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.     

The influence of colonization in nested species subsets

Biotic communities inhabiting collections of insular habitat patches often exhibit compositional patterns described as nested subsets. In nested biotas, the assemblages of species in relatively depauperate sites comprise successive subsets of species in relatively richer sites. In theory, nestedness may result from selective extinction, selective colonization, or other mechanisms, such as nested habitats. Allopatric speciation is expected to reduce nestedness. Previous studies, based largely on comparisons between land-bridge and oceanic archipelagos, have emphasized the role of selective extinction. However, colonization could also be important in generating strong patterns of nestedness. We apply a recently published index of nestedness to more than 50 island biogeographic data sets, and examine the roles of colonization, extinction, endemism, and, to a limited extent, habitat variability on the degree on nestedness. Most data sets exhibit a significant degree of nestedness, although there is no general tendency for land-bridge biotas to appear more nested than oceanic ones. Endemic species are shown to generally reduce nestedness. Comparisons between groups of non-endemic species differing in overwater or inter-patch dispersal ability indicate that superior dispersers generally exhibit a greater degree of nestedness than poorer dispersers, a result opposite that expected if colonization were a less predictable process than extinction. These results suggest that frequent colonization is likely to enhance nestedness, thereby increasing the compositional overlap among insular biotas. The prevalence of selective extinction in natural communities remains in question. The importance of colonization in generating and maintaining nested subsets suggests that (1) minimum critical areas will be difficult to determine from patterns of species distributions on islands; (2) multiple conservation sites are likely to be required to preserve communities in subdivided landscapes; and (3) management of dispersal processes may be as important to preserving species and communities as is minimizing extinctions.     

Mammalian biogeography of the Alexander Archipelago of Alaska: a north temperate nested fauna

Aim A large number of studies have analysed the distribution of mammals within archipelagos, yet few have focused on islands that were heavily glaciated and subsequently colonized following deglaciation. Location We explored the relative effects of island area and isolation on faunal composition based on twenty-three mammalian taxa of twenty-four islands of the Alexander Archipelago, Southeast Alaska. Methods We used regression of log-transformed variables and several indices of nestedness. Results These faunas showed significant nested structure using tests of nestedness and regression models. Unlike most landbridge and mainland archipelagos studied previously, isolation appears to be the primary factor determining patterns of species richness. Main conclusions Colonization ability of particular taxa, rather than extinction, has determined this nested structure. We suggest that other higher latitude archipelagos may show similar historical patterns.     

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.     

Investigating causality of nestedness of insular communities: selective immigrations or extinctions?

Abstract. Most archipelagos are comprised of a nested set of communities, with species on depauperate islands representing proper subsets of those on richer islands. The causality for this common intra-archipelago pattern, however, remains elusive. Here, I present a Monte Carlo approach for investigating whether nestedness results from selective extinctions, selective immigrations, or both. Results for mammal communities of three nearshore archipelagos and two montane forest archipelagos suggested that nestedness may result from both forces. Nestedness was significantly associated with area (extinctions) for all archipelagos studied, and with isolation (immigrations) for islands of Lake Huron and the American Southwest. The degree of nestedness is influenced by the degree of variations among species and islands. In addition, our ability to assess causality of nestedness is influenced by our ability to calculate biologically relevant measures of isolation, and by the potentially confounding effects of selective immigrations and extinctions on community nestedness.     

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     

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.     

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.     

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.     

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。     

The measure of order and disorder in the distribution of species in fragmented habitat

Species distribution patterns within naturally fragmented habitat have been found to often exhibit patterns of pronounced nestedness. Highly predictable extinction sequences are implied by these nested species distribution patterns, thus the patterns are important to both the philosophy and practice of conservation biology. A simple thermodynamic measure of the order and disorder apparent in the nested patterns is described. The metric offers (i) a measure of the uncertainty in species extinction order, (ii) a measure of relative populational stabilities, (iii) a means of identifying minimally sustainable population sizes, and (iv) an estimate of the historical coherence of the species assemblage. Four presumptions govern the development of the metric and its theory: (i) the fragmented habitat was once whole and originally populated by a single common source biota, (ii) the islands were initially uniform in their habitat heterogeneity and type mix, and have remained so throughout their post-fragmentation history, (iii) no significant clinal (latitudinal) gradation exists across the archipelago so as to promote species turnover across the archipelago, and (iv) all species of interest are equally isolated on all islands. The violation of these conditions promotes species distributions which are idiosyncratic to the general extinction order expected in fragmentation archipelagos. While some random variation in extinction order is to be expected, idiosyncratic distributional patterns differ from randomness and are readily segregatable from such noise. A method of identifying idiosyncratic species and sites is described.     

