Vascular plant species richness in relation to habitat diversity and island area in the Finnish Archipelago

Aim The aim of this study is to explore the interrelationships between island area, species number and habitat diversity in two archipelago areas. Location The study areas, Brunskär and Getskär, are located in an archipelago in south‐western Finland. Methods The study areas, 82 islands in Brunskär and 78 in Getskär, were classified into nine habitat types based on land cover. In the Brunskär area, the flora (351 species) was surveyed separately for each individual habitat on the islands. In the Getskär area, the flora (302 species) was surveyed on a whole‐island basis. We used standard techniques to analyse the species–area relationship on a whole‐island and a habitat level. We also tested our data for the small island effect (SIE) using breakpoint and path analysis models. Results Species richness was significantly associated with both island area and habitat diversity. Vegetated area in particular, defined as island area with the rock habitat subtracted, proved to be a strong predictor of species richness. Species number had a greater association with island area multiplied by the number of habitats than with island area or habitat number separately. The tests for a SIE in the species–area relationship showed the existence of a SIE in one of the island groups. No SIE could be detected for the species–vegetated area relationship in either of the island groups. The strength of the species–area relationship differed considerably between the habitats. Main conclusions The general principles of island biogeography apply well to the 160 islands in this study. Vascular plant diversity for small islands is strongly influenced by physiographic factors. For the small islands with thin and varying soil cover, vegetated area was the most powerful predictor of species richness. The species–area curves of various habitats showed large variations, suggesting that the measurement of habitat areas and establishment of habitat‐based species lists are needed to better understand species richness on islands. We found some evidence of a SIE, but it is debatable whether this is a ‘true’ SIE or a soil cover/habitat characteristics feature.     

Ecological traits modulate bird species responses to forest fragmentation in an Amazonian anthropogenic archipelago

Aim

We assessed patterns of avian species loss and the role of morpho‐ecological traits in explaining species vulnerability to forest fragmentation in an anthropogenic island system. We also contrasted observed and detectability‐corrected estimates of island occupancy, which are often used to infer species vulnerability.

Location

Tucuruí Hydroelectric Reservoir, eastern Brazilian Amazonia.

Methods

We surveyed forest birds within 36 islands (3.4–2,551.5 ha) after 22 years of post‐isolation history. We applied species–area relationships to assess differential patterns of species loss among three data sets: all species, forest specialists and habitat generalists. After controlling for phylogenetic non‐independence, we used observed and detectability‐corrected estimates of island occupancy separately to build competing models as a function of species traits. The magnitude of the difference between these estimates of island occupancy was contrasted against species detectability.

Results

The rate of species loss as a function of island area reduction was higher for forest specialists than for habitat generalists. Accounting for the area effect, forest fragmentation did not affect the overall number of species regardless of the data set. Only the interactive model including natural abundance, habitat breadth and geographic range size was strongly supported for both estimates of island occupancy. For 30 species with detection probabilities below 30%, detectability‐corrected estimates were at least tenfold higher than those observed. Conversely, differences between estimates were negligible or non‐existent for all 31 species with detection probabilities exceeding 45.5%.

Main conclusions

Predicted decay of avian species richness induced by forest loss is affected by the degree of habitat specialisation of the species under consideration, and may be unrelated to forest fragmentation per se. Natural abundance was the main predictor of species island occupancy, although habitat breadth and geographic range size also played a role. We caution against using occupancy models for low‐detectability species, because overestimates of island occupancy reduce the power of species‐level predictions of vulnerability.

     

Quantifying island isolation – insights from global patterns of insular plant species richness

