

Aim: The General Dynamic Model of island biogeography (GDM) explains patterns of species richness on volcanic hotspot archipelagos with respect to island age (T) and area (A), most commonly employing what we here call semi‐log ATT2 and LogATT2 models. These can be considered direct extensions of semi‐log and power law species‐area relationships (SARs), with a hump‐shaped function of T modelling drivers of biodiversity over and above area per se. We compare these and related models and investigate their conceptual and mathematical connections. Location: The Azores, Galapagos, Hawaii, Marquesas, Society Islands, Cape Verde Islands, Canary Islands and Aeolian archipelagos. Taxa: Vascular plants, birds, bryophytes, arthropods and molluscs. Methods: We gathered species richness data from databases and published sources, resulting in 34 datasets for various taxa. We used the Akaike information criterion corrected for small sample size (AICc) to compare the relative performance of models. We analysed the data at archipelago level and across archipelagos by using mixed effect models. We also calculated the relative time of expected peak diversity (Tpeak) from parameter estimates. Results: In almost three‐fourth of the cases the best model overall was one that contained only A or T rather than both predictors. There was greater variance between taxa than between archipelagos. Better dispersed taxa tended to reach peak diversity earlier. The most generally applicable of the models including T were the LogATT2 and LogTT2 models. These have important, previously unrecognized mathematical properties. Main Conclusions: SARs and species–time relationships should always be considered as potentially more parsimonious options. Considering the LogATT2 model as a modification of the power law S = cAz, where a lognormal function in T replaces the constant, c allows the construction of an extended framework for the GDM with incorporation of additional predictors such as measures of environmental heterogeneity and application beyond hotspot archipelagos. [ABSTRACT FROM AUTHOR]

Aim: The General Dynamic Model (GDM) links island biogeographical processes to island geological history. A key premise of the GDM implies that environmental factors shaping the ecology and evolution of biota on oceanic islands follow a hump‐shaped trend over the island's life span and drive dynamics in carrying capacity, species diversity and endemism. An important component of the GDM is environmental heterogeneity (EH), but its effects on insular diversity remain poorly understood. Here, we first quantified EH, tested whether EH follows the expected hump‐shaped trend along island ontogeny and evaluated how EH relates to plant diversity. Location: 135 oceanic islands of volcanic origin. Taxon: Vascular plants. Methods: We calculated 20 EH metrics focusing on topographic and climatic components of EH, and compared whole‐island metrics (e.g. range) and moving‐window metrics (e.g. roughness). Using linear mixed‐effects models, we evaluated the trends of EH with island age and the EH–plant diversity relationship expected based on the GDM. Results: Our analysis revealed some EH components to be collinear, for example, elevation and temperature heterogeneity but also that EH metrics capture different aspects of EH, for example, climatic gradients versus climatic complexity. EH generally followed a hump‐shaped trend with island age, peaking early during island ontogeny. Among the EH components, climatic heterogeneity had the strongest effect on plant species richness and elevational heterogeneity on endemism. Lastly, including EH metrics in GDMs (traditionally, only island age and area were included) improved their predictive power. Main conclusions: The EH metrics compared here captured various attributes of the environment that influence insular plant diversity. In line with the GDM, our results strongly support a hump‐shaped relationship between EH and island age, suggesting that islands become highly heterogeneous early in their ontogeny. Finally, the contribution of EH to GDM‐based models of species richness and endemism suggests that EH is a main driver of the diversity of oceanic island biotas. [ABSTRACT FROM AUTHOR]

