The effect of habitat type on speciation rates and range movements in aquatic beetles: inferences from species-level phylogenies

Most aquatic beetles in the family Dytiscidae are tightly associated either with running (lotic) or stagnant (lentic) water bodies. The range size of lotic species is known to be, on average, much smaller than that of lentic species, presumably as a result of differences in dispersal strategies in each habitat type. We explored possible effects of these differences on clade evolution and speciation rates by comparing species-level phylogenies based on cytochrome oxidase I (COI) and 16S rRNA mitochondrial genes for two genera, the lentic Ilybius and the lotic Deronectes. The expectation that species turnover is higher in lotic lineages due to their lower dispersal propensity compared to lentic species was not strongly supported. Deronectes displays a higher frequency of recent splits than Ilybius, consistent with the hypothesis, but the difference was not significant compared to expected patterns under a constant speciation rate null model. Similarly, when the degree of sympatry was plotted against relative node age, more allopatric splits were evident in the lentic Deronectes, suggesting a slower rate of range movement since speciation, but the differences were not significant. We discuss two explanations for our failure to detect differences between the two clades. First, current methods for analysing species-level phylogenies may be sensitive to taxonomic and sampling artefacts. Second, lentic and lotic clades may indeed display similar levels of species turnover despite occupying very different habitats at different spatial scales. More work is needed to investigate the effects of population level processes and spatial scale on macroevolutionary dynamics.     

Parallel habitat-driven differences in the phylogeographical structure of two independent lineages of Mediterranean saline water beetles

It has been hypothesized that species living in small lentic water bodies, because of the short-term geological persistence of their habitat, should show higher dispersal ability, with increased gene flow among populations and a less pronounced phylogeographical structure. Conversely, lotic species, living in more geologically stable habitats, should show reduced dispersal and an increased phylogeographical structure at the same geographical scales. In this work we tested the influence of habitat type in two groups of aquatic Coleoptera ( Nebrioporus ceresyi and Ochthebius notabilis groups, families Dytiscidae and Hydraenidae respectively), each of them with closely related species typical of lotic and lentic saline Western Mediterranean water bodies. We used mitochondrial cox1 sequence data of 453 specimens of 77 populations through the range of nine species to compare a lotic vs. a lentic lineage in each of the two groups. Despite the differences in biology (predators vs. detritivorous) and evolutionary history, in both lotic lineages there was a higher proportion of nucleotide diversity among than within groups of populations, and a faster rate of accumulation of haplotype diversity (as measured by rarefaction curves) than in the lentic lineages. Similarly, lotic lineages had a higher absolute phylogenetic diversity, more remarkable considering their smaller absolute geographical ranges. By comparing closely related species, we were able to show the effect of contrasting habitat preferences in two different groups, in agreement with predictions derived from habitat stability.     

Dispersal ability rather than ecological tolerance drives differences in range size between lentic and lotic water beetles (Coleoptera: Hydrophilidae)

Aim In aquatic ecosystems, standing (lentic) and running (lotic) waters differ fundamentally in their stability and persistence, shaping the comparative population genetic structure, geographical range size and speciation rates of lentic versus lotic lineages. While the drivers of this pattern remain incompletely understood, the suite of traits making up the ability of a species to establish new populations is instrumental in determining such differences. Here we explore the degree to which the association between habitat type and geographical range size results from differences in dispersal ability or fundamental niche breadth in the members of the Enochrus bicolor complex, an aquatic beetle clade with species across the lentic–lotic divide. Location Western Mediterranean, with a special focus on North Africa, the Iberian Peninsula and Sicily. Methods DNA sequences for four loci were obtained from species of the E. bicolor complex and analysed using phylogenetic inference. Dispersal and establishment abilities were assessed in lentic–lotic species pairs of the complex, using flight wing morphometrics and thermal tolerance ranges as surrogates, respectively. Results There were clear differences in range size between the lotic and lentic taxa of the complex, which appears to have had a lotic origin with two transitions to standing waters. Only small differences were observed in temperature tolerance and acclimation ability between the two lotic–lentic sister species studied. By contrast, wing morphometrics revealed clear, consistent differences between lotic and lentic Enochrus species pairs, the latter having a higher dispersal capacity. Main conclusions We hypothesize that there have been two habitat shifts from lotic to lentic waters, which have allowed marked expansions in geographical range size in western Mediterranean species of the E. bicolor complex. Differences in dispersal rather than in establishment ability appear to underlie differences in geographical range extent, as transitions to lentic waters were associated with changes in wing morphology, but not in thermal tolerance range. In this lineage of water beetles, selection for dispersal in geologically short‐lived lentic systems has driven the evolution of larger range sizes in lentic taxa compared with those of their lotic relatives.     