Terrestrial bird community patterns on the coralline islands of the Dahlak Archipelago, Red Sea, Eritrea

Aim This study aims to explain the patterns of species richness and nestedness of a terrestrial bird community in a poorly studied region. Location Twenty‐six islands in the Dahlak Archipelago, Southern Red Sea, Eritrea. Methods The islands and five mainland areas were censused in summer 1999 and winter 2001. To study the importance of island size, isolation from the mainland and inter‐island distance, I used constrained null models for the nestedness temperature calculator and a cluster analysis. Results Species richness depended on island area and isolation from the mainland. Nestedness was detected, even when passive sampling was accounted for. The nested rank of islands was correlated with area and species richness, but not with isolation. Idiosyncrasies appeared among species‐poor and species‐rich islands, and among common and rare species. Cluster analysis showed differences among species‐rich islands, close similarity among species‐poor and idiosyncratic islands, and that the compositional similarity among islands decreased with increasing inter‐island distance. Thus, faunas of species‐poor, smaller islands were more likely to be subsets of faunas of species‐rich, larger islands if the distance between the islands was short. Main conclusions Species richness and nestedness were related to island area, and nestedness also to inter‐island distances but not to isolation from the mainland. Thus, nestedness and species richness are not affected in the same way by area and distance. Moreover, idiosyncrasies may have been the outcome of species distributions among islands being influenced also by non‐nested distributions of habitats, inter–specific interactions, and differences in species distributions across the mainland. Idiosyncrasies in nested patterns may be as important as the nested pattern itself for conservation – and conservation strategies based on nestedness and strong area effects (e.g. protection of only larger islands) may fail to preserve idiosyncratic species/habitats.     

Extinctions of montane mammals reconsidered: putting a global-warming scenario on ice

McDonald and Brown developed a model based on island biogeographic principles to predict the magnitude and composition of small mammal extinctions from isolated boreal habitats atop mountains of the Great Basin following global warming. The model predicts that three of 14 boreal mammals will go extinct regionally and that four of 19 mountain ranges will lose upwards of 50% of their present faunas. Here, we re-examine the model on the basis of its underlying assumptions, on the statistical and biogeographic protocols used, and on its predictive power. A key assumption, that populations of these small mammals are isolated by absolute barriers to dispersal, is challenged by published field observations and by extensive trapping records. Statistical procedures used to construct the model are questionable and the model itself yields imprecise estimates. The biogeographic principle used to identify extinction-prone species, nested subsets of species, makes predictions that are at odds with available autecological information. The demonstration of a nested pattern of species occurrences does not provide definitive evidence in resolving SLOSS-or whether a single large island or several small islands of equivalent total area will contain more species. We conclude that the model is not a reliable method for forecasting species extinctions following global warming. The final resolution of the biogeography of montane mammals (and predictive models of extinction) in the Great Basin must await a full and accurate accounting of past and present species distributions.     

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.     

Biogeography of Puerto Rican ants: a non-equilibrium case?

Ants were studied on Puerto Rico and 44 islands surrounding Puerto Rico. Habitat diversity was the best predictor of the number of species per island and the distributions of species followed a nested subset pattern. The number of extinctions per island was low, approximately 1–2 extinctions per island in a period of 18 years, and the rates of colonization seem to be greater than the extinction rates. Ant dynamics on these islands do not seem to support the basic MacArthur and Wilson model of island biogeography. The MacArthur and Wilson equilibrium is based on the notion that species are interchangeable, but some extinctions and colonizations can change the composition and number of species drastically.     

