Direct and indirect effects of area,energy and habitat heterogeneity on breeding bird communities

Aim To compare the ability of island biogeography theory, niche theory and species–energy theory to explain patterns of species richness and density for breeding bird communities across islands with contrasting characteristics. Location Thirty forested islands in two freshwater lakes in the boreal forest zone of northern Sweden (65°55′ N to 66°09′ N; 17°43′ E to 17°55′ E). Methods We performed bird censuses on 30 lake islands that have each previously been well characterized in terms of size, isolation, habitat heterogeneity (plant diversity and forest age), net primary productivity (NPP), and invertebrate prey abundance. To test the relative abilities of island biogeography theory, niche theory and species–energy theory to describe bird community patterns, we used both traditional statistical approaches (linear and multiple regressions) and structural equation modelling (SEM; in which both direct and indirect influences can be quantified). Results Using regression‐based approaches, area and bird abundance were the two most important predictors of bird species richness. However, when the data were analysed by SEM, area was not found to exert a direct effect on bird species richness. Instead, terrestrial prey abundance was the strongest predictor of bird abundance, and bird abundance in combination with NPP was the best predictor of bird species richness. Area was only of indirect importance through its positive effect on terrestrial prey abundance, but habitat heterogeneity and spatial subsidies (emerging aquatic insects) also showed important indirect influences. Thus, our results provided the strongest support for species–energy theory. Main conclusions Our results suggest that, by using statistical approaches that allow for analyses of both direct and indirect influences, a seemingly direct influence of area on species richness can be explained by greater energy availability on larger islands. As such, animal community patterns that seem to be in line with island biogeography theory may be primarily driven by energy availability. Our results also point to the need to consider several aspects of habitat quality (e.g. heterogeneity, NPP, prey availability and spatial subsidies) for successful management of breeding bird diversity at local spatial scales and in fragmented or insular habitats.     

BIODIVERSITY RESEARCH: Nestedness for different reasons: the distributions of birds,lizards and small mammals on islands of an inundated lake

Aim We examined whether the community compositions of birds, lizards and small mammals were nested in a fragmented landscape in the Thousand Island Lake, China. We also assessed whether the mechanisms influencing nestedness differed among these taxonomic groups. Location Thousand Island Lake, China. Methods Presence/absence matrices were compiled for birds (42 islands) and lizards (42 islands) using line‐transect methods, and for small mammals (14 islands) using live‐trapping methods from 2006 to 2009. Nestedness was analysed using BINMATNEST, and statistical significance was assessed using the conservative null model 3. We used Spearman rank correlations and partial Spearman rank correlations to examine associations of nestedness and habitat variables (area, isolation, habitat diversity and plant richness) as well as life‐history traits (body size, habitat specificity, geographical range size and area requirement) related to species extinction and immigration tendencies. Results The community compositions of birds, lizards and small mammals were all significantly nested, but the causal factors underlying nestedness differed among taxonomic groups. For birds, island area, habitat specificity and area requirement were significantly correlated with nestedness after controlling for other independent variables. For lizards, habitat heterogeneity was the single best correlate of nestedness. For small mammals, island area, habitat heterogeneity and habitat specificity were significantly correlated with nestedness. The nested patterns of birds, lizards and small mammals were not attributable to passive sampling or selective colonization. Main conclusions The processes influencing nested patterns differed among taxonomic groups. Nestedness of bird assemblages was driven by selective extinction, and lizard assemblage was caused by habitat nestedness, while nestedness of small mammals resulted from both selective extinction and habitat nestedness. Therefore, we should take taxonomic differences into account when analysing nestedness to develop conservation guidelines and refrain from using single taxa as surrogates for others.     

