Diversity and similarity of butterfly communities in five different habitat types at Tam Dao National Park, Vietnam

Diversity and similarity of butterfly communities were assessed in five different habitat types (from natural closed forest to agricultural lands) in the mountains of Tam Dao National Park, Vietnam for 3 years from 2002 to 2004. The line transect count was used to record species richness and abundance of butterfly communities in the different habitat types. For each habitat, the number of species and individuals, and indices of species richness, evenness and diversity of butterfly communities were calculated. The results indicated that species richness and abundance of butterfly communities were low in the natural closed forest, higher in the disturbed forest, highest in the forest edge, lower in the shrub habitat and lowest in the agricultural lands. The indices of species richness, evenness and diversity of butterfly communities were low in agricultural lands and natural closed forest but highest in the forest edge and shrub habitats. The families Satyridae and Amathusiidae have the greatest species richness and abundance in the natural closed forest, with a reduction in their species richness and abundance from the natural closed forest to the agricultural lands. Species composition of butterfly communities was different among five different habitat types (40%), was similar in habitats outside the forest (68%) and was similar in habitats inside the forest (63%). Diversity and abundance of butterfly communities are not different between the natural closed forest and the agriculture lands, but species composition changed greatly between these habitat types. A positive correlation between the size of species geographical distribution range and increasing habitat disturbance was found. The most characteristic natural closed forest species have the smallest geographical distribution range.     

Consequences of organic farming and landscape heterogeneity for species richness and abundance of farmland birds

It has been suggested that organic farming may benefit farmland biodiversity more in landscapes that have lost a significant part of its former landscape heterogeneity. We tested this hypothesis by comparing bird species richness and abundance during the breeding season in organic and conventional farms, matched to eliminate all differences not directly linked to the farming practice, situated in either homogeneous plains with only a little semi-natural habitat or in heterogeneous farmland landscapes with abundant field borders and semi-natural grasslands. The effect of farm management on species richness interacted with landscape structure, such that there was a positive relationship between organic farming and diversity only in homogeneous landscapes. This pattern was mainly dependent on the species richness of passerine birds, in particular those that were invertebrate feeders. Species richness of non-passerines was positively related to organic farming independent of the landscape context. Bird abundance was positively related to landscape heterogeneity but not to farm management. This was mainly because the abundance of passerines, particularly invertebrate feeders, was positively related to landscape heterogeneity. We suggest that invertebrate feeders particularly benefit from organic farming because of improved foraging conditions through increased invertebrate abundances in otherwise depauperate homogeneous landscapes. Although many seed-eaters also benefit from increased insect abundance, they may also utilize crop seed resources in homogeneous landscapes and conventional farms. The occurrence of an interactive effect of organic farming and landscape heterogeneity on bird diversity will have consequences for the optimal allocation of resources to restore the diversity of farmland birds.     

寒温带牧林交错区生境复杂度对啮齿类物种多样性的影响

采用铗日法对嫩江流域牧林交错区5个生境梯度中啮齿动物的多样性水平进行了研究,以探讨栖息地复杂程度对地面啮齿类物种多样性的影响.结果表明,黑线姬鼠、黑线仓鼠和大林姬鼠是嫩江流域牧林交错区中的优势种和常见种,不同物种在不同生境类型中呈明显的不均匀分布,反映了它们对不同生境的选择倾向性.同时,地面草本植物的覆盖度和丰富度对地面活动的小型哺乳动物的物种组成和群落结构具有较大的影响,啮齿动物的捕获率和生物量与草本植物的多样性水平呈现出显著的相关性关系,其物种丰富度随草本植物多样性指数的增加呈递增的趋势.此外,各生境中啮齿动物的物种多样性指数与草本植物多样性的相关关系并不显著,可能是因为灌丛区和林-灌交错区内大量的低矮灌丛与萌生丛改善了生境的隐蔽条件与食物资源,为更多物种的共存创造了良好的微环境,使得两生境中啮齿动物的物种多样性水平尤为突出.研究表明,小型啮齿动物的物种组成与栖息地的复杂程度有关,但地面植被对小型兽类生物量和物种多样性的影响力度并不是等同的,对地面小型兽类物种多样性影响的研究,统计生境复杂度或异质性时应考虑不同植被型/生境类型距地面的高度,不同植被型对小型兽类物种多样性的贡献程度随其距地面高度的增加而降低.     

