Response of bird assemblages to windstorm and salvage logging — Insights from analyses of functional guild and indicator species

Natural disturbances, such as fire, windstorms and insect outbreaks, are important drivers of biodiversity in forest ecosystems, but at the same time cause large economic losses. Among the natural disturbances in Europe, windstorms cause the highest economic loss. After such storms, damaged forest stands are commonly salvage logged to restore economic value. However, such interventions could affect species assemblages of various taxonomic groups, including breeding birds. Despite these potential effects, investigations of the impacts of post-storm logging are largely lacking. We thus investigated assemblages of breeding birds in 21 logged and 21 unlogged windstorm-disturbed forest plots and 18 undisturbed, control forest plots using fixed-radius point-stop counts three, five and seven years after a windstorm within the Bavarian Forest National Park as part of the European Long-Term Ecosystem Research Network. We recorded 2100 bird individuals of 55 bird species. Bird assemblages were predominantly altered by the consequences of the windstorm and affected only to a minor degree by subsequent logging of storm-felled trees. Nevertheless, bird species richness was significantly reduced by post-storm logging within the first season. In general, the windstorm led to a shift in bird assemblage composition from typical forest species towards open- and shrub-land species. Assemblages of logged and unlogged disturbed plots consisted mainly of long-distance migrants and ground-foraging bird species, whereas assemblages of undisturbed control plots consisted of resident species that forage within vegetation. Both unlogged and logged storm-felled coniferous forest stands were inhabited by endangered or declining bird species, such as Water Pipit (Anthus spinoletta) on logged plots and Eurasian Redstart (Phoenicurus phoenicurus) on unlogged plots. Indicator species analyses suggested that species of unlogged storm disturbed plots depended on storm-created legacies, such as pits and snags, for foraging and nesting. Hence, we recommend reducing post-storm logging of these legacies to support species restricted to unlogged disturbed forest. To increase the diversity of breeding birds on the landscape scale, diverse logged and unlogged post-disturbance stands should be provided, which could be gained by a partial benign neglect strategy of storm disturbed forest stands.     

High Bird Species Diversity in Structurally Heterogeneous Farmland in Western Kenya

Tropical ecosystems are globally important for bird diversity. In many tropical regions, land‐use intensification has caused conversion of natural forests into human‐modified habitats, such as secondary forests and heterogeneous agricultural landscapes. Despite previous research, the distribution of bird communities in these forest‐farmland mosaics is not well understood. To achieve a comprehensive understanding of bird diversity and community turnover in a human‐modified Kenyan landscape, we recorded bird communities at 20 sites covering the complete habitat gradient from forest (near natural forest, secondary forest) to farmland (subsistence farmland, sugarcane plantation) using point counts and distance sampling. Bird density and species richness were on average higher in farmland than in forest habitats. Within forest and farmland, bird density and species richness increased with vegetation structural diversity, i.e., were higher in near natural than in secondary forest and in subsistence farmland than in sugarcane plantations. Bird communities in forest and farmland habitats were very distinct and very few forest specialists occurred in farmland habitats. Moreover, insectivorous bird species declined in farmland habitats whereas carnivores and herbivores increased. Our study confirms that tropical farmlands can hardly accommodate forest specialist species. Contrary to most previous studies, our findings show that structurally rich tropical farmlands hold a surprisingly rich and distinct bird community that is threatened by conversion of subsistence farmland into sugarcane plantations. We conclude that conservation strategies in the tropics must go beyond rain forest protection and should integrate structurally heterogeneous agroecosystems into conservation plans that aim at maintaining the diverse bird communities of tropical forest‐farmland mosaics.     

Effects of disturbances on scuttle flies (Diptera: Phoridae) in Pine Forests

I investigated the ecological consequences of disturbances (anthropogenic and natural) on the scuttle fly communities in four large Pine Forests in Poland. I used data on 17,547 male individuals representing 183 species. Communities found in pine plantations (established in clear-cut areas) and in differently treated post-windstorm (with windthrow logs being left or removed) were less diverse than those found in old-growth forest. The communities recorded in the same habitat types in different forest complexes (ca. 300 km apart) were found to display greater similarity than those recorded on adjacent plots in a given forest (ca. 1 km apart), but covering different habitats. The species-specific preference for habitats after disturbances (clear-cuts and post-windstorm areas) was highly correlated between the forests. The abundance of the species with saprophagous larvae was distinctly higher in the disturbed areas than in the old-growth stands. Also, the body length of the scuttle flies was significantly related to their preference for disturbed or undisturbed habitats: smaller species preferred clear-cuts and post-windstorm areas, whereas larger species were related to intact stands.     

