Attempted tests of the surrogacy value of the ED environmental diversity measures highlight the need for corroboration assessment of surrogacy hypotheses

A recent paper in this journal (Hortal et al., 2009) claimed to have evaluated the ED biodiversity surrogates methods of Faith and Walker, 1994, Faith and Walker, 1996, and to have provided evidence for poor performance of the continuous ED method. In fact, their study neither used nor evaluated the continuous ED method. Here, I document their misrepresentation. I then discuss some constructive lessons emerging from their study and other recent studies that have attempted tests of ED surrogacy value. The need to consider the actual degree of support that observed evidence provides for a hypothesis about surrogacy raises general issues for evaluations of indicators’ performance, and suggests a greater role for corroboration assessments. Guidelines for achieving this cover three aspects of surrogates testing: experimental design of tests, ongoing corroboration assessment of evidence produced by tests, and accumulation of lessons learned from multiple test studies over time.     

An illustrated key to adult males of neotropical Fannia Robineau-Desvoidy belonging to pusio sub-group (Diptera, Fanniidae).

The 11 neotropical species of Fannia belonging to the pusio sub-group (canicularis group), are keyed F. dodgei Seago; F. femoralis (Stein); F. pamplonae Couri & Araújo; F. parafemoralis Araújo & Couri; F. paraisensis Araújo & Couri; F. punctivervis Malloch; F. pusio (Wiedemann); F. sabroskyi Seago; F. snyderi Seago; F. trimaculata (Stein); F. trimaculatoides Couri & Pamplona. The key is addressed only to the males and the illustrations help in the visualization of the characters.     

Male and female association in Trichomyia Haliday in Curtis, 1839 using a molecular approach (Diptera,Psychodidae, Trichomyiinae), and description of new species from Brazil

A new species of Trichomyia from the state of Bahia, Brazil, is described and illustrated, and male and female are associated using DNA barcoding. Additionally, fragments of the COI of two other species, Trichomyia cerdosa Araújo & Bravo, 2016 and Trichomyia ituberensis Araújo & Bravo, 2016, and the females of two unidentified species, are sequenced.     

<Emphasis Type="Italic">Scleria rubrostriata,</Emphasis> a new species of Cyperaceae from the Brazilian Atlantic Forest

A new species is here described: Scleria rubrostriata A. C. Araújo & N. A. Brummitt, a herb from the Atlantic forest of Espírito Santo State, south-eastern Brazil. A key to the sections of Scleria and to the American species of Scleria sect. Schizolepis is provided, and an illustration of the new species and comparison to similar taxa are given.     

Best Practicable Aggregation of Species: a step forward for species surrogacy in environmental assessment and monitoring

The available taxonomic expertise and knowledge of species is still inadequate to cope with the urgent need for cost‐effective methods to quantifying community response to natural and anthropogenic drivers of change. So far, the mainstream approach to overcome these impediments has focused on using higher taxa as surrogates for species. However, the use of such taxonomic surrogates often limits inferences about the causality of community patterns, which in turn is essential for effective environmental management strategies. Here, we propose an alternative approach to species surrogacy, the “Best Practicable Aggregation of Species” (BestAgg), in which surrogates exulate from fixed taxonomic schemes. The approach uses null models from random aggregations of species to minimizing the number of surrogates without causing significant losses of information on community patterns. Surrogate types are then selected in order to maximize ecological information. We applied the approach to real case studies on natural and human‐driven gradients from marine benthic communities. Outcomes from BestAgg were also compared with those obtained using classic taxonomic surrogates. Results showed that BestAgg surrogates are effective in detecting community changes. In contrast to classic taxonomic surrogates, BestAgg surrogates allow retaining significantly higher information on species‐level community patterns than what is expected to occur by chance and a potential time saving during sample processing up to 25% higher. Our findings showed that BestAgg surrogates from a pilot study could be used successfully in similar environmental investigations in the same area, or for subsequent long‐term monitoring programs. BestAgg is virtually applicable to any environmental context, allowing exploiting multiple surrogacy schemes beyond stagnant perspectives strictly relying on taxonomic relatedness among species. This prerogative is crucial to extend the concept of species surrogacy to ecological traits of species, thus leading to ecologically meaningful surrogates that, while cost effective in reflecting community patterns, may also contribute to unveil underlying processes. A specific R code for BestAgg is provided.     

