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Abstract: Aim: To determine the relationships between the functional trait composition of forest communities and environmental gradients across scales and biomes and the role of species relative abundances in these relationships. Location: Global. Time period: Recent. Major taxa studied: Trees. Methods: We integrated species abundance records from worldwide forest inventories and associated functional traits (wood density, specific leaf area and seed mass) to obtain a data set of 99,953 to 149,285 plots (depending on the trait) spanning all forested continents. We computed community‐weighted and unweighted means of trait values for each plot and related them to three broad environmental gradients and their interactions (energy availability, precipitation and soil properties) at two scales (global and biomes). Results: Our models explained up to 60% of the variance in trait distribution. At global scale, the energy gradient had the strongest influence on traits. However, within‐biome models revealed different relationships among biomes. Notably, the functional composition of tropical forests was more influenced by precipitation and soil properties than energy availability, whereas temperate forests showed the opposite pattern. Depending on the trait studied, response to gradients was more variable and proportionally weaker in boreal forests. Community unweighted means were better predicted than weighted means for almost all models. Main conclusions: Worldwide, trees require a large amount of energy (following latitude) to produce dense wood and seeds, while leaves with large surface to weight ratios are concentrated in temperate forests. However, patterns of functional composition within‐biome differ from global patterns due to biome specificities such as the presence of conifers or unique combinations of climatic and soil properties. We recommend assessing the sensitivity of tree functional traits to environmental changes in their geographic context. Furthermore, at a given site, the distribution of tree functional traits appears to be driven more by species presence than species abundance.

Tropical cyclones are common disturbances on many Pacific islands and affect forest structure and community composition. However, we know little about the process of succession after disturbances in the tropical South Pacific. We utilize published data of vegetation surveys in a lowland tropical rainforest reserve in Samoa that were undertaken within 1 year, and after 6 and 15 years, after two cyclones and a fire that occurred in 1990 and 1991. We combine these surveys with functional trait data from literature and the field. Community and functional composition differed little from inferred pre-cyclone conditions soon after the disturbances but had changed considerably 6 years after the disturbance. Early successional species with functional trait characteristics relating to resources acquisition and faster growth (lower wood density, larger leaf area, shorter maximum height, smaller seeds) had become dominant 6 years after the disturbances, but had declined considerably by 15 years. No clear differences in community-weighted means of functional traits were detected between burned and unburned forest, but community composition differed considerably. In particular, the introduced rubber tree, Funtumia elastica, which was functionally intermediate between early and late successional species, had become very abundant in burned forest. Our results suggest that ecological functions may be more resilient to cyclone disturbance than community composition, but this requires further study. Our findings highlight the impact of cyclones on community composition and functioning, the importance of long-term data for investigating the recovery after disturbances, and the potential of multiple disturbances to facilitate the proliferation of invasive species. Keywords: alien invasive species, fire, Funtumia elastica, multiple disturbances, Samoa, secondary succession, tropical cyclone, wood density, TROPICAL CYCLONES have been shown to cause various levels of structural damage and increased tree mortality rates (Elmqvist et al. 1994, Franklin et al. 2004, Uriarte et al. 2019, Taylor [...]

There is a debate concerning the definition and extent of tropical dry forest biome and vegetation type at a global spatial scale. We identify the potential extent of the tropical dry forest biome based on bioclimatic definitions and climatic data sets to improve global estimates of distribution, cover, and change. We compared four bioclimatic definitions of the tropical dry forest biome-Murphy and Lugo, Food and Agriculture Organization (FAO), DryFlor, aridity index-using two climatic data sets: WorldClim and Climatologies at High-resolution for the Earth's Land Surface Areas (CHELSA). We then compared each of the eight unique combinations of bioclimatic definitions and climatic data sets using 540 field plots identified as tropical dry forest from a literature search and evaluated the accuracy of World Wildlife Fund tropical and subtropical dry broadleaf forest ecoregions. We used the definition and climate data that most closely matched field data to calculate forest cover in 2000 and change from 2001 to 2020. Globally, there was low agreement (< 58%) between bioclimatic definitions and WWF ecoregions and only 40% of field plots fell within these ecoregions. FAO using CHELSA had the highest agreement with field plots (81%) and was not correlated with the biome extent. Using the FAO definition with CHELSA climatic data set, we estimate 4,931,414 km2 of closed canopy (≥ 40% forest cover) tropical dry forest in 2000 and 4,369,695 km2 in 2020 with a gross loss of 561,719 km2 (11.4%) from 2001 to 2020. Tropical dry forest biome extent varies significantly based on bioclimatic definition used, with nearly half of all tropical dry forest vegetation missed when using ecoregion boundaries alone, especially in Africa. Using site-specific field validation, we find that the FAO definition using CHELSA provides an accurate, standard, and repeatable way to assess tropical dry forest cover and change at a global scale.