人类休闲活动对森林鸟类群落无有害影响

我们对西班牙马德里森林深处(未受到人类干扰)、森林小路沿线(间或有徒步旅行者经过)以及森林野餐区(有较持久干扰)的鸟类进行了调查,以查明人类休闲活动对该地区鸟类群落的分布和多度的影响.结合样方法对以上3类区域鸟类群落的总体特征(如:鸟类的多度、物种的丰富度和多样性)、不同鸟类群体的特征(如:鸦科鸟类的多度,营巢或者取食集团的特征)、各鸟种的多度等参数进行了比较.结果表明:小路沿线和野餐区样方内鸟类的多度和物种丰富度比森林深处更高,已查明14种鸟类的多度在人为干扰区域比森林深处高,只有5种鸟类的多度在森林深处更高一些;在树冠层筑巢的鸟类、在树干和地面觅食的鸟类及鸦科鸟类在休闲活动区有更高的多度;地巢性鸟类在休闲区的多度较低,这种差别尤以阔叶落叶林中加明显.而森林小路的鸟类群落结构与森林内部非常相似,说明人类休闲活动对这些林区鸟类群落的影响更多缘自野餐.原因是野餐区常常具有以下特点:具有高大的乔木、中层植被稀疏、有较多的洞和缝隙以及野餐后会遗留下一些垃圾.以上结论与研究的预期目标正好相反,人类的这些休闲活动可能会降低地巢性鸟类的适宜度,但是对大多数鸟的类栖息地没有不利影响.     

Effects of forest modification on bird community composition and seed removal in a heterogeneous landscape in South Africa

Human disturbance threatens and modifies forest ecosystems worldwide. Previous studies have investigated the effects of human impact on local bird communities in disturbed forests, but we still lack information on how bird species richness and ecological processes respond to different forest modifications present at a landscape scale. In a heterogeneous South African landscape, we chose six types of indigenous scarp forest, differing in the intensity of human disturbance: continuous natural forests and natural forest fragments in nature reserves, forest fragments in eucalyptus plantations, fragments in the agricultural matrix, forest gardens and secondary forests in game reserves. In 36 study sites, we investigated the bird community using point counts and observed the seed removal of birds at the native tree species Celtis africana. Species richness did not differ among the forest types, but abundance varied significantly with most birds observed in fragments in the agricultural matrix, forest gardens, and secondary forests. The higher bird abundance in these forests was mainly due to forest generalists, shrubland and open country species whereas forest specialists were rarely present. Changes in species composition were also confirmed by multivariate analysis which clearly separated bird communities by forest type. Frugivore abundance in C. africana was highest in natural forest fragments, fragments in the agricultural matrix, forest gardens and secondary forests. The same trend was found for the estimated total number of fruits removed per C. africana tree, though the differences among forest types were not significant. Consequently, modified forests seem to maintain important ecological functions as they provide food sources for generalist species which may, due to their mobility, enhance natural plant regeneration. However, we could show that protected forest habitats are important refugees for specialist species sensitive to human disturbance.     

四川老君山自然保护区不同生境鸟类多样性研究

2006年4~5月采用"点样带法"(point transects)对四川老君山自然保护区的鸟类物种组成和种群数量进行了调查,共记录到85种鸟类,分属7目22科。其中,国家级保护鸟类5种,我国特有种13种,优势种4种。把调查区域划分为3种生境:原始林、次生林和人工林,并对各生境鸟类群落的鸟类数量级、鸟类群落的物种多样性、鸟类群落间的相似性进行了比较分析,其结果表明:1)国家级保护鸟类在原始林中最多,次生林中最少;我国特有种则是次生林中最多,人工林中最少。2)鸟类群落Shannon-Wiener物种多样性指数原始林最高,人工林最低。3)原始林和次生林之间的B ray-Curtis相似性指数最高,原始林和人工林之间相似性指数最低。     