Temperature‐driven color lightness and body size variation scale to local assemblages of European Odonata but are modified by propensity for dispersal

1. Previous macrophysiological studies suggested that temperature‐driven color lightness and body size variations strongly influence biogeographical patterns in ectotherms. However, these trait–environment relationships scale to local assemblages and the extent to which they can be modified by dispersal remains largely unexplored. We test whether the predictions of the thermal melanism hypothesis and the Bergmann's rule hold for local assemblages. We also assess whether these trait–environment relationships are more important for species adapted to less stable (lentic) habitats, due to their greater dispersal propensity compared to those adapted to stable (lotic) habitats.
2. We quantified the color lightness and body volume of 99 European dragon‐ and damselflies (Odonata) and combined these trait information with survey data for 518 local assemblages across Europe. Based on this continent‐wide yet spatially explicit dataset, we tested for effects temperature and precipitation on the color lightness and body volume of local assemblages and assessed differences in their relative importance and strength between lentic and lotic assemblages, while accounting for spatial and phylogenetic autocorrelation.
3. The color lightness of assemblages of odonates increased, and body size decreased with increasing temperature. Trait–environment relationships in the average and phylogenetic predicted component were equally important for assemblages of both habitat types but were stronger in lentic assemblages when accounting for phylogenetic autocorrelation.
4. Our results show that the mechanism underlying color lightness and body size variations scale to local assemblages, indicating their general importance. These mechanisms were of equal evolutionary significance for lentic and lotic species, but higher dispersal ability seems to enable lentic species to cope better with historical climatic changes. The documented differences between lentic and lotic assemblages also highlight the importance of integrating interactions of thermal adaptations with proxies of the dispersal ability of species into trait‐based models, for improving our understanding of climate‐driven biological responses.

     

Not going with the flow: a comprehensive time‐calibrated phylogeny of dragonflies (Anisoptera: Odonata: Insecta) provides evidence for the role of lentic habitats on diversification

Ecological diversification of aquatic insects has long been suspected to have been driven by differences in freshwater habitats, which can be classified into flowing (lotic) waters and standing (lentic) waters. The contrasting characteristics of lotic and lentic freshwater systems imply different ecological constraints on their inhabitants. The ephemeral and discontinuous character of most lentic water bodies may encourage dispersal by lentic species in turn reducing geographical isolation among populations. Hence, speciation probability would be lower in lentic species. Here, we assess the impact of habitat use on diversification patterns in dragonflies (Anisoptera: Odonata). Based on the eight nuclear and mitochondrial genes, we inferred species diversification with a model‐based evolutionary framework, to account for rate variation through time and among lineages and to estimate the impact of larval habitat on the potentially nonrandom diversification among anisopteran groups. Ancestral state reconstruction revealed lotic fresh water systems as their original primary habitat, while lentic waters have been colonized independently in Aeshnidae, Corduliidae and Libellulidae. Furthermore, our results indicate a positive correlation of speciation and lentic habitat colonization by dragonflies: speciation rates increased in lentic Aeshnidae and Libellulidae, whereas they remain mostly uniform among lotic groups. This contradicts the hypothesis of inherently lower speciation in lentic groups and suggests species with larger ranges are more likely to diversify, perhaps due to higher probability of larger areas being dissected by geographical barriers. Furthermore, larger range sizes may comprise more habitat types, which could also promote speciation by providing additional niches, allowing the coexistence of emerging species.     

不同生境间龟鳖类体型的差异

体型是动物重要的形态特征,影响动物的生境利用。为揭示龟鳖类体型与生境之间的关系,通过文献收集331种龟鳖(龟鳖目Tesudines总物种数的98.8%)的最大背甲长及其生境信息,将生境分为海洋、淡水、岛屿性陆地和大陆性陆地4种类型,再将淡水生境分为大静水、大流水、小静水、小流水和所有水域5种亚类型,大陆性陆地生境分为高地、平地和荒漠3种亚类型,从而比较不同生境类型或亚类型之间龟鳖类体型的差异。广义线性混合模型分析结果显示:1)海龟体型最大,岛屿性陆龟次之,淡水龟鳖和大陆性陆龟体型最小,且后两者差异无统计学意义。2)淡水龟鳖类的体型在5种亚类型生境间存在差异,大静水和大流水水域的体型均显著大于小静水和小流水水域,而体型在大静水与大流水水域、小静水与小流水水域之间的差异均无统计学意义,表明淡水龟鳖类体型与水域面积有关,而与水域是静水或流水无关。广布所有水域的淡水龟鳖类体型趋于中间型,且与其他4种亚类型生境中的体型之间的差异均无统计学意义。3)大陆性陆龟的体型从高地到平地再到荒漠有逐渐变大的趋势,但差异无统计学意义。本研究揭示龟鳖类的保护对策需要考虑其体型和生境面积的相关性。     

Phylogeography and population genomics of a lotic water beetle across a complex tropical landscape