The integration and synthesis of data from varied sources is becoming increasingly common in biogeography and ecology. Although analyses of large, global datasets from multiple, varied sources only became common recently, such studies trace their origin to the early 19th century. Alexander von Humboldt (1769-1859) was an eminent and much-celebrated scientist, who introduced the concept of collecting high-quality quantitative data across continents. More importantly, he championed inter-disciplinary research, data-driven studies, and global comparative syntheses. With these studies, he helped to lay the foundations for the fields of biogeography and ecology, and his ideas, concepts, and scientific approach remain at the forefront of exciting recent developments in both disciplines. Remembering Humboldt’s 250th birthday, we show how his scientific philosophy and visionary approach to multi-disciplinary data integration and synthesis have recently been revitalized due to increasing computing power and the numerous complex environmental challenges facing our planet.

BackgroundRefugia are island-like habitats that are linked to long-term environmental stability and, as a result, high endemism. Conservation of refugia and endemism hotspots should be based on a deep ecological and evolutionary understanding of their functioning, which remains limited. Although functional traits can provide such insights, a corresponding, coherent framework is lacking.Proposed frameworkPlant communities in refugia and endemism hotspots should, due to long-term environmental stability, display unique functional characteristics linked to distinct phylogenetic patterns. Therefore, such communities should be characterized by a functional signature that exhibits: (1) distinct values and combinations of traits, (2) higher functional diversity and (3) a prevalence of similar traits belonging to more distantly related lineages inside, compared to outside, of endemism hotspots and refugia. While the limited functional trait data available from refugia and endemism hotspots do not allow these predictions to be tested rigorously, three potential applications of the functional signature in biogeography and conservation planning are highlighted. Firstly, it allows the functional characteristics of endemism hotspots and refugia to be identified. Secondly, the strength of the functional signature can be compared among these entities, and with the surrounding landscape, to provide an estimate of the capacity of endemism hotspots and refugia to buffer environmental changes. Finally, the pattern of the functional signature can reveal ecological and evolutionary processes driving community assembly and functioning, which can assist in predicting the effect of environmental changes (e.g. climate, land-use) on communities in endemism hotspots and refugia.ConclusionThe proposed functional signature concept allows the systematic integration of plant functional traits and phylogeny into the study of endemism hotspots and refugia, but more data on functional traits in these entities are urgently needed. Overcoming this limitation would facilitate rigorous testing of the proposed predictions for the functional signature, advancing the eco-evolutionary understanding of endemism hotspots and refugia.

Experimental approaches to measure territory defence can be used to interpret the relative magnitude of intraspecific aggression in different ecological contexts. The aim of this study is to examine the response of Fijian forest bird species to conspecific intruders using playback of previously recorded songs along an elevational gradient. We address two questions: Do different foraging guilds in Fiji's forest birds show different patterns of territorial defence behaviour towards conspecific intruders? Secondly, do Fiji's forest birds show stronger territorial defence at higher elevations? We explore these questions across two foraging guilds (insectivore and omnivore) on Viti Levu Island, Fiji. All four focal species had a stronger response at the highest elevation, and the endemic insectivore had the strongest response. Perhaps high elevation insects are less abundant and hence increased aggressiveness is favoured to defend the territory, and/or perhaps in general more aggressive birds colonise higher elevation areas, both ideas that require further testing. This study provides first insights into response to conspecifics in Fiji's forest birds that can be explored in relation to avian behaviour also in the context of human activity, human disturbance and threats to the persistence of birds across elevational gradients. [ABSTRACT FROM AUTHOR]