Dynamic species co‐occurrence networks require dynamic biodiversity surrogates

In conservation it is inevitable that surrogates be selected to represent the occurrence of hard‐to‐find species and find priority locations for management. However, species co‐occurrence can vary over time. Here we demonstrate how temporal dynamics in species co‐occurrence influence the ability of managers to choose the best surrogate species. We develop an efficient optimisation formulation that selects the optimal set of complementary surrogate species from any co‐occurrence network. We apply it to two Australian datasets on successional bird responses to disturbances of revegetation and fire. We discover that a surprisingly small number of species are required to represent the majority of species co‐occurrences at any one time. Because co‐occurrence patterns are temporally dynamic, the optimal set of surrogates, and the number of surrogates required to achieve a desired surrogacy power, depend on sampling effort and the successional state of a system. Overlap in optimal sets of surrogates for representing 70% of co‐occurring species ranges from zero to 57% depending on when the surrogacy decision is made. Surrogate sets representing early successional communities over‐estimate the power of surrogacy decisions at later times. Our results show that in dynamic systems, optimal surrogates might be selected in different ways: 1) use short‐term monitoring to choose a larger number of static less‐informative surrogates; 2) use long‐term monitoring to choose a smaller number of static high‐power surrogates that may poorly represent early successional co‐occurrence; 3) develop adaptive surrogate selection frameworks with high short‐term and long‐term surrogacy power that update surrogate sets and capture temporal dynamics in species co‐occurrence. Our results suggest vigilance is needed when selecting surrogates for other co‐occurring species in dynamic landscapes, as selected surrogates from one time may have reduced effectiveness at a different time. Ultimately, decisions that fail to acknowledge dynamic species co‐occurrence will lead to uninformative or redundant surrogates.     

Shortcuts in biodiversity research: What determines the performance of higher taxa as surrogates for species?

Biodiversity research is often impeded by the time and resources required to identify species. One possible solution is to use higher taxa to predict species richness and community composition. However, previous studies have shown that the performance of higher taxa as surrogates for species is highly variable, making it difficult to predict whether the method will be reliable for a particular objective. Using 8 independent datasets, I tested whether higher taxa accurately characterize the responses of beetle and ant communities to environmental drivers. For each dataset, ordinations were carried out using species and higher taxa, and the two compared using the Procrustes m² statistic (a scale‐independent variant of Procrustes sum of squares). I then modelled the relationship between five hypothesised explanatory variables and 1) Procrustes m², and 2) the coefficient of determination (R²) for the correlation between richness of species and higher taxa. The species to higher taxon ratio, community structure, beta diversity, completeness of sampling, and taxon (beetles or ants) were all significant predictors of m², together explaining 88% of the variance. The only significant predictor of R² was the species to higher taxon ratio, which explained 45% of the variance. When using higher taxa to predict community composition, better performance is expected when the ratio of species to higher taxa is low, in communities with high evenness and high species turnover, and when there is niche conservation within higher taxa. When using higher taxa to predict species richness, effective surrogacy can be expected when the species to higher taxon ratio is very low. When it is not, surrogacy performance may be strongly influenced by stochastic factors, making predictions of performance difficult.     