Aim: We mapped global patterns of tree phylogenetic endemism (PE) to identify hotspots and test hypotheses about possible drivers. Specifically, we tested hypotheses related to current climate, geographical characteristics and historical conditions and assessed their relative importance in shaping PE patterns. Location: Global. Time period: We used the present distribution of trees, and predictors covering conditions from the mid‐Miocene to present. Major taxa studied: All seed‐bearing trees. Methods: We compiled distributions for 58,542 tree species across 463 regions worldwide, matched these to a recent phylogeny of seed plants and calculated PE for each region. We used a suite of predictor variables describing current climate (e.g., mean annual temperature), geographical characteristics (e.g., isolation) and historical conditions (e.g., tree cover at the Last Glacial Maximum) in a spatial regression model to explain variation in PE. Results: Tree PE was highest on islands, and was higher closer to the equator. All three groups of predictor variables contributed substantially to the PE pattern. Isolation and topographic heterogeneity promoted high PE, as did high current tree cover. Among mainland regions, temperature seasonality was strongly negatively related to PE, while mean annual temperature was positively related to PE on islands. Some relationships differed among the major floristic regions. For example, tree cover at the Last Glacial Maximum was a positive predictor of PE in the Palaeotropics, while tree cover at the Miocene was a negative predictor of PE in the Neotropics. Main conclusions: Globally, PE can be explained by a combination of geographical, historical and current factors. Some geographical variables appear to be key predictors of PE. However, the impact of historic and current climate variables differs considerably among the major floristic regions, reflecting their unique histories. Hence, the current distribution of trees is the result of globally relevant geographical drivers and regional climatic histories. [ABSTRACT FROM AUTHOR]

Understanding how species diversity is related to sampling area and spatial scale is central to ecology and biogeography. Small islands and small sampling units support fewer species than larger ones. However, the factors influencing species richness may not be consistent across scales. Richness at local scales is primarily affected by small‐scale environmental factors, stochasticity and the richness at the island scale. Richness at whole‐island scale, however, is usually strongly related to island area, isolation and habitat diversity. Despite these contrasting drivers at local and island scales, island species–area relationships (SARs) are often constructed based on richness sampled at the local scale. Whether local scale samples adequately predict richness at the island scale and how local scale samples influence the island SAR remains poorly understood. We investigated the effects of different sampling scales on the SAR of trees on 60 small islands in the Raja Ampat archipelago (Indonesia) using standardised transects and a hierarchically nested sampling design. We compared species richness at different grain sizes ranging from single (sub)transects to whole islands and tested whether the shape of the SAR changed with sampling scale. We then determined the importance of island area, isolation, shape and habitat quality at each scale on species richness. We found strong support for scale dependency of the SAR. The SAR changed from exponential shape at local sampling scales to sigmoidal shape at the island scale indicating variation of species richness independent of area for small islands and hence the presence of a small‐island effect. Island area was the most important variable explaining species richness at all scales, but habitat quality was also important at local scales. We conclude that the SAR and drivers of species richness are influenced by sampling scale, and that the sampling design for assessing the island SARs therefore requires careful consideration. [ABSTRACT FROM AUTHOR]

Determining patterns of plant diversity on granite inselbergs is an important task for conservation biogeography due to mounting threats. However, beyond the tropics there are relatively few quantitative studies of floristic diversity, or consideration of these patterns and their environmental, biogeographic, and historical correlates for conservation. We sought to contribute broader understanding of global patterns of species diversity on granite inselbergs and inform biodiversity conservation in the globally significant Southwest Australian Floristic Region (SWAFR). We surveyed floristics from 16 inselbergs (478 plots) across the climate gradient of the SWAFR stratified into three major habitats on each outcrop. We recorded 1,060 species from 92 families. At the plot level, local soil and topographic variables affecting aridity were correlated with species richness in herbaceous (HO) and woody vegetation (WO) of soil‐filled depressions, but not in woody vegetation on deeper soils at the base of outcrops (WOB). At the outcrop level, bioclimatic variables affecting aridity were correlated with species richness in two habitats (WO and WOB) but, contrary to predictions from island biogeography, were not correlated with inselberg area and isolation in any of the three habitats. Species turnover in each of the three habitats was also influenced by aridity, being correlated with bioclimatic variables and with interplot geographic distance, and for HO and WO habitats with local site variables. At the outcrop level, species replacement was the dominant component of species turnover in each of the three habitats, consistent with expectations for long‐term stable landscapes. Our results therefore highlight high species diversity and turnover associated with granite outcrop flora. Hence, effective conservation strategies will need to focus on protecting multiple inselbergs across the entire climate gradient of the region. [ABSTRACT FROM AUTHOR]

