Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

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.     

Spatial and temporal organisation of the pre-dispersal seed predator guild in a perennial legume, Vicia tenuifolia

Abstract.  1. The variability of species combinations and membership assembly patterns in an insect pre-dispersal seed predator guild were studied at various spatial and temporal resolutions using presence/absence and abundance data in null models. The guild consists of seven beetle species (four bruchids and three curculionids) and one moth species that live on a perennial vetch, Vicia tenuifolia Roth (Leguminosae).
2. The seed predator guild varied considerably in the number of members and species combinations in space and time, and, contrary to expectations, there was no evidence of interspecific competition among guild members, supporting the view that other processes, including chance events, could contribute to guild formation.
3. It is concluded that, apart from the possibility of stochastic co-occurrence, it is the narrow host specialisation that constrains seed predator members to participate in the guild, and small differences in habitat preference can also lead to spatial and temporal variation.     

How does habitat diversity affect the species–area relationship?

Aim   To examine the way in which 'area' and 'habitat diversity' interact in shaping species richness and to find a simple and valid way to express this interaction.
Location   The Natura 2000 network of terrestrial protected areas in Greece, covering approximately 16% of the national territory.
Methods   We used the Natura 2000 framework, which provides a classification scheme for natural habitat types, to quantify habitat heterogeneity. We analysed data for the plant species composition in 16,143 quadrats in which 5044 species and subspecies of higher plants were recorded. We built a simple mathematical model that incorporates the effect of habitat diversity on the species–area relationship (SAR).
Results   Our analysis showed that habitat diversity was correlated with area. However, keeping habitat diversity constant, species richness was related to area; while keeping area constant, species richness was related to habitat diversity. Comparing the SAR of the 237 sites we found that the slope of the species–area curve was related to habitat diversity.
Main conclusions   Discussion of the causes of the SAR has often focused on the primacy of area per se versus habitat heterogeneity, even though the two mechanisms are not mutually exclusive and should be considered jointly. We find that increasing habitat diversity affects the SAR in different ways, but the dominant effect is to increase the slope of the SAR. While a full model fit typically includes a variety of terms involving both area and habitat richness, we find that the effect of habitat diversity can be reduced to a linear perturbation of the slope of the species accumulation curve.     

Null model analyses of temporal patterns of bird assemblages and their foraging guilds revealed the predominance of positive and random associations

Patterns of species associations have been commonly used to infer interactions among species. If species positively co‐occur, they may form predominantly neutral assemblages, and such patterns suggest a relatively weak role for compensatory dynamics. The main objective of this study was to test this prediction on temporal samples of bird assemblages (n = 19, 10–57 years) by the presence/absence and quantitative null models on assemblage and guild levels. These null model outcomes were further analyzed to evaluate the effects of various data set characteristics on the outcomes of the null models. The analysis of two binary null models in combination with three association indices revealed 20% with significant aggregations, 61% with random associations, and only 19% with significant segregations (n = 95 simulations). The results of the quantitative null model simulations detected more none‐random associations: 61% aggregations, 6% random associations, and 33% segregations (n = 114 simulations). Similarly, quantitative analyses on guild levels showed 58% aggregations, 20% segregations, and 22% random associations (n = 450 simulations). Bayesian GLMs detected that the outcomes of the binary and quantitative null models applied to the assemblage analyses were significantly related to census plot size, whereas the outcomes of the quantitative analyses were also related to the mean population densities of species in the data matrices. In guild‐level analyses, only 9% of the GLMs showed a significant influence of matrix properties (plot size, matrix size, species richness, and mean species population densities) on the null model outcomes. The results did not show the prevalence of negative associations that would have supported compensatory dynamics. Instead, we assume that a similar response of the majority of species to climate‐driven and stochastic factors may be responsible for the revealed predominance of positive associations.     