Landslides, resulting from disturbances in slope equilibrium, pose a significant threat to landscapes, infrastructure, and human life. Triggered by factors such as intense precipitation, seismic activities, or volcanic eruptions, these events can cause extensive damage and endanger nearby communities. A comprehensive understanding of landslide characteristics, including spatio-temporal patterns, dimensions, and morphology, is vital for effective landslide disaster management. Existing remote sensing approaches mostly use either optical or synthetic aperture radar sensors. Integrating information from both these types of sensors promises greater accuracy for identifying and locating landslides. This study proposes a novel approach, the ML-LaDeCORsat (Machine Learning-based coseismic Landslide Detection using Combined Optical and Radar Satellite Imagery), that integrates freely available Sentinel-1, Palsar-2, and Sentinel-2 imagery data in Google Earth Engine (GEE). The approach also integrates relevant spectral indices and suitable bands used in a machine learning-based classification of coseismic landslides. The approach includes a robust and reproducible training and validation strategy and allows one to choose between five classifiers (CART, Random Forest, GTB, SVM, and Naive Bayes). Using landslides from four different earthquake case studies, we demonstrate the superiority of our approach over existing solutions in coseismic landslide identification and localization, providing a GTB-based detection accuracy of 87–92%. ML-LaDeCORsat can be adapted to other landslide events (GEE script is provided). Transfer learning experiments proved that our model can be applied to other coseismic landslide events without the need for additional training data. Our novel approach therefore facilitates quick and reliable identification of coseismic landslides, highlighting its potential to contribute towards more effective disaster management. [ABSTRACT FROM AUTHOR]

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]

Introduction: Seagrass roots harbour diverse assemblages of microorganisms that likely benefit the growth and survival of meadows. Yet, restoration efforts rarely consider their effect on developing seagrass seedlings. Sediment origin should determine the types of rhizosphere and root-colonising (rhizoplane) microorganisms and thus the performance of seedlings during restoration, particularly for slow growing climax species like Posidonia. Recent Posidonia restoration attempts in South Australia used commercially sourced 'play pit sand' for seedling propagation and planting, but have been impacted by high mortality. More natural substrates like seagrass meadow sediment have not been previously investigated for suitability over conventional substrates with regard to seedling growth and survival. Methods: To assess the relevance of seagrass associated microorganisms in the growth of Posidonia angustifolia seedlings, we investigated the bacterial communities from tank-raised seedlings propagated in autoclave treated and untreated 'play pit sand' and meadow mix (comprising a 1:3 ratio of natural meadow sediment and beach sand) over a 12-week period. Autoclave treatment was adopted in order to diminish the bacterial load prior to planting and thus inform their contribution (if any) on early seedling growth. Samples for bacterial community analysis and seedling growth measurements (numbers and total length of roots/leaves, root diameter, seedling weight, starch reserves) were taken at 4 and 12 weeks. Bacterial assemblages were surveyed from DNA extracts from bulk and rhizosphere sediments and root tissues, as well as from swabs of P. angustifolia fruit, established meadow sediment and water samples prior to seedling propagation, by constructing Illumina 16S rRNA gene libraries. Results: While most growth measurements did not vary significantly between sediment type or treatment, proportional growth of roots versus leaves (as expressed as a pseudo root:shoot ratio) was significantly related to treatment, sediment type and seed length. Seedlings from meadow mix invested more in leaves, regardless of treatment, when compared to play sand. Autoclave treatment increased investment in roots for play sand but increased the investment in leaves for meadow mix. Bacterial communities differed significantly between sediments and between sample types (bulk, rhizosphere and roots), with the roots from meadow mix seedlings containing an increased abundance of various potentially beneficial bacterial taxa. Discussion: While such changes appear to affect the early development of seedlings, bacterial community dynamics are also likely coupled to changes in nutrient availability. Further research is thus required to disentangle host seedling growth-nutrient-bacterial community dynamics with the view to identifying microbes that may support the growth and vigour of seedlings under different nutrient conditions as part of future restoration efforts. [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]

A disjunct population of red stringybark (Eucalyptus macrorhyncha) trees in South Australia is experiencing increasing amounts of dieback. Because the population is considered vulnerable to extinction, we investigated spatiotemporal vegetation changes, quantified the extent of dieback, and determined how topography influences dieback using aerial and satellite imagery. Classification of vegetation health status using hyperspectral aerial imagery indicated that 37% (accuracy = 0.87 Kappa) of the population was unhealthy and potentially experiencing dieback. When correlating this classification with a digital terrain model (DTM), the aspect and amount of solar radiation had the strongest relationship with the presence of unhealthy vegetation. PlanetScope satellite-derived, and spectral index-based analysis indicated that 7% of the red stringybark population experienced negative vegetation health changes during a five-year period (2017–2022), with positive vegetation health changes (9.5%) noted on pole-facing slopes. Therefore, our integrated remote sensing approach documented the extent and spatiotemporal dynamics of dieback, suggesting it could be applied in other studies. Topographical aspects exposed to high-solar radiation were particularly vulnerable to dieback, and pole-facing aspects demonstrated some recovery between droughts. The influence of topography and maps of vegetation health can be used to guide future management and restoration of the population. [ABSTRACT FROM AUTHOR]

