Nested bird and micro-habitat assemblages in a peatland archipelago

Biotic assemblages of insular habitats are nested when poor assemblages are subsets of richer ones. Nestedness of species assemblages is frequent and may result from selective extinction or frequent colonization in insular habitats. It may also be created by a nested distribution of habitats among islands or by sampling bias. We sampled 67 isolated peatlands (7–843 ha) in southern Quebec, Canada, to measure nestedness of bird species assemblages among peatlands and assess the habitat nestedness hypothesis. Species and microhabitat assemblages were both strongly nested among peatlands. Whether sites were ranked by species richness, microhabitat richness or peatland area had no effect on nestedness. However, microhabitat nestedness was significantly reduced when sites were sorted by area rather than by microhabitat richness. As expected, if bird-microhabitat associations are responsible for the nested pattern of distribution, we found a positive correlation between the contributions of bird species and microhabitats to individual site nestedness. Nevertheless, microhabitat assemblages were significantly less nested than bird species assemblages, possibly because of frequent recolonization by birds or uneven sampling among sites. Received: 12 June 1998 / Accepted: 20 September 1998     

Compositional dissimilarity patterns of reptiles and amphibians in insular systems around the world

Several studies have shown that taxa with poor dispersal ability have a higher level of compositional dissimilarity than good dispersers. However, compositional dissimilarity patterns between islands with respect to dispersal ability of taxa have never been investigated before. In this study, we investigated compositional dissimilarity patterns of three taxonomic groups, namely amphibians, lizards, and snakes, differing in their dispersal abilities, in various insular systems around the world. We compiled presence–absence matrices, based on which we calculated several metacommunity indices to check for differences among taxonomic groups and island types (oceanic and continental shelf) using classical statistical tests and generalized linear mixed-effects models. According to our results, compositional dissimilarity was positively affected by the isolation of the insular system, in accordance to theory. In particular, oceanic systems were characterized by a high level of compositional dissimilarity between islands and subsequently by a low level of nestedness. SIEs may be generating these patterns causing distortions from expected levels of nestedness. Similar to our predictions, compositional dissimilarity patterns were also dependent on taxon-specific dispersal ability, with good dispersers showing lower levels of between-island compositional dissimilarity than poor dispersers in continental shelf systems. However, this pattern was not observed in oceanic systems. In conclusion, compositional dissimilarity in insular systems is dependent on both taxon and island type.     

Nestedness and niche‐based species loss in moorland plant communities

Communities in isolated habitat patches surrounded by inhospitable matrices often form a nested subset pattern. However, the underlying causal mechanisms and conservation implications of nestedness in regional communities remain controversial. The nested ranks of species in a nested species‐by‐site matrix may reflect a gradient of species vulnerability to extinction or of colonization ability. However, nestedness analysis has rarely been used to explore determinants of species rank; consequently, little is known of underpinning mechanisms. In this study, we examined nestedness in moorland plant communities widely interspersed within the subalpine zone of northern Japan. Moorland sites differed in area (1000–160 000 m²) and were naturally isolated from one another to various extents within an inhospitable forest matrix. We also determined whether site characteristics (physical and morphometric measures) and species characteristics (niche position and breadth, based on species’ traits) are related to nestedness. Moorland plant communities in the study area were significantly nested. The pH and moorland kernel density (proxy for spatial clustering of moorlands around the focal site) were the most important predictors of moorland site nested rank in a nestedness matrix. Niche breadths of species (measured as variation in leaf mass area and height) predicted species’ nested ranks. Selective environmental tolerances imposed by environmental harshness and selective extinction caused by declines in site carrying capacities probably account for the nested subset pattern in moorland plant communities. The nested rank of species in the nestedness matrix can therefore be translated into the potential order of species loss explainable by species niche breadths (based on variation in functional traits). Complementary understanding of the determinants of site ranking and species ranking in the nestedness matrix provides powerful insight into ecological processes underlying nestedness and into the ways by which communities are assembled or disassembled by such processes.     