Non-interactive fish communities in the coastal streams of North-western France

1. Spatial patterns of freshwater fish species at regional and local scales were investigated to explore the possible role of interspecific interactions in influencing distribution and abundance within communities occupying coastal streams of North-Western France.
2. Nine sites from nine streams situated in the same biogeographical region were sampled annually over the 6-year period from 1990 to 1995.
3. Similar habitats (sites) with richer regional colonization pools exhibited proportionally richer local communities in terms of number of species, total density and total biomass of individuals. Furthermore, no negative relationships were found between density and biomass of each of the most common species and local species richness.
4. Results of dynamic regression models (applied to the above-mentioned species) suggest an absence of strong competition between all pairs of species.
5. The evidence on lack of density compensation for species-poor communities and absence of perceptible interspecific competition between species suggest that the communities studied are non-interactive.
6. Two main explanations can be advanced. First, the local abundance of species in the communities studied could be determined through differential responses to unpredictable environmental changes, rather than through biological interactions. Second, as a result of historical events, the communities studied are reduced in congeneric species which can limit, in turn, the influence of interspecific competition in structuring these communities.
7. These results underline the strong influence of regional processes in shaping local riverine fish communities and minimize the possible influence of species interactions in governing these communities.     

Random sampling, abundance–extinction dynamics and niche-filtering immigration constraints explain the generation of species richness gradients

Aim  The paradigm that species' patterns of distribution, abundance and coexistence are the result of adaptations of the species to their niches has recently been challenged by evidence that similar patterns may be generated by simple random processes. We argue here that a better understanding of macroecological patterns requires an integration of both ecological and neutral stochastic approaches. We demonstrate the utility of such an integrative approach by testing the sampling hypothesis in a species–energy relationship of forest bird species.
Location  A Mediterranean biome in Catalonia, Spain.
Methods  To test the sampling hypothesis we designed a metacommunity model that reproduces the stochastic sampling from a regional pool to predict local species richness variation. Four conceptually different sampling procedures were evaluated.
Results  We showed that stochastic sampling processes predicted a substantial part (over 40%) of the observed variation in species richness, but left considerable variation unexplained. This remaining variation in species richness may be better understood as the result of alternative ecological processes. First, the sampling model explained more variation in species richness when the probability that a species colonises a new locality was assumed to increase with its niche width, suggesting that ecological differences between species matter when it comes to explaining macroecological patterns. Second, extinction risk was significantly lower for species inhabiting high-energy regions, suggesting that abundance–extinction processes play a significant role in shaping species richness patterns.
Main conclusions  We conclude that species–energy relationships may not simply be understood as a result of either ecological or random sampling processes, but more likely as a combination of both.     

Latitudinal variation of diversity in European freshwater animals is not concordant across habitat types

Aim   We analysed the variation of species richness in the European freshwater fauna across latitude. In particular, we compared latitudinal patterns in species richness and β-diversity among species adapted to different habitat types.
Location   Europe.
Methods   We compiled data on occurrence for 14,020 animal species across 25 pre-defined biogeographical regions of European freshwaters from the Limnofauna Europaea . Furthermore, we extracted information on the habitat preferences of species. We assigned species to three habitat types: species adapted to groundwater, lotic (running water) and lentic (standing water) habitats. We analysed latitudinal patterns of species richness, the proportion of lentic species and β-diversity.
Results   Only lentic species showed a significant species–area relationship. We found a monotonic decline of species richness with latitude for groundwater and lotic habitats, but a hump-shaped relationship for lentic habitats. The proportion of lentic species increased from southern to northern latitudes. β-Diversity declined from groundwater to lentic habitats and from southern to northern latitudes.
Main conclusions   The differences in the latitudinal variation of species richness among species adapted to different habitat types are in part due to differences in the propensity for dispersal. Since lentic habitats are less persistent than lotic or groundwater habitats, lentic species evolved more efficient strategies for dispersal. The dispersal propensity of lentic species facilitated the recolonization of central Europe after the last glaciation. Overall, we stress the importance of considering the history of regions and lineages as well as the ecological traits of species for understanding patterns of biodiversity.     