Isolation is a driving factor of species richness and other island community attributes. Most empirical studies have investigated the effect of isolation measured as distance to the nearest continent. Here we expanded this perspective by comparing the explanatory power of seventeen isolation metrics in sixty‐eight variations for vascular plant species richness on 453 islands worldwide. Our objectives were to identify ecologically meaningful metrics and to quantify their relative importance for species richness in a globally representative data set. We considered the distances to the nearest mainland and to other islands, stepping stone distances, the area of surrounding landmasses, prevailing wind and ocean currents and climatic similarity between source and target areas. These factors are closely linked to colonization and maintenance of plant species richness on islands. We tested the metrics in spatial multi‐predictor models accounting for area, climate, topography and island geology. Besides area, isolation was the second most important factor determining species richness on the studied islands. A model including the proportion of surrounding land area as the isolation metric had the highest predictive power, explaining 86.1% of the variation. Distances to large islands, stepping stone distances and distances to climatically similar landmasses performed slightly better than distance to the nearest mainland. The effect of isolation was weaker for large islands suggesting that speciation counteracts the negative effect of isolation on immigration on large islands. Continental islands were less affected by isolation than oceanic islands. Our results suggest that a variety of immigration mechanisms influence plant species richness on islands and we show that this can be detected at macro‐scales. Although the distance to the nearest mainland is an adequate and easy‐to‐calculate measure of isolation, accounting for stepping stones, large islands as source landmasses, climatic similarity and the area of surrounding landmasses increases the explanatory power of isolation for species richness.     

Ecological correlates of mammal β‐diversity in Amazonian land‐bridge islands: from small‐ to large‐bodied species

Aim

Mega hydroelectric dams have become one of the main drivers of biodiversity loss in the lowland tropics. In these reservoirs, vertebrate studies have focused on local (α) diversity measures, whereas between‐site (β) diversity remains poorly assessed despite its pivotal importance in understanding how species diversity is structured and maintained. Here, we unravel the patterns and ecological correlates of mammal β‐diversity, including both small (SM) and midsized to large mammal species (LM) across 23 islands and two continuous forest sites within a mega hydroelectric reservoir.

Location

Balbina Hydroelectric Dam, Central Brazilian Amazonia.

Methods

Small mammals were sampled using live and pitfall traps (48,350 trap‐nights), and larger mammals using camera traps (8,160 trap‐nights). β‐diversity was examined for each group using multiplicative diversity decomposition of Hill numbers, which considers the importance of rare, common and dominant species, and tested to what extent those were related to a set of environmental characteristics measured at different spatial scales.

Results

β‐diversity for both mammal groups was higher when considering species presence–absence. When considering species abundance, β‐diversity was significantly higher for SM than for LM assemblages. Habitat variables, such as differences in tree species richness and percentage of old‐growth trees, were strong correlates of β‐diversity for both SMs and LMs. Conversely, β‐diversity was weakly related to patch and landscape characteristics, except for LMs, for which β‐diversity was correlated with differences in island sizes.

Main conclusions

The lower β‐diversity of LMs between smaller islands suggests subtractive homogenization of this group. Although island size plays a major role in structuring mammal α‐diversity in several land‐bridge islands, local vegetation characteristics were additional key factors determining β‐diversity for both mammal groups. Maintaining the integrity of vegetation characteristics and preventing the formation of a large set of small islands within reservoirs should be considered in long‐term management plans in both existing and planned hydropower development in lowland tropical forests.

     

Species richness, environmental heterogeneity and area: a case study based on land snails in Skyros archipelago (Aegean Sea, Greece)

Aim To test the performance of the choros model in an archipelago using two measures of environmental heterogeneity. The choros model is a simple, easy‐to‐use mathematical relationship which approaches species richness as a combined function of area and environmental heterogeneity. Location The archipelago of Skyros in the central Aegean Sea (Greece). Methods We surveyed land snails on 12 islands of the archipelago. We informed the choros model with habitat data based on natural history information from the land snail species assemblage. We contrast this with habitat information taken from traditional vegetation classification to study the behaviour of choros with different measures of environmental heterogeneity. R² values and Akaike's information criterion (AIC) were used to compare the choros model and the Arrhenius species–area model. Path analysis was used to evaluate the variance in species richness explained by area and habitat diversity. Results Forty‐two land snail species were recorded, living in 33 different habitat types. The choros model with habitat types had more explanatory power than the classic species–area model and the choros model using vegetation types. This was true for all islands of the archipelago, as well as for the small islands alone. Combined effects of area and habitat diversity primarily explain species richness in the archipelago, but there is a decline when only small islands are considered. The effects of area are very low both for all the islands of the archipelago, and for the small islands alone. The variance explained by habitat diversity is low for the island group as a whole, but significantly increases for the small islands. Main conclusions The choros model is effective in describing species‐richness patterns of land snails in the Skyros Archipelago, incorporating ecologically relevant information on habitat occupancy and area. The choros model is more effective in explaining richness patterns on small islands. When using traditional vegetation types, the choros model performs worse than the classic species–area relationship, indicating that use of proxies for habitat diversity may be problematic. The slopes for choros and Arrhenius models both assert that, for land snails, the Skyros Archipelago is a portion of a larger biogeographical province. The choros model, informed by ecologically relevant habitat measures, in conjunction with path analysis points to the importance of habitat diversity in island species richness.     