舟山群岛蝶类群落嵌套分布格局及其影响因素

片断化生境中群落的物种组成常呈现嵌套分布格局。2013年7-8月, 我们在浙江舟山群岛采用截线法对28个岛屿上的蝴蝶群落进行了野外调查, 探讨了岛屿物种嵌套分布格局及其影响因素。通过测量采集标本获得蝶类的生活史特征(最小需求面积、翅展和体重), 查阅文献资料获得蝶类的栖息地特征(岛屿面积、距最近大陆距离和距最近大岛距离), 分析了影响蝶类群落嵌套结构的因素。研究结果显示: (1)舟山群岛蝶类群落符合嵌套分布格局; (2)岛屿面积和物种最小需求面积对嵌套格局的形成有显著影响; (3)舟山群岛蝶类群落嵌套格局的形成支持选择性灭绝假说; (4)随机检验零模型结果显示该嵌套分布格局并非采样偏差造成的。因此, 在制定舟山群岛区域蝶类保护措施时, 应优先考虑那些分布在面积较大岛屿的和最小需求面积较大的物种。     

Nestedness of butterfly assemblages in the Zhoushan Archipelago,China: area effects,life-history traits and conservation implications

The nested subset pattern (nestedness) of faunal assemblages has been a research focus in the fields of island biogeography and conservation biology in recent decades. However, relatively few studies have described nestedness in butterfly assemblages in oceanic archipelago systems. Moreover, previous studies often quantified nestedness using inappropriate nestedness metrics and random fill algorithms with high Type I errors. The aims of this study are to examine the existence of nestedness and underlying causal mechanisms of butterfly assemblages in the Zhoushan Archipelago, China. We used the line-transect method to determine butterfly occupancy and abundance on 42 study islands from July to August 2014. We obtained butterfly life-history traits (wingspan, body weight and minimum area requirement) by field work and island geographical features (area and isolation) from the literature. We used the recently developed metric WNODF to estimate nestedness. Partial Spearman rank correlation was used to evaluate the associations of nestedness and island geographical features as well as butterfly life-history traits related to species extinction risk and colonization ability. The butterfly assemblages were significantly nested. Island area and minimum area requirement of butterflies were significantly correlated with nestedness after controlling for other independent variables. In contrast, the nestedness of butterflies did not appear to result from passive sampling or selective colonization. However, multi-year studies are needed to confirm that target effects are not muddling these results. Our results indicate that selective extinction may be the main driver of nestedness of butterfly assemblages in our study system. From a conservation viewpoint, we should protect both large islands and species with large area requirement to maximize the number of species preserved.     

Effects of Fragmentation and Sea-Level Changes upon Frog Communities of Land-Bridge Islands off the Southeastern Coast of Brazil

We investigate the composition of anuran communities of land-bridge islands off the southeastern coast of Brazil. These islands provide natural long-term experiments on the effects of fragmentation in the Brazilian Atlantic Forest (BAF). We hypothesize that Pleistocene sea-level changes, in combination with other abiotic variables such as area and habitat diversity, has affected anuran species richness and community composition on these islands. Data from the literature and collections databases were used to produce species lists for eight land-bridge islands and for the mainland adjacent to the islands. We assess the effects of area, number of breeding habitats and distance to the mainland upon anuran species richness on land-bridge islands. Additionally we use nestedness analysis to quantify the extent to which the species on smaller and less habitat-diverse islands correspond to subsets of those on larger and more diverse ones. We found that area has both direct and indirect effects on anuran species richness on land-bridge islands, irrespective of distance to the mainland. However, on islands with comparable sizes, differences in species richness can be attributed to the number and quality of breeding habitats. Anuran communities on these islands display a nested pattern, possibly caused by selective extinction related to habitat loss. Common lowland pond-breeders were conspicuous by their absence. In the BAF, the conservation of fragments with a high diversity of breeding habitats could compensate for the generally negative effect of small area upon species richness. We suggest that sea-level changes have an important role in shaping composition of anuran species on coastal communities.     