Modelling patterns of coexistence of three congeneric headwater fishes

1. Mechanisms driving patterns of occurrence and co-occurrence among North American freshwater fishes are poorly understood. In particular, the influence of biotic interactions on coexistence among stream reaches and their effects on regional species distribution patterns is not well understood for congeneric headwater fishes.
2. Occupancy models provide a useful framework for examining patterns of co-occurrence while also accounting for imperfect detection. Occupancy models may be extended to test for evidence that a dominant species influences the occurrence of a subordinate species and thus evaluate support for the hypothesis that species interactions drive patterns of coexistence.
3. We examined patterns of occurrence and co-occurrence at the stream-reach scale among three species of darters (Percidae: Etheostomatinae) that occupy headwater streams within a Gulf Coastal Plain drainage in the south-eastern U.S.A. We assessed species occurrences at 97 sites in first- to third-order streams on one occasion each and used data from four sub-reaches sampled with equal effort at each site to estimate species-specific detection probabilities. Following sampling, a suite of habitat variables was collected at three equidistant points along each of the three transects established within a sub-reach. Coarse (stream-segment, catchment, network) scale variables were also incorporated using geospatial data. Single-species and two-species occupancy models were used to examine patterns of occupancy and coexistence.
4. The occupancy of each species was influenced by distinct habitat variables. Goldstripe darters (Etheostoma parvipinne) were constrained by a stream size gradient, groundwater input appeared to influence the occurrence of Yazoo darters (Etheostoma raneyi), and local habitat heterogeneity (e.g. variation in depth and current velocity) appeared to influence the occupancy of redspot darters (Etheostoma artesiae).
5. We found no evidence that the presence of one species influenced the occurrence of another within a stream-reach based on two-species occupancy models. Rather, species co-occurrences were best explained as independent occurrences within a stream-reach according to species-specific habitat associations.
6. Occupancy modelling may provide a suitable framework for evaluating the influence of biotic interactions among congeneric stream fishes along species-specific habitat gradients at the stream reach scale. Our study offers insight into how habitat variation can influence coexistence of potential competitors across a large river system.

     

The small‐island effect: fact or artefact?

Positive relationships between species richness and sampling area are perhaps the most pervasive patterns in nature. However, the shape of species–area relationships is often highly variable, for reasons that are poorly understood. One such source of variability is the "small-island effect", which refers to a decrease in the capacity of sampling area to predict species richness on small islands. Small-island effects have been attributed to a variety of processes, including spatial subsidies, habitat characteristics and ocean-born disturbances. Here, we show that small-island effects can be generated by logarithmic data transformations, which are commonly applied to both axes of species–area relationships. To overcome this problem, we derive several null models to test for non-random variability in the capacity of island area to predict species richness and apply them to data sets on island plant communities in Canada and New Zealand. Both archipelagos showed evidence for small-island effects using traditional breakpoint regression techniques on log-log axes. However, null model analyses revealed different results. The capacity of sampling area to predict species richness in the Canadian archipelago was actually lowest at intermediate island size classes. In the New Zealand archipelago, island area was similarly capable of predicting species richness across the full range of island sizes, indicating the small-island effect detected by breakpoint regression is an artifact of logarithm data transformation. Overall results show that commonly used regression techniques can generate spurious small-island effects and that alternative analytic procedures are needed to detect non-random patterns in species richness on small islands.     

Habitat diversity, ecological requirements of species and the Small Island Effect

Aim  To explore the causal factors leading to a significant Small Island Effect (SIE), that is, the absence of the commonly found species–area relationships below an island size, on the terrestrial isopod communities from a large number of islands.
Location  Ninety islands of the Aegean Sea (Greece).
Methods  The detection of a significant SIE is assessed through the application of all three methods available in the literature. Species are divided into generalists and specialists. We tested if the minimum area and the area range of each species' occurrences differ between generalists and specialists. Next, we searched for differences in the ratios of specialists to generalists above and below the SIE threshold, and tested their cumulative ratios when islands are arranged according to increasing area, altitude or habitat diversity in order to identify the threshold where they become statistically indistinguishable from the ratio of the total set of islands.
Results  Our results indicate a strong effect of habitat availability on the SIE. Communities of islands within the SIE range, host a higher percentage of generalists. An analysis of the specific habitat requirements shows that, for isopods, the crucial factor is the lack of habitats related to inland waters from small islands.
Main conclusions  The distribution of habitats on islands of different size is of major importance for the occurrence of a SIE. The relative representation of specialist and generalist species on islands of different size plays an important role in shaping SIE-related patterns. Conservation efforts should pay special attention on freshwater habitats, especially on small Aegean islands. Identifying the causal factors of SIE, combined with a thorough knowledge of the ecological requirements of species can offer insights into identifying habitat types and groups of species that are more vulnerable to alterations of the environment.     