塔里木荒漠河岸林异质生境物种多样性比较与其测度指标筛选及评价

选择合适的物种多样性测度指标与多样性指数是进行群落多样性研究的基础工作。依据塔里木河上游荒漠河岸林样地调查资料,分别采用重要值、盖度和多度为测度指标比较了反映群落物种丰富度、多样性、均匀度和优势度12种多样性指数与异质生境群落多样性特征,并对多样性指数进行了相关分析与评价。结果表明,荒漠河岸林异质生境群落物种组成种类差异明显,轮南镇胡杨群落物种丰富度与多样性指数最高,水工三连灰胡杨群落多样性最低,土壤水盐的空间异质性是引起荒漠植被空间分布与群落多样性差异的主导因子。表征荒漠群落多样性以重要值和盖度为测度指标优于多度指标,其中以重要值为测度指标来反映群落多样性更为合理。相关与主成分分析表明,均匀度与多样性指数间的相关性高于丰富度与多样性指数,且多样性指数受均匀度、优势度指数受丰富度影响较大,反映出荒漠河岸林群落多样性主要决定于物种分布的均匀程度。12种多样性指数中Margalef丰富度指数（Ma）、Shannon-Weiner多样性指数（H）与Simpson多样性指数（D）能客观真实地反映异质生境荒漠植物群落多样性。同时,针对高度生境异质性的荒漠植物群落,还应综合考虑群落物种组成与生境特征,选择合适的多样性指数组合可更客观地反映荒漠河岸林群落多样性变化。     

Forest Fragment and Breeding Habitat Characteristics Explain Frog Diversity and Abundance in Singapore

Habitat loss and fragmentation can have severe negative and irreversible effects on biodiversity. We investigated the effects of forest fragmentation on frog diversity in Singapore because of its high rates of deforestation and the demonstration that frogs are some of the most sensitive species to habitat degradation. We surveyed frog species in 12 forest fragments varying from 11 to 935 ha. We compared differences in species richness, abundance, and Shannon's index in relation to forest fragment size, connectivity (distance between fragments), and breeding habitat heterogeneity. A total of 20 species from 12 genera and five families were encountered in 12 fragments. Larger fragments and those closer to larger fragments had higher species richness. Abundance, however, was not correlated with forest area or connectivity, but we found fewer individual frogs in the larger fragments. We also found that breeding habitat heterogeneity best explained frog species diversity and abundance in forest fragments. Fragments with a high diversity of breeding habitats had more species. We found no evidence to suggest that abundance and diversity are strongly correlated, particularly in disturbed areas, but that breeding habitat heterogeneity is an under-appreciated factor that should be considered when prioritizing areas for anuran conservation. Enriching breeding habitat heterogeneity, creating corridors between fragments, and reforesting degraded areas are some of the most beneficial strategies for preserving urban frog biodiversity.     

Fine-scale habitat structure complexity determines insectivorous bird diversity in a tropical forest

Habitat complexity in reforested stands has been acknowledged as a key factor that influences habitat use by birds, being especially critical for habitat disturbance-sensitive species such as tropical understory insectivorous birds. Most studies regarding the relationship between forest structure and species diversity were conducted at the landscape scale, but different diversity patterns may emerge at a finer scale (i.e., within a habitat patch). We examined a tropical reforested area (State of Caldas, Colombia), hypothesizing that insectivorous bird richness, abundance, and foraging guild abundance would increase as intra-habitat complexity increases. We established 40 monitoring plots within a reforested area, measured their structural features, and determined their relationships with species richness, total abundance, and foraging guild abundance, using Generalized Additive Models. We found that the increasing variation in basal area, stem diameter, and number of stems was positively correlated with species richness, total abundance, and foraging guild abundance. Relationships between richness or abundance and structural features were not lineal, but showing curvilinear responses and thresholds. Our results show that heterogeneity on basal area, stem diameter, and the number of stems was more correlated to insectivorous bird richness and abundance than the average of those structural features. Promoting structural variation on reforested areas by planting species with different growth rates may contribute to increase the richness and abundance of a tropical vulnerable group of species such as the understory insectivorous birds.     