Nestedness theory suggests wetland fragments with large areas and macrophyte diversity benefit waterbirds

Many artificial wetland constructions are currently underway worldwide to compensate for the degradation of natural wetland systems. Researchers face the responsibility of proposing wetland management and species protection strategies to ensure that constructed wetlands positively impact waterbird diversity. Nestedness is a commonly occurring pattern for biotas in fragmented habitats with important implications for conservation; however, only a few studies have focused on seasonal waterbird communities in current artificial wetlands. In this study, we used the nestedness theory for analyzing the annual and seasonal community structures of waterbirds in artificial wetlands at Lake Dianchi (China) to suggest artificial wetland management and waterbird conservation strategies. We carried out three waterbird surveys per month for one year to observe the annual, spring, summer, autumn, and winter waterbird assemblages in 27 lakeside artificial wetland fragments. We used the NeD program to quantify nestedness patterns of waterbirds at the annual and seasonal levels. We also determined Spearman partial correlations to examine the associations of nestedness rank and habitat variables to explore the factors underlying nestedness patterns. We found that annual and all four seasonal waterbird compositions were nested, and selective extinction and habitat nestedness were the main factors governing nestedness. Further, selective colonization was the key driver of nestedness in autumn and winter waterbirds. We suggest that the area of wetland fragments should be as large as possible and that habitat heterogeneity should be maximized to fulfill the conservation needs of different seasonal waterbirds. Furthermore, we suggest that future studies should focus on the least area criterion and that vegetation management of artificial wetland construction should be based on the notion of sustainable development for humans and wildlife.     

Geographic variation in patterns of nestedness among local stream fish assemblages in Virginia

Nestedness of faunal assemblages is a multi-scale phenomenon, potentially influenced by a variety of factors. Prior small-scale studies have found freshwater fish species assemblages to be nested along stream courses as a result of either selective colonization or extinction. However, within-stream gradients in temperature and other factors are correlated with the distributions of many fish species and may also contribute to nestedness. At a regional level, strongly nested patterns would require a consistent set of structuring mechanisms across streams, and correlation among species tolerances of the environmental factors that influence distribution. Thus, nestedness should be negatively associated with the spatial extent of the region analyzed and positively associated with elevational gradients (a correlate of temperature and other environmental factors). We examined these relationships for the freshwater fishes of Virginia. Regions were defined within a spatial hierarchy and included whole river drainages, portions of drainages within physiographic provinces, and smaller subdrainages. In most cases, nestedness was significantly stronger in regions of smaller spatial extent and in regions characterized by greater topographic relief. Analysis of hydrologic variability and patterns of faunal turnover provided no evidence that inter-annual colonization/extinction dynamics contributed to elevational differences in nestedness. These results suggest that, at regional scales, nestedness is influenced by interactions between biotic and abiotic factors, and that the strongest nestedness is likely to occur where a small number of organizational processes predominate, i.e., over small spatial extents and regions exhibiting strong environmental gradients.The Virginia Cooperate Fish and Wildlife Research Unit is jointly sponsored by United States Geological Survey, Virginia Polytechnic Institute and State University, and Virginia Department of Game and Inland Fisheries.     

Effects of site and species characteristics on nested patterns of species composition in sedge meadows

Biotas of both geographical islands and habitat islands are often nested subsets of the biotas of successively more species-rich islands within the same system. The life history characteristics of a species may determine how that species contributes to the general pattern of species nestedness. Here, I investigate the floras of 56 sedge meadow wetlands in northern Illinois (USA) in order to characterize the degree of nestedness in these communities, determine which individual plant species contribute to the nested pattern, and investigate species characteristics that might be related to nonrandom patterns of distribution in individual plant species. The entire assemblage of species at all sedge meadows was significantly nested. Species richness and area were significantly correlated, and the nested pattern was closely related to site area, suggesting that species drop out of the assemblage in a predictable order as site area decreases. Some individual species exhibited nonrandom distributions across the sites, occurring more often in large, species-rich sites. Large sites were more likely than smaller sites to contain conservative species, i.e., those typical of pristine natural habitat, whereas nonconservative species were distributed more randomly among sites. Nested patterns of distribution of conservative species with respect to site area may result from their high probability of extinction on small sites or from a tendency for required habitats to co-occur on the same large sites. This revised version was published online in June 2006 with corrections to the Cover Date.     