Natural Windbreaks Sustain Bird Diversity in a Tea-Dominated Landscape

Windbreaks often form networks of forest habitats that improve connectivity and thus conserve biodiversity, but little is known of such effects in the tropics. We determined bird species richness and community composition in windbreaks composed of remnant native vegetation amongst tea plantations (natural windbreaks), and compared it with the surrounding primary forests. Fifty-one, ten-minute point counts were conducted in each habitat type over three days. Despite the limited sampling period, our bird inventories in both natural windbreaks and primary forests were nearly complete, as indicated by bootstrap true richness estimator. Bird species richness and abundance between primary forests and windbreaks were similar, however a difference in bird community composition was observed. Abundances of important functional groups such as frugivores and insectivores did not vary between habitat types but nectarivores were more abundant in windbreaks, potentially as a result of the use of windbreaks as traveling routes, foraging and nesting sites. This preliminary study suggests that natural windbreaks may be important habitats for the persistence of bird species in a production landscape. However, a better understanding of the required physical and compositional characteristics for windbreaks to sustain bird communities is needed for effective conservation management.     

Moderate Human Disturbance of Rain Forest Alters Composition of Fruiting Plant and Bird Communities

Forests worldwide are experiencing rapid environmental change due to human activity. We aimed to increase understanding of anthropogenic impacts on community composition and species interactions. In a natural experiment, we asked whether subsistence human land use has altered the community composition of a Neotropical rain forest on the island of Tobago, in the West Indies. We surveyed fruiting plants and birds in three adjacent habitat types that varied in level of disturbance, and used multivariate analyses to determine whether changes in the plant community were associated with differences in avifauna composition. The three forest habitats had similar plant and bird diversities, yet markedly different species compositions and abundances. Primary forest had the most diverse plant community, while disturbed habitats had a more homogeneous plant composition. Primary and disturbed forest had distinct community compositions, with canopy cover and the relative abundance of plant types explaining 83 percent of the variation in bird species assemblages. Seemingly moderate human disturbance has led to substantial changes in the plant and bird assemblages of Tobago's rain forest, outside of a protected reserve. Our study highlights the direct links between human disturbance and the structure of rain forests, underscoring the impact of even moderate activity on community composition.     

Tree recovery and seed dispersal by birds: Comparing forest, agroforestry and abandoned agroforestry in coastal Ecuador

We used a highly replicated study to examine vegetation characteristics between patches of intervened forest, abandoned agroforestry systems with coffee and actively managed agroforestry systems with coffee in a tropical landscape. In all habitats, plant structural characteristics, individual abundance, species richness and composition were recorded for the three plant size classes: adult trees, saplings and seedlings. Furthermore, bird species richness and composition, and seeds dispersed by birds were recorded. Tree abundance was higher in forest habitats while saplings and seedlings were more abundant in abandoned coffee sites. Although species richness of adult trees was similar in the three habitats, species richness of saplings and seedlings was much higher in forest and abandoned coffee than in managed coffee sites. However, in spite of their relatively low species richness, managed coffee sites are an important refuge for tree species common to the almost disappeared mature forest in the area. Floristic similarity for adult trees was relatively low between land use types, but clearly higher for seedlings, indicating homogenizing processes at the landscape level. More than half of the saplings and seedling were not represented by adults in the canopy layer, suggesting the importance of seed dispersal by birds between habitats. Our results show that each of the studied ecosystems plays a unique and complementary role as seed source and as habitat for tree recovery and tree diversity.     

Are post-fire residual forest patches refugia for boreal bryophyte species? Implications for ecosystem based management and conservation

Residual patches of forest remaining after natural or anthropogenic disturbance may facilitate regeneration of fragmented forest. However, residual patch function remains unclear, especially after natural wildfire. We investigate the role of residual boreal forest patches as refugia for bryophytes and ask the question, do they house bryophyte communities similar to those encountered in undisturbed forests? Bryophytes were sampled in three habitat types in black spruce boreal forests illustrating a gradient of disturbance severity: undisturbed forests, residual patches and burned matrices. Temporal, disturbance severity, spatial and structural variables of habitats were also recorded. Bryophyte community composition differed among habitat types with residual patches characterized by higher species richness, the loss of forest specialists and the addition of disturbance-prone species. The bryophyte community found in residual patches is at the interface between the communities of undisturbed forests and burned matrices. As residual patches did not conserve all species, particularly forest specialists, they were not refugia. However, we identify temporal, spatial and structural characteristics that can maintain bryophyte communities most similar to undisturbed forests and enhance residual patch “refugia potential”. Residual patches enhance bryophyte diversity of the landscape housing species that cannot survive in the burned matrix. As conclusion we discuss the use of retention patches in harvested stands, together with the preservation of undisturbed stands that house singular bryophyte communities and especially sensitive forest specialists.     

