Reach- and catchment-scale determinants of the distribution of freshwater mussels (Bivalvia: Unionidae) in south-eastern Michigan, U.S.A.

1. We investigated the diversity and distribution of freshwater mussels at 40 sites in an agricultural catchment, the River Raisin in south‐eastern Michigan, to relate mussel assemblages and individual taxa to reach and catchment‐scale variables. Unionids were surveyed by timed searches in 100‐m reaches, and in‐stream and riparian habitat were quantified as well as flow, water chemistry and channel morphology. Land use/cover and surficial geology were determined for site subcatchments and riparian buffers. 2. Some 21 mussel species were found overall; richness ranged from 0 to 12 living species per site. From the upper to middle to lower catchment, the number of individuals, number of species, Shannon–Weaver diversity and relative abundance of intolerant unionids all declined significantly. 3. Four groupings based on overall mussel diversity and abundance were significantly related to reach‐scale habitat variables. The richest mussel assemblages were associated with sites with higher overall habitat quality, greater flow stability, less fine substratum, and lower specific conductance. 4. Stepwise multiple regressions revealed that the distribution and abundance of the total mussel assemblage, as well as the most common species, could be predicted from a combination of reach‐ and catchment‐scale variables (R² = 0.63 for total mussels, R² = 0.51–0.86 for individual species). 5. Flow stability, substratum composition and overall reach habitat quality were the most commonly identified reach‐scale variables, and measures of surficial geology were the most effective catchment‐scale variables. The spatial pattern of geology is likely to be responsible for the diversity gradient from the upper to the lower catchment. 6. Prior studies, attempting to explain mussel distributions from local habitat features alone, have found relatively weak relationships. By employing a combination of reach‐ and catchment‐scale habitat variables, this study was able to account for a substantial amount of the spatial variability in mussel distributions.     

Linkages among land-use, water quality, physical habitat conditions and lotic diatom assemblages: A multi-spatial scale assessment

We assessed the importance of spatial scales (catchment, stream network, and sample reach) on the effects of agricultural land-use on lotic diatom assemblages along a land-use gradient in the agricultural Willamette Valley Ecoregion of Oregon. Periphyton, water chemistry, and physical habitat conditions were characterized for 25 wadeable streams during a dry season (July to September, 1997). Additional water chemistry samples were collected in the following wet season (February 1998) to assess seasonal effects of land-use on stream water chemistry. Percent agricultural land-use in the study catchments ranged from 10% to 89% with an average of 52%. Partial canonical correspondence analysis (CCA) with the first axis constrained by % agricultural land-use showed that % agricultural land-use at 3 spatial scales explained between 3.7%–6.3% of variability in the diatom species dataset. Monte Carlo Permutation tests indicated that the variance explained by % agricultural land-use was only significant at the spatial scale of the stream network with 10- and 30-m band width (p<0.05, 999 permutations). In addition to the effects of % agricultural land-use, partial CCAs with a forward selection option showed that water chemistry (e.g., SiO₂), reach-scale stream channel dimensions (e.g., width, depth, and slope), reach-scale in-stream habitats (substrates and filamentous algal cover in stream beds), and riparian vegetative buffer were all important with relation to diatom species assemblages. Percent of obligately nitrogen-heterotrophic taxa was the only diatom autecological metric that showed a significant but weak correlation with % agricultural land-use along the stream network (r=0.50), but not at catchment or sample reach scale. Correlation between % agricultural land-use and water chemistry variables varied among the spatial scales and between seasons. Physical habitat variables (log₁₀ erodible substrate diameters and stream reach slope) were significantly correlated with % agricultural land-use along the stream network but not at catchment or sample reach scale. Our data suggest that spatial scales are important in assessing effects of land-use on stream conditions but the spatial scale effects may vary between seasons. Direct linkages between agricultural land-use and lotic diatom assemblages were weak during summer base-flow time regardless of the spatial scales. Summer sampling may underestimate the effects of catchment land-use on stream conditions in areas where seasonal patterns are so distinctive as in the Willamette Valley.     