Refugia are key environments in biogeography and conservation. Because of their unique eco‐evolutionary formation and functioning, they should display distinct functional trait signatures. However, comparative trait‐based studies of plants in refugia and non‐refugia are lacking. Here, we provide a comparison between resource‐rich (putative microrefugia for species preferring mesic habitats under increasing aridity) and resource‐impoverished woodlands (non‐refugia) around two granite outcrops in south‐western Australia. We measured and compared six functional traits (bark thickness, foliar δ13C, foliar C:N, leaf dry matter content, plant height, specific leaf area) in four woody species. We performed multiple‐trait, multiple‐species and single‐trait, within‐species analyses to test whether plants in resource‐rich habitats were functionally distinct and more diverse than those in the surrounding resource‐impoverished woodlands. We found that species in resource‐rich woodlands occupied larger and distinct multiple‐trait functional spaces and showed distinct single‐trait values (for specific leaf area and bark thickness). This suggests that plants in resource‐rich woodlands can deploy unique and more diverse ecological strategies, potentially making these putative microrefugia more resilient to environmental changes. These findings suggest that species in microrefugia may be characterised by unique functional signatures, illustrating the utility of comparative trait‐based approaches to improve understanding of the functioning of refugia. [ABSTRACT FROM AUTHOR]

Tropical cyclones (TCs) are large‐scale disturbances that regularly impact tropical forests. Although long‐term impacts of TCs on forest structure have been proposed, a global test of the relationship between forest structure and TC frequency and intensity is lacking. We test on a pantropical scale whether TCs shape the structure of tropical and subtropical forests in the long term.We compiled forest structural features (stem density, basal area, mean canopy height and maximum tree size) for plants ≥10 cm in diameter at breast height from published forest inventory data (438 plots ≥0.1 ha, pooled into 250 1 × 1‐degree grid cells) located in dry and humid forests. We computed maps of cyclone frequency and energy released by cyclones per unit area (power dissipation index, PDI) using a high‐resolution historical database of TCs trajectories and intensities. We then tested the relationship between PDI and forest structural features using multivariate linear models, controlling for climate (mean annual temperature and water availability) and human disturbance (human foot print).Forests subject to frequent cyclones (at least one TCs per decade) and high PDI exhibited higher stem density and basal area, and lower canopy heights. However, the relationships between PDI and basal area or canopy height were partially masked by lower water availability and higher human foot print in tropical dry forests.Synthesis. Our results provide the first evidence that tropical cyclones have a long‐term impact on the structure of tropical and subtropical forests in a globally consistent way. The strong relationship between power dissipation index and stem density suggests that frequent and intense tropical cyclones reduce canopy cover through defoliation and tree mortality, encouraging higher regeneration and turnover of biomass. The projected increase in intensity and poleward extension of tropical cyclones due to anthropogenic climate change may therefore have important and lasting impacts on the structure and dynamics of forests in the future. We show that tropical cyclones have a long‐term impact on the structure of tropical and subtropical forests in a globally consistent way. Forests subject to frequent and intense cyclones exhibit higher stem density and lower canopy heights. The projected changes in tropical cyclone regime may therefore have important and lasting impacts on the structure of forests in the future. [ABSTRACT FROM AUTHOR]

Abstract: Plants respond to changing environmental conditions, and their ability to adjust intra‐specifically to such shifts represents an ecological and evolutionary advantage. We studied seven plant traits for two common, rhizomatous granite outcrop species (the fern Cheilanthes austrotenuifolia, and the herb Stypandra glauca) with seasonal foliage during the cooler, wetter winter months at seven sites across an aridity gradient in southwestern Australia. We investigated trait patterns at regional and habitat scale, by investigating changes in trait values along the aridity gradient, and by comparing two different habitats types (sun‐exposed and sheltered). We expected plants occurring in more arid sites and highly irradiated, shallow‐soil (sun‐exposed) habitats, to exhibit traits indicative of more conservative resource acquisition, retention and use strategies. At the habitat scale, we found support for our prediction, with plants in more stressful, sun‐exposed habitats showing traits’ values associated with more conservative strategies (especially for water), such as smaller plants, denser leaves, higher foliar δ13C and C/N. However, at the regional scale many traits displayed the opposite pattern, suggesting less conservative resource acquisition in more arid sites. This evidence was particularly pronounced for specific leaf area (SLA), which exhibited a significant, positive relationship with increasing aridity. We suggest that the unexpected regional trends in foliar traits relate to shorter lived, faster growing leaves linked to highly efficient resource acquisition and use strategies during the shorter growing season in the more arid regions. These highly exploitative strategies may enable plants to avoid climate extremes, that is, hot and dry periods in the more arid sites. Our findings of contrasting foliar traits responses at different scales support the importance of multi‐scale approaches to quantify the role of intraspecific trait variability. [ABSTRACT FROM AUTHOR]