Context. Habitat loss is a global problem and in Fiji >50% of the land area once covered by forests has been converted to grasslands and agricultural land. About 99% of Fiji's endemic biodiversity and 80% of the land bird species have been identified as forest species. Aims. In this study, we compare forest and grassland sites and test for consistency in avian diversity, abundance, foraging guild, and distribution status (endemic, native, introduced to Fiji) over a 5-year period (2016-2020). Methods. We surveyed bird communities using the point count method with a 100 m radius and 7-min observation period per site. Key results. A one-way analysis of similarities (ANOSIM) analysis showed significant differences in species composition and bird abundance between the forested habitats and grassland habitats. A general linear model test showed significant differences in foraging guild composition and distribution status between forested and grassland habitats. There were no significant differences between the three forested sites (primary montane forest, secondary old-growth forest, old-growth mahogany plantations with regenerating native species), while grassland sites had stronger annual change in species composition. Implications. Forest cover, irrespective of whether these forests are of primary or secondary nature, therefore plays an important role in maintaining the native and endemic land bird species and other biodiversity in oceanic island ecosystems such as Viti Levu Island, Fiji. [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]

Expert knowledge is widely used for assessing the conservation status of little-known species. Pterocymbium oceanicum is a large, emergent tree endemic to Fiji. Experts classified the species as 'Critically Endangered' (IUCN Red List) in 2016 and estimated the population size at <250 mature individuals with the largest subpopulation consisting of a single stand of 100 individuals. We surveyed this stand and counted 133 individuals, which validated the expert estimate. However, we discovered an additional three stands nearby, increasing the total subpopulation size to 433 individuals. Therefore, the expert estimate reliably and conservatively estimated population size but missed important information. Field surveys suggest that P. oceanicum has a narrow ecological niche as a long-lived early pioneer at mid-elevations (400–800 m) in seasonally dry forests, one of Fiji's most degraded ecosystems. The introduced African Tulip Tree, Spathodea campanulata , is invading the sampled subpopulation and shares the characteristics of being a large, fast-growing and early successional tree. Therefore, the narrow niche of P. oceanicum may now be threatened by S. campanulata. Our study highlights that expert knowledge can provide valuable information about threatened species but also has limitations. IUCN Red List assessments based solely on expert assessments should therefore be cautiously interpreted. The invasion by S. campanulata poses a serious threat to P. oceanicum and needs to be carefully monitored, with the impacts of S. campanulata on the establishment and growth of P. oceanicum requiring urgent investigation. Pterocymbium oceanicum is a large, emergent tree endemic to Fiji that was classified as 'Critically Endangered' (IUCN Red List) based on expert knowledge. Our field survey quadruples the known size of the largest subpopulation by discovering three previously unknown stands, highlighting the value and limitations of expert knowledge for conservation assessments. Furthermore, we found that the species has a narrow ecological niche and is potentially threatened by the invading African Tulip Tree, Spathodea campanulata , which should be carefully monitored. [ABSTRACT FROM AUTHOR]

Charles Darwin proposed his 'principle of divergence' to account for changes in traits that could promote speciation and coexistence of diverse forms through occupation of different niches to reduce interspecific competition. We explore interspecific foraging behaviour overlap in Fiji's forest birds, and address two main questions: (1) Is there vertical stratification of foraging behavior? and (2) Is there evidence of interspecific competition driving the differences in foraging behaviour? We explore these questions across three foraging guilds, nectarivores (three species), insectivores (two species), and omnivores (two species), and find vertical portioning of foraging in each group. To investigate the effect of interspecific competition, we compared foraging heights of the Orange-breasted Myzomela (Myzomela jugularis) honeyeater on Viti Levu Island (where it coexists with two other honeyeater species) and Leleuvia Island (no other honeyeater species). On the main island Viti Levu, we found evidence for vertical niche partitioning within each foraging guild. On Leleuvia, with the 'one-species only foraging guild', Orange-breasted Myzomela occupied broader vertical foraging niche than on Viti Levu with two other competitor honeyeater species. This result supports the idea that vertical foraging height can be shaped by interspecific competition. The findings of this study support Darwin's principle of divergence in Fiji's forest birds for every foraging guild measured and adds to our understanding of the significance of interspecific competition and niche divergence for patterns of ecological speciation on islands. [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]