Effects of polyculture and monoculture farming in oil palm smallholdings on tropical fruit‐feeding butterfly diversity

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Responses of tropical fruit bats to monoculture and polyculture farming in oil palm smallholdings

The oil palm industry is one of the main economic drivers in Southeast Asia. The industry has caused tropical deforestation on a massive scale in producing countries, and this forest conversion to oil palm agriculture has decimated the habitat of numerous native species. Monoculture and polyculture practices are two distinctive oil palm production systems. We hypothesize that polyculture farming hosts a greater diversity of species than monoculture farming. Habitat complexity in smallholdings is influenced by multiple farming practices (i.e. polyculture and monoculture). However, little is known about the effects of such farming practices in smallholdings on mammalian biodiversity, and particularly frugivorous bats. Our study aimed to find the best farming practice to reconcile oil palm production with biodiversity conservation. Mist-nets were used to trap frugivorous bats at 120 smallholdings in Peninsular Malaysia. We compared species richness and the abundance of frugivorous bats between monoculture and polyculture smallholdings. We investigated their relationships with vegetation structure characteristics. Our results revealed that species richness and abundance of frugivorous bats were significantly greater in polyculture smallholdings than monoculture smallholdings. We also found that 28.21% of the variation in species richness was explained by in situ habitat characteristics, including the number of dead standing oil palms and immature oil palms, non-grass cover, height of non-grass cover, and farming practices. The in situ habitat quality was closely associated with oil palm farming management. Commercial growers should implement polyculture rather than monoculture farming because polyculture farming has positive effects on the abundance and species richness of bats in oil palm production landscapes.     

里海西北部Caspian地区草原扩大与啮齿类群落结构的变化

于1997-1999年检测了俄联邦卡尔梅克(Kalmyki)共和国ChernieZemly半干旱草原由10个物种组成的啮齿类群落。该区域的植被特点是家畜大量减少后形成的恢复植被。我们的任务之一就是了解不同啮齿动物的生境需求,以预测不同物种对环境变化的特异性反应。我们也检测了群落空间结构、生态位参数和物种多样性。逐步回归分析表明,各种啮齿动物的空间分布仅部分地决定于已知环境因子(2-6个变量)。虽然方程式具有强显著性,但决定系数R2很低,不超过18%。对于大多数种类,这种结果可由最近草原扩大而来的生境异质性降低解释。物种生境选择条件和资源的明显低水平可能是由物种对环境变化的惯性反应造成的。判别函数分析和主分量分析结果表明,群落结构以及单个物种的生态位参数在不同年间并不稳定。空间分层结构在种间中等和低重叠的高物种多样性年度表现明显。多样性、生态位组成和生态位宽度分析结果表明,不同物种对环境条件和资源变化的反应具有不同的个性化方式。同时,物种对环境动态的明显个性化反应成为物种多样性定向变化的原因。限定空间和相对单一条件下的结果表明,啮齿类α多样性与生境结构复杂性呈正相关,而与生境生产力特征呈负相关。观察表明,卡尔梅克国草原扩大伴随着初级生产力的升高和生境异质性的降低。随着时间的推移会导致大多数草原和半干旱区域啮齿类多样性的降低,并形成动物区系核心.     

The effects of land-use change on arthropod richness and abundance on Santa Maria Island (Azores): unmanaged plantations favour endemic beetles

We study how endemic, native and introduced arthropod species richness, abundance, diversity and community composition vary between four different habitat types (native forest, exotic forest of Cryptomeria japonica, semi-natural pasture and intensive pasture) and how arthropod richness and abundance change with increasing distance from the native forest in adjacent habitat types in Santa Maria Island, the Azores. Arthropods were sampled in four 150 m long transects in each habitat type. Arthropods were identified to species level and classified as Azorean endemic, single-island endemic (SIE), native, or introduced. The native forest had the highest values for species richness of Azorean endemics, SIEs and natives; and also had highest values of Azorean endemic diversity (Fisher’s alpha). In contrast, the intensive pasture had the lowest values for endemic and native species richness and diversity, but the highest values of total arthropod abundance and introduced species richness and diversity. Arthropod community composition was significantly different between the four habitat types. In the semi-natural pasture, the number of SIE species decreased with increasing distance from the native forest, and in the exotic forest the abundance of both Azorean endemics and SIEs decreased with increasing distance from the native forest. There is a gradient of decreasing arthropod richness and abundance from the native forest to the intensive pasture. Although this study demonstrates the important role of the native forest in arthropod conservation in the Azores, it also shows that unmanaged exotic forests have provided alternative habitat suitable for some native species of forest specialist arthropods, particularly saproxylic beetles.     