Levels of endemism are not necessarily biased by the co-presence of species with different range sizes: a case study of Vilenkin and Chikatunov's models

Aim  To illustrate problems in the methods proposed by B. Vilenkin and V. Chikatunov to study levels of endemism and species–area relationships.
Location  The study used data on the distribution of tenebrionid beetles (Coleoptera, Tenebrionidae) on the Aegean Islands (Greece).
Methods  A total of 32 islands and 170 taxa (species and subspecies) were included in this study. Levels of endemism were evaluated both as the proportion of endemic taxa, and according to the methods proposed by Vilenkin and Chikatunov, which are based on the number of non-endemic taxa and various relationships with area. A model of the species–area relationship proposed by these authors was also analysed.
Results  The number of endemic taxa was positively correlated with the number of taxa with different distribution types, but this positive correlation did not influence the estimation of the level of endemism. In fact, the commonly used estimate of endemicity as a percentage was strongly correlated with the endemism values calculated according to the method of Vilenkin and Chikatunov. The usual power function fitted the species–area relationship as well as the most complicated method of Vilenkin and Chikatunov.
Main conclusions  As hypothesized by Vilenkin and Chikatunov, the number of endemic taxa was influenced both by the number of taxa of other biogeographical ranks, and by an island's area. However, explanations for the positive relationship between the number of endemic taxa and taxa of different biogeographical ranks are equivocal. Importantly, this relationship did not necessarily influence the level of endemism, which could be expressed adequately by percentages. The method proposed by Vilenkin and Chikatunov to estimate the species–area relationship cannot be clearly justified on theoretical grounds and is of questionable practical utility.     

Co‐occurrence patterns in a diverse arboreal ant community are explained more by competition than habitat requirements

A major goal of community ecology is to identify the patterns of species associations and the processes that shape them. Arboreal ants are extremely diverse and abundant, making them an interesting and valuable group for tackling this issue. Numerous studies have used observational data of species co‐occurrence patterns to infer underlying assembly processes, but the complexity of these communities has resulted in few solid conclusions. This study takes advantage of an observational dataset that is unusually well‐structured with respect to habitat attributes (tree species, tree sizes, and vegetation structure), to disentangle different factors influencing community organization. In particular, this study assesses the potential role of interspecific competition and habitat selection on the distribution patterns of an arboreal ant community by incorporating habitat attributes into the co‐occurrence analyses. These findings are then contrasted against species traits, to explore functional explanations for the identified community patterns. We ran a suite of null models, first accounting only for the species incidence in the community and later incorporating habitat attributes in the null models. We performed analyses with all the species in the community and then with only the most common species using both a matrix‐level approach and a pairwise‐level approach. The co‐occurrence patterns did not differ from randomness in the matrix‐level approach accounting for all ant species in the community. However, a segregated pattern was detected for the most common ant species. Moreover, with the pairwise approach, we found a significant number of negative and positive pairs of species associations. Most of the segregated associations appear to be explained by competitive interactions between species, not habitat affiliations. This was supported by comparisons of species traits for significantly associated pairs. These results suggest that competition is the most important influence on the distribution patterns of arboreal ants within the focal community. Habitat attributes, in contrast, showed no significant influence on the matrix‐wide results and affected only a few associations. In addition, the segregated pairs shared more biological characteristic in common than the aggregated and random ones.     

Associations between avian functional guild response and regional landscape properties for conservation planning

This project facilitates a regional approach to conservation planning in Pennsylvania based on avian breeding habitat selection. The objectives were to: (1) determine the sensitivity of spatial pattern in avian diversity to changing thresholds of intra-guild species richness and (2) relate change of spatial pattern in avian diversity with landscape characteristics of bird Atlas blocks. Two state-wide spatial data layers, based on Landsat satellite data were constructed for this study. These regional landscape data were compared to Breeding Bird Atlas data from 1983 to 1989 using a geographic information system. Breeding bird data were recorded from 4928 blocks that form a grid covering Pennsylvania. Correlation analysis reduced landscape variables to 12 originally derived from forest, urban, roads, streams, and topographic data.Avian functional response guilds were used to analyze associations between breeding bird data and landscape variables. Functional response guilds were created by grouping organisms based on shared habitat preferences or behavioral characteristics. Most of the 18 avian guilds identified for this study were based on shared structural resource characteristics of preferred breeding habitat. Preferred structural resources frequently included the amount and type of forest. For this study, guilds separate resource characteristics by: (1) primary habitat (i.e. forest interior, forest edge), (2) area sensitivity (i.e. forest and grassland), (3) migratory status (i.e. resident, temperate, and tropical), and (4) nest placement (i.e. canopy nester, forest ground nester). Wetland obligate species were treated as a separate guild. Breeding Bird Atlas blocks were tabulated with respect to the number of species present from each guild. For a given guild, the number of its species in a block is termed guild-specific species richness. Sample blocks having high species richness for a given guild often occur adjacent or in close proximity forming spatial clusters in the landscape. Spatial coherence (adjacency/proximity) among the blocks forming these islands is shared guild-specific richness. Spatial clustered blocks of each guild represent areas that presumably possess required resources for members of that guild. Blocks having high intra-guild richness were evaluated with a group of block-level continuous variables using multiple logistic regressions. Logistic regression results indicate that a convincing connection exists between landscape properties of Breeding Bird Atlas blocks and habitat selection characteristics of guild members. Percent of forest cover and mean elevation were the most important habitat characteristics influencing intra-guild richness for most of the guilds tested. Concordance values from logistic regression were used to determine the strength of each guild model. Concordance, the proportion that represents the percent of correct guild richness predictions versus incorrect predictions, suggests a relationship between guild-rich clusters and habitat resources required by each guild. The highest concordance was for the exotics guild at 76.3% and the next highest was 74.8% for the grassland area sensitive guild. This signifies a 75% certainty that landscape variables could predict occurrence of a guild-rich block. Eight more guilds had concordance values greater than 65%.By using a guild approach, this study goes beyond total diversity to the more informative structural and functional diversity of guilds. Spatially clustered blocks of high species richness for a particular guild are more indicative of habitat availability and quality than would be the case for overall species richness. Clusters of blocks having high intra-guild species richness become candidate areas for conservation efforts.     