<Emphasis Type="Italic">Cyperaceae Raddianae</Emphasis>: a nomenclatural and taxonomic study of the Cyperaceae published in G. Raddi’s <Emphasis Type="Italic">Agrostografia brasiliensis</Emphasis>

A study of the 26 Cyperaceae published in G. Raddi’s Agrostografia brasiliensis is presented. Several new synonyms and a re-evaluation of some taxa described by G. Raddi are proposed. The first citation of Fimbristylis pilosa Vahl for Brazil is reported. The new combination, Pleurostachys pilulifera (Bertol.) Longhi-Wagner, Baldini & A. C. Araújo is made.     

SURROGACY: DONOR CONCEPTION REGULATION IN JAPAN

As of 2008, surrogacy is legal and openly practised in various places; Japan, however, has no regulations or laws regarding surrogacy. This paper reports the situation of surrogacy in Japan and in five other regions (the USA, the UK, Taiwan, Korea and France) to clarify the pros and cons of prohibiting surrogacy, along with the problems and issues relating to surrogacy compensation. Not only in a country such as France that completely prohibits surrogacy within the country, but also in a country such as the UK that allows non‐commercial surrogacy, infertile couples travel overseas for the purpose of surrogacy. In addition, some couples might seek underground surrogacy if the government prohibits surrogacy. If an intended parent couple and a surrogate make an agreement among themselves and then a problem occurs, they cannot ask for support from professionals or bring a case to court, as can be observed in South Korea and Taiwan. We also conclude that there is little difference between commercial surrogacy and non‐commercial surrogacy in the absence of a clear definition of ‘reasonable expenses.’ In the UK, the law does not allow surrogates to receive compensation. However, in reality, there may be little difference between the amounts paid to surrogates for profit in the US and those paid to surrogates for reasonable expenses in the UK. We conclude that the issue of surrogacy demands further discussion in Japan.     

Species traits and environmental characteristics together regulate ant‐associated biodiversity

Host‐associated organisms (e.g., parasites, commensals, and mutualists) may rely on their hosts for only a portion of their life cycle. The life‐history traits and physiology of hosts are well‐known determinants of the biodiversity of their associated organisms. The environmental context may strongly influence this interaction, but the relative roles of host traits and the environment are poorly known for host‐associated communities. We studied the roles of host traits and environmental characteristics affecting ant‐associated mites in semi‐natural constructed grasslands in agricultural landscapes of the Midwest USA. Mites are frequently found in ant nests and also riding on ants in a commensal dispersal relationship known as phoresy. During nonphoretic stages of their development, ant‐associated mites rely on soil or nest resources, which may vary depending on host traits and the environmental context of the colony. We hypothesized that mite diversity is determined by availability of suitable host ant species, soil detrital resources and texture, and habitat disturbance. Results showed that that large‐bodied and widely distributed ant species within grasslands support the most diverse mite assemblages. Mite richness and abundance were predicted by overall ant richness and grassland area, but host traits and environmental predictors varied among ant hosts: mites associated with Aphaenogaster rudis depended on litter depth, while Myrmica americana associates were predicted by host frequency and grassland age. Multivariate ordinations of mite community composition constructed with host ant species as predictors demonstrated host specialization at both the ant species and genus levels, while ordinations with environmental variables showed that ant richness, soil texture, and grassland age also contributed to mite community structure. Our results demonstrate that large‐bodied, locally abundant, and cosmopolitan ant species are especially important regulators of phoretic mite diversity and that their role as hosts is also dependent on the context of the interaction, especially soil resources, texture, site age, and area.     

A test for assessment of saproxylic beetle biodiversity using subsets of “monitoring species”