Abstract: Aim: To determine the role of regional forcing on plot‐level species diversity and composition, and to quantify the relative importance of biogeographical and climatic factors in explaining woody plant diversity and composition at the local‐, island‐ and archipelago‐scale. Location: Forty‐one tropical islands of the Indo‐Pacific region from Madagascar to Hawai‘i Island. Methods: We analysed the diversity and composition of tropical woody plant communities located across 113 plots, 41 islands and 19 archipelagos. We used generalized linear mixed‐effects models and generalized dissimilarity models to determine the role of regional forcing at the island and archipelago scale and to assess the relative importance of biogeographical (area and isolation of islands or archipelagos, geographical distance between plots) and climatic factors in explaining differences in local diversity and composition (species turnover). Analyses were conducted at different geographical scales (local, island and archipelago) and taxonomic levels (species, genus and family). Results: Variation in local (plot‐level) diversity (as species density, the number of species per 100 woody plants) was primarily explained by island and archipelago identity. Maximum species density was positively correlated with the area of an island (or archipelago) and negatively correlated with the isolation of an archipelago. Local climatic variability was also a significant predictor of species density, but less important than regional forcing. Climate variables explained < 20% of the variation in species turnover across all plots. The importance of geographical distance between plots relative to climate in driving species turnover decreased from the species to family level, and from the regional to island level. Main conclusions: Regional forcing was the key driver of local diversity and composition on islands. Island area and archipelago isolation are likely driving local diversity through their effects on the pool of island species. Geographical distance between plots is the main factor explaining species turnover, while at higher taxonomic levels, climatic factors and niche conservatism are the main drivers. [ABSTRACT FROM AUTHOR]

Complex processes related to biotic and abiotic forces can impose limitations to assembly and composition of plant communities. Quantifying the effects of these constraints on plant functional traits across environmental gradients, and among communities, remains challenging. We define ecological constraint ( C i) as the combined, limiting effect of biotic interactions and environmental filtering on trait expression (i.e., the mean value and range of functional traits). Here, we propose a set of novel parameters to quantify this constraint by extending the trait-gradient analysis (TGA) methodology. The key parameter is ecological constraint, which is dimensionless and can be measured at various scales, for example, on population and community levels. It facilitates comparing the effects of ecological constraints on trait expressions across environmental gradients, as well as within and among communities. We illustrate the implementation of the proposed parameters using the bark thickness of 14 woody species along an aridity gradient on granite outcrops in southwestern Australia. We found a positive correlation between increasing environmental stress and strength of ecological constraint on bark thickness expression. Also, plants from more stressful habitats (shrublands on shallow soils and in sun-exposed locations) displayed higher ecological constraint for bark thickness than plants in more benign habitats (woodlands on deep soils and in sheltered locations). The relative ease of calculation and dimensionless nature of C i allow it to be readily implemented at various scales and make it widely applicable. It therefore has the potential to advance the mechanistic understanding of the ecological processes shaping trait expression. Some future applications of the new parameters could be investigating the patterns of ecological constraints (1) among communities from different regions, (2) on different traits across similar environmental gradients, and (3) for the same trait across different gradient types. [ABSTRACT FROM AUTHOR]

Aim Refugia under past climates have been important in structuring current patterns in diversity, while refugia under anthropogenic climate change will likely be important in retaining this diversity and shaping new patterns. However, few studies have examined the congruence of past, present and future refugia, or the spatiotemporal connectivity of these refugia. Our aim was to test the extent of overlap of refugia under Last Glacial Maximum ( LGM), present (2015) and likely future climates (2100), for Tasmania's palaeoendemic flora. We then aimed to identify areas of high spatiotemporal refugia connectivity, as priority areas for conservation and management. Location Tasmania, Australia. Methods We developed and applied a new community-level approach to identifying refugia, based on generalized dissimilarity modelling of compositional turnover and a set of reference sites with known biodiversity value. Using these projections of palaeoendemic plant refugia for past, present and future climates, we developed and applied a second approach to quantify the level of connectivity of these refugia over space and time. Results Although there was large overlap (85%) between current and future climates in the distribution of the highest value palaeoendemic refugia, the small congruence of these areas with refugia at the LGM resulted in only a small area ( c. 9 km2) of persistent high value refugia over all three time periods. Despite this, our spatiotemporal analysis identified several areas of high connectivity in refugial environments for Tasmania's palaeoendemic flora over time. Main conclusions The community-level approaches we demonstrate here to quantify refugia and their spatiotemporal connectivity have the potential to advance our understanding of biodiversity dynamics, particularly for taxonomic groups that are species-rich, poorly studied or comprised of many rare species, where species-level approaches are less suitable. [ABSTRACT FROM AUTHOR]