Dung beetle assemblages in primary forest and disturbed habitats in a tropical dry forest landscape in western Mexico

Dung beetles (Scarabaeinae) are conspicuous components in most terrestrial ecosystems, performing important ecological functions and services. Being sensitive to several types of disturbance, they have been successfully used as indicators of habitat change. Dung beetle communities in tropical rainforests have been well studied, but considerably less information is available for tropical dry forests. In this study I sampled dung beetles in two undisturbed habitats, deciduous forest and semideciduous forest, and two disturbed habitats, secondary forest and open area habitat, in the Chamela-Cuixmala region of western Mexico. Dung beetle species with high indicator value for each habitat were identified. Beetle abundance, observed species richness and estimated species richness were similar in the three forest habitats, but significantly lower in the open area habitat. A more detailed analysis of species-specific abundances in the three forest habitats revealed some differences. Transects of one of the undisturbed habitats, the deciduous forest, were more similar to the non-adjacent transects of disturbed secondary forest, than to the adjacent undisturbed semideciduous forest transects. Unlike studies in other tropical sites that have found a decrease in equitability in Scarabaeinae assemblages between undisturbed forest and disturbed habitat (particularly open habitats), in the Chamela-Cuixmala region all four habitats showed similar low equitability in community structure, with two or three very dominant species.     

Animal-habitat relationships in the Knysna Forest,South Africa: discrimination between forest types by birds and invertebrates

Summary Effects of forest plant species composition and physiognomy on bird and invertebrate communities were investigated in three discrete, relatively undisturbed forest types along a dry-wet soil moisture gradient. Using discriminant function analysis, a 100% floristic and a 78% vegetation structural discrimination were obtained between the three forest types. However, the bird communities of these different forest types were very similar in species composition, and had much lower densities than those normally encountered in other, superficially similar forests. Although an 81% discrimination between forest types was attained through analysis of ground surface invertebrates, measures of litter and aerial invertebrate abundance were also of limited use as discriminators. Historical and biogeographic factors, as well as the low nutritional levels in the soil and vegetation may be the causes of low bird and invertebrate density and diversity. It is concluded that floristics and vegetation structure have, at best, a minor influence on bird community structure, and possibly also on invertebrate community structure in the Knysna Forest.     

Bird Populations in Rustic and Planted Shade Coffee Plantations of Eastern Chiapas, México

Much of the remaining “forest” vegetation in eastern Chiapas, Mexico is managed for coffee production. In this region coffee is grown under either the canopy of natural forest or under a planted canopy dominated by Inga spp. Despite the large differences in diversity of dominant plant species, both planted and rustic shade coffee plantations support a high overall diversity of bird species; we recorded approximately 105 species in each plantation type on fixed radius point counts. We accumulated a combined species list of 180 species on repeatedly surveyed transects through both coffee plantation types. These values are exceeded regionally only by moist tropical forest. Of the habitats surveyed, shade coffee was second only to acacia groves in the abundance and diversity of Nearctic migrants. The two plantation types have similar bird species lists and both are similar in composition to the dominant woodland—mixed pine-oak. Both types of shade coffee plantation habitats differ from other local habitats in supporting highly seasonal bird populations. Survey numbers almost double during the dry season—an increase that is found in omnivorous migrants and omnivorous, frugivorous, and nectarivorous resident species. Particularly large influxes were found for Tennessee warblers (Vermivora peregrina) and northern orioles (Icterus galbula) in Inga dominated plantations.     