In European forests, large scale biodiversity monitoring networks need to be implemented – networks which include components such as taxonomical groups that are at risk and that depend directly on forest stand structure. In this context, monitoring the species-rich group of saproxylic beetles is challenging. In the absence of sufficient resources to comprehensively survey a particular group, surrogates of species richness can be meaningful tools in biodiversity evaluations. In search of restricted subsets of species to use as surrogates of saproxylic beetle richness, we led a case study in Western Europe.Beetle data were compiled from 67 biodiversity surveys and ecological studies carried out from 1999 to 2010 with standardised trapping methods in France and Belgium. This large-scale dataset contains 642 forest plots, 1521 traps and 856 species. Twenty-two simplified species subsets were identified as potential surrogates, as well as the number of genera, a higher taxonomic level, taking into account, for each surrogate, the effort required for species identification, the practical monitoring experience necessary, the species conservation potential or the frequency of species occurrence. The performance of each surrogate was analyzed based on the following parameters: overall surrogacy (correlation between subset richness and total species richness), surrogacy vs. identification cost balance, surrogacy variation over a wide range of ecological conditions (forest type, altitude, latitude and bio-geographical area) and consistency with spatial scale. Ecological representativeness and ability to monitor rare species were supplementary criteria used to assess surrogate performance.The subsets consisting of the identifiable (or only easy-to-identify species) could easily be applied in practice and appear to be the best performing subsets, from a global point of view.The number of genera showed good prediction at the trap level and its surrogacy did not vary across wide environmental gradients. However, the subset of easy-to-identify species and the genus number were highly sensitive to spatial scale, which limits their use in large-scale studies. The number of rare species or the species richness of single beetle families (even the best single-family subset, the Cerambycidae) was very weak surrogates for total species richness. Conversely, the German list of monitoring species had high surrogacy, low identification costs and was not strongly influenced by the main geographical parameters, even with our French and Belgian data.In European-wide monitoring networks, such internationally validated subsets could be very useful with regard to the timing and cost-efficiency of field inventories.     

Spatial conservation priorities are highly sensitive to choice of biodiversity surrogates and species distribution model type

Pressure to conserve biodiversity with limited resources has led to increasing use of species distribution models (SDMs) for spatial conservation prioritization. Published spatial prioritization exercises often focus on well‐studied groups, with data compiled from on‐line databases of ad‐hoc collections. Conservation plans generally aim to protect all components of biodiversity, and it is implied that the species used in prioritization act as surrogates. Here, we assess the sensitivity of spatial priorities to model and surrogate choice using a case study from a fragmented agricultural area of south eastern Australia that is poorly represented in the national reserve system. We model the distributions of 30 species of bird, microbat and bee using two types of SDM; generalised linear models based on systematic surveys that yield presence and absence observations, and MaxEnt models based on biodiversity database records. Eight prioritization scenarios were tested using Zonation software, and were based on either the presence–background or presence–absence SDMs and combinations of surrogacy among the three taxa. We found low correlations between SDMs generated for the same species using different modelling frameworks (μ = 0.18, n = 26). Area under the receiver operating characteristic curve (AUC) estimates generated by MaxEnt were optimistic; on average 1.36 times higher than when tested against the systematic survey data. Conservation priorities were sensitive to the choice of surrogate and type of data used to fit SDMs, and though bats and birds formed moderately good surrogates for each other, there was less compelling evidence of surrogacy for bees. Because valid surrogacy is unlikely with most existing data sets, investment in high quality data for less‐surveyed groups prior to planning should still be a priority. If this is not possible, then it is advisable to analyse the sensitivity of conservation plans to the assumed surrogacy and quality of data available.     

High‐severity fire corroborated in historical dry forests of the western United States: response to Fulé et al.

Accurate assessment of changing fire regimes is important, since climatic change and people may be promoting more wildfires. Government wildland fire policies and restoration programmes in dry western US forests are based on the hypothesis that high‐severity fire was rare in historical fire regimes, modern fire severity is unnaturally high and restoration efforts should focus primarily on thinning forests to eliminate high‐severity fire. Using General Land Office (GLO) survey data over large dry‐forest landscapes, we showed that the proportion of historical forest affected by high‐severity fire was not insignificant, fire severity has not increased as a proportion of total fire area and large areas of dense forest were present historically (Williams & Baker, Global Ecology and Biogeography, 21 , 1042–1052, 2012; W&B). In response, Fulé et al. (Global Ecology and Biogeography, 2013, doi: 10.1111/geb.12136; FE) suggest that our inferences are unsupported and land management based on our research could be damaging to native ecosystems. Here, we show that the concerns of FE are unfounded. Their criticism comes from misquoting W&B, mistaking W&B's methods, misusing evidence (e.g. from Aldo Leopold) and missing substantial available evidence. We also update corroboration for the extensive historical high‐severity fire shown by W&B. We suggest that restoration programmes are misdirected in seeking to reduce all high‐severity fire in dry forests, given findings from spatially extensive GLO data and other sources.     