Aim To determine if habitat diversity, as estimated by climatic and topographic variables, can predict patterns of orchid diversity on different islands and archipelagos with similar explanatory power to biogeographical variables, such as area, isolation and age of an island. Location Sixty-three islands on eight archipelagos (Solomon Islands, Vanuatu, New Caledonia, Fiji, Samoa, Tonga, Niue and the Cook Islands) in the south-west Pacific. Methods For each island, we determined the orchid species present, age, area, isolation, and indicators of topographic heterogeneity and climatic variability. We then determined the power of various biogeographical, climatic and topographic variables to predict the number of indigenous and endemic species on archipelagos, and on islands within archipelagos, using generalized linear models ( GLMs) and generalized linear mixed models ( GLMMs) respectively. Results We identified a total of 552 species in 110 genera. Area was the only significant biogeographical variable for predicting patterns of orchid species diversity on archipelagos and islands. However, climatic and topographic predictors of habitat diversity performed similarly well. The range in curvature was the best indicator of species richness from the topographic variables, while the range of temperature was the best climatic predictor. These key variables were often strongly correlated with area. Main conclusions Climatic and topographic variables are useful indicators of habitat diversity. The high explanatory power of area and climatic and topographic predictors, and the strong correlation among these variables, suggests that increasing habitat diversity with increasing area may be the major driver of the species-area relationship. Using climatic and topographic variables as predictors of species richness therefore allows determining the key environmental factors and processes driving species diversity. [ABSTRACT FROM AUTHOR]

Refugia - areas that may facilitate the persistence of species during large-scale, long-term climatic change - are increasingly important for conservation planning. There are many methods for identifying refugia, but the ability to quantify their potential for facilitating species persistence (ie their "capacity") remains elusive. We propose a flexible framework for prioritizing future refugia, based on their capacity. This framework can be applied through various modeling approaches and consists of three steps: (1) definition of scope, scale, and resolution; (2) identification and quantification; and (3) prioritization for conservation. Capacity is quantified by multiple indicators, including environmental stability, microclimatic heterogeneity, size, and accessibility of the refugium. Using an integrated, semi-mechanistic modeling technique, we illustrate how this approach can be implemented to identify refugia for the plant diversity of Tasmania, Australia. The highest- capacity climate-change refugia were found primarily in cool, wet, and topographically complex environments, several of which we identify as high priorities for biodiversity conservation and management. [ABSTRACT FROM AUTHOR]

Identifying refugia is a critical component of effective conservation of biodiversity under anthropogenic climate change. However, despite a surge in conceptual and practical interest, identifying refugia remains a significant challenge across diverse continental landscapes. We provide an overview of the key properties of refugia that promote species' persistence under climate change, including their capacity to (i) buffer species from climate change; (ii) sustain long-term population viability and evolutionary processes; (iii) minimize the potential for deleterious species interactions, provided that the refugia are (iv) available and accessible to species under threat. Further, we classify refugia in terms of the environmental and biotic stressors that they provide protection from (i.e. thermal, hydric, cyclonic, pyric and biotic refugia), but ideally refugia should provide protection from a multitude of stressors. Our systematic characterization of refugia facilitates the identification of refugia in the Australian landscape. Challenges remain, however, specifically with respect to how to assess the quality of refugia at the level of individual species and whole species assemblages. It is essential that these challenges are overcome before refugia can live up to their acclaim as useful targets for conservation and management in the context of climate change. [ABSTRACT FROM AUTHOR]

Aim Granite outcrops may be able to act as refugia for species during adverse climate change, owing to their topographic complexity. We assessed this hypothesis by examining phylogeographical patterns in a common, geographically widespread granite endemic, Stypandra glauca (Hemerocallidaceae). Location Granite outcrops of the Southwest Australian Floristic Region, Western Australia. Methods Twenty-four tetraploid individuals of the granite endemic Stypandra glauca were sampled from each of 12 granite outcrops: 7 from a mesic environment and 5 from the semi-arid region. Phylogenetic reconstruction and divergence-dating was achieved using Bayesian and parsimony analyses of chloroplast haplotypes from 90 individuals. Nuclear diversity and population differentiation were analysed across all individuals using 10 microsatellite loci. Results Stypandra glauca exhibited high (chloroplast) or moderate (nuclear) levels of divergence among, and low diversity within, outcrops. Haplotype diversity was high in both sampling regions, and each haplotype was unique to one outcrop. There was little correlation between geographical and genetic distance. Both nuclear and chloroplast diversity were higher in southern (mesic) outcrops than in northern (semi-arid) outcrops, although the level of chloroplast divergence among outcrops was similar for both climatic regions. Main conclusions The levels of divergence and low diversity revealed in S. glauca support a scenario of prolonged isolation and persistence on granite outcrops in both mesic and semi-arid climatic regions, with no evidence of contraction-expansion dynamics across the outcrop network. The higher levels of diversity in the southern populations may result from the maintenance of a larger effective population size in southern regions, which retained more mesic climates during drier glacial periods. Although the climatic conditions differ between outcrops in this study, our results indicate that outcrops in both regions have harboured S. glauca throughout climatic changes, accentuating the value of these habitats to biodiversity conservation under future changing climate. [ABSTRACT FROM AUTHOR]

Aim We examine variation in woody plant species richness and endemism within tropical dry forest on oceanic islands and determine what climatic and biogeographic metrics best explain native species richness and endemism across archipelagos, islands and plots. Location Oceanic islands in the Pacific. Methods Stand-level sampling (0.1 ha) at 35 different dry forest sites across 16 islands, and five archipelagos (New Caledonia, Fiji, Marquesas, Marianas and Hawaii). Descriptors of native species richness and endemism were calculated at the plot, island and archipelago level. Biogeographic and climate metrics at the archipelago, island and plot level were drawn from the literature and computer databases. The effects of biogeographic and climate metrics were investigated using linear mixed-effects models. Results Dry forests of New Caledonia and Fiji had the highest native species richness, while New Caledonia and Hawaii had the highest endemism. Native species richness and endemism within tropical dry forests on oceanic islands are primarily influenced by biogeographic metrics, especially isolation of the archipelago, and not climatic metrics. Most variance in native species richness and endemism (60% and 64%) is at the archipelago level compared with the island (8%, 16%) and plot (32%, 15%) level. At the island level, species richness in tropical dry forest is affected by precipitation, while island area significantly affects endemism. The area of forest fragments is an important predictor of native species richness and endemism in plots. Main conclusions Although dry forests in the Pacific have been exceptionally deforested and degraded, high native species richness and endemism remains in a number of forest fragments. Biogeographic metrics explain most of the variance in native species richness and endemism across scales, while climatic metrics are important at the island level. First-order assessments of native richness and endemism at the archipelago, island and stand-level are possible for forest types on oceanic archipelagos. [ABSTRACT FROM AUTHOR]

Aim Regional patterns of forest composition in West Polynesia were analysed to determine the relative importance of dispersal limitation, speciation, environmental filtering and biotic interactions. Location West Polynesia (Fiji, Tonga, Samoa). Methods We applied clustering and indicator species analysis to presence/absence data for 397 tree taxa in 338 forest plots to detect groups of plots sharing similar species composition. Patterns of plot composition and co-occurrence of congeneric species were compared between archipelagos for evidence of dispersal limitation, speciation, and environmental filtering versus competitive displacement or niche differentiation across the three archipelagos at the local (plot) and regional scales. Results Dispersal limitation was evident in the compositional segregation of inland rain forests across the three archipelagos, whereas compositional overlap was widespread for coastal forests, which are composed of species dispersed by water. Within genera, species numbers attenuated from Fiji to Tonga to Samoa, but speciation also contributed to compositional differences in all three island groups. Environmental filtering was suggested by the distinct composition of both secondary and mature inland rain forest versus coastal forest. At the local scale, however, clumping of congeners, which might suggest fine-scale environmental filtering, was observed for only a minority of inland forest genera. Most genera showed either overdispersion, suggesting competitive displacement or niche differentiation, or patterns not significantly different from random. Main conclusions Well-established biogeographical patterns of dispersal limitation and speciation contribute to West Polynesian tree diversity at the community scale. Environmental filtering and competitive displacement further structure forest communities; the evidence from this study suggests that environmental filtering is more important at a landscape scale, while competitive displacement may be more important at a local scale, although their relative importance varies considerably among taxa. [ABSTRACT FROM AUTHOR]

The independent island nations of the South Pacific have a rich and threatened terrestrial biota. Despite considerable investment of resources into conservation over the last three decades, biodiversity is dwindling and protected area systems remain inadequate. This lack of success is caused by important differences in cultural, economic, landownership, and social factors in developing Pacific Island countries, compared to developed nations that often fund conservation programs and plans. Despite the obvious need for capacity building and information exchange among stakeholders, little collaboration and development has eventuated. A coordinated and integrated approach, focusing conservation resources on national priorities, is essential to achieve efficient conservation. This will need to include active involvement of landowners, a good sociocultural understanding of target communities, improved collaboration between the various stakeholders, provision of sustainable alternative economic activities, and a commitment to long funding cycles for projects. [ABSTRACT FROM AUTHOR]

ABSTRACT Aim Identifying and protecting refugia is a priority for conservation under projected anthropogenic climate change, because of their demonstrated ability to facilitate the survival of biota under adverse conditions. Refugia are habitats that components of biodiversity retreat to, persist in and can potentially expand from under changing environmental conditions. However, the study and discussion of refugia has often been ad hoc and descriptive in nature. We therefore: (1) provide a habitat-based concept of refugia, and (2) evaluate methods for the identification of refugia. Location Global. Methods We present a simple conceptual framework for refugia and examine the factors that describe them. We then demonstrate how different disciplines are contributing to our understanding of refugia, and the tools that they provide for identifying and quantifying refugia. Results Current understanding of refugia is largely based on Quaternary phylogeographic studies on organisms in North America and Europe during significant temperature fluctuations. This has resulted in gaps in our understanding of refugia, particularly when attempting to apply current theory to forecast anthropogenic climate change. Refugia are environmental habitats with space and time dimensions that operate on evolutionary time-scales and have facilitated the survival of biota under changing environmental conditions for millennia. Therefore, they offer the best chances for survival under climate change for many taxa, making their identification important for conservation under anthropogenic climate change. Several methods from various disciplines provide viable options for achieving this goal. Main conclusions The framework developed for refugia allows the identification and description of refugia in any environment. Various methods provide important contributions but each is limited in scope; urging a more integrated approach to identify, define and conserve refugia. Such an approach will facilitate better understanding of refugia and their capacity to act as safe havens under projected anthropogenic climate change. [ABSTRACT FROM AUTHOR]

1. Testing the comparative strength and influence of age and area of islands, proximity of source propagules and disturbances on community assembly, species diversity and vegetation structure has proved difficult at large scales. The little-studied rain forests in the Tropical South Pacific (TSP) provide a unique study area to investigate determinants of community dynamics, with islands varying in age, isolation, area and cyclone frequency. We tested the effects of biogeographical factors and cyclone frequency on the species composition, species diversity and forest structure of old-growth rain forest using 1-ha inventory plots on 12 islands between New Guinea and the Solomon Islands. 2. As predicted by the General Dynamic Model of Oceanic Island Biogeography, the biogeographical variables of archipelago age and island area are the most important factors affecting species richness and diversity, with older and larger islands having higher richness and diversity. There is no significant effect of cyclone frequency on species diversity. 3. The theory that diversity drives endemism is not supported in this system as endemism is not correlated with species diversity. Instead, age and isolation of an island best explain patterns of endemism, with the latter suggesting dispersal limitations between archipelagos. 4. Proximity to source area influences species composition of lowland tropical rain forests in the TSP, which is also supported by a strong correlation between geographic distance and floristic similarity. Vector-fitting onto non-metric multidimensional scaling suggests that archipelago age and cyclone frequency may, in addition to proximity to source area, influence species composition. This implies that a species’ tolerance to cyclones affects its abundance at different cyclone frequencies. 5. Synthesis. Both biogeographical variables (island area and isolation) and cyclone frequency appear to affect community assembly in lowland rain forests in the TSP. While species are hence not ecologically equivalent, interspecific ecological differences do not seem to affect the overall patterns of species diversity, which are mostly determined by biogeographical variables, as predicted by the neutral theory. [ABSTRACT FROM AUTHOR]