The habitat template concept applied to a freshwater system indicates that lotic species, or those which occupy permanent habitats along stream courses, are less dispersive than lentic species, or those that occur in more ephemeral aquatic habitats. Thus, populations of lotic species will be more structured than those of lentic species. Stream courses include both flowing water and small, stagnant microhabitats that can provide refuge when streams are low. Many species occur in these microhabitats but remain poorly studied. Here, we present population genetic data for one such species, the tropical diving beetle Exocelina manokwariensis (Dytiscidae), sampled from six localities along a ~300 km transect across the Birds Head Peninsula of New Guinea. Molecular data from both mitochondrial (CO1 sequences) and nuclear (ddRAD loci) regions document fine‐scale population structure across populations that are ~45 km apart. Our results are concordant with previous phylogenetic and macroecological studies that applied the habitat template concept to aquatic systems. This study also illustrates that these diverse but mostly overlooked microhabitats are promising study systems in freshwater ecology and evolutionary biology. With the advent of next‐generation sequencing, fine‐scale population genomic studies are feasible for small nonmodel organisms to help illuminate the effect of habitat stability on species’ natural history, population structure and geographic distribution.     

Evolution of territoriality in Hylinae treefrogs: Ecological and morphological correlates and lineage diversification

Given the diverse nature of traits involved in territorial defence, they may respond to different selective pressures and then exhibit distinct patterns of evolution. These selective pressures also may cause territorial behaviour to be associated with environmental and morphological variables. Such associations, however, have mostly been studied at the intraspecific level, being phylogenetic analyses of territoriality in a broad taxonomic framework rare in the literature. We used the anuran subfamily Hylinae to test (1) whether two territorial-behaviour traits with different levels of aggression—territorial call and physical combat—are evolutionarily more labile than a morphological trait used in physical combat—the spine-shaped prepollex; (2) whether reproduction in lentic waters and phytotelmata, as well as resource scarcity, might favour the occurrence of territoriality; (3) if physical combat is more important than territorial call for the evolution of body size and sexual size dimorphism and (4) the relationships between territorial-behaviour traits and lineage diversification. We mainly used the literature to build two datasets with different levels of certainty. Territorial-behaviour traits exhibited intermediate levels of phylogenetic signal in Hylinae, whereas the phylogenetic signal for the presence of the spine-shaped prepollex was strong. We found support for the hypothesis that reproduction in lentic water favours the occurrence of territorial behaviour, because the expression of territorial-behaviour traits was more associated with reproduction in lentic than in lotic waters. Territorial-behaviour traits were not correlated with annual precipitation nor with habitat complexity. Body size and sexual size dimorphism were not correlated with the presence of territorial call nor with physical combat. We identified negative correlations between diversification rates and physical combat. Relationships of territorial call and physical combat with diversification rates suggest that these territorial behaviours influence evolutionary processes in different ways.     

Habitat availability does not explain the species richness patterns of European lentic and lotic freshwater animals

Aim In Europe, the relationships between species richness and latitude differ for lentic (standing water) and lotic (running water) species. Freshwater animals are highly dependent on suitable habitat, and thus the distribution of available habitat should strongly influence large‐scale patterns of species richness. We tested whether habitat availability can account for the differences in species richness patterns between European lentic and lotic freshwater animals. Location Europe. Methods We compiled occurrence data of 1959 lentic and 2445 lotic species as well as data on the amount of lentic and lotic habitats across 25 pre‐defined biogeographical regions of European freshwaters. We used the range of elevation of each region as a proxy for habitat diversity. We investigated the relationships between species richness, habitat availability and habitat diversity with univariate and multiple regression analyses. Results Species richness increased with habitat availability for lentic species but not for lotic species. Species richness increased with elevational range for lotic species but decreased for lentic species. For both groups, neither habitat availability nor diversity could account for previously reported latitudinal patterns in species richness. For lotic species, richness declined with latitude, whereas there was no relationship between habitat availability and latitude. For lentic species, richness showed a hump‐shaped relationship with latitude, whereas available habitat increased with latitude. Main conclusions Habitat availability and diversity are poor predictors of species richness of the European freshwater fauna across large scales. Our results indicate that the distributions of European freshwater animals are probably not in equilibrium and may still be influenced by history, namely the recurrent European glaciations and possible differences in post‐glacial recolonization. The distributions of lentic species appear to be closer to equilibrium than those of lotic species. This lends further support to the hypothesis that lentic species have a higher propensity for dispersal than lotic species.     

An interspecific test of Bergmann's rule reveals inconsistent body size patterns across several lineages of water beetles (Coleoptera: Dytiscidae)

1. Bergmann's rule sensu lato, the ecogeographic pattern relating animals' body size with environmental temperature (or latitude), has been shown to be inconsistent among insect taxa. Body size clines remain largely unexplored in aquatic insects, which may show contrasting patterns to those found in terrestrial groups because of the physiological or mechanical constraints of the aquatic environment. 2. Bergmann's rule was tested using data on body size, phylogeny and distribution for 93 species belonging to four lineages of dytiscid water beetles. The relationship between size and latitude was explored at two taxonomic resolutions – within each independent lineage, and for the whole dataset – employing phylogenetic generalised least‐squares to control for phylogenