Additive partitioning of butterfly diversity in a fragmented landscape: importance of scale and implications for conservation

Aim Most of the Atlantic Forest in Brazil occurs in fragments of various sizes. Previous studies indicate that forest fragmentation affects fruit‐feeding butterflies. Conservation strategies that seek to preserve organisms that are distributed in high‐fragmented biomes need to understand the spatial distribution of these organisms across the landscape. In view of the importance of understanding the fauna of these forest remnants, the objective of the present work is to investigate the extent to which the diversity of this group varies across spatial scales ranging from within‐forest patches to between landscapes. Location South America, south‐eastern Brazil, São Paulo State. Methods We used bait traps to sample fruit feeding butterflies at 50 points in 10 fragments in two different landscapes during a period of 12 months. Total species richness and Shannon index were partitioned additively in diversity at trap level, and beta diversity was calculated among traps, among forest patches, and between landscapes. We used permutation tests to compare these values to the expected ones under the null hypothesis that beta diversity is only a random sampling effect. Results There was significant beta diversity at the smallest scale examined; however, the significance at higher scales depends on the diversity measurement used. Beta diversity with Shannon index was smaller than expected by chance among fragments, whereas species richness was not. Among landscapes, only beta diversity in richness was higher than expected by chance. Main conclusions The results observed occur because there is great variability in species composition among forest patches in the same landscape, changing this diversity even though the communities are formed from the same pool of species. At the largest scale evaluated (between landscapes), these pattern changes and differences in beta diversity in richness were detectable. This difference is probably caused by the presence of rare species. Thus, a conservation strategy that seeks to preserve as many species as possible per unit of area in high‐fragmented biomes should give priority to protecting fragments in different landscapes, rather than more fragments in the same landscape.     

Partitioning multiple-site tree-like beta diversity into turnover and nestedness components without pairwise comparisons

In this letter, I extend a species-level partitioning framework for a multiple-site dissimilarity index established by Ricotta and Pavoine (2015) to account for tree-like information (i.e., evolutionary history, taxonomic classification or functional divergence of species). This novel framework can be applied to evaluate the relative contribution of turnover versus nestedness to total multiple-site tree-like beta diversity in empirical settings. The feature of the framework is to account for expectedly and unexpectedly absence of species in the sites without pairwise comparisons between sites. Simple examples with step-by-step calculation details and corresponding R computing code are provided to better understand and apply the proposed framework.     

Inventory, differentiation, and proportional diversity: a consistent terminology for quantifying species diversity

Almost half a century after Whittaker (Ecol Monogr 30:279–338, 1960) proposed his influential diversity concept, it is time for a critical reappraisal. Although the terms alpha, beta and gamma diversity introduced by Whittaker have become general textbook knowledge, the concept suffers from several drawbacks. First, alpha and gamma diversity share the same characteristics and are differentiated only by the scale at which they are applied. However, as scale is relative––depending on the organism(s) or ecosystems investigated––this is not a meaningful ecological criterion. Alpha and gamma diversity can instead be grouped together under the term “inventory diversity.” Out of the three levels proposed by Whittaker, beta diversity is the one which receives the most contradictory comments regarding its usefulness (“key concept” vs. “abstruse concept”). Obviously beta diversity means different things to different people. Apart from the large variety of methods used to investigate it, the main reason for this may be different underlying data characteristics. A literature review reveals that the multitude of measures used to assess beta diversity can be sorted into two conceptually different groups. The first group directly takes species distinction into account and compares the similarity of sites (similarity indices, slope of the distance decay relationship, length of the ordination axis, and sum of squares of a species matrix). The second group relates species richness (or other summary diversity measures) of two (or more) different scales to each other (additive and multiplicative partitioning). Due to that important distinction, we suggest that beta diversity should be split into two levels, “differentiation diversity” (first group) and “proportional diversity” (second group). Thus, we propose to use the terms “inventory diversity” for within-sample diversity, “differentiation diversity” for compositional similarity between samples, and “proportional diversity” for the comparison of inventory diversity across spatial and temporal scales. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effect of landscape structure on the spatial distribution of Mediterranean dung beetle diversity