Effectiveness of biological surrogates for predicting patterns of marine biodiversity: a global meta-analysis

The use of biological surrogates as proxies for biodiversity patterns is gaining popularity, particularly in marine systems where field surveys can be expensive and species richness high. Yet, uncertainty regarding their applicability remains because of inconsistency of definitions, a lack of standard methods for estimating effectiveness, and variable spatial scales considered. We present a Bayesian meta-analysis of the effectiveness of biological surrogates in marine ecosystems. Surrogate effectiveness was defined both as the proportion of surrogacy tests where predictions based on surrogates were better than random (i.e., low probability of making a Type I error; P) and as the predictability of targets using surrogates (R(2)). A total of 264 published surrogacy tests combined with prior probabilities elicited from eight international experts demonstrated that the habitat, spatial scale, type of surrogate and statistical method used all influenced surrogate effectiveness, at least according to either P or R(2). The type of surrogate used (higher-taxa, cross-taxa or subset taxa) was the best predictor of P, with the higher-taxa surrogates outperforming all others. The marine habitat was the best predictor of R(2), with particularly low predictability in tropical reefs. Surrogate effectiveness was greatest for higher-taxa surrogates at a <10-km spatial scale, in low-complexity marine habitats such as soft bottoms, and using multivariate-based methods. Comparisons with terrestrial studies in terms of the methods used to study surrogates revealed that marine applications still ignore some problems with several widely used statistical approaches to surrogacy. Our study provides a benchmark for the reliable use of biological surrogates in marine ecosystems, and highlights directions for future development of biological surrogates in predicting biodiversity.     

Measuring the effectiveness of regional conservation assessments at representing biodiversity surrogates at a local scale: A case study in Réunion Island (Indian Ocean)

In a context of scarce financial and human resources, the allocation of conservation efforts needs to be optimized. Our analysis attempts to draw conclusions on the integration of regional and local conservation assessments, specifically, with regard to the acquisition of fine‐scale data to complement the regional assessment. This study undertaken in Réunion Island (Indian Ocean) assessed how biodiversity surrogates targeted at a regional scale represented other biodiversity surrogates at a local scale. Biodiversity surrogates at both scales consisted of species, habitats and processes. Habitats and processes at regional scale were defined using a coarser scale of thematic resolution than at local scale. The surrogacy was tested in terms of incidental representation of local‐scale features in the regional assessments, and correlation of irreplaceability values between scales. Near‐minimum sets and irreplaceability values were generated using MARXAN software. Our results revealed that conservation targets for processes at local scale were never met incidentally, while threatened species and fragmented habitats were also usually under‐represented. More specifically, requiring only 12% of the local planning domain, the application of species as surrogates at regional scale was the least effective option at representing biodiversity features at local scale. In contrast, habitats at a coarse scale of thematic resolution achieved a significant proportion of conservation targets incidentally (67%) and their irreplaceability values were well correlated with the irreplaceability values of surrogates at local scale. The results highlighted that all three types of biodiversity surrogates are complementary for assessing overall biodiversity. Because of the cost of data acquisition, we recommended that the most efficient strategy to develop nested regional/local conservation plans is to apply habitats and processes at a coarse scale of thematic resolution at regional scale, and threatened species and degraded habitats at local scale, with their fine‐scale mapping limited to highly transformed areas.     

Higher taxa as surrogates of species richness of spiders in insect‐resistant transgenic rice

Abstract Biodiversity assessments can often be time‐ and resource‐consuming. Several alternative approaches have been proposed to reduce sampling efforts, including indicator taxa and surrogates. In this study, we examine the reliability of higher taxon surrogates to predict species richness in two experimental rice fields of Fujian Province, southeastern China during 2005 and 2009. Spider samples in transgenic and nontransgenic plots were collected using a suction sampler. Both the genus and family surrogates had significant and positive linear relationships with species richness in the transgenic and nontransgenic rice fields. The rice varieties did not significantly influence the linear relationships. Our findings suggest that higher‐taxon surrogacy could be a useful alternative to complete species inventory for risk assessments of transgenic rice.     

New frameworks for species surrogacy in monitoring highly variable coastal ecosystems: Applying the BestAgg approach to Mediterranean coastal lagoons

Benthic invertebrates are good indicators of aquatic ecosystem health. Yet, environmental monitoring and assessment of community changes in relation to both natural and human sources of disturbance involves considerable efforts for sample processing and time-consuming identifications of organisms, which make challenging large-scale and continuous monitoring programs required under the current regulatory frameworks at European scale. The use of higher taxa (e.g. families) as surrogates for species is a mainstream approach to reduce cost and time associated to fine taxonomic resolution in environmental studies, especially concerning macro-invertebrate communities. However, this approach of ‘taxonomic sufficiency’ simply relies on the static grouping of organisms in taxa belonging to a single higher taxonomic level irrespective of their ecological relevance or difficulties in their taxonomic identification, leading to unnecessary losses of taxonomic detail or ecological information. A new approach, namely the Best Practicable Aggregation of Species (BestAgg), has been recently developed as an alternative procedure for species surrogacy. BestAgg is based on aggregating species in the minimum number of surrogates sufficient to depict species-level community patterns consistently, while capitalizing on ecological information. Although the approach has been successfully applied to marine and freshwater invertebrate assemblages, its effectiveness in transitional water systems, where the complex and highly variable environmental conditions may affect the performance of surrogacy methods, still remain untested. Here, we applied the BestAgg approach to quantifying spatio-temporal patterns of variability of macro-invertebrate assemblages from Mediterranean coastal lagoons (Northern Adriatic Sea). Surrogates were defined using species-level data from a representative lagoon system, which served as pilot study. Then, they were used to analyze macro-invertebrate assemblages in two independent lagoons in the same geographic area. Results showed that BestAgg surrogates were effective in depicting multivariate patterns of macro-invertebrate assemblages as at species level over and beyond potential variations among the investigated lagoons. The use of families, following a more classic approach based on taxonomic sufficiency, also provided results comparable to those obtained using species. However, with respect to families, BestAgg surrogates allowed an estimated timesaving of 40% higher while still retaining an equivalent amount of information on species-level patterns. More importantly, BestAgg allowed exploiting different criteria of species aggregation, leading to a set of surrogates more aligned with ecological/functional characteristics of organisms, suggesting that BestAgg approach may provide a fresh perspective for optimizing trade-offs between pragmatism and the need for relevant ecological information in environmental assessment and monitoring.     

Effectiveness of Biodiversity Surrogates for Conservation Planning: Different Measures of Effectiveness Generate a Kaleidoscope of Variation