Plants on small islands revisited: the effects of spatial scale and habitat quality on the species–area relationship

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.     

Dung beetle assemblages on tropical land‐bridge islands: small island effect and vulnerable species

Aim Using dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) in a tropical land‐bridge island system, we test for the small island effect (SIE) in the species–area relationship and evaluate its effects on species richness and community composition. We also examine the determinants of species richness across island size and investigate the traits of dung beetle species in relation to their local extinction vulnerability following forest fragmentation. Location Lake Kenyir, a hydroelectric reservoir in north‐eastern Peninsular Malaysia. Methods We sampled dung beetles using human dung baited pitfall traps on 24 land‐bridge islands and three mainland sites. We used regression tree analyses to test for the SIE, as well as species traits related to local rarity, as an indication of extinction vulnerability. We employed generalized linear models (GLMs) to examine determinants for species richness at different scales and compared the results with those from conventional linear and breakpoint regressions. Community analyses included non‐metric multidimensional scaling, partial Mantel tests, nestedness analysis and abundance spectra. Results Regression tree analysis revealed an area threshold at 35.8 ha indicating an SIE. Tree basal area was the most important predictor of species richness on small islands (<35.8 ha). Results from GLMs supported these findings, with isolation and edge index also being important for small islands. The SIE also manifested in patterns of dung beetle community composition where communities on small islands (<35.8 ha) departed from those on the mainland and larger islands, and were highly variable with no significant nestedness, probably as a result of unexpected species occurrences on several small islands. The communities exhibited a low degree of spatial autocorrelation, suggesting that dispersal limitation plays a part in structuring dung beetle assemblages. Species with lower baseline density and an inability to forage on the forest edge were found to be rarer among sites and hence more prone to local extinction. Main conclusions We highlight the stochastic nature of dung beetle community composition on small islands and argue that this results in reduced ecosystem functionality. A better understanding of the minimum fragment size required for retaining functional ecological communities will be important for effective conservation management and the maintenance of tropical forest ecosystem stability.     

Patterns of invertebrate density and taxonomic richness across gradients of area,isolation, and vegetation diversity in a lake‐island system

Over the past half century, ecologists have tried to unravel the factors that drive species richness patterns in ecological communities. One influential theory is island biogeography theory (IBT), which predicts that island or habitat area and isolation are drivers of species richness. However, relatively few studies testing IBT have considered invertebrate or belowground communities, and it is unclear as to whether the predictions made by IBT hold for these communities. Other theories predict that habitat characteristics such as vegetation diversity may be important drivers of invertebrate species richness. To investigate patterns of invertebrate density and species richness across gradients of area, isolation, and vegetation diversity, we used a system of 30 lake islands in the boreal zone of northern Sweden. We assessed density and taxonomic richness of ground‐dwelling spiders, web‐building spiders, beetles, collembolans, mites, and nematodes, for all islands during two consecutive summers. For all invertebrate groups, both density and taxonomic richness were either neutrally or negatively related to island size, and either neutrally or positively related to island isolation. Meanwhile the density and taxonomic richness for several groups was positively related to vegetation diversity (i.e. habitat heterogeneity). In multiple regression analyses, island size was often the single best predictor for both invertebrate density and taxonomic richness, but in some cases island size and isolation in combination explained more variation than each factor considered singly. Contrary to IBT predictions, invertebrate density and richness was never positively related to island size or negatively related to island isolation. Instead, our results suggest that plant diversity (and thus habitat heterogeneity) was the main driver of the patterns that we found, although other factors could have some influence. We conclude that several factors, but not necessarily those predicted as important by IBT, are important in determining invertebrate abundance and species richness in island systems.     