Aim The biogeographical patterns and drivers of diversity on oceanic islands in the tropical South Pacific (TSP) are synthesized. We use published studies to determine present patterns of diversity on TSP islands, the likely sources of the biota on these islands and how the islands were colonized. We also investigate the effect of extinctions. Location We focus on oceanic islands in the TSP. Methods We review available literature and published molecular studies. Results Examples of typical island features (e.g. gigantism, flightlessness, gender dimorphism) are common, as are adaptive radiations. Diversity decreases with increasing isolation from mainland sources and with decreasing size and age of archipelagos, corresponding well with island biogeographical expectations. Molecular studies support New Guinea/Malesia, New Caledonia and Australia as major source areas for the Pacific biota. Numerous studies support dispersal-based scenarios, either over several 100 km (long-distance dispersal) or over shorter distances by island-hopping (stepping stones) and transport by human means (hitch-hiking). Only one vicariance explanation, the eastward drift of continental fragments (shuttles) that may have contributed biota to Fiji from New Caledonia, is supported by some geological evidence, although there is no evidence for the transport of taxa on shuttle fragments. Another vicariance explanation, the existence of a major continental landmass in the Pacific within the last 100 Myr (Atlantis theory), receives little support and appears unlikely. Extinction of lineages in source areas and persistence in the TSP has probably occurred many times and has resulted in misinterpretation of biogeographical data. Main conclusions Malesia has long been considered the major source region for the biota of oceanic islands in the TSP because of shared taxa and high species diversity. However, recent molecular studies have produced compelling support for New Caledonia and Australia as alternative important source areas. They also show dispersal events, and not vicariance, to have been the major contributors to the current biota of the TSP. Past extinction events can obscure interpretations of diversity patterns. [ABSTRACT FROM AUTHOR]

Aim Cycads constitute an ancient plant group that is generally believed to disperse poorly. However, one group of cycads (subsection Rumphiae) is thought to have dispersed relatively recently from a Malesian source area westwards to East Africa and eastwards into the Pacific, using a floatation-facilitating layer in their seeds. We use morphological and allozyme characters to investigate the relationships among the species within this group and to deduce whether the wide distribution was achieved by recent dispersal (as evidenced by high genetic similarity) or more distant vicariance events (high genetic differentiation). Location We examined specimens collected throughout the range of subsection Rumphiae, from East Africa through Southeast Asia to Tonga in the South-west Pacific. Methods We investigated relationships within subsection Rumphiae of the genus Cycas by analysing 18 variable (11 informative) morphological characters and 22 allozyme loci for seven of the 10 species currently assigned to this taxon. Results Distinctive morphological characters are few and fail to resolve relationships within the group. Allozyme data show that species within this subsection are closely related and suggest that there are two groups within the subsection, one comprising Cycas thouarsii (East Africa) and C. edentata (the Philippines), and the other the remaining species (from Malesia and the Pacific). The Australian species C. silvestris is sister to subsection Rumphiae in the morphological analysis but is closely allied to C. rumphii (nested within the subsection) in the allozyme analysis, suggesting that Rumphiae may be paraphyletic and that characters thought to be taxonomically important may need to be re-evaluated. Main conclusions Cycads within subsection Rumphiae are closely related, and the wide distribution of this group was probably achieved through relatively recent oceanic dispersal events. Separate events probably account for the dispersal of these cycads into the Pacific and to Africa. The origin and distribution of C. silvestris (Australia) could be explained by a dispersal event from New Guinea or may have resulted from a former land connection between Australia and New Guinea. [ABSTRACT FROM AUTHOR]

A letter to the editor is presented regarding the refugia as a key to climatic change management.

The article discusses impact of ocean warming and acidification on refugia for coral reefs. Topics discussed include importance of conserving refugia for conservation planning in terrestrial ecosystems, negative impact of ocean acidification water pollution, overexploitation and coral-limiting environments, and identification of refugia for coral reefs for safeguarding the persistence of these communities under anthropogenic climate change.