Evidence for community assembly constraints during succession in dune slack plant communities

Community assembly during succession can be constrained by both local and regional factors. Despite an increasing regional species pool size during succession, we found a limit on the number of species in 1 × 1 m plots in dune slacks. Three alternative hypotheses (habitat heterogeneity, dispersal limitation and niche limitation) explaining this community saturation were tested. A null model analysis showed that species richness in the plots had an unusually low variance suggesting that beta habitat diversity was not likely to explain the limitation on species richness. Because we did not find a correlation between the distribution of species over the slack and their dispersal capacity, we also excluded the dispersal limitation hypothesis. Finally, a guild proportionality analysis revealed that the abundances of forb, graminoid and ruderal species showed low an unusually low variance over all age classes involved. This provides evidence for nonrandom community assembly during succession, likely to be determined by competitive exclusion among species of the same guild.     

Bird diversity and environmental gradients in Britain: a test of the species–energy hypothesis

1.  We tested the species diversity–energy hypothesis using the British bird fauna. This predicts that temperature patterns should match diversity patterns. We also tested the hypothesis that the mechanism operates directly through effects of temperature on thermoregulatory loads; this further predicts that seasonal changes in temperature cause matching changes in patterns of diversity, and that species' body mass is influential.
2.  We defined four assemblages using migration status (residents or visitors) and season (summer or winter distribution). Records of species' presence/absence in a total of 2362, 10 × 10-km, quadrats covering most of Britain were used, together with a wide selection of habitat, topographic and seasonal climatic data.
3.  We fitted a logistic regression model to each species' distribution using the environmental data. We then combined these individual species models mathematically to form a diversity model. Analysis of this composite model revealed that summer temperature was the factor most strongly associated with diversity.
4.  Although the species–energy hypothesis was supported, the direct mechanism, predicting an important role for body mass and matching seasonal patterns of change between diversity and temperature, was not supported.
5.  However, summer temperature is the best overall explanation for bird diversity patterns in Britain. It is a better predictor of winter diversity than winter temperature. Winter diversity is predicted more precisely from environmental factors than summer diversity.
6.  Climate change is likely to influence the diversity of different areas to different extents; for resident species, low diversity areas may respond more strongly as climate change progresses. For winter visitors, higher diversity areas may respond more strongly, while summer visitors are approximately neutral.     

Additive partitioning of rarefaction curves and species-area relationships: unifying alpha-, beta- and gamma-diversity with sample size and habitat area

Additive partitioning of species diversity is widely applicable to different kinds of sampling regimes at multiple spatial and temporal scales. In additive partitioning, the diversity within and among samples ( α and β ) is expressed in the same units of species richness, thus allowing direct comparison of α and β . Despite its broad applicability, there are few demonstrated linkages between additive partitioning and other approaches to analysing diversity. Here, we establish several connections between diversity partitions and patterns of habitat occupancy, rarefaction, and species–area relationships. We show that observed partitions of species richness are equivalent to sample-based rarefaction curves, and expected partitions from randomization tests are approximately equivalent to individual-based rarefaction. Additive partitions can also be applied to species–area relationships to determine the relative contributions of factors influencing the β -diversity among habitat fragments.     