Do forest plants conform to the theory of island biogeography: the case study of bog islands

Biodiversity conservation is confronted with increasing risk of extinction in isolated small-area remnants and the limitation of species to colonize recently formed habitats. We hypothesized that the equilibrium pattern of forest herb layer in long-term fragmented landscape should comply with the theory of island biogeography. Forests on mineral soil islands located in large mires of western Estonia were considered as dispersal target habitats, and forests on mainland and peninsulas in mires as sources. Species richness was the lowest in mainland forests and the effect was confounded by habitat structure, suggesting a negative effect of silvicultural management in easily accessible forests. We observed the ‘small island effect’, i.e. greater overall species richness in small-area habitats, which was determined by the habitat preference of shade tolerant generalists. The average species richness of common mainland forest specialists varied little, but capitalizing on the traditional approach and analyzing only island data, weak effects of distance and habitat quality were detected. At single species level, unexpectedly, many habitat specialists were observed to have successfully dispersed to islands, indicating insufficient knowledge of the long-distance dispersal mechanisms of forest-dwelling plants. In fragmented forest landscapes the theory of island biogeography can be applied to habitat specialist plant species, but only regarding the effect of isolation and in conditions of persistent forest structural quality. In the light of global changes, optimized conservation planning should primarily target on (i) the conservation of ancient habitat fragments independent of their current area, and (ii) the promotion of diversity of potential dispersal vectors in the landscape.     

From plots to islands: species diversity at different scales

Aim To investigate how plant diversity of whole islands (‘gamma’) is related to alpha and beta diversity patterns among sampling plots within each island, thus exploring aspects of diversity patterns across scales. Location Nineteen islands of the Aegean Sea, Greece. Methods Plant species were recorded at both the whole‐island scale and in small 100 m² plots on each island. Mean plot species richness was considered as a measure of alpha diversity, and six indices of the ‘variation’‐type beta diversity were also applied. In addition, we partitioned beta diversity into a ‘nestedness’ and a ‘replacement’ component, using the total species richness recorded in all plots of each island as a measure of ‘gamma’ diversity. We also applied 10 species–area models to predict the total observed richness of each island from accumulated plot species richness. Results Mean alpha diversity was not significantly correlated with the overall island species richness or island area. The range of plot species richness for each island was significantly correlated with both overall species richness and area. Alpha diversity was not correlated with most indices of beta diversity. The majority of beta diversity indices were correlated with whole‐island species richness, and this was also true for the ‘replacement’ component of beta diversity. The rational function model provided the best prediction of observed island species richness, with Monod’s and the exponential models following closely. Inaccuracy of predictions was positively correlated with the number of plots and with most indices of beta diversity. Main conclusions Diversity at the broader scale (whole islands) is shaped mainly by variation among small local samples (beta diversity), while local alpha diversity is not a good predictor of species diversity at broader scales. In this system, all results support the crucial role of habitat diversity in determining the species–area relationship.     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.     

Low Primate Diversity and Abundance in Northern Amazonia and its Implications for Conservation

Large areas of the Rio Negro basin in Amazonia are covered by continuous tracts of tropical forest, but have few primate species. This is anomalous considering the general relationship between area and number of species. One possibility is that much of the forest is unsuitable habitat for most primates and the area of suitable habitat is much less than the forested area. This has important consequences for the design of reserves and predictions of the consequences of climate change, which tend to be based on broad categories based on satellite images, and not on information of species distributions within those broad categories. The study was conducted through diurnal and nocturnal line‐transect surveys in the Biodiversity Research Program 25‐km² permanent grid in Viruá National Park, which has vegetation associations typical of much of northern Amazonia. The highest primate diversity and abundances occurred in tall terra firme forests (58%), whereas inundated forests and scrublands, which cover 42 percent of the survey grid and 90.8 percent of the Viruá National Park, have virtually no primates. This suggests that parks and reserves in northwestern Amazonia will have to be very large to maintain viable populations of most primates and their ecological interactions, and that very broad habitat categories are not sufficient to make predictions about actual and future suitability of areas for primate conservation.     

千岛湖库区岛屿面积对植物分布的影响及植物物种多样性保护研究

修建大坝形成水库蓄水是造成景观破碎化的原因和形式之一.本研究选取了浙江千岛湖库区景观破碎化的典型区域小金山林场,调查了74个大小不一的岛屿,研究了面积对植物物种分布的影响,并比较了使物种多样性最大化的几种植物物种保护方案:小岛组合、中岛组合、大岛组合、大中小组合岛.调查共记录到乔木物种56种,灌木物种79种.将岛屿按照面积由大到小排序,将相对应分布的乔木、灌木绘制成植物物种分布表.结果显示:乔木中分布不受面积影响的物种有15种,受面积影响的有11种,介于两者之间的有30种;灌木物种中不受面积影响的有24种,受面积影响的有16种,介于两者之间的有39种.将乔木、灌木和总物种数分别累加,同时将相对应的岛屿面积按不同组合累加,绘制累加物种和面积关系图,发现在特定的相同累加面积下,组合岛的累加乔木、灌木、总物种数最多,小岛组合次之,大岛组合最少.因此,建立相同面积的保护区时,组合岛有利于保护更多的物种.     