千岛湖陆桥岛屿地表蚂蚁群落物种多样性空间格局及其影响因素

为了探讨千岛湖岛屿景观参数对地表蚂蚁群落物种α和β多样性空间格局的影响, 作者分别于2017和2018年的5-8月, 采用陷阱法、凋落物分拣法和手捡法调查了千岛湖33个岛屿上的地表蚂蚁群落, 并依据食性将其划分为捕食性蚂蚁和杂食性蚂蚁。利用回归模型分析了全部蚂蚁、捕食性蚂蚁和杂食性蚂蚁群落α和β多样性与岛屿景观参数的关系。结果表明, 岛屿面积对全部蚂蚁、捕食性蚂蚁和杂食性蚂蚁的物种丰富度均有显著的正向影响, 而隔离度则无显著作用。蚂蚁群落的β多样性由空间周转组分主导。岛屿面积差对全部蚂蚁、捕食性蚂蚁和杂食性蚂蚁群落β多样性的嵌套组分有正向影响, 隔离度差只对杂食性蚂蚁的总体β多样性有正向影响。因此, 岛屿面积是影响千岛湖地表蚂蚁群落物种丰富度的主要因素, 并且岛屿面积通过嵌套组分来影响蚂蚁群落的β多样性, 表现出选择性灭绝过程。此外, 不同食性蚂蚁可能因为扩散能力的差异对岛屿景观参数产生不同的响应。     

千岛湖陆桥岛屿鸟类集团对栖息地片段化的敏感性及其季节变化

为探究千岛湖陆桥岛屿不同鸟类集团对栖息地片段化敏感性的差异和季节变化,于2009年4月—2012年1月鸟类繁殖季(4、5、6月)和冬季(11、12、1月)对千岛湖41个陆桥岛屿鸟类集团进行了研究。结果表明,冬季杂食鸟对片段化敏感性高于食虫鸟,繁殖季时二者无显著差异,繁殖季和冬季时下层鸟对片段化敏感性均高于林冠鸟,冬季留鸟对片段化敏感性高于候鸟,繁殖季则无显著差异。杂食鸟和留鸟对片段化敏感性存在季节差异,而食虫鸟、林冠鸟、下层鸟和候鸟对片段化敏感性均无季节差异。不同鸟类集团对栖息地片段化敏感性的差异和季节变化规律,有助于人们在栖息地管理和保护区设计时采取更有针对性的鸟类保护措施。     

Hot,dry and different: Australian lizard richness is unlike that of mammals,amphibians and birds

Aims (1) To map the species richness of Australian lizards and describe patterns of range size and species turnover that underlie them. (2) To assess the congruence in the species richness of lizards and other vertebrate groups. (3) To search for commonalities in the drivers of species richness in Australian vertebrates. Location Australia. Methods We digitized lizard distribution data to generate gridded maps of species richness and β‐diversity. Using similar maps for amphibians, mammals and birds, we explored the relationship between species richness and temperature, actual evapotranspiration, elevation and local elevation range. We used spatial eigenvector filtering and geographically weighted regression to explore geographical patterns and take spatial autocorrelation into account. We explored congruence between the species richness of vertebrate groups whilst controlling for environmental effects. Results Lizard richness peaks in the central deserts (where β‐diversity is low) and tropical north‐east (where β‐diversity is high). The intervening lowlands have low species richness and β‐diversity. Generally, lizard richness is uncorrelated with that of other vertebrates but this low congruence is strongly spatially structured. Environmental models for all groups also show strong spatial heterogeneity. Lizard richness is predicted by different environmental factors from other vertebrates, being highest in dry and hot regions. Accounting for environmental drivers, lizard richness is weakly positively related to richness of other vertebrates, both at global and local scales. Main conclusions Lizard species richness differs from that of other vertebrates. This difference is probably caused by differential responses to environmental gradients and different centres of diversification; there is little evidence for inter‐taxon competition limiting lizard richness. Local variation in habitat diversity or evolutionary radiations may explain weak associations between taxa, after controlling for environmental variables. We strongly recommend that studies of variation in species richness examine and account for non‐stationarity.     

黄淮平原农业景观不同生境鸟类多样性特征及年际动态

鸟类是地球生物多样性中的重要组成部分,在生态系统功能和服务中发挥着重要作用,是生态环境变化的重要指示物种。农业景观中的食虫鸟类提供了重要的虫害控制服务。当前,农业景观中鸟类多样性丧失加剧,为探讨鸟类多样性在各生境以及年际间的变化,以黄淮平原为研究区,在河流、湖泊、农田、村庄等生境中共设置20个样点。于2016-2019年连续4年在繁殖期采用样线法对鸟类进行多样性调查。调查结果显示：（1）共发现22922只个体,分属14目,38科,53属。从区系分布来看,各生境各年间均以广布种为主;从生态类群来看,鸣禽占绝对优势;从居留型来看,留鸟所占比例最高。（2）在食性组成上,从物种丰富度看,食虫鸟类有57种,约占总物种数的77%;从个体数来看,杂食性鸟类占比超52%。（3）物种丰富度、多样性和均匀度指数最高值均出现在湖泊或河流生境中。（4）鸟类群落相似性分析显示,各生境间鸟类群落均为中等相似程度;鸟类物种丰富度波动幅度在农田和村庄中呈逐年上升趋势。（5）物种多样性加性拆分分析显示,在生境尺度上,局地的α多样性是生物多样性的最重要组成,而从整个研究区来看,生境间的差异则更为重要。造成鸟类多样性时空差异的原因复杂多样,而生境异质性的增加和水域的存在对提高鸟类多样性是具有积极作用的。调查中超过77%的物种和40%的个体均为食虫鸟类,应当充分重视鸟类为区域农业景观提供的虫害控制服务。本研究可为区域鸟类多样性保护及鸟类提供的生态系统服务的提升管理提供理论基础和科学依据。     

Winter bird communities in the Åland archipelago: an island biogeographic point of view

The distribution of wintering land birds was examined in the archipelago of Åland (60°N, 20°E) along two ecological dimensions: (1) the island continuum from small skerries ((0.5 ha) to the mainland of Åland (97000 ha), and (2) the habitat range of the mainland of Åland. In the species-area relationship, both the exponential and power function models fitted well. The increase of the number of species with island size was interpreted as a consequence of increasing habitat diversity on larger islands. The smallest islands supported only few seed-eating species utilizing the islands in a fine-grained fashion; none of these birds belonged to the breeding fauna of the islands. With increasing island size and habitat diversity, insectivores and species tied to the human culture were added. On the mainland of Åland, more than half of the birds were observed near human habitations: bird communities wintering in pine forests were richer than in deciduous forests. Proportion of wintering species was greatest in the breeding communities of pine forests. The proportion of wintering species was presumably regulated by the degree of structural change in the habitat between summer and winter, this change being most drastic in deciduous habitats The species observed could be broadly classified into three feeding categories: opportunists ., omnivores and scavengers, dependent on the diverse food items provided by man; food specialists , species éating seeds and berries, ranging widely in the islands and habitats; and insectivorous habitat specialists , inhabiting coniferous habitats, and islands with coniferous (pine) forests. Opportunists, in particular, gel increasing wintering opportunities by the impact of man.     

湿地公园鸟类栖息地营建研究——以北京琉璃河湿地公园为例

北京地区处于全球候鸟东亚-澳大利西亚的迁徙路线上,是候鸟重要的迁徙路线,近些年,随着人为活动的影响,该区生境破碎化问题愈发突出,直接威胁着本地鸟种和过境迁徙鸟类的生存。为达到保护鸟类多样性的目的,需开展相应的栖息地恢复工作。不同生态类群的鸟类对栖息地有着不同的要求,相同鸟种在不同空间、季节和生活期对栖息地的选择也有着不同的特点。因而,鸟类栖息地恢复应针对目标鸟种根据其繁殖特点、巢位空间分布、食性特点、活动空间特点等进行规划营造。以北京房山琉璃河湿地公园为例,针对项目所在区域的鸟类分布特征,确定目标恢复鸟种,结合项目区现场条件,围绕目标鸟种对于栖息地水系、植被等方面的需求,从岸线重塑、水深设计、植物配置、生态鸟岛等方面规划设计鸟类栖息地修复措施。     

Bird assemblages on Amazonian river islands: Patterns of species diversity and composition

The principles of island biogeography are rarely applied to the animal assemblages of Amazonian river islands. Here, we compare bird assemblages of Amazonian river islands with a variety of mainland habitats. We also examine how bird species diversity and composition are related to island physical attributes. Birds were sampled with mist nets and qualitative censuses on 11 river islands and 24 mainland sites on the lower reaches of the Rio Negro in the Brazilian Amazon. Island bird assemblages were characterized by lower species richness and a higher abundance of a few dominant species. Additionally, the species composition of the islands was distinct from that of the mainland, including the nearby floodplain habitats. The number of bird species increased with island size and habitat diversity, and decreased with degree of isolation. In addition, small islands tended to harbor an impoverished subset of the species present on larger ones. Bird species diversity and composition on Amazonian river islands are likely influenced by the ecological succession and historical events affecting island formation. Considering their small total area across the Amazon basin, these insular fluvial communities could be disproportionately threatened by river channel disturbances related to climate change or hydroelectric dam development. Abstract in Portughese is available with online material.     

Non‐random patterns of invasion and extinction reduce phylogenetic diversity in island bird assemblages

Anthropogenically driven changes in bird communities on oceanic islands exemplify the biotic upheaval experienced by island floras and faunas. While the influence of invasions and extinctions on species richness and beta‐diversity of island bird assemblages has been explored, little is known about the impact of these invasions and extinctions on phylogenetic diversity. Here we quantify phylogenetic diversity of island bird assemblages resulting from extinctions alone, invasions alone, and the combination of extinctions and invasions in the historic time period (1500 CE to the current), and compare it to the expected phylogenetic diversity that would result if these processes involved randomly selected island bird species. We assessed phylogenetic diversity and structure at the scale of the island (n = 152), the archipelago containing the islands (n = 22), and the four oceans containing the archipelagos using three measures. We found that extinction, invasion, and the combination of invasion and extinction generally resulted in lower phylogenetic diversity than expected, regardless of the spatial scale examined. We conclude that extinction and invasion of birds on islands are non‐random with respect to phylogeny and that these processes generally leave bird assemblages with lower phylogenetic diversity than we would expect under random invasion or extinction.     

Host phylogeography and beta diversity in avian haemosporidian (Plasmodiidae) assemblages of the Lesser Antilles

1. We estimated the correlation between host phylogeographical structure and beta diversity of avian haemosporidian assemblages of passerine birds to determine the degree to which parasite communities change with host evolution, expressed as genetic divergence between island populations, and we investigated whether differences among islands in the haemosporidia of a particular host species reflect beta diversity in the entire parasite assemblage, beta diversity in vectors, turnover of bird species and/or geographical distance. 2. We used Mantel tests to assess the significance of partial correlations between host nucleotide difference (based on cytochrome b) and haemosporidian (Haemoproteus spp. and Plasmodium spp.) mitochondrial lineage beta diversity within a given host species and between Plasmodium mitochondrial lineage beta diversity and mosquito and bird species beta diversity (or turnover). Three abundant and widespread host species (Tiaris bicolor, Coereba flaveola and Loxigilla noctis/barbadensis) were included in the study. Haemosporidian lineage beta diversity among nine islands was assessed using the Chao-Jaccard, Chao-S?rensen and Morisita-Horn indices of community similarity. Beta diversity indices of mosquito species and turnover of bird species were calculated from data in published records and field guides. 3. In Loxigilla spp., we found a positive correlation with geographical distance and an unexpected negative correlation between haemosporidian beta diversity and host genetic distance. Tiaris bicolor exhibited a significant positive correlation between haemosporidian beta diversity and beta diversity within the entire parasite assemblage. We did not find significant correlations between parasite beta diversity and mosquito beta diversity or bird species turnover. 4. Host phylogeographical structure does not appear to drive within-host beta diversity of haemosporidian lineages. Instead, the array of parasites on one host can reflect the haemosporidian assemblage on other hosts.     

Temporal Beta Diversity of Bird Assemblages in Agricultural Landscapes: Land Cover Change vs. Stochastic Processes