The role of stochastic processes in producing nested patterns of species distributions

Nestedness has received considerable attention in community ecology and conservation biology from both theoretical and empirical perspectives. This has lead to the creation of various metrics and null models to analyze nested subsets, all of which rely on the random placement of species to assess significance. However, if immigration and extinction are the processes that underlie species distributions on island systems, then null models might be better determined on the basis of randomly placed individuals. Consequently, we examined the effects of species–abundance distributions (uniform, dominance–decay, random–assortment, and dominance–preemption), island–size distributions (uniform and linear decrease), and total abundances (128, 256, 512, 1024, 2048, 4096 and 8192) on the degree of nestedness and its significance. Generally, matrices of species presence and absence created from the random placement of individuals were nested significantly according to null models based on the random placement of species. Island size and abundance had less of an effect on nestedness in systems dominated by only a few species than in systems in which abundances were distributed more evenly. Stochastic processes, such as the random placement of individuals, predispose systems to evince patterns of nestedness at the species level, which may account, in part, for the ubiquity of nestedness in nature.     

Invasive species and land bird diversity on remote South Atlantic islands

Invasive species pose significant threats to biodiversity, especially on islands. They cause extinctions and population declines, yet little is known about their consequences on the emergent, metacommunity-level patterns of native species in island assemblages. We investigated differences in species–area relationships, nestedness, and occupancy of 9 species of native land birds between island assemblages with and without invasive Norway rats (Rattus norvegicus) in the Falkland Archipelago. We found that species–area curves, nestedness, and individual species’ occurrences differed between island assemblages with and without rats. Rat-free islands had, on average, 2.1 more land bird species than rat-infested islands of similar size. Passerine bird communities on islands with and without rats were significantly nested, but nestedness was significantly higher on rat-free islands than on rat-infested islands. The presence of rats was associated with differences in the incidence of many, but not all bird species. On rat free islands the occurrence of all species increased with island area. The occurrence of most, albeit not all, bird species was lower on islands with than on islands without rats. Two species of conservation concern, Troglodytes aedon cobbi and Cinclodes antarcticus, were abundant on rat-free islands, but absent or found at very low frequencies on islands with rats. The occurrence of three species was not associated with the presence of rats. The patterns presented here can be used to evaluate the consequences of ongoing rat eradications for passerine diversity, distribution, and abundance.     

Effects of habitat size and isolation on species immigration–extinction dynamics and community nestedness in a desert river system

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Regionally nested patterns of fish assemblages in floodplain lakes of the Magdalena river (Colombia)

We investigated if fish assemblages in neotropical floodplain lakes (cienagas) exhibit nestedness, and thus offer support to the managers of natural resources of the area for their decision making. The location was floodplain lakes of the middle section of the Magdalena river, Colombia. We applied the nested subset analysis for the series of 30 cienagas (27 connected to the main river and three isolated). All fish were identified taxonomically in the field and the matrix for presence-absence in all the lakes was used for the study of the pattern of nestedness. The most diverse order was Characiformes (20 species), followed by Siluriformes (19 species). Characidae and Loricaridae were the richest families. The species found in all the lakes studied were migratory species (17), and sedentary species (33). Two species (Caquetaia kraussii and Cyphocharax magdalenae) were widespread across the cienagas archipelago (100% of incidence). Nestedness analysis showed that the distribution of species over the spatial gradient studied (840 km) is significantly nested. The cienagas deemed the most hospitable were Simiti, El Llanito, and Canaletal. Roughly, 13 out of the 50 species caught show markedly idiosyncratic distributions. The resulting dataset showed a strong pattern of nestedness in the distribution of Magdalenese fishes, and differed significantly from random species assemblages. Out of all the measurements taken in the cienagas, only the size (area) and local richness are significantly related to the range of order of nested subset patterns (r=-0.59 and -0.90, respectively, at p < 0.01). Differential species extinction is suggested as the cause of a nested species assemblage, when the reorganized matrix of species occurring in habitat islands is correlated with the island area. Our results are consistent with this hypothesis.