Assemblage-level responses of phyllostomid bats to tropical forest fragmentation: land-bridge islands as a model system

Aim Working within a system of high structural contrast between fragments and the surrounding matrix, we assessed patterns of species loss and changes in species composition of phyllostomid bats on artificial land‐bridge islands relative to mainland assemblages, and evaluated the responses of bats to forest edges. We further examined the relative influence of local‐scale characteristics (e.g. vegetation structure, island area) versus landscape attributes (e.g. forest cover, patch density) and the importance of spatial scale in determining phyllostomid species richness and composition on islands. Location Islands in Gatún Lake and adjacent mainland peninsulas in the Barro Colorado Nature Monument, Panama. Methods Bats were sampled over a 2‐year period on 11 islands as well as at forest‐edge and interior sites on adjacent mainland, resulting in > 8400 captures. Results The islands harboured a less diverse and structurally simplified phyllostomid bat fauna. Islands far from the mainland were especially species‐poor. This decline in species richness was associated with compositional shifts towards assemblages strongly dominated by frugivores with good dispersal abilities. Members of other ensembles, most importantly gleaning animalivores, were much less common or absent. Although overall species composition was not significantly altered, species richness at continuous forest‐edge sites was significantly lower compared with that at interior sites. Distance from the mainland and amount of forest cover in the landscape were the best predictors of species richness and assemblage composition. Responses were scale‐dependent. At the local scale, species richness was independent of island area but was correlated positively with distance from the mainland. In contrast, area effects became more important at larger spatial scales, suggesting that many species use multiple fragments. Main conclusions Our results underline the conservation value of small habitat remnants, which, even when embedded in a hostile matrix, can support a relatively diverse bat fauna, provided that there is a low degree of patch isolation and spatial proximity to larger tracts of continuous forest. Although the results at the assemblage level were inconclusive, we demonstrate that certain bat species and ensembles, particularly gleaning animalivores, exhibit high edge‐sensitivity. Our results point to habitat loss rather than changes in landscape configuration as the main process after isolation underlying phyllostomid bat responses, suggesting that conservation efforts should focus on habitat preservation instead of trying to minimize fragmentation per se at the expense of habitat amount.     

Sampling design may obscure species–area relationships in landscape-scale field studies

We investigated 1) the role of area per se in explaining anuran species richness on reservoir forest islands, after controlling for several confounding factors. We also assessed 2) how sampling design affects the inferential power of island species–area relationships (ISARs) aiming to 3) provide guidelines to yield reliable estimates of area-induced species losses in patchy systems. We surveyed anurans with autonomous recording units at 151 plots located on 74 islands and four continuous forest sites at the Balbina Hydroelectric Reservoir landscape, central Brazilian Amazonia. We applied semi-log ISAR models to assess the effect of sampling design on the fit and slope of species–area curves. To do so, we subsampled our surveyed islands following both a 1) stratified and 2) non-stratified random selection of 5, 10, 15, 20 and 25 islands covering 1) the full range in island size (0.45–1699 ha) and 2) only islands smaller than 100 ha, respectively. We also compiled 25 datasets from the literature to assess the generality of our findings. Island size explained ca half of the variation in species richness. The fit and slope of species–area curves were affected mainly by the range in island size considered, and to a very small extent by the number of islands surveyed. In our literature review, all datasets covering a range of patch sizes larger than 300 ha yielded a positive ISAR, whereas the number of patches alone did not affect the detection of ISARs. We conclude that 1) area per se plays a major role in explaining anuran species richness on forest islands within an Amazonian anthropogenic archipelago; 2) the inferential power of island species–area relationships is severely degraded by sub-optimal sampling designs; 3) at least 10 habitat patches spanning three orders of magnitude in size should be surveyed to yield reliable species–area estimates in patchy systems.     

