Network properties of arboreal plants: Are epiphytes,mistletoes and lianas structured similarly?

Network analyses provide a unified framework to evaluate different types of species interactions. We used a network approach to comparatively evaluate three types of arboreal plant metacommunities. Interactions between mistletoes, lianas and epiphytes and their host trees were quantified in two New Zealand forests and individual-based null models were used to test for non-random patterns in network degree, nestedness and negative co-occurrence patterns. Arboreal plants were simulated to randomly occur on individual host trees to derive ‘null’ interaction matrices, which were then compared to the observed matrix. Results showed that mistletoes, lianas and epiphytes had very different network properties. Mistletoe and liana degree distributions exhibited fewer links than expected under the null model, indicating strong host preferences. Conversely, degree distributions for epiphytes were consistent with randomised expectations. Mistletoes and lianas were less nested than null model expectations and instead showed support for negative co-occurrence patterns, meaning mistletoe and liana species tended to have mutually exclusive host preferences. Conversely, epiphytes were more nested than expected by chance and showed positive co-occurrence patterns. Overall results indicate that plant–plant interactions exhibited by different types of arboreal plants have very different network properties. We hypothesize that these differences result from (1) differences in coevolutionary dynamics between arboreal plants and their hosts, which range from parasitic (mistletoes) to commensal (epiphytes), and (2) biotic interactions among arboreal plant species for access to host trees.     

Patterns in the assembly of an island plant community

Aim  To test for patterns in the assembly of an island plant community.
Location  Islands off the west coast of Vancouver Island, British Columbia, Canada.
Methods  Twenty-seven islands were visited by boat, and the abundance of six woody angiosperm species was quantified. Null models were then used to test whether: (1) some species co-occur less than expected by chance (i.e. co-occurrence assembly rule), (2) the incidence and abundance of some species are inversely related to the abundance of other species (i.e. incidence assembly rule), and (3) support for assembly rules precludes evidence for nestedness, which refers to a pattern in species composition in which the species present on depauperate islands form regular subsets of those occurring on progressively more diverse islands.
Results  Most species co-occurred with other species at frequencies expected by chance. However, one species ( Sambucus racemosa ) co-occurred with other species less frequently than randomized expectations. The observed incidence and abundance patterns of most species were also consistent with randomized patterns. However, the incidence and abundance of S. racemosa declined with the abundance of other plant species. Weak, variable support was found for nestedness of the total plant community. However, stronger, consistent support was found after removing S. racemosa from the matrix prior to analyses.
Main conclusions  Most species were assembled on islands in a manner consistent with randomized expectations. However, non-random distributional patterns were observed in one species whose distribution was consistent with the hypothesis that competition limits the assembly of island communities.     

Scaling coexistence and assemblage patterns of desert small mammals

Scaling biodiversity patterns has been recognized lately as a very important issue in the search of global processes; however coexistence and assemblage patterns are typically approached at a single spatial scale. Here, we examined coexistence and co-occurrence patterns of desert small mammal communities across different spatial scales in the search of general community patterns. We sampled small mammals in Monte desert (Argentina, South America) for small spatial scales and reviewed published papers from other worldwide deserts for large spatial scale analyses. We used classic community estimators (Shannon, Richness), rank abundance curves and fitting distributions to analyze species coexistence and co-occurrence patterns. Assemblage patterns were analyzed evaluating nestedness across spatial scales and among deserts. Worldwide desert small mammal assemblages are characterized mainly by low species richness and high variation in species composition. The central Monte desert of Argentina showed a consistent assemblage pattern across spatial scales, with a generalist species being the most abundant and widely distributed, accompanied by other subordinate and more narrowly distributed species. All Monte desert communities were significantly nested, with nestedness increasing with scale from patch to regional. Assemblage and coexistence patterns were similar when comparing worldwide deserts despite differences in total richness and faunal singularity. The degree of nestedness varied among worldwide deserts; however all of them showed a consistent nested pattern. Differences in the degree of nestedness could be a result of different regulating factors depending on the desert and scale. These results highlight the importance of including multiscale approaches when dealing with processes that structure desert communities.     

Species co-occurrence, nestedness and guild–environment relationships in stream macroinvertebrates

1. Describing species distribution patterns and the underlying mechanisms is at the heart of ecological research. A number of recent studies have used null model approaches to explore mechanisms behind spatial variation in community structure.
2. However, unexplored questions are the degree to which single guilds of potentially competing stream macroinvertebrate species show: (i) interspecific segregation among-stream sites (i.e. occur together less often than expected by chance), suggesting competitive interactions; (ii) interspecific aggregation (i.e. occur together more often than expected by chance), suggesting similar responses to the environment; (iii) comply with nestedness, suggesting the existence of selective extinctions or colonisations and (iv) show similar environmental relationships.
3. The present analyses showed that guilds of stream macroinvertebrates exhibit non-random co-occurrence patterns that were generally contingent on the weighting of sites by stream size. Despite significant segregation of species, each guild also showed significantly nested patterns. Species richness was correlated with different environmental factors between the guilds, although these correlations were relatively low. By contrast, correlations between the major ordination axes and key environmental variables were slightly stronger in canonical correspondence analysis, and generally the same factors were most strongly correlated with variation in the species composition of each guild.
4. The present findings are the first to show that species within each stream macroinvertebrate guild show significant negative co-occurrence at the among-stream riffle scale. These findings present challenges for future studies that aim to disentangle whether these patterns comply with the habitat checkerboard or the competitive checkerboard explanations.     

Species co-occurrences and neutral models: reassessing J. M. Diamond's assembly rules

The question whether species co-occurrence patterns are non-random has intrigued ecology for more than two decades. Recently Gotelli and McCabe used meta-analysis to show that natural assemblages indeed tend to have non-random species co-occurrence patterns and that these patterns are in line with the predictions of Diamond's assembly rule model. Here I show that neutral ecological drift models are able to generate patterns in line with Diamond's assembly rules and very similar to the empirical results in Gotelli and McCabe. Ecological drift shifted species co-occurrence patterns (measured by C-scores, checkerboard scores and species combination scores) of model species placed into a grid of the 25 cells (sites; metacommunity sizes 5 to 25 species with 100 individuals each) significantly from an initial random pattern towards a pattern predicted by the assembly rule model of Diamond. These findings imply that instead of asking whether natural communities are structured according to some assembly rules we should ask whether these non-random patterns are generated by species interactions or by stochastic drift processes.     

Are ectoparasite communities structured? Species co-occurrence, temporal variation and null models

1. We studied temporal variation in the structure of flea communities on small mammalian hosts from eastern Slovakia using null models. We asked (a) whether flea co-occurrences in infracommunities (in the individual hosts) in different hosts as well as in the component communities (in the host species) demonstrate a non-random pattern; (b) whether this pattern is indicative of either positive or negative flea species interactions; (c) whether this pattern varies temporally; and (d) whether the expression of this pattern is related to population size of either fleas or hosts or both. 2. We constructed a presence/absence matrix of flea species for each temporal sample of a host species and calculated four metrics of co-occurrence, namely the C-score, the number of checkerboard species pairs, the number of species combinations and the variance ratio (V-ratio). Then we compared these metrics with the respective indices calculated for 5000 null matrices that were assembled randomly using two algorithms, namely fixed-fixed (FF) and fixed-equiprobable (FE). 3. Most co-occurrence metrics calculated for real data did not differ significantly from the metrics calculated for simulated matrices using the FF algorithm. However, the indices observed for 42 of 75 presence/absence matrices differed significantly from the null expectations for the FE models. Non-randomness was detected mainly by the C-score and V-ratio metrics. In all cases, the direction of non-randomness was the same, namely the aggregation, not competition, of flea species in host individuals and host species. 4. The inclusion or exclusion of the uninfested hosts in the FE models did not affect the results for individual host species. However, exclusion of the uninfested host species led to the acceptance of the null hypothesis for only six of 13 temporal samples of the component flea communities for which non-randomness was detected when the uninfested hosts were included in the analysis. 5. In most host species, the absolute values of the standardized size effect of both the C-score and V-ratio increased with an increase in host density and a concomitant decrease in flea abundance and prevalence. 6. Results of this study demonstrated that (a) flea assemblages on small mammalian hosts were structured at some times, whereas they appeared to be randomly assembled at other times; (b) whenever non-randomness of flea co-occurrences was detected, it suggested aggregation but never segregation of flea species in host individuals or populations; and (c) the expression of structure in flea assemblages depended on the level of density of both fleas and hosts.     