荒漠啮齿动物群落对开垦干扰的响应及其种群生态对策

开垦对功能相对脆弱的荒漠生态系统是重要的干扰,这种干扰往往导致栖息地破碎化,并对动植物群落产生强烈影响。作为荒漠生态系统的重要成分,啮齿动物群落受到开垦干扰后对环境的响应及其群落中种群的生态对策,是荒漠生态系统生物多样性及其功能维持稳定的重要基础。2006～2011年,采用标志重捕法对内蒙古阿拉善荒漠区未开垦和开垦草地啮齿动物群落格局及其不同生态对策种群的数量组成进行了专门研究。目的（1）明确开垦干扰下啮齿动物群落格局时间尺度变化;（2）验证假设：人为干扰区（开垦区）啮齿动物群落中r?对策者占据优势,未干扰区（未开垦区）以K?对策者为主;（3）依据啮齿动物群落中不同生态对策种群的数量组成来判断群落所受干扰的程度。结果表明,开垦干扰降低了啮齿动物群落多样性,改变了群落中不同生态对策种群的数量组成,种群以r?对策者为优势;未开垦区啮齿动物群落中以K?对策种群为主。开垦区啮齿动物群落受到严重干扰。     

Relative Importance of Coral Cover,Habitat Complexity and Diversity in Determining the Structure of Reef Fish Communities

The structure of coral reef habitat has a pronounced influence on the diversity, composition and abundance of reef-associated fishes. However, the particular features of the habitat that are most critical are not always known. Coral habitats can vary in many characteristics, notably live coral cover, topographic complexity and coral diversity, but the relative effects of these habitat characteristics are often not distinguished. Here, we investigate the strength of the relationships between these habitat features and local fish diversity, abundance and community structure in the lagoon of Lizard Island, Great Barrier Reef. In a spatial comparison using sixty-six 2m² quadrats, fish species richness, total abundance and community structure were examined in relation to a wide range of habitat variables, including topographic complexity, habitat diversity, coral diversity, coral species richness, hard coral cover, branching coral cover and the cover of corymbose corals. Fish species richness and total abundance were strongly associated with coral species richness and cover, but only weakly associated with topographic complexity. Regression tree analysis showed that coral species richness accounted for most of the variation in fish species richness (63.6%), while hard coral cover explained more variation in total fish abundance (17.4%), than any other variable. In contrast, topographic complexity accounted for little spatial variation in reef fish assemblages. In degrading coral reef environments, the potential effects of loss of coral cover and topographic complexity are often emphasized, but these findings suggest that reduced coral biodiversity may ultimately have an equal, or greater, impact on reef-associated fish communities.     

Scale-specific correlations between habitat heterogeneity and soil fauna diversity along a landscape structure gradient

Habitat heterogeneity contributes to the maintenance of diversity, but the extent that landscape-scale rather than local-scale heterogeneity influences the diversity of soil invertebrates—species with small range sizes—is less clear. Using a Scottish habitat heterogeneity gradient we correlated Collembola and lumbricid worm species richness and abundance with different elements (forest cover, habitat richness and patchiness) and qualities (plant species richness, soil variables) of habitat heterogeneity, at landscape (1 km²) and local (up to 200 m²) scales. Soil fauna assemblages showed considerable turnover in species composition along this habitat heterogeneity gradient. Soil fauna species richness and turnover was greatest in landscapes that were a mosaic of habitats. Soil fauna diversity was hump-shaped along a gradient of forest cover, peaking where there was a mixture of forest and open habitats in the landscape. Landscape-scale habitat richness was positively correlated with lumbricid diversity, while Collembola and lumbricid abundances were negatively and positively related to landscape spatial patchiness. Furthermore, soil fauna diversity was positively correlated with plant diversity, which in turn peaked in the sites that were a mosaic of forest and open habitat patches. There was less evidence that local-scale habitat variables (habitat richness, tree cover, plant species richness, litter cover, soil pH, depth of organic horizon) affected soil fauna diversity: Collembola diversity was independent of all these measures, while lumbricid diversity positively and negatively correlated with vascular plant species richness and tree canopy density. Landscape-scale habitat heterogeneity affects soil diversity regardless of taxon, while the influence of habitat heterogeneity at local scales is dependent on taxon identity, and hence ecological traits, e.g. body size. Landscape-scale habitat heterogeneity by providing different niches and refuges, together with passive dispersal and population patch dynamics, positively contributes to soil faunal diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