Landscape variables influence taxonomic and trait composition of insect assemblages in Neotropical savanna streams

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The influence of scale and geography on relationships between stream community composition and landscape variables: description and prediction

SUMMARY 1. We analysed an existing database of macroinvertebrates and fish in the context of a newly established geographical information system (GIS) of physical features to determine the relationships between stream community composition and physical factors measured at three landscape scales – catchment, reach and bedform. Both an exploratory (concordance analysis) and a predictive ( ausrivas ) approach were used.
2. The environmental variables that most successfully accounted for variation in macroinvertebrate assemblages were mainly 'natural' and at the catchment-scale (relief ratio, basin diameter, etc.) but the human-induced physical setting of percentage of pasture in the riparian zone was also influential. For fish, 'natural' variables were also dominant, but these were mostly at the bedform or reach scales and land use featured strongly.
3. Geographic location accounted for some of the variation in invertebrate assemblages, partly because geography and influential conditions/resources are correlated but also because different species may have evolved in different places and have not colonised every 'ecologically appropriate' location. Geographic location was not influential in accounting for variation in assemblages of strongly flying invertebrates, supporting the hypothesis that organisms having high dispersal potential can be expected to break down geographic barriers more readily than those with poor dispersal powers. In accord with what is known about the local evolution and restricted distributions of native and exotic species, history (reflected in geography) appeared to account for some variation in fish assemblages.
4. Given their different mathematical bases, the fact that exploratory and predictive analyses yielded similar results provides added confidence to our conclusions.     

Implications of community concordance for assessing stream integrity at three nested spatial scales in Minnesota,U.S.A.

1. Fish and invertebrate assemblage data collected from 670 stream sites in Minnesota (U.S.A.) were used to calculate concordance across three nested spatial scales (statewide, ecoregion and catchment). Predictive taxa richness models, calibrated using the same data, were used to evaluate whether concordant communities exhibited similar trends in human‐induced taxa loss across all three scales. Finally, we evaluated the strength of the relationship between selected environmental variables and the composition of both assemblages at all three spatial scales. 2. Significant concordance between fish and invertebrate communities occurred at the statewide scale as well as in six of seven ecoregions and 17 of the 21 major catchments. However, concordance was not consistently indicative of significant relationships between rates of fish and invertebrate taxa loss at those same scales. 3. Fish and invertebrate communities were largely associated with different environmental variables, although the composition of both communities was strongly correlated with stream size across all three scales. 4. Predictive taxa‐loss models for fish assemblages were less sensitive and precise than models for invertebrate assemblages, likely because of the relatively low number of common fish taxa in our data set. Both models, however, distinguished reference from non‐reference sites. 5. The importance of concordance, geographic context and scale are discussed in relation to the design and interpretation of stream integrity indicators. In particular, our findings suggest that community concordance should not be viewed as a substitute for an evaluation of how assemblages respond to environmental stressors.     

The influence of different spatial‐scale variables on caddisfly assemblages in Flemish lowland streams

1. Patterns in species assemblages are the result of the combined influence of processes acting on different spatial scales. Various studies describe the distribution of macroinvertebrate communities and their relationship with environmental factors at different geographical scales, but only a few of these studies concentrate on Western European lowlands. 2. Using Flanders as representative for the densely populated Western‐European lowlands, the specific aims of this study are: (i) to identify the different trichopteran species assemblages and to characterise them biologically using indicator species; (ii) to determine which environmental gradients most influence the observed species assemblages; and (iii) to analyse the relative importance of different spatial scale variables in constraining the Trichoptera distributions. 3. Assessment of the main environmental gradients suggested that the absence of Trichoptera from certain locations was mainly due to elevated nutrient concentrations and lower oxygen contents, confirming their sensitivity to anthropogenic disturbance. 4. Five Trichoptera species assemblages were distinguished based on Bray–Curtis dissimilarity coefficients. These assemblages did not differ significantly in species richness, but a shift in stream zonation preference was observed. In the ordination analysis 11 variables that were selected using a stepwise model building function manifested themselves as upstream–downstream and size‐related gradients. The Trichoptera assemblages in lowland streams thus appear to follow a longitudinal succession pattern that corresponds with the species‐specific preferences. 5. Partitioning the variance over the different spatial scales indicated that the reach‐scale variables were far more important in explaining the variation in species composition. The study design, which limited the minimum–maximum range of catchment‐scale characteristics, however, may have led to an overestimation of the impact of the local‐scale variables.     

Relative influence of variables at multiple spatial scales on stream macroinvertebrates in the Northern Lakes and Forest ecoregion, U.S.A.