Aim  Landscape structure influences the distribution of animals, altering their movements and their ability to reach habitat patches. We analysed the spatial patterns of dung beetle species diversity in three differently structured natural landscapes in a Mediterranean protected area in the centre of the Iberian Peninsula.
Location  Cabañeros National Park, Central Spain.
Methods  Diversity components within (α) and among (β) the three main vegetation types in the reserve were compared by using a hierarchical nested design. These were forests, scrublands and grasslands embedded in three different landscapes, where each was the most dominant and structurally connected habitat.
Results  Species richness of grassland habitat did not vary across landscapes, but forest habitat showed lower species richness in the grassland-dominated landscape. Scrubland was the least species-rich habitat, but here again there was no significant variation across landscapes. However, in all cases, there was a significant influence of habitat context (configuration of habitat patches within landscape matrix) on similarity of species composition. These tended to be more similar to the dominant landscape matrix where they were embedded, rather than to the same habitat type in other landscapes. Additive partitioning of diversity showed higher than expected values of β in all landscapes, which indicated a structured response. Highest values of β in the grassland-dominated landscape suggest that this was the least connected landscape for dung beetles.
Main conclusions  Our results suggest that in homogeneous conditions of climate and trophic resources, landscape structure may well be more important than habitat type as a determinant of dung beetle distribution in the Mediterranean.     

Fine‐scale Beta‐diversity Patterns Across Multiple Arthropod Taxa Over a Neotropical Latitudinal Gradient

Documenting how diversity patterns vary at fine‐ and broad scales may help answer many questions in theoretical and applied ecology. However, studies tend to compare diversity patterns at the same scale and within the same taxonomic group, which limits the applicability and generality of the results. Here, we have investigated whether vegetation‐dwelling arthropods from different trophic ranks and with distinct life histories (i.e., ants, caterpillars, cockroaches, and spiders) have different beta‐diversity patterns at multiple scales. Specifically, we compared their beta diversity across architecturally distinct plant species (fine‐scale process) and a latitudinal gradient of sites (broad‐scale process) along 2040 km of coastal restinga vegetation in the Neotropics. Over 50 percent of the compositional changes (β‐diversity) in ants, caterpillars, and spiders and 41 percent of those in cockroaches were explained by plant identity within each site. Even groups that do not feed on plant tissues, such as omnivores and predators, were strongly affected by plant identity. Fine‐scale variation was more important than large‐scale processes for all studied groups. Performing a cross‐scale comparison of diversity patterns of groups with distinct life histories helps elucidate how processes that act at regional scales, such as dispersal, interact with local processes to assemble arthropod communities.     

Bryophyte and lichen diversity: A comparative study

Abstract We describe the regional species richness, variation in species richness and species turnover of bryophytes and lichens from 36 sites in lowland forests of southeastern Australia. The analyses subdivided the two major taxa into their constituent sub-groups: mosses, liverworts, and crustose, fruticose and foliose lichens. They also explored correlations between selected environmental variables and patterns of diversity. On a regional scale, there were 77 species of bryophytes and 69 species of lichens, giving a total of approximately one-third of the total number of vascular plant species in the region. Mean species richness was higher for lichens than bryophytes. Also, the two taxa were negatively correlated because lichens favoured dry sites and bryophytes favoured moist ones. Species turnover was greater for bryophytes than lichens, largely due to the distribution of liverwort species. Foliose lichens showed higher levels of turnover than crustose lichens. Multiple regression and canonical correspondence analysis showed that both taxa and all sub-groups responded to the same three variables: vascular plant cover, time since last fire and topographic position. Other variables, including time since logging and intensity of logging, explained little variation in bryophyte or lichen diversity. The data suggest that the strategies for the conservation of bryophyte and lichen biodiversity will be different, to reflect the different patterns of species richness and species turnover.     

Management defines species turnover of bees and flowering plants in vineyards

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Partitioning plant spectral diversity into alpha and beta components

Plant spectral diversity – how plants differentially interact with solar radiation – is an integrator of plant chemical, structural, and taxonomic diversity that can be remotely sensed. We propose to measure spectral diversity as spectral variance, which allows the partitioning of the spectral diversity of a region, called spectral gamma (γ) diversity, into additive alpha (α; within communities) and beta (β; among communities) components. Our method calculates the contributions of individual bands or spectral features to spectral γ‐, β‐, and α‐diversity, as well as the contributions of individual plant communities to spectral diversity. We present two case studies illustrating how our approach can identify 'hotspots’ of spectral α‐diversity within a region, and discover spectrally unique areas that contribute strongly to β‐diversity. Partitioning spectral diversity and mapping its spatial components has many applications for conservation since high local diversity and distinctiveness in composition are two key criteria used to determine the ecological value of ecosystems.     

Headwaters are critical reservoirs of microbial diversity for fluvial networks

Streams and rivers form conspicuous networks on the Earth and are among nature''s most effective integrators. Their dendritic structure reaches into the terrestrial landscape and accumulates water and sediment en route from abundant headwater streams to a single river mouth. The prevailing view over the last decades has been that biological diversity also accumulates downstream. Here, we show that this pattern does not hold for fluvial biofilms, which are the dominant mode of microbial life in streams and rivers and which fulfil critical ecosystem functions therein. Using 454 pyrosequencing on benthic biofilms from 114 streams, we found that microbial diversity decreased from headwaters downstream and especially at confluences. We suggest that the local environment and biotic interactions may modify the influence of metacommunity connectivity on local biofilm biodiversity throughout the network. In addition, there was a high degree of variability in species composition among headwater streams that could not be explained by geographical distance between catchments. This suggests that the dendritic nature of fluvial networks constrains the distributional patterns of microbial diversity similar to that of animals. Our observations highlight the contributions that headwaters make in the maintenance of microbial biodiversity in fluvial networks.     

From plots to islands: species diversity at different scales

Aim To investigate how plant diversity of whole islands (‘gamma’) is related to alpha and beta diversity patterns among sampling plots within each island, thus exploring aspects of diversity patterns across scales. Location Nineteen islands of the Aegean Sea, Greece. Methods Plant species were recorded at both the whole‐island scale and in small 100 m² plots on each island. Mean plot species richness was considered as a measure of alpha diversity, and six indices of the ‘variation’‐type beta diversity were also applied. In addition, we partitioned beta diversity into a ‘nestedness’ and a ‘replacement’ component, using the total species richness recorded in all plots of each island as a measure of ‘gamma’ diversity. We also applied 10 species–area models to predict the total observed richness of each island from accumulated plot species richness. Results Mean alpha diversity was not significantly correlated with the overall island species richness or island area. The range of plot species richness for each island was significantly correlated with both overall species richness and area. Alpha diversity was not correlated with most indices of beta diversity. The majority of beta diversity indices were correlated with whole‐island species richness, and this was also true for the ‘replacement’ component of beta diversity. The rational function model provided the best prediction of observed island species richness, with Monod’s and the exponential models following closely. Inaccuracy of predictions was positively correlated with the number of plots and with most indices of beta diversity. Main conclusions Diversity at the broader scale (whole islands) is shaped mainly by variation among small local samples (beta diversity), while local alpha diversity is not a good predictor of species diversity at broader scales. In this system, all results support the crucial role of habitat diversity in determining the species–area relationship.     

Spatial scale and the diversity of macroinvertebrates in a Neotropical catchment

1. Lotic ecosystems can be studied on several spatial scales, and usually show high heterogeneity at all of them in terms of biological and environmental characteristics. Understanding and predicting the taxonomic composition of biological communities is challenging and compounded by the problem of scale. Additive diversity partitioning is a tool that can show the diversity that occurs at different scales. 2. We evaluated the spatial distribution of benthic macroinvertebrates in a tropical headwater catchment (S.E. Brazil) during the dry season and compared alpha and beta diversities at the scales of stream segments, reaches, riffles and microhabitats (substratum types: gravels, stones and leaf litter). We used family richness as our estimate of diversity. Sampling was hierarchical, and included three stream segments, two stream reaches per segment, three riffles per reach, three microhabitats per riffle and three Surber sample units per microhabitat. 3. Classification analysis of the 53 families found revealed groups formed in terms of stream segment and microhabitat, but not in terms of stream reaches and riffles. Separate partition analyses for each microhabitat showed that litter supported lower alpha diversity (28%) than did stones (36%) or gravel (42%). In all cases, alpha diversity at the microhabitat scale was lower than expected under a null model that assumed no aggregation of the fauna. 4. Beta diversity among patches of the microhabitats in riffles depended on substratum type. It was lower than expected in litter, similar in stone and higher in gravel. Beta diversities among riffles and among reaches were as expected under the null model. On the other hand, beta diversity observed was higher than expected at the scale of stream segments for all microhabitat types. 5. We conclude that efficient diversity inventories should concentrate sampling in different microhabitats and stream sites. In the present study, sampling restricted to stream segments and substratum types (i.e. excluding riffles and stream reaches) would produce around 75% of all observed families using 17% of the sampling effort employed. This finding indicates that intensive sampling (many riffles and reaches) in few stream segments does not result in efficient assessment of diversity in a region.     

Additive diversity partitioning as a guide to regional montane reserve design in Asia: an example from Yunnan Province,China

Aim Data on spatial and temporal turnover in species composition within a region is essential to design regional protected areas. Montane systems are often recognized as biodiversity hotspots. The primary objective of this study is to identify patterns of montane bird diversity across multiple spatial and temporal scales using an additive diversity partitioning framework. Location The Ailao Mountains, central Yunnan Province, China. Methods We used point counts to sample bird communities in four elevational zones, on eastern and western slopes, during both the breeding and the non‐breeding seasons. Diversity (richness and Shannon) was partitioned across space (points, elevational zones and slopes) and time (seasons). We used permutation tests to compare observed values to values expected by random chance. A complementary cluster analysis was also used to evaluate beta diversity. Results Overall, the gamma diversity was attributed to significantly higher beta diversity (relative to that of randomization tests) among elevational zones and, to a lesser extent, between slopes. For Shannon–Wiener Index, beta diversity between seasons was significantly higher than expected and had a similar contribution to the gamma diversity as with beta diversity between slopes. Hierarchical cluster analysis supported the findings for Shannon–Wiener Index. The contribution of beta diversity among points to gamma diversity within each elevational zone generally lessened with increasing elevation. Main conclusions Our results show significantly high levels of beta diversity among elevational zones and between slopes, as well as between seasons for Shannon diversity, in a small area of the Ailao Mountain range. Thus, a regional montane reserve system should cover the entire elevational gradient and multiple slopes, rather than only the montane crest. Furthermore, higher pattern diversity in lower elevational zones suggests that larger areas should be preserved at lower elevational zones. Finally, the design of regional reserve systems require more studies conducted at multiple seasons at a regional scale.     

BIODIVERSITY RESEARCH: Conserving macroinvertebrate diversity in headwater streams: the importance of knowing the relative contributions of α and β diversity

Aim We investigated partitioning of aquatic macroinvertebrate diversity in eight headwater streams to determine the relative contributions of α and β diversity to γ diversity, and the scale dependence of α and β components. Location Great Dividing Range, Victoria, Australia. Methods We used the method of Jost (Ecology, 2007, 88, 2427–2439) to partition γ diversity into its α and β components. We undertook the analyses at both reach and catchment scales to explore whether inferences depended on scale of observation. Results We hypothesized that β diversity would make a large contribution to the γ diversity of macroinvertebrates in our dendritic riverine landscape, particularly at the larger spatial scale (among catchments) because of limited dispersal among sites and especially among catchments. However, reaches each had relatively high taxon richness and high α diversity, while β diversity made only a small contribution to γ diversity at both the reach and catchment scales. Main conclusions Dendritic riverine landscapes have been thought to generate high β diversity as a consequence of limited dispersal and high heterogeneity among individual streams, but this may not hold for all headwater stream systems. Here, α diversity was high and β diversity low, with individual headwater stream reaches each containing a large portion of γ diversity. Thus, each stream could be considered to have low irreplaceability since losing the option to use one of these sites in a representative reserve network does not greatly diminish the options available for completing the reserve network. Where limited information on individual taxonomic distributions is available, or time and money for modelling approaches are limited, diversity partitioning may provide a useful ‘first‐cut’ for obtaining information about the irreplaceability of individual streams or subcatchments when establishing representative freshwater reserves.