The relationship between nested subsets,habitat subdivision,and species diversity

Biotic assemblages are said to be nested when the species making up relatively species-poor biotas comprise subsets of the species present at richer sites. Because species number and site area are often correlated, previous studies have suggested that nestedness may be relevant to questions of how habitat subdivision affects species diversity, particularly with respect to the question of whether a single large, contiguous patch of habitat will generally contain more species than collections of smaller patches having the same total combined area. However, inferences from analyses of nestedness are complicated by (1) variability in degrees of nestedness measured in natural communities, (2) variability in species-area relationships, and (3) the fact that nestedness statistics do not account for the size of habitat patches, only in the degree of overlap among sites with different numbers of species. By comparing various indices of nestedness with a saturation index that more directly measures the effect of habitat subdivision, it is shown that the first two of these factors are not as important as the third. Whether a single large site or several smaller ones having the same total combined area maximizes species diversity is dependent on (1) overlap in species composition among sites and (2) the number of species per unit area in the different sites. Because nestedness indices do not account for species number at a site, they cannot accurately predict how habitat subdivision affects species diversity patterns. Still, nestedness analyses are important in that they indicate the degree to which rare species tend to be found in the largest, or the most species-rich, sites, patterns not revealed by the saturation index. Both types of analysis are important in order to obtain a more complete picture of how species richness and compositional patterns are influenced by habitat subdivision.     

On the meaning and measurement of nestedness of species assemblages

Nestedness of species assemblages occurs when thebiotas of sites with lower numbers of species tend to be subsets of the biotas at richer sites. We develop new quantitative and statistical techniques for measuring, testing, and comparing nestedness, and apply these methods to data from the literature. Significantly nonrandom nestedness was present in all 27 assemblages examined, and tended to be stronger in systems dominated by extinction, such as landbridge islands. Sets of assemblages that were very strongly nested were more likely to have greater species richness on one or a few large sites than on several smaller sites of equivalent total area — that is, to fall toward the single large side of the Single Large Or Several Small (SLOSS) continuum. Our analysis indicates that nestedness, when quantified as a single number for a presence-absence matrix, measures community-wide differences in incidence (the frequency of occurrence or distribution of species). Factors that lead to consistent differences among species in immigration or extinction rates cause strong patterns of nestedness of species assemblages. Nestedness is negatively related to beta diversity: nestedness is low when beta diversity is high, and vice versa. Conservation managers will thus seek to minimize nestedness and the development of nested structure in systems of nature reserves.     

Can the biotic nestedness matrix be used predictively?

The biotas of a suite of neighboring patches of remnant vegetation often form a series of nested sub-sets, in which the species present in species-poor patches are non-random sub-sets of those present in richer patches. There has been recent interest in ways in which this knowledge may be used to aid conservation. We focus here on whether nested patterns can be used predictively. If nestedness in a fragmented system increases over time through biotic relaxation, locations where particular species may become extinct or are likely to colonize might be predictable and this could be useful in threatened-species management. We used the Temperature Calculator of Atmar and Patterson (1995) to arrange a matrix of bird species' occurrences in a series of buloke Allocasuarina leuhmannii woodland remnants so that nestedness was maximized. Probability bands generated by the calculator were used to predict possible colonization and extinction events. We then re-surveyed the avifauna of the fragments after a seven-year interval to test these predictions. Although nestedness increased between the two survey periods, there was no linear relationship between the generated probability of extinctions or colonizations and the accuracy of the predictions. The predictions derived from the calculator were no more accurate than a second set of predictions generated by use of a simple non-nested model. Despite the increase in nestedness, the arrangement of sites in each of the two maximally packed matrices was substantially different. For the nestedness matrix to generate accurate predictions, an increase in nestedness must be due to a minimization of unexpected species presences and absences rather than an extensive redistribution of species among remnants, as we found. The potential utility of nested patterns in predicting systematic colonization and extinction events should be further evaluated in other, less dynamic, fragmented systems such as those undergoing biotic relaxation.     