Nestedness in island faunas: novel insights into island biogeography through butterfly community profiles of colonization ability and migration capacity

Aim To relate variation in the migration capacity and colonization ability of island communities to island geography and species island occupancy. Location Islands off mainland Britain and Ireland. Methods Mean migration (transfer) capacity and colonization (establishment) ability (ecological indices), indexed from 12 ecological variables for 56 butterfly species living on 103 islands, were related to species nestedness, island and mainland source geography and indices using linear regression models, RLQ analysis and fourth‐corner analysis. Random creation of faunas from source species, rank correlation and rank regression were used to examine differences between island and source ecological indices, and relationships to island geography. Results Island butterfly faunas are highly nested. The two ecological indices related closely to island occupancy, nestedness rank of species, island richness and geography. The key variables related to migration capacity were island area and isolation; for colonization ability they were area, isolation and longitude. Compared with colonization ability, migration capacity was found to correlate more strongly with island species occupancy and species richness. For island faunas, the means for both ecological indices decreased, and variation increased, with increasing island species richness. Mean colonization ability and migration capacity values were significantly higher for island faunas than for mainland source faunas, but these differences decreased with island latitude. Main conclusions The nested pattern of butterfly species on islands off mainland Britain and Ireland relates strongly to colonization ability but especially to migration capacity. Differences in colonization ability among species are most obvious for large, topographically varied islands. Generalists with abundant multiple resources and greater migration capacity are found on all islands, whereas specialists are restricted to large islands with varied and long‐lived biotopes, and islands close to shore. The inference is that source–sink dynamics dominate butterfly distributions on British and Irish islands; species are capable of dispersing to new areas, but, with the exception of large and northern islands, facilities (resources) for permanent colonization are limited. The pattern of colonization ability and migration capacity is likely to be repeated for mainland areas, where such indices should provide useful independent measures for assessing the conservation status of faunas within spatial units.     

Species‐habitat relationships and ecological correlates of butterfly abundance in a transformed tropical landscape

Tropical butterfly conservation strategies often focus on total and/or common species richness to assess the conservation value of a patch or habitat. However, such a strategy overlooks the unique dynamics of rare species. We evaluated the species‐habitat relationships of 209 common, intermediate, and rare butterfly species (including morphospecies) across four habitat types (mature, degraded, or fragmented forest, and urban parks) and two patch sizes (<400 ha, ≥400 ha) in Singapore. Common species richness was consistent across habitat types. Intermediate species richness declined by more than 50 percent in urban parks (relative to all forest habitats), and rare species richness was reduced by 50 percent in degraded and fragmented forest and by 90 percent in urban parks (relative to mature forest). Large patches had comparable overall richness to small patches, but they supported more rare species and three times as many habitat‐restricted species over a similar area. Importantly, a number of rare species were confined to single small patches. Mixed‐effects regression models were constructed to identify habitat and ecological/life history variables associated with butterfly abundance. These models revealed that species with greater habitat specialization, rare larval host plants, few larval host plant genera, and narrow global geographic ranges were more likely to be rare species. Overall, these results demonstrate that the richness of habitat‐restricted and rare species do not follow the same spatial distribution patterns as common species. Therefore, while conserving mature forests is key, effective butterfly conservation in a transformed landscape should take into account rare and habitat‐restricted species.     

Influence of damming on anuran species richness in riparian areas: A test of the serial discontinuity concept

Almost all large rivers worldwide are fragmented by dams, and their impacts have been modeled using the serial discontinuity concept (SDC), a series of predictions regarding responses of key biotic and abiotic variables. We evaluated the effects of damming on anuran communities along a 245‐km river corridor by conducting repeated, time‐constrained anuran calling surveys at 42 locations along the Broad and Pacolet Rivers in South Carolina, USA. Using a hierarchical Bayesian analysis, we test the biodiversity prediction of the SDC (modified for floodplain rivers) by evaluating anuran occupancy and species diversity relative to dams and degree of urbanized land use. The mean response of the anuran community indicated that occupancy and species richness were maximized when sites were farther downstream from dams. Sites at the farthest distances downstream of dams (47.5 km) had an estimated ~3 more species than those just below dams. Similarly, species‐specific occupancy estimates showed a trend of higher occupancy downstream from dams. Therefore, using empirical estimation within the context of a 245‐km river riparian landscape, our study supports SDC predictions for a meandering river. We demonstrate that with increasing distance downstream from dams, riparian anuran communities have higher species richness. Reduced species richness immediately downstream of dams is likely driven by alterations in flow regime that reduce or eliminate flows which sustain riparian wetlands that serve as anuran breeding habitat. Therefore, to maintain anuran biodiversity, we suggest that flow regulation should be managed to ensure water releases inundate riparian wetlands during amphibian breeding seasons and aseasonal releases, which can displace adults, larvae, and eggs, are avoided. These outcomes could be achieved by emulating pre‐dam seasonal discharge data, mirroring discharge of an undammed tributary within the focal watershed, or by basing real‐time flow releases on current environmental conditions.     