• 1 We used 94 sites within the Northern Lakes and Forests ecoregion spanning Minnesota, Wisconsin and Michigan to identify environmental variables at the catchment, reach and riparian scales that influence stream macroinvertebrates. Redundancy analyses (RDA) found significantly influential variables within each scale and compared their relative importance in structuring macroinvertebrate assemblages.
• 2 Environmental variables included landcover, geology and groundwater delivery estimates at the catchment scale, water chemistry, channel morphology and stream habitat at the reach scale, and landcover influences at three distances perpendicular to the stream at the riparian scale. Macroinvertebrate responses were characterised with 22 assemblage attributes, and the relative abundance and presence/absence of 66 taxa.
• 3 Each scale defined macroinvertebrates along an erosional to depositional gradient. Wisconsin's macroinvertebrate index of biotic integrity, Ephemeroptera–Plecoptera–Trichoptera taxa and erosional taxa corresponded with forest streams, whereas organic pollution tolerant, Chironomidae and depositional taxa corresponded with wetland streams. Reach scale analyses defined the gradient similarly as dissolved oxygen and wide, shallow channels (erosional) opposed instream macrophytes and pool habitats (depositional). Riparian forests within 30 m of the stream coincided with an erosional assemblage and biotic integrity.
• 4 Next, we combined all significant environmental variables across scales to compare the relative influence of each spatial scale on macroinvertebrates. Partial RDA procedures described how much of the explained variance was attributable to each spatial scale and each interrelated scale combination.
• 5 Our results appeared consistent with the concept of hierarchical functioning of scale in which large‐scale variables restrict the potential for macroinvertebrate traits or taxa at smaller spatial scales. Catchment and reach variables were equally influential in defining assemblage attributes, whereas the reach scale was more influential in determining relative abundance and presence/absence.
• 6 Ultimately, comprehending the relative influence of catchment and reach scale properties in structuring stream biota will assist prioritising the scale at which to rehabilitate, manage and derive policies for stream ecosystem integrity.

     

The influence of stream invertebrate composition at neighbouring sites on local assemblage composition

1. The composition of freshwater invertebrate assemblages at a location is determined by a range of physico‐chemical and biotic factors in the local environment, as well as larger‐scale spatial factors such as sources of recruits. We assessed the relative importance of the species composition of local neighbourhoods and proximal environmental factors on the composition of invertebrate assemblages. 2. Macroinvertebrate assemblages were sampled at 188 running‐water sites in the catchment of the River Rede, north‐east England. A total of 176 species were recorded. 3. Environmental data, in the form of 13 biotic and abiotic measurements that described stream physical structure, aquatic vegetation and water characteristics, were recorded for each site. Detrended correspondence analysis was then used to simplify nine of these stream environmental variables to create an index of stream structure. 4. The species composition of the invertebrate assemblages was related to the environmental variables, using an information theoretic approach. The impact of the species composition of neighbouring sites on each site was determined using Moran's I and autoregressive modelling techniques. 5. Species composition was primarily associated with water pH and stream structure. The importance of the species composition of neighbouring sites in determining local species assemblages differed markedly between taxa. The autoregressive component was low for Coleoptera, intermediate for Trichoptera and Plecoptera, and high for Ephemeroptera. 6. We hypothesise that the observed differences in the autoregressive component amongst these orders reflects variation in their dispersal abilities from neighbouring sites.     

Stressor prioritisation in riverine ecosystems: Which environmental factors shape benthic invertebrate assemblage metrics?