Conservation planners represent many aspects of biodiversity by using surrogates with spatial distributions readily observed or quantified, but tests of their effectiveness have produced varied and conflicting results. We identified four factors likely to have a strong influence on the apparent effectiveness of surrogates: (1) the choice of surrogate; (2) differences among study regions, which might be large and unquantified (3) the test method, that is, how effectiveness is quantified, and (4) the test features that the surrogates are intended to represent. Analysis of an unusually rich dataset enabled us, for the first time, to disentangle these factors and to compare their individual and interacting influences. Using two data-rich regions, we estimated effectiveness using five alternative methods: two forms of incidental representation, two forms of species accumulation index and irreplaceability correlation, to assess the performance of ‘forest ecosystems’ and ‘environmental units’ as surrogates for six groups of threatened species—the test features—mammals, birds, reptiles, frogs, plants and all of these combined. Four methods tested the effectiveness of the surrogates by selecting areas for conservation of the surrogates then estimating how effective those areas were at representing test features. One method measured the spatial match between conservation priorities for surrogates and test features. For methods that selected conservation areas, we measured effectiveness using two analytical approaches: (1) when representation targets for the surrogates were achieved (incidental representation), or (2) progressively as areas were selected (species accumulation index). We estimated the spatial correlation of conservation priorities using an index known as summed irreplaceability. In general, the effectiveness of surrogates for our taxa (mostly threatened species) was low, although environmental units tended to be more effective than forest ecosystems. The surrogates were most effective for plants and mammals and least effective for frogs and reptiles. The five testing methods differed in their rankings of effectiveness of the two surrogates in relation to different groups of test features. There were differences between study areas in terms of the effectiveness of surrogates for different test feature groups. Overall, the effectiveness of the surrogates was sensitive to all four factors. This indicates the need for caution in generalizing surrogacy tests.     

Spatial and temporal variation in the diet of the sandy beach gastropod Olivella minuta

Marine gastropods exert a strong influence on the structure of marine ecosystems through their foraging activities, yet little is known about how environmental differences in space and time may affect their feeding behavior. To enhance the knowledge about the trophic ecology of coastal benthic species, we investigated temporal and spatial variations in the diet of the sandy beach gastropod Olivella minuta. We collected individuals of O. minuta seasonally, over 1 year, at two sandy beaches with different morphodynamic characteristics (Araçá Bay, an ultra‐dissipative tidal flat, and Barequeçaba Beach, a dissipative beach) and analyzed the stomach contents of specimens of O. minuta. We identified approximately 15,000 food items, which we grouped into 14 categories. Thirteen food categories were found at each area, but significant differences in diet were observed between sites and sampling periods. Individuals from Araçá consumed an overall larger quantity and diversity of food items, which consisted mainly of foraminiferans, polychaetes, and crustaceans. At Barequeçaba, diatoms were the most important item, followed by crustaceans and polychaetes. Temporal variation depended on sites: the number of food items in the stomachs of individuals of O. minuta and dietary composition was roughly constant over time at Araçá, whereas larger variations were observed at Barequeçaba. Although O. minuta has been previously characterized as a carnivore/scavenger, our results show that the diet of this species varies in space and time and comprises a large variety of food resources. The species thus plays a role in the cycling of both vegetal and animal organic matter. This generalist behavior may allow individuals to shift their diet according to habitat alterations and food availability, thereby enhancing the persistence and resilience of O. minuta in different environments or future climate change scenarios.     

The need for richness‐independent measures of turnover when delineating biogeographical regions

Delineating biogeographical regions is one of the primary steps when analysing biogeographical patterns. In their proposed quantitative framework, Kreft & Jetz (2010, Journal of Biogeography, 37 , 2029–2053) recommended the use of the β_sim index to delineate biogeographical regions because this turnover measure is weakly affected by differences in species richness between localities. A recent study by Carvalho et al. (2012, Global Ecology and Biogeography, 21 , 760–771) critiziced the use of β_sim in ecological and biogeographical studies, and proposed the β_‐3 index. Here we used simple numerical examples and an empirical case study (European freshwater fishes) to highlight potential pitfalls associated with the use of β_‐3 for bioregionalization. We show that β_‐3 is not a richness‐independent measure of species turnover. We also show that this index violates the ‘complementarity’ property, namely that localities without species in common have the largest dissimilarity, which is an essential prerequisite for beta diversity studies.