Species richness, composition and abundance of fruit-feeding butterflies in the Brazilian Atlantic Forest: comparison between a fragmented and a continuous landscape

Aim  To evaluate frugivorous butterflies as indicators of forest disturbance in the Brazilian Atlantic Forest.
Location  The study area is located in the municipality of Cotia, São Paulo State, south-eastern Brazil (23°35'−23°50' S, 46°45'−47°15' W).
Methods  Sampling was done at four sites inside a large forest block, the Morro Grande State Reserve, and in five forest fragments in an adjacent fragmented landscape. Butterflies were sampled with portable traps, baited with a fermented mixture of banana and sugar cane juice. Sampling was carried out during the period most favourable for the capture of frugivorous butterflies in south-eastern Brazil.
Results  All richness-related results indicated no effect of forest fragmentation on the frugivorous butterfly guild, concurring with the suggestion of appreciable resistance of Atlantic Forest butterflies to habitat modification. However, species composition discriminated between the two landscapes, indicating that fragmentation may have effects beyond simple species richness. When species were analysed individually, clear patterns of distribution emerged, with some species that were very abundant in the fragmented landscape being practically absent in the continuous landscape, and vice versa. This pattern seems consistent even for some subfamilies.
Main conclusion  Our findings support the usefulness of the frugivorous butterfly guild as a biological indicator of forest disturbance effects in one of the world's most threatened ecosystems.     

Inferring species abundance distribution across spatial scales

A long-standing problem in ecology is to understand how the species–abundance distribution (SAD) varies with sampling scale. The problem was first characterized by Preston as the veil line problem. Although theoretical and empirical studies have now shown the nonexistence of the veil line, this problem has generated much interest in scaling biodiversity patterns. However, research on scaling SAD has so far exclusively focused on the relationship between the SAD in a smaller sampling area and a known SAD assumed for a larger area. An unsolved challenge is how one may predict species–abundance distribution in a large area from that of a smaller area. Although upscaling biodiversity patterns (e.g. species–area curve) is a major focus of macroecological research, upscaling of SAD across scale is, with few exceptions, ignored in the literature. Methods that directly predict SAD in a larger area from that of a smaller area have just started being developed. Here we propose a Bayesian method that directly answers this question. Examples using empirical data from tropical forests of Malaysia and Panama are employed to demonstrate the use of the method and to examine its performance with increasing sampling area. The results indicate that only 10-15% of the total census area is needed to adequately predict species abundance distribution of a region. In addition to species abundance distributions, the method also predicts well the regional species richness.     

Effects of habitat characteristics and interspecific interactions on co-occurrence patterns of saproxylic beetles breeding in tree boles after forest fire: null model analyses

It is often suggested that habitat attributes and interspecific interactions can cause non-random species co-occurrence patterns, but quantifying their contributions can be difficult. Null models that systematically exclude and include habitat effects can give information on the contribution of these factors to community assembly. In the boreal forest, saproxylic beetles are known to be attracted to recently burned forests where they breed in dead and dying trees. We examined whether species co-occurrences of saproxylic beetles that develop in, and emerge from, boles of recently burned trees show non-random patterns. We also estimated the extent to which both the post-fire habitat attributes and interspecific interactions among beetles contribute to such patterns. We sampled tree boles encompassing key attributes (tree species, tree size/dbh and burn severity) that are thought to characterize species–habitat associations of saproxylic beetles, a proposition that we tested using indicator species analysis. Two null models with no habitat constraints (“unconstrained”) indicated that a total of 29.4% of the species pairs tested had significant co-occurrence patterns. Habitat-constrained null models indicated that most of the detected species aggregations (72%) and segregations (59%) can be explained by shared and distinct species–habitat relationships, respectively. The assembly pattern was also driven by interspecific interactions, of which some were modulated by habitat; for example, predator and prey species tended to co-occur in large-sized trees (a proxy of available bark/wood food resource primarily for the prey). In addition, some species segregation suggesting antagonistic, competitive, or prey–predator interactions were evident after accounting for the species’ affinities for the same tree species. Overall, our results suggest that an intimate link between habitat and interspecific interactions can have important roles for community assembly of saproxylic assemblages even following disturbance by fire. We also show that a systematic application of null models can offer insight into the mechanisms behind the assembly of ecological communities.