Aquatic ecosystems are amongst the most heavily altered ecosystems and exhibit a disproportional loss of biodiversity. Numerous stressors, such as nutrient enrichment, contaminant pollution, sedimentation and alterations in stream hydrology and habitat structure, account for these losses. Understanding these forces is of utmost importance to prevent riverine ecosystems from further deterioration and to provide helpful insights for restoration practices. In the present study, we analyse the response of biological indicators to a large number of environmental factors. For this, benthic invertebrate assemblages from 83 sites in Germany were described based on 25 metrics from four different metric types. The condition of the sites was described using 27 environmental factors: 13 for water quality, 4 for land use in the catchment and 10 for local scale habitat structure. The relative importance of single environmental predictors or predictor combinations for benthic invertebrate assemblages was analysed with single and multiple linear regression models. The results for the latter models were statistically supported via a bootstrap approach. The models revealed the importance of water quality and catchment-scale land use in explaining benthic invertebrate assemblages; in particular, chloride, oxygen, total organic carbon and the amounts of artificial surfaces and arable land were the most important predictors. Models including solely structural variables such as plan form, bank structures and substrate diversity had lower goodness of fit values than those for other variables. Regarding the four different assemblage metric types, functional metrics had on average lower goodness of fit values than composition/abundance, richness/diversity and sensitivity/tolerance metrics. Among the richness/diversity metrics, however, the model results for the Shannon–Wiener and Simpson diversity indices and evenness were poor. Our results show that catchment-related factors and water quality were of overriding importance in shaping biodiversity patterns and causing species loss. In contrast, structural degradation at a local scale was not the most significant stressor. This finding might explain why structural restoration at a reach scale often yields a low benefit–cost ratio and may be considered to represent inappropriate investment prioritisation.     

Biome transitions as centres of diversity: habitat heterogeneity and diversity patterns of West African bat assemblages across spatial scales

It is widely accepted that species diversity is contingent upon the spatial scale used to analyze patterns and processes. Recent studies using coarse sampling grains over large extents have contributed much to our understanding of factors driving global diversity patterns. This advance is largely unmatched on the level of local to landscape scales despite being critical for our understanding of functional relationships across spatial scales. In our study on West African bat assemblages we employed a spatially explicit and nested design covering local to regional scales. Specifically, we analyzed diversity patterns in two contrasting, largely undisturbed landscapes, comprising a rainforest area and a forest‐savanna mosaic in Ivory Coast, West Africa. We employed additive partitioning, rarefaction, and species richness estimation to show that bat diversity increased significantly with habitat heterogeneity on the landscape scale through the effects of beta diversity. Within the extent of our study areas, habitat type rather than geographic distance explained assemblage composition across spatial scales. Null models showed structure of functional groups to be partly filtered on local scales through the effects of vegetation density while on the landscape scale both assemblages represented random draws from regional species pools. We present a mixture model that combines the effects of habitat heterogeneity and complexity on species richness along a biome transect, predicting a unimodal rather than a monotonic relationship with environmental variables related to water. The bat assemblages of our study by far exceed previous figures of species richness in Africa, and refute the notion of low species richness of Afrotropical bat assemblages, which appears to be based largely on sampling biases. Biome transitions should receive increased attention in conservation strategies aiming at the maintenance of ecological and evolutionary processes.     

Additive partitioning of plant diversity with respect to grassland management regime,fertilisation and abiotic factors

Understanding patterns of vascular plant diversity in managed temperate grasslands and the processes that determine them requires analyses at multiple spatial scales. In this study, we applied additive partitioning to plant species richness data of two contrasting management regimes (meadow vs. pasture) collected from a nested sampling design that consisted of two hierarchical scales. At the local scale, we quantified additive diversity components in 180 vegetation plots, and at the regional scale in 60 grassland parcels. Total observed regional species richness (γ_r) was partitioned into its additive components within (α_l) and among vegetation plots (β_l) and among grassland parcels (β_r). We used the same approach in a comparison of common and infrequent plant species. Partitioning analyses revealed that the relative contributions of diversity components to total observed regional species richness changed as a function of spatial scale. We found that species richness among grassland parcels (β_r) of both meadows and pastures contributed most to total observed regional species richness (γ_r) of all and infrequent plant species (up to 81% and 96%, respectively), whereas for common species only up to 51% of γ_r were attributable to species richness among grassland parcels (β_r). To gain insight into the processes that may affect local patterns of species richness in grasslands, we analysed the observed local species diversity components with respect to management regime, nitrogen fertilisation and abiotic environmental factors (slope angle and soil quality). Our results show that grazing at a low-to-moderate stocking density promotes the β-diversity of all plant species at the local scale due to increased within-habitat heterogeneity. Low application rates of nitrogen fertilisers and abiotic environmental conditions such as steep slopes and soils with a low nutrient status generally benefited local species diversity components. We conclude that the observed patterns of plant species diversity are shaped by processes at multiple spatial scales.     

Taxonomic and functional homogenisation of macroinvertebrate communities in recently intermittent Alpine watercourses

1. Mountain streams in southwestern European Alps are currently shifting from perennial to intermittent flow due to the combined effects of climate change and local anthropogenic pressures. Given that flow intermittency is a recently documented phenomenon in the Alps, only scattered studies have investigated functional and taxonomical diversity of benthic invertebrate communities in recently intermittent Alpine streams.
2. We used a hierarchical sampling design to investigate patterns in taxonomic and functional diversity of benthic invertebrate communities in 13 recently intermittent Alpine streams in north-west Italy. in April 2017, we sampled benthic communities in two reaches of each stream with different hydrological conditions: a control reach, with permanent flow; and an intermittent reach, which recently experienced non-flow periods in summer.
3. We tested for the response of taxonomic richness at multiple spatial scales by partitioning total diversity into the average richness of local communities and the richness due to variation among local communities both within and among reaches. By partitioning total diversity (γ) into its local (α) and turnover (β) components we showed a decrease in local and regional species richness both within and among reaches, whereas variation among communities was significantly lower in intermittent reaches at the reach scale only.
4. The analysis of multidimensional trait space of macroinvertebrate communities in reaches with different hydrological conditions revealed a significant reduction of functional diversity, dispersion, and evenness in intermittent reaches. There was trait overdispersion in intermittent reaches, as these hosted both typical Alpine taxa and organisms adapted to flow intermittency. In particular, we observed the replacement of taxa with aquatic respiration and those preferring medium- to fast-flowing oligotrophic waters by taxa adapted to lentic habitats, air breathing and with larval dormancy phases.
5. These results indicate that recent flow intermittency has caused drastic changes in benthic invertebrate communities in Alpine streams. Our work highlights the importance of integrating taxonomic and functional diversity to thoroughly assess the impacts of flow intermittency.

     

Influences of catchment and corridor imperviousness on urban stream macroinvertebrate communities at multiple spatial scales

Resolving land cover hierarchy relationships in urban settings is important for defining the scale and type of management required to enhance stream health. We investigated associations between macroinvertebrate assemblages in urban streams of Hamilton, New Zealand, and environmental variables measured at multiple spatial scales comprising (i) local-scale physicochemical conditions, (ii) impervious area in multiple stream corridor widths (30, 50 and 100 m) along segments (sections of stream between tributary nodes) and for entire upstream networks, and (iii) total impervious area in stream segment sub-catchments and upstream catchments. Imperviousness was higher for stream segment sub-catchments than for entire catchments because of the agricultural headwaters of some urban streams. Imperviousness declined as corridor width declined at both segment and catchment scales reflecting the vegetated cover along most urban stream gullies. Upstream catchment imperviousness was strongly and inversely correlated with dissolved organic carbon concentration, whereas segment and upstream corridor scales were correlated with water temperature and pH. Corridor imperviousness appeared to be a stronger predictor than catchment imperviousness of Ephemeroptera, Plecoptera and Trichoptera taxa richness and the Quantitative Urban Community Index specifically developed to assess impacts of urbanisation. In contrast, imperviousness at all measured scales added only marginal improvement in assemblage-based models over that provided by the local-scale physicochemical variables of reach width, habitat quality, macrophyte cover, pH and dissolved oxygen concentration. These findings infer variable scales of influence affecting macroinvertebrate communities in urban streams and suggest that it may be important to consider local and corridor factors when determining mechanisms of urbanisation impacts and potential management options.     

Functional diversity and trait–environment relationships of stream fish assemblages in a large tropical catchment

1. The species composition of stream fish assemblages changes across the longitudinal fluvial gradient of large river basins. These changes may reflect both zonation in species distributions and environmental filtering of fish traits as stream environments change from the uplands to the lowlands of large catchments. Previous research has shown that taxonomic diversity generally increases in larger, lowland streams, and the River Continuum Concept, the River Habitat Template and other frameworks have provided expectations for what functional groups of fishes should predominate in certain stream types. However, studies addressing the functional trait composition of fish assemblages across large regions are lacking, particularly in tropical river basins. 2. We examined functional trait–environment relationships and functional diversity of stream fish assemblages in the Río Grijalva Basin in southern Mexico. Traits linked to feeding, locomotion and life history strategy were measured in fishes from streams throughout the catchment, from highland headwaters to broad, lowland streams. Relationships between functional traits and environmental variables at local and landscape scales were examined using multivariate ordination, and the convex hull volume of trait space occupied by fish assemblages was calculated as a measure of functional diversity. 3. Although there were a few exceptions, functional diversity of assemblages increased with species richness along the gradient from uplands to lowlands within the Grijalva Basin. Traits related to swimming, habitat preference and food resource use were associated with both local (e.g. substratum type, pool availability) and landscape‐scale (e.g. forest cover) environmental variables. 4. Along with taxonomic structure and diversity, the functional composition of fish assemblages changed across the longitudinal fluvial gradient of the basin. Trait–environment relationships documented in this study partially confirmed theoretical expectations and revealed patterns that may help in developing a better understanding of general functional responses of fish assemblages to environmental change.     

Additive partitioning of aquatic invertebrate species diversity across multiple spatial scales

1. Additive partitioning of three measures of diversity (species richness, Shannon's diversity index H and Simpson's diversity D) was used to study the relationship between local and regional diversity of benthic macroinvertebrate communities of boreal lakes (littoral habitats) and streams (riffle habitats) across three spatial scales (sampling sites, ecoregions and biogeographic regions). 2. Alpha (α) and beta (β) diversity are defined as within‐habitat and between‐habitat diversity, respectively. According to the concept of additive partitioning, diversity can be partitioned across multiple spatial scales such that the total (γ) diversity on one spatial scale becomes within‐habitat (α) diversity at the next higher scale. Hence, the total diversity at one scale is determined by the α diversity and the between‐habitat diversity (β) at the next lower scale. Consequently, one of the advantages of additive partitioning is that it is possible to study simultaneously β diversity and the regional‐local species relationship and the scale dependence of α and β components. 3. For both lakes and streams α diversity was low for sites and ecoregions, whereas β diversity was high, indicating that among‐site factors are important in describing the variability among the lakes and streams studied here. 4. Weak, albeit significant, evidence was found for regional and local species saturation patterns. Multiple stepwise regression indicated that local processes might be more important in structuring lake‐littoral and stream‐riffle species assemblages than regional processes. From these results we conclude that environmental heterogeneity may act as an important factor contributing to species coexistence, resulting in the observed saturation patterns. 5. Our study supports the use of additive partitioning for identifying specific patterns of macroinvertebrate diversity on multiple spatial scales and the underlying processes generating these patterns. This information is needed to improve understanding of the relation between patterns and processes affecting (decreasing) trends in aquatic biodiversity.     

Modelling and mapping the distribution,diversity and abundance of freshwater mussels (Family Unionidae) in wadeable streams of Illinois,U.S.A.

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Global patterns of specialization and coexistence in bird assemblages

Aim Increased specialization has been hypothesized to facilitate local coexistence and thus high species richness, but empirical evaluations of the richness–specialization relationships have been relatively scant. Here, we provide a first assessment of this relationship for terrestrial bird assemblages at global extent and from fine to coarse grains. Location World‐wide. Methods We use two indices of specialization that describe species‐level resource use: diet and habitat specialization. The relationship between richness and mean assemblage‐level specialization was independently assessed at realm, biome‐realm, 12,100 km² equal‐area grid cells and fine‐grained scales. To identify assemblages that are diverse relative to environmental conditions we: (1) applied quantile regressions, (2) statistically accounted for other environmental variables which may constrain richness, and (3) parsed the data according to the residuals of a model relating species richness to the environmental variables. Results Assemblage species richness increases with both measures of specialization at all scales. Statistically, richness appears constrained by levels of specialization, with the highest richness values only found in specialized assemblages. Richness is positively associated with specialization even after accounting for gradients in resource availability. Net primary productivity and assemblage specialization have complementary statistical effects on assemblage species richness. Contrary to expectations based on niche partitioning of local resources, the relationship between specialization and richness is steep even at coarse scales. Main conclusions The results demonstrate that for an entire clade, totalling > 9000 species, specialization and species richness are related, at least for diverse assemblages. The strong patterns observed across scales suggest that this relationship does not solely originate from (1) limits on coexistence in present‐day assemblages, or (2) increased specialization in richer assemblages imposed by species’ abilities to partition ecological space. Instead, regional‐scale influences on the species pool may determine much of the observed relationship between richness and specialization. Although causal attribution is not straightforward, these findings support the idea that, for the scale of our analysis, specialization may be related to the past origination of high‐diversity assemblages, rather than their contemporary assembly.