区域变动尺度上不同生境类型及环境因子对西双版纳蝴蝶多样性的影响

【目的】生境类型和环境因子对物种分布和维持具有重要的影响。本研究通过分析不同生境类型对蝴蝶群落多样性及其群落结构影响的差异,以及环境因子对蝴蝶物种丰富度和多度的影响,为区域变动尺度蝴蝶多样性维持机制的研究奠定基础。【方法】于2019年8月和10月,在西双版纳地区采用样线法,调查了天然林、次生林、复合生境、人工林和农田5种生境中蝴蝶的物种,分析了蝴蝶群落多样性、群落结构相似性及物种丰富度和多度与环境因子的关系。【结果】2019年从西双版纳共采集蝴蝶2 226头,隶属于11科98属175种,在西双版纳州级尺度上蝴蝶物种丰富度高于县域尺度。在西双版纳州级尺度上,蝴蝶的物种丰富度和多度在5种生境间存在显著差异,而在县域尺度上,物种丰富度、多度和Chao 1物种丰富度估计值没有一致性规律。群落结构相似性结果显示,在西双版纳州级尺度上,蝴蝶群落结构在不同生境类型间存在极显著差异,在县域尺度上,仅勐腊区域蝴蝶群落结构在不同生境类型间存在显著差异。蝴蝶物种丰富度和多度不仅受到生境类型的影响,还受到温度、年均降水和海拔的影响。【结论】本研究结果表明,在区域变动尺度上,生境类型对西双版纳蝴蝶的多样性的影响较大,而温度、年均降水和海拔是维持蝴蝶物种多样性的重要因素。这些发现对当前人类导致的生境丧失和气候变化时代生物多样性的保护具有重要意义。     

Habitat structure components are effective predictors of trap-nesting Hymenoptera diversity

Habitat heterogeneity can be the major factor affecting species diversity in a community and measuring bee and wasp community habitat preferences in natural systems may provide insights for biodiversity management and conservation. In the present study, we investigate the effects of habitat structure components on solitary bee and wasp species richness and abundance. The research was conducted in an urban forest remnant in southeast Brazil. Our main questions were: (1) is similarity in habitat structure mirrored by similarity in Aculeate assemblage composition? and (2) what are the vegetation features that could be used as predictors of solitary bee and wasp richness and abundance? Aculeate bees and wasps were sampled using trap nests from February to November 2004. Trap nests were placed in sampling units located in 6 ha of secondary mesophytic forest. One hundred and thirty-seven trap nests were occupied by four species of wasps and seven species of bees. Altogether, our sampling units had a mean capture rate (relative to expected richness) of 72% during all the study period. The more similar sampling units were in terms of vegetation structure, the more similar they were in solitary bee and wasp species composition. The variance of tree abundance, shrub height and the abundance of wood logs were good predictors of solitary bee and wasp species richness and abundance in the study area. We demonstrate that even in a small scale it is possible to detect significant influences of habitat features on alpha diversity and that some of them are effective as predictors of trap-nesting Hymenoptera richness and abundance.     

The stability–diversity relationship in stream macroinvertebrates: influences of sampling effects and habitat complexity

1. Theory predicts that the stability of a community should increase with diversity. However, despite increasing interest in the topic, most studies have focused on aggregate community properties (e.g. biomass, productivity) in small‐scale experiments, while studies using observational field data on realistic scales to examine the relationship between diversity and compositional stability are surprisingly rare. 2. We examined the diversity–stability relationship of stream invertebrate communities based on a 4‐year data set from boreal headwater streams, using among‐year similarity in community composition (Bray–Curtis coefficient) as our measure of compositional stability. We related stability to species richness and key environmental factors that may affect the diversity–stability relationship (stream size, habitat complexity, productivity and flow variability) using simple and partial regressions. 3. In simple regressions, compositional stability was positively related to species richness, stream size, productivity and habitat complexity, but only species richness and habitat complexity were significantly related to stability in partial regressions. There was, however, a strong relationship between species richness and abundance. When abundance was controlled for through re‐sampling, stability was unrelated to species richness, indicating that sampling effects were the predominant mechanism producing the positive stability–diversity relationship. By contrast, the relationship between stability and habitat complexity (macrophyte cover) became even stronger when the influence of community abundance was controlled for. Habitat complexity is thus a key factor enhancing community stability in headwater streams.     

Local and landscape drivers of biodiversity of four groups of ants in coffee landscapes

Agriculture of varying management intensity dominates fragmented tropical areas and differentially impacts organisms across and within taxa. We examined impacts of local and landscape characteristics on four groups of ants in an agricultural landscape in Chiapas, Mexico comprised of forest fragments and coffee agroecosystems varying in habitat quality. We sampled ground ants found in leaf litter and rotten logs and arboreal ants found in hollow coffee twigs and on tree trunks. Then using vegetation and agrochemical indices and conditional inference trees, we examined the relative importance of local (e.g. vegetation, elevation, agrochemical) and landscape variables (e.g. distance to and amount of nearby forest and rustic coffee) for predicting richness and abundance of ants. Leaf litter ant abundance increased with vegetation complexity; richness and abundance of ants from rotten logs, twig-nests, and tree trunks were not affected by vegetation complexity. Agrochemical use did not affect species richness or abundance of any ant group. Several local factors (including humus mass, degree of decay of logs, number of hollow twigs, tree circumference, and absence of fertilizers) were significant positive predictors of abundance and richness of some ant groups. Two landscape factors (forest within 200 m, and distance from forest) predicted richness and abundance of twig-nesting and leaf litter ants. Thus, different ant groups were influenced by different characteristics of agricultural landscapes, but all responded primarily to local characteristics. Given that ants provide ecosystem services (e.g. pest control) in coffee farms, understanding ant responses to local and landscape characteristics will likely inform farm management decisions.     

Forest complexity drives dung beetle assemblages along an edge-interior gradient in the southwest Amazon rainforest

1. The habitat heterogeneity hypothesis predicts that heterogeneous habitats may provide more niches and diverse ways of exploiting environmental resources, thereby allowing more species to coexist, persist and diversify. 2. We aimed to investigate how an edge-interior gradient related to forest complexity influences species composition, abundance and richness of dung beetles in the western Amazon rainforest. We expected dung beetle abundance and richness to increase along the forest edge-interior gradient, in accordance with the habitat heterogeneity hypothesis. We also expected strong changes in species composition driven by species turnover in the forest interior and nestedness along the forest edges. We sampled dung beetles using baited pitfall traps across an edge-interior gradient. We also assessed the variation in forest features along the edge-interior gradient to identify changes in forest complexity. 3. Both species richness and abundance of dung beetles increased along the forest edge-interior, following the gradient of forest complexity. The Sorensen dissimilarity of dung beetle assemblages was higher among sampling units placed near the forest edge, although neither turnover, nor nestedness was different between the extremes of the forest edge-interior gradient. There was a clear compositional change along the edge-interior gradient mostly driven by species turnover. Individual indicator value analysis revealed that species were strongly associated with the forest interior conditions. 4. The simplification of the Amazon rainforest near clearings causes compositional changes in dung beetle assemblages. These changes are characterised by species-poor and low-abundance assemblages and may impair dung beetle ecological functions and therefore forest recovery.     

Vegetation patterns at the alpine treeline ecotone: the influence of tree cover on abrupt change in species composition of alpine communities

Aims: The upper elevation limit of forest vegetation in mountain ranges (the alpine treeline ecotone) is expected to be highly sensitive to global change. Treeline shifts and/or ecotone afforestation could cause fragmentation and loss of alpine habitat, and are expected to trigger considerable alterations in alpine vegetation. We performed an analysis of vegetation structure at the treeline ecotone to evaluate whether distribution of the tree population determines the spatial pattern of vegetation (species composition and diversity) across the transition from subalpine forest to alpine vegetation. Location: Iberian eastern range of the Pyrenees. Methods: We studied 12 alpine Pinus uncinata treeline ecotones. Rectangular plots ranging from 940 to 1900 m² were placed along the forest‐alpine vegetation transition, from closed forest to the treeless alpine area. To determine community structure and species distribution in the treeline ecotone, species variation along the forest‐alpine vegetation transition was sampled using relevés of 0.5 m² set every 2 m along the length of each plot. Fuzzy C‐means clustering was performed to assess the transitional status of the relevés in terms of species composition. The relation of P. uncinata canopy cover to spatial pattern of vegetation was evaluated using continuous wavelet transform analysis. Results: Vegetation analyses revealed a large degree of uniformity of the subalpine forest between all treeline ecotone areas studied. In contrast, the vegetation mosaic found upslope displayed great variation between sites and was characterized by abrupt changes in plant community across the treeline ecotone. Plant richness and diversity significantly increased across the ecotone, but tree cover and diversity boundaries were not spatially coincident. Conclusions: Our results revealed that no intermediate communities, in terms of species composition, are present in the treeline ecotone. Ecotone vegetation reflected both bedrock type and fine‐scale heterogeneity at ground level, thereby reinforcing the importance of microenvironmental conditions for alpine community composition. Tree cover did not appear to be the principal driver of alpine community changes across the treeline ecotone. Microenvironmental heterogeneity, together with effects of past climatic and land‐use changes on ecotone vegetation, may weaken the expected correlation between species distribution and vegetation structure.     

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.