Temporal dynamics and nestedness of an oceanic island bird fauna

Aim To examine temporal variation in nestedness and whether nestedness patterns predict colonization, extinction and turnover across islands and species. Location Dahlak Archipelago, Red Sea. Method The distributions of land birds on 17 islands were recorded in two periods 30 years apart. Species and islands were reordered in the Nestedness Temperature Calculator, software for assessing degrees of nestedness in communities. The occupancy probability of each cell, i.e. species–island combinations, was calculated in the nested matrix and an extinction curve (boundary line) was specified. We tested whether historical and current nested ranks of species and islands were correlated, whether there was a relationship between occupancy probability (based on the historical data) and number of extinctions or colonizations (regression analyses) and whether the boundary line could predict extinctions and colonizations (chi‐square analyses). Results Historical and current nested ranks of islands and species were correlated but changes in occupancy patterns were common, particularly among bird species with intermediate incidence. Extinction and turnover of species were higher for small than large islands, and colonization was negatively related to isolation. As expected, colonizations were more frequent above than below the boundary line. Probability of extinction was highest at intermediate occupancy probability, giving a quadratic relationship between extinction and occupancy probability. Species turnover was related to the historical nested ranks of islands. Colonization was related negatively while extinction and occupancy turnover were related quadratically to historical nested ranks of species. Main conclusions Some patterns of the temporal dynamics agreed with expectations from nested patterns. However, the accuracy of the predictions may be confounded by regional dynamics and distributions of idiosyncratic, resource‐limited species. It is therefore necessary to combine nestedness analysis with adequate knowledge of the causal factors and ecology of targeted species to gain insight into the temporal dynamics of assemblages and for nestedness analyses to be helpful in conservation planning.     

嵌套性:研究方法、形成机制及其对生物保护的意义

岛屿或者“生境岛”中的生物区系常常显示出一种嵌套结构 ,即物种较贫乏的岛屿中的物种是物种较丰富的岛屿中的物种的一个适当的子集 ,如果将各个岛屿中的生物区系排列起来就形成一个嵌套的序列。与种 面积关系一样 ,嵌套结构在很多生境类型和生物类群中也都存在。嵌套性对生物保护也有一定的意义 ,特别是与SLOSS争论 (是单个大的还是几个小的保护区能保护更多的物种 )有一定关系。在过去的十几年中 ,已经提出了一些方法 ,可以对嵌套性进行定量刻画和统计检验。同时 ,对嵌套性的形成机制也进行了大量的研究 ,其中选择性的迁移和选择性的灭绝是两个主要的原因。由于嵌套性分析只需要物种的存在 /不存在数据 ,使得很多调查数据都能够利用起来 ,因此 ,这是一个值得深入研究的领域     

Can human disturbance promote nestedness? A case study with breeding birds in urban habitat fragments