Temporal variation in the composition of species assemblages could be the result of deterministic processes driven by environmental change and/or stochastic processes of colonization and local extinction. Here, we analyzed the relative roles of deterministic and stochastic processes on bird assemblages in an agricultural landscape of southwestern France. We first assessed the impact of land cover change that occurred between 1982 and 2007 on (i) the species composition (presence/absence) of bird assemblages and (ii) the spatial pattern of taxonomic beta diversity. We also compared the observed temporal change of bird assemblages with a null model accounting for the effect of stochastic dynamics on temporal beta diversity. Temporal assemblage dissimilarity was partitioned into two separate components, accounting for the replacement of species (i.e. turnover) and for the nested species losses (or gains) from one time to the other (i.e. nestedness-resultant dissimilarity), respectively. Neither the turnover nor the nestedness-resultant components of temporal variation were accurately explained by any of the measured variables accounting for land cover change (r²<0.06 in all cases). Additionally, the amount of spatial assemblage heterogeneity in the region did not significantly change between 1982 and 2007, and site-specific observed temporal dissimilarities were larger than null expectations in only 1% of sites for temporal turnover and 13% of sites for nestedness-resultant dissimilarity. Taken together, our results suggest that land cover change in this agricultural landscape had little impact on temporal beta diversity of bird assemblages. Although other unmeasured deterministic process could be driving the observed patterns, it is also possible that the observed changes in presence/absence species composition of local bird assemblages might be the consequence of stochastic processes in which species populations appeared and disappeared from specific localities in a random-like way. Our results might be case-specific, but if stochastic dynamics are generally dominant, the ability of correlative and mechanistic models to predict land cover change effects on species composition would be compromised.     

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.     

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.     

Island theory, matrix effects and species richness patterns in habitat fragments

Island biogeography theory, created initially to study diversity patterns on islands, is often applied to habitat fragments. A key but largely untested assumption of this application of theory is that landscape matrix species composition is non‐overlapping with that of the islands. We tested this assumption in successional old field patches in a closely mowed matrix, and because our patches are appropriately viewed as sets of contiguous habitat units we studied patterns of species richness per unit area. Previous studies at our site did not find that diversity patterns on patch ‘islands’ conformed to predictions of island biogeography theory. Our results indicate that when matrix species are removed from the patch samples, diversity patterns conform better to theory. We suggest that classical island theory remains an appropriate tool to study diversity patterns in fragmented habitats, but that allowances should be made for spill‐over colonization of ‘islands’ from the ‘sea’.     

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.     

The contribution of different habitat types to species diversity of Baltic uplift islands

Islands are vulnerable ecosystems worldwide, increasingly exposed to human pressure, global climate change and invasive species. Thus, understanding island species diversity is key for nature conservation. Recent studies on insular plant communities indicated that habitat-specific species composition and richness might largely affect diversity patterns observed at the island scale. In consequence, habitat-based approaches are needed to (i) estimate how environmental changes at the habitat scale may affect island diversity, and to (ii) estimate the contribution of different patches of the same habitat to island diversity with respect to habitat-specific environmental constraints.In the present study, we tested these habitat-to-island diversity relationships for shoreline habitats (brackish reeds, salt marsh, rocky shore, tall herbs) and island interior habitats (rocks, semi-natural grassland, pioneer forest, coniferous forest, mixed forest) using 108 islands of three Baltic archipelagos in Sweden. These islands differed in terms of island-scale variables describing effects of island configuration and distance, and habitat-scale variables representing the effects of habitat area, abiotic environment and land-use.The studied habitats differed in their contribution to island species diversity, called habitat specificity. Shoreline habitats shared many common specialist species adapted to extreme conditions like sea salt or bird grazing, while habitats of the island interior harbored mainly species adapted to the specific conditions of a single habitat. We found high variability in habitat specificity as a consequence of habitat-specific environmental factors. Variability was highest for grasslands, where it was related to abandonment and soil fertility, stressing the importance of grassland management for maintaining island biodiversity. Habitats with high habitat specificity through either high species richness or many habitat-specific specialists should be the primary targets for biodiversity management.     

Small island biogeography in the Gulf of California: lizards, the subsidized island biogeography hypothesis, and the small island effect

Aim We used insular lizard communities to test the predictions of two hypotheses that attempt to explain patterns of species richness on small islands. We first address the subsidized island biogeography (SIB) hypothesis, which predicts that spatial subsidies may cause insular species richness to deviate from species–area predictions, especially on small islands. Next, we examine the small island effect (SIE), which suggests small islands may not fit the traditional log‐linear species–area curve. Location Islands with arthropodivorous lizard communities throughout the Gulf of California. Methods To evaluate the SIB hypothesis, we first identified subsidized and unsubsidized islands based on surrogate measures of allochthonous productivity (i.e. island size and bird presence). Subsequently, we created species–area curves from previously published lizard species richness and island area data. We used the residuals and slopes from these analyses to compare species richness on subsidized and unsubsidized islands. To test for an SIE, we used breakpoint regression to model the relationship between lizard species richness and island area. We compared results from this model to results from the log‐linear regression model. Results Subsidized islands had a lower slope than unsubsidized islands, and the difference between these groups was significant when small islands were defined as < 1 km². In addition to comparing slopes, we tested for differences in the magnitude of the residuals (from the species–area regression of all islands) for subsidized vs. unsubsidized islands. We found no significant patterns in the residual values for small vs. large islands, or between islands with and without seabirds. The SIE was found to be a slightly better predictor of lizard species richness than the traditional log‐linear model. Main conclusions Predictions of the SIB hypothesis were partially supported by the data. The absence of a significant SIE may be a result of spatial subsidies as explained by the SIB hypothesis and data presented here. We conclude by suggesting potential scenarios to test for interactions between these two small island hypotheses. Future studies considering factors affecting species richness should examine the possible role of spatial subsidies, an SIE, or a synergistic effect of the two in data sets with small islands.     

Birds,Cattle, and Bracken Ferns: Bird Community Responses to a Neotropical Landscape Shaped by Cattle Grazing Activities

Large areas of tropical moist forests have been converted to cattle pastures, generating complex landscapes where different habitats are represented by small patches with an uneven spatial distribution. Here, we describe how bird communities respond to the different elements present in a livestock landscape that was originally dominated by tropical moist forest. We surveyed six habitats: open pastures, pastures with shrubs, early‐ and middle‐secondary forests, mature forest, and pastures invaded by bracken ferns (Pteridium aquilinum). Bird diversity was high in secondary and mature forests, and low in fern‐invaded sites and open pastures. Fern‐dominated sites had the lowest bird species richness, and trophic guild diversity of all habitats. Habitat structure affected both bird species richness and densities in similar ways. Tree species richness was the habitat attribute that had a bigger positive effect on bird species richness. Bird community structure varied among sampled habitats, separating habitats in two major groups (forests and pastures). Our data indicate that bracken fern‐invaded pastures were the worst habitat condition for avian communities. To increase bird diversity, we recommend to eliminate or manage bracken fern and to increase shrub and tree cover in open pastures to provide food resources and shelter for birds. Finally, we encourage the maintenance of secondary and mature forest remnants as a strategy to conserve resident birds within a landscape dominated by livestock activities.     

Heterogeneous Landscapes on Steep Slopes at Low Altitudes as Hotspots of Bird Diversity in a Hilly Region of Nepal in the Central Himalayas

Understanding factors determining the distribution of species is a key requirement for protecting diversity in a specific area. The aim of this study was to explore the factors affecting diversity and distribution of species of birds on different forested hills in central Nepal. The area is rich in species of birds. Because the area is characterized by steep gradients, we were also interested in the importance of altitude in determining the diversity and species composition of the bird communities. We assessed bird diversity and species composition based on point observations along a gradient of increasing altitude in two valleys (Kathmandu and Palung) in central Nepal. Data on environmental variables were also collected in order to identify the main determinants of bird diversity and species composition of the bird communities. We recorded 6522 individual birds belonging to 146 species, 77 genera and 23 families. Resident birds made up 80% (117 species) of the total dataset. The study supported the original expectation that altitude is a major determinant of species richness and composition of bird communities in the area. More diverse bird communities were found also in areas with steeper slopes. This together with the positive effect of greater heterogeneity suggests that forests on steep slopes intermixed with patches of open habitats on shallow soil at large spatial scales are more important for diverse bird communities than more disturbed habitats on shallow slopes. In addition, we demonstrated that while different habitat characteristics such as presence of forests edges and shrubs play an important role in driving species composition, but they do not affect species richness. This indicates that while habitat conditions are important determinants of the distribution of specific species, the number of niches is determined by large scale characteristics, such as landscape level habitat heterogeneity and altitude. Thus, to protect bird diversity in the mid-hills of central Nepal, we should maintain diverse local habitats (viz. forest, shrubs, open land, etc.) but also make sure the natural habitats on steeper slopes with large scale heterogeneity are maintained.