Patterns of association, nestedness, and species co-occurrence of helminth parasites in the greater kudu, Tragelaphus strepsiceros, in the Kruger National Park, South Africa, and the Etosha National Park, Namibia

The helminth parasites of the greater kudu from the Kruger National Park (KNP), South Africa, and the Etosha National Park (ENP), Namibia, were examined to determine the major patterns of spatial and demographic variation in community structure and to evaluate nonrandomness in parasite community assembly. Nonmetric multidimensional scaling ordination procedures were used to test for differences in parasite community composition between hosts of the 2 parks and between hosts of different demographic groups within KNP. Infracommunities within KNP were also examined for patterns of nonrandomness using 2 null models, i.e., nestedness and species co-occurrence. Infracommunities of KNP and ENP were significantly different from each other, as were infracommunities of different host demographic groups within KNP. Parasite species in the greater kudu from KNP displayed significant levels of nestedness and were found to co-occur less frequently than expected by chance; however, this lack of co-occurrence was significant only when all demographic groups were considered. When restricted to any particular age class, co-occurrence patterns could not be distinguished from random. Overall, these data suggest that biogeography and host demographics are important factors in determining community organization of helminth parasites in the greater kudu.     

Metacommunity patterns of highly diverse stream midges: gradients,chequerboards, and nestedness,or is there only randomness?

Abstract.  1. Several non-random patterns in the distribution of species have been observed, including Clementsian gradients, Gleasonian gradients, nestedness, chequerboards, and evenly spaced gradients. Few studies have examined these patterns simultaneously, although they have often been studied in isolation and contrasted with random distribution of species across sites.
2. This study examined whether assemblages of chironomid midges exhibit any of the idealised distribution patterns as opposed to random distribution of species across sites within the metacommunity context in a boreal drainage system. Analyses were based on stream surveys conducted during three consecutive years. Analytical approaches included ordinations, cluster analysis, null models, and associated randomisation methods.
3. Midge assemblages did not conform to Clementsian gradients, which was evidenced by the absence of clearly definable assemblage types with numerous species exclusive to each assemblage type. Rather, there were signs of continuous Gleasonian variability of assemblage composition, as well as significant nested subset patterns of species distribution.
4. Midge assemblages showed only weak relationships with any of the measured environmental variables, and even these weak environmental relationships varied among years.
5. Midge assemblages did not appear to be structured by competition. This finding was somewhat problematic, however, because the two indices measuring co-occurrence provided rather different signs of distribution patterns. This was probably a consequence of how they actually measure co-occurrence.
6. Although midge assemblages did not show a perfect match with any of the idealised distribution patterns, they nevertheless showed a resemblance to the empirical patterns found previously for several plant and animal groups.     

人工红松林步行虫(Coleoptera:Carabidae)群落物种共存格局动态分析

揭示群落物种共存格局是群落生态学研究的重点内容之一,零模型的应用极大的促进了群落物种共存格局及其调控机制的进展,然而针对地下生物群落共存格局动态特征的研究并不多见。在帽儿山森林生态站的人工红松林内,通过5次调查取样基于零模型模拟分析小尺度空间(20 m×20 m)步行虫群落物种共存格局的动态特征。结果表明:(1)共捕获步行虫20种,2278只个体,其中Carabus billergi maoershanensis为所有调查季节数量最具优势且分布最广泛的物种,步行虫群落结构具明显的时间变异性;(2)2013年6月步行虫群落为集群性共存格局,而2014年8、10月为竞争性共存格局,基于目前的零模型指标和法则难以准确揭示其他月份是集群性还是竞争性的共存格局,但所有季节的群落均表现为明显的非随机性共存格局,这些共存格局的发现并不完全支持Diamond的群落构建机制理论;(3)所有调查季节均发现很少的显著物种对,基于更严格的检验表明群落中集群性物种对多于隔离性物种对,那些表现为显著的非随机性共存关系的物种对往往是群落内数量较大且分布广泛的优势和常见物种。表明非随机性共存格局可能是帽儿山人工红松林小尺度空间步行虫群落的常见格局,这种非随机性格局具一定的短期动态稳定性,但不同季节这种非随机性共存格局类型表现不同,群落内这些较少的显著物种对可能对群落物种共存格局具有一定的贡献。     

Co-occurrence of ectoparasites on rodent hosts: null model analyses of data from three continents

We studied patterns of species co-occurrence in communities of ectoparasitic arthropods (ixodid ticks, mesostigmate mites and fleas) harboured by rodent hosts from South Africa ( Rhabdomys pumilio ), South America ( Scapteromys aquaticus and Oxymycterus rufus ) and west Siberia ( Apodemus agrarius , Microtus gregalis , Microtus oeconomus and Myodes rutilus ) using null models. We compared frequencies of co-occurrences of parasite species or higher taxa across host individuals with those expected by chance. When non-randomness of parasite co-occurrences was detected, positive but not negative co-occurrences of parasite species or higher taxa prevailed (except for a single sample of mesostigmate mites from O. rufus ). Frequency of detection of non-randomness of parasite co-occurrences differed among parasite taxa, being higher in fleas and lower in mites and ticks. This frequency differed also among host species independent of parasite taxon, being highest in Microtus species and lowest in O. rufus and S. aquaticus . We concluded that the pattern of species co-occurrence in ectoparasite communities on rodent hosts is predominantly positive, depends on life history of parasites and may be affected to a great extent by life history of a host.     

Factors contributing to non-randomness in species Co-occurrences on Islands

Summary There has been dispute whether patterns of species co-occurrence on islands are largely random. We present a new method for testing this question; this method lets one not only examine whether a whole fauna is non-randomly structured, but also identify in which direction and by how much each particular species combination deviates from expectations based on randomness. Application of this method to the whole Bismarck and New Hebridean avifaunas, and to two particular guilds, shows that some pairs of species have more exclusive distributions than expected for random placement of species, because of competition, differing distributional strategies, or different geographical orgins. Other pairs of species have more coincident distributions than expected, because of shared habitat, single-island endemism, shared distributional strategies, or shared geographical origins. Much of the information about non-random co-occurrence is contained in the incidence graphs for occurrence of individual species. Finally, our present understanding of assembly rules is summarized.     

Nestedness, anti-nestedness, and the relationship between prevalence and intensity in ectoparasite assemblages of marine fish: a spatial model of species coexistence

Nested species subset patterns consist in a hierarchical structure of species composition in related assemblages, with the species found in depauperate assemblages representing non-random subsets of progressively richer ones. This pattern has been found at the infracommunity level in about a third of the fish ectoparasite assemblages studied to date. Here we present evidence for another non-random structural pattern in assemblages of fish ectoparasites, anti-nestedness, which corresponds to situations in which parasite species are always absent from infracommunities richer than the most depauperate one in which they occur. We show that this pattern is exactly as common as nestedness, and that anti-nested assemblages are characterised by significantly lower prevalence and mean intensities of parasites than nested assemblages. In addition, we found a positive relationship between the prevalence and the mean intensity of parasites across the different assemblages. We propose a link between the nestedness/anti-nestedness continuum and the prevalence-intensity relationship that may involve colonisation-extinction processes. The results presented here suggest that, although nestedness may not be common in parasite communities, other departures from random species assembly are possible, and that some form of structure may be present in many communities. The continuum between nestedness and anti-nestedness also has implications for recent models of species coexistence in communities.     

Biogeography of Aphodiinae dung beetles based on the regional composition and distribution patterns of genera

Aim  To examine current biogeographical patterns of Aphodiinae dung beetles (Coleoptera: Scarabaeoidea: Scarabaeidae) in order to reveal relationships among regions and their potential impact on the diversification of this group.
Location  Worldwide.
Methods  Information about all Aphodiinae genera was obtained from the literature. An occurrence matrix was built for the six worldwide biogeographical regions, and their faunas were characterized through simple statistics. Regional variations and similarities were further explored using co-occurrence and nestedness analyses, sequential agglomerative, hierarchical and nested clustering (SAHN), and a parsimony analysis of endemicity (PAE). Mantel tests were also employed to assess the relationships between several characteristics of the regions and their faunas.
Results  The Palaearctic and Palaeotropical regions showed the highest total numbers of Aphodiinae genera and the greatest generic endemism. Both these regions and the Oriental also showed higher numbers of genera than would be expected according to their size. Co-occurrence and nestedness analyses confirmed the non-randomness of the distribution of genera. Clustering and PAE showed that the Palaearctic and Oriental regions are the most similar, followed by the Palaeotropical region. Regional dissimilarity in genera composition was related to biological and historical traits, but not to ecoregions.
Main conclusions  A structured geographical pattern for Aphodiinae was confirmed. Land continuity and proximity in the long term could have played a unifying role in regional faunas. We suggest that the different biogeographical regions have acted as either macroevolutionary sources (basically the Palaearctic and the Palaeotropical regions) or sink regions, according to their role as diversification centres. We review the processes and events that could account for current patterns of Aphodiinae diversity.     

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.     

Simultaneous modelling of distributional patterns in a guild of eastern-Australian cicadas

Summary The distributions, with respect to habitat structure, of nine species of eastern-Australian cicadas have been shown to be non-random. The most striking consequence of this non-randomness is a marked inverse relationship between habitat breadth and habitat position (terms defined in text). Eight basic models and 12 derived models were used in conjunction with a canonical space to try to account for the ways in which the species of cicadas were distributed with respect to habitat. Several models produced results that were in reasonable agreement with the observed data. The most parsimonious of these corresponds to analytical results of other workers, such as Diamond's (1975) incidence curves, occurrence sequences (Schoener and Schoener 1983), and probability functions (Adler and Wilson 1985). The distributions of cicadas can be modelled by assuming that the species occupy sites independently of one another. These species of cicadas are unlikely to engage in interspecific competition, which is consistent with independence of distributions.     

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.     

From sampling stations to archipelagos: investigating aspects of the assemblage of insular biota

Aim To investigate the formation of nestedness and species co‐occurrence patterns at the local (sampling station), the intermediate (island group), and the archipelago scale. Location The study used data on the distribution of terrestrial isopods on 20 islands of the central Aegean (Greece). These islands are assigned to two distinct subgroups (Kyklades and Eastern islands). Methods The Nestedness Temperature Calculator was used to obtain nestedness values and maximally nested matrices, the EcoSim7 software and a modified version of Sanderson (2000 ) method were used for the analysis of species co‐occurrences. Idiosyncratic temperatures of species and the order of species placement in the maximally nested matrices were used for further comparisons among spatial scales. The relationships of nestedness values with beta‐diversity, habitat diversity and a number of ecological factors recorded for each sampling station were also investigated. Results Significant nestedness was found at all spatial scales. Levels of nestedness were not related to beta‐diversity or habitat diversity. Nestedness values were similar among spatial scales, but they were affected by matrix size. The species that contributed most to the nested patterns within single islands were not the same as those that produce nestedness at the archipelago scale. There was significant variation in the frequency of species occurrence among islands and among spatial scales. There was no direct effect of ecological factors on the shaping of patterns of nestedness within individual islands, but habitat heterogeneity was crucial for the existence of such patterns. Positive associations among species prevailed at all scales when species per station were considered, while negative associations prevailed in the species per island matrices. All associations resulted from the habitat structure of sampling stations and from particularities of geographical distributions. Conclusions There was no clear‐cut distinction between nestedness patterns among spatial scales, even though different species, and partially different factors, contributed to the formation of these patterns in each case. There was a core of species that contributed to the formation of nested patterns at all spatial scales, while the patterns of species associations suggested that biotic interactions are not an important causal factor. The results of this study suggest that locally rare species cannot be widespread at a higher spatial scale, while locally common species can have a restricted distribution.     

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.     

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.     

Climate drives temporal replacement and nested‐resultant richness patterns of Scottish coastal vegetation

Beta diversity quantifies spatial and/or temporal variation in species composition. It is comprised of two distinct components, species replacement and nestedness, which derive from opposing ecological processes. Using Scotland as a case study and a β‐diversity partitioning framework, we investigate temporal replacement and nestedness patterns of coastal grassland species over a 34‐yr time period. We aim to 1) understand the influence of two potentially pivotal processes (climate and land‐use changes) on landscape‐scale (5 × 5 km) temporal replacement and nestedness patterns, and 2) investigate whether patterns from one β‐diversity component can mask observable patterns in the other. We summarised key aspects of climate driven macro‐ecological variation as measures of variance, long‐term trends, between‐year similarity and extremes, for three important climatic predictors (minimum temperature, water‐balance and growing degree‐days). Shifts in landscape‐scale heterogeneity, a proxy of land‐use change, was summarised as a spatial multiple‐site dissimilarity measure. Together, these climatic and spatial predictors were used in a multi‐model inference framework to gauge the relative contribution of each on temporal replacement and nestedness patterns. Temporal β‐diversity patterns were reasonably well explained by climate change but weakly explained by changes in landscape‐scale heterogeneity. Climate was shown to have a greater influence on temporal nestedness than replacement patterns over our study period, linking nestedness patterns, as a result of imbalanced gains and losses, to climatic warming and extremes respectively. Important climatic predictors (i.e. growing degree‐days) of temporal β‐diversity were also identified, and contrasting patterns between the two β‐diversity components revealed. Results suggest climate influences plant species recruitment and establishment processes of Scotland's coastal grasslands, and while species extinctions take time, they are likely to be facilitated by climatic perturbations. Our findings also highlight the importance of distinguishing between different components of β‐diversity, disentangling contrasting patterns than can mask one another.