Several factors, singly or in combination, have proven relevant in promoting nestedness in animal communities (area, isolation, habitat structure, etc.); however, little empirical evidence exists about the role of human disturbance. The goal of this paper was to assess whether human disturbance (pedestrians) may affect bird species composition in such a way as to generate a nested distributional pattern. The study was conducted in an urban fragmented landscape, the city of Madrid, where wooded parks were suitable fragments for many bird species, and had different levels of human visitation. At the community level, the distribution of species was significantly nested in two consecutive breeding seasons. Using two different procedures ("temperature" and "departures" methods) to analyse causality in nestedness, I found that pedestrian rate, fragment size and the diversity of stems were significantly correlated to the nested pattern. When analysed simultaneously, and controlling for their independent effects, these factors still accounted for nestedness. Pedestrian rate was the only factor significantly associated with changes in species composition between years. At the individual species level, 74 and 41% of species were significantly nested in relation to pedestrian rate in 1997 and 1998, respectively; however, these percentages were independent of foraging substrate and body size. Besides the classic area and habitat diversity effects, human disturbance can also promote nestedness: locally, by restraining the time and space of foraging and breeding opportunities, thus reducing fragment suitability, and regionally, by increasing extinction and decreasing colonization probabilities in highly disturbed fragments.     

Distributions of forest birds and butterflies in the Andaman islands, Bay of Bengal: nested patterns and processes

The distributional patterns of forest birds and butterflies in the Andaman islands, an oceanic chain located off SE Asia, were tested for nestedness. Both taxa were highly nested. Nestedness could be due to colonization or extinction processes, area or distance effects or nestedness of habitats. Nestedness in forest bird distributions were strongly influenced by area and habitat related factors. Habitats were significantly nested in all three island groups, however most strongly for the North Andamans. However forest bird distributions in the North Andamans, as indicated by row order in the packed matrix, was not correlated with habitat diversity, suggesting that habitat related factors alone cannot account for these patterns. Other causal influences could be passive sampling, where common and abundant species and habitats are more likely to have a widespread distribution than rare species and habitats. The nested subset pattern seen in two unrelated taxa suggests that the Andamans are extinction dominated and that the protection of forests on large islands is critical for the conservation of its biodiversity.     

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.     

千岛湖陆桥岛屿繁殖鸟类的扩散能力差异 对群落动态的影响

岛屿生物地理学理论的核心过程是岛屿物种的周转, 包括迁入与灭绝。本研究旨在探讨扩散能力差异对岛屿繁殖鸟类群落动态的影响。2007年4月至2013年6月, 采用样线法调查了千岛湖36个陆桥岛屿的繁殖鸟类, 依据扩散能力强弱将其划分为两类, 结合陆桥岛屿参数, 并运用逻辑斯蒂回归模型和最大似然法, 来研究鸟类扩散能力的不同对其周转率的影响。结果表明, 千岛湖繁殖鸟类扩散能力强的物种具有较高周转率且受岛屿参数约束较小, 而扩散能力弱的物种周转率较低且对岛屿参数变化更敏感。因此, 千岛湖陆桥岛屿繁殖鸟类的扩散能力显著影响其群落动态。     

Nestedness of desert bat assemblages: species composition patterns in insular and terrestrial landscapes

Nested patterns of community composition exist when species at depauperate sites are subsets of those occurring at sites with more species. Nested subset analysis provides a framework for analyzing species occurrences to determine non-random patterns in community composition and potentially identify mechanisms that may shape faunal assemblages. We examined nested subset structure of desert bat assemblages on 20 islands in the southern Gulf of California and at 27 sites along the Baja California peninsula coast, the presumable source pool for the insular faunas. Nested structure was analyzed using a conservative null model that accounts for expected variation in species richness and species incidence across sites (fixed row and column totals). Associations of nestedness and island traits, such as size and isolation, as well as species traits related to mobility, were assessed to determine the potential role of differential extinction and immigration abilities as mechanisms of nestedness. Bat faunas were significantly nested in both the insular and terrestrial landscape and island size was significantly correlated with nested structure, such that species on smaller islands tended to be subsets of species on larger islands, suggesting that differential extinction vulnerabilities may be important in shaping insular bat faunas. The role of species mobility and immigration abilities is less clearly associated with nestedness in this system. Nestedness in the terrestrial landscape is likely due to stochastic processes related to random placement of individuals and this may also influence nested patterns on islands, but additional data on abundances will be necessary to distinguish among these potential mechanisms.