The effect of windthrow, wild fire, and management practices on epigeic Collembola in windthrown forest stands of the High Tatra Mts (Slovakia)

Epigeic communities of Collembola were studied after a windthrow in differently managed spruce forest stands in the High Tatras, Slovakia. Investigations were carried out in following treatments: reference forest stands (REF), windthrown stands left to natural regeneration (NEX), windthrown stands from which fallen wood had been extracted (EXT), and windthrown stands from which fallen wood had been removed and which had been affected by fire (FIR). A Kruskal-Wallis nonparametric ANOVA revealed significant differences of Collembola community composition and activity between the treatments studied, which was also further supported by the results from NMS ordination and cluster analysis using indicator species analysis. EXT stands had communities with remarkably high activity and diversity of Collembola, whereas FIR stands had low species richness and diversity indices compared to other treatments (REF, NEX, EXT). Windthrown stands had greater collembolan activity and diversity (NEX, EXT) than intact forest stands. However, forest specialists and bryophilous species were sensitive to forest clearing and fire disturbance and were replaced by xeroresistant and generalist species, or species preferring open habitats. The collembolan community most similar to that found in REF stands, in terms of community homogeneity and the magnitude of diversity indices, was found in NEX stands.     

Composition and diversity of bird communities in Swedish farmland–forest mosaic landscapes

CapsuleThe amount of forest (at local and landscape scales) and occurrence of residual habitats at the local scale are shown to be the major factors influencing bird community composition in farmland–forest landscapes in central Sweden.

Aims To investigate the importance of local habitat and landscape structure for breeding birds in farmland–forest landscapes in central Sweden.

Methods Breeding birds were censused at 292 points. A detailed habitat mapping was made within 300 m of the points. Within a 300–600 m radius only two major habitats (forests and arable fields) were identified.

Results Cluster analyses of bird communities identified three site types that also differed in habitat composition: (i) partially forested sites in forested landscapes; (ii) heterogeneous sites with residual habitats in mosaic landscapes; and (iii) field-dominated farmland sites in open landscapes. A total of 19 of 25 farmland bird species (restricted to farmland or using both farmland and forest) had the highest abundance in farmland sites with mosaics of forest and farmland, while only six farmland species had the highest abundance in field-dominated sites. The bird community changed from being dominated by farmland species to being dominated by forest species (common in forest landscapes without farmland) at small proportions (10–20%) of forest at the local scale. A major difference in habitat composition between heterogeneous and field-dominated sites was the occurrence of different residual habitats (e.g. shrubby areas and seminatural grasslands). These habitats seemed to influence bird community composition more than land-use, despite covering <10% of the area. Seminatural grasslands were important for bird community composition and species-richness, but grazing seemed to be less important. Among different land-use types, cereal crops were the least preferred fields. Set-asides with tall vegetation and short rotation coppices were positively associated with species-richness of farmland birds.

Conclusion In general, the composition of the landscape was important for bird community composition, although amount and distribution of forests, occurrence of residual habitats and land-use of fields at the local scale had the strongest influence on bird community composition. The possible implications of these patterns for managing farmland–forest landscapes are discussed.     

Spatial Variation of Bird Species in Landscapes Dominated by Old-growth Forests in Northern Boreal Finland

An important issue in conservation planning is to study the distribution and abundance patterns of species in natural landscapes. Information of concordant and clumped species distribution pattertns can enhance attempts to focus conservation efforts in sites/areas preferred by various species. We compared bird species abundance, community composition and species distribution patterns in three large old-growth forest areas (size 40–120 km²) in northern boreal Finland on the basis of quantitative bird censuses. Total bird density and species composition, and mean density and variances of the most abundant species were highly similar between the three areas. Most of the bird species were distributed randomly within the areas and species preferring old-growth forests showed compensatory density variation without preference for any sub-area. Mean density of species preferring old-growth forests was significantly higher in the studied large old-growth forests than in smaller old-growth forests and was about threefold higher than regional density of species in a predominantly managed landscape matrix. Bird species preferring old-growth forests are probably not dependent on a certain specific part of a virgin boreal forest landscape but rather on the overall size of the high-quality and diverse old-growth forest area.     

Forest edges have high conservation value for bird communities in mosaic landscapes

A major conservation challenge in mosaic landscapes is to understand how trait‐specific responses to habitat edges affect bird communities, including potential cascading effects on bird functions providing ecosystem services to forests, such as pest control. Here, we examined how bird species richness, abundance and community composition varied from interior forest habitats and their edges into adjacent open habitats, within a multi‐regional sampling scheme. We further analyzed variations in Conservation Value Index (CVI), Community Specialization Index (CSI) and functional traits across the forest‐edge‐open habitat gradient. Bird species richness, total abundance and CVI were significantly higher at forest edges while CSI peaked at interior open habitats, i.e., furthest from forest edge. In addition, there were important variations in trait‐ and species‐specific responses to forest edges among bird communities. Positive responses to forest edges were found for several forest bird species with unfavorable conservation status. These species were in general insectivores, understorey gleaners, cavity nesters and long‐distance migrants, all traits that displayed higher abundance at forest edges than in forest interiors or adjacent open habitats. Furthermore, consistently with predictions, negative edge effects were recorded in some forest specialist birds and in most open‐habitat birds, showing increasing densities from edges to interior habitats. We thus suggest that increasing landscape‐scale habitat complexity would be beneficial to declining species living in mosaic landscapes combining small woodlands and open habitats. Edge effects between forests and adjacent open habitats may also favor bird functional guilds providing valuable ecosystem services to forests in longstanding fragmented landscapes.     

Breakdown of the species-area relationship in exotic but not in native forest patches

We studied the pattern of bird species richness in native and exotic forest patches in Hungary. We hypothesized that species-area relationship will depend on forest naturalness, and on the habitat specialization of bird species. Therefore, we expected strong species-area relationship in native forest patches and forest bird species, and weaker relationship in exotic forest patches containing generalist species. We censused breeding passerine bird communities three times in 13 forest patches with only native tree species, and 14 with only exotic trees in Eastern Hungary in 2003. Although most bird species (92%) of the total of 41 species occurred in both exotic and native forests, the species-area relationship was significant for forest specialist, but not for generalist species in the native forests. No relationship between bird species and area was found for either species group in the forest with exotic tree species. The comparison of native versus exotic forest patches of similar sizes revealed that only large (>100 ha) native forests harbor higher bird species richness than exotic forests for the forest specialist bird species. There is no difference between small and medium forest patches and in richness of generalist species. Thus, the species-area relationship may diminish in archipelago of exotic habitat patches and/or for habitat generalist species; this result supports the warning that the extension of exotic habitats have been significantly contributing to the decline of natural community patterns.     

Assessing avian diversity and community composition along a successional gradient in traditional Lacandon Maya agroforests

Evidence regarding the ability of agroforests to conserve biological diversity has been mixed; they tend to maintain avian communities with species richness similar to that of undisturbed forest ecosystems but generally do not completely preserve community composition. Using a combination of occupancy modeling and non-metric multidimensional scaling on point-count data, we assessed changes in avian community diversity and composition along a successional gradient in traditional Lacandon Maya agroforests and compared them to protected areas in the region. Bird species richness and diversity in Lacandon agroforests peaked in early secondary forest stages. These agroforests' mean Shannon–Weiner diversity was 5% higher than that of nearby protected areas, but their species richness was similar. Community composition in Lacandon agroforests changed throughout succession, with earlier stages supporting communities distinctly characterized by generalist species, while subsequent, less-intensively managed stages tended to support more forest-dwellers. The bird community observed in even the most mature secondary forest stages in Lacandon agroforests differed from that of undisturbed rain forest ecosystems. These results demonstrate the potential of traditional Lacandon agroforestry management to conserve avian biodiversity while ensuring food sovereignty for farmers. However, because the community composition of early-successional stages was different than later stages, shortening fallow cycles and reducing forest cover to increase agricultural production will limit the species this system can support. This study illustrates the value of incorporating traditional agroecosystems into conservation planning as well as maintaining protected areas, because the latter serve as refugia for species that require undisturbed forest habitat in an agroecological matrix.     

浙江古田山自然保护区鸟类群落生态研究

对古田山地区鸟类进行的调查表明:该地区鸟类群落的主要鸟类共计48种,其中优势种有14种,占29.17％。根据鸟类生境的分布系数分析。中性分布型有(25—100％)28种,占58.33％。不同生境内鸟类群落的种类数目和群体密度存在差异,结构复杂的生境,其鸟类群落由三个分布群组成,如农田村落、农田河滩、混交林和阔叶林;结构简单的生境,如竹林、针叶林和迹地灌丛,其鸟类群落由二个分布群所组成。各种鸟类分布群在鸟类群落内的比例随栖息地结构的变化而改变。由于栖息地结构的变化同样也导致鸟类群落其他参数的变化,如Shannon-Wiener多样性指数、Simpson优势度、种间相遇机率PIE等。