Orchids of the West Indies: predictability of diversity and endemism

Aim We examined phytogeographical patterns of West Indian orchids, and related island area and maximum elevation with orchid species richness and endemism. We expected strong species–area relationships, but that these would differ between low and montane island groups. In so far as maximum island elevation is a surrogate for habitat diversity, we anticipated a strong relationship with maximum elevation and both species richness and endemism for montane islands. Location The West Indies. Methods Our data included 49 islands and 728 species. Islands were classified as either montane (≥ 300 m elevation) or low (< 300 m). Linear and multivariate regression analyses were run to detect relationships between either area or maximum island elevation and species richness or the number of island endemic species. Results For all 49 islands, the species–area relationship was strong, producing a z‐value of 0.47 (slope of the regression line) and explaining 46% of the variation. For 18 relatively homogeneous, low islands we found a non‐significant slope of z = −0.01 that explained only 0.1% of the variation. The 31 montane islands had a highly significant species–area relationship, with z = 0.49 and accounting for 65% of the variation. Species numbers were also strongly related to maximum island elevation. For all islands < 750 km², we found a small‐island effect, which reduced the species–area relationship to a non‐significant z = 0.16, with only 5% of the variation explained by the model. Species–area relationships for montane islands of at least 750 km² were strong and significant, but maximum elevation was the best predictor of species richness and accounted for 79% of the variation. The frequency of single‐island endemics was high (42%) but nearly all occurred on just nine montane islands (300 species). The taxonomic distribution of endemics was also skewed, suggesting that seed dispersability, while remarkable in some taxa, is very limited in others. Montane island endemics showed strong species–area and species–elevation relationships. Main conclusions Area and elevation are good predictors of orchid species diversity and endemism in the West Indies, but these associations are driven by the extraordinarily strong relationships of large, montane islands. The species richness of low islands showed no significant relationship with either variable. A small‐island effect exists, but the montane islands had a significant relationship between species diversity and maximum elevation. Thus, patterns of Caribbean orchid diversity are dependent on an interplay between area and topographic diversity.     

Species–area relationships in the Andaman and Nicobar Islands emerge because rarer species are disproportionately favored on larger islands

The island species–area relationship (ISAR) describes how the number of species increases with increasing size of an island (or island‐like habitat), and is of fundamental importance in island biogeography and conservation. Here, we use a framework based on individual‐based rarefaction to infer whether ISARs result from passive sampling, or whether some processes are acting beyond sampling (e.g., disproportionate effects and/or habitat heterogeneity). Using data on total and relative abundances of four taxa (birds, butterflies, amphibians, and reptiles) from multiple islands in the Andaman and Nicobar archipelago, we examine how different metrics of biodiversity (total species richness, rarefied species richness, and abundance‐weighted effective numbers of species emphasizing common species) vary with island area. Total species richness increased for all taxa, as did rarefied species richness controlling for a given sampling effort. This indicates that the ISAR did not result because of passive sampling, but that instead, some species were disproportionately favored on larger islands. For birds, frogs, and lizards, this disproportionate effect was only associated with species that were rarer in the samples, but for butterflies, both more common and rarer species were affected. Furthermore, for the two taxa for which we had plot‐level data (reptiles and amphibians), within‐island β‐diversity did not increase with island size, suggesting that within‐island compositional effects were unlikely to be driving these ISARs. Overall, our results indicate that the ISARs of these taxa are most likely driven by disproportionate effects, that is, where larger islands are important sources of biodiversity beyond a simple sampling expectation, especially through their influence on rarer species, thus emphasizing their role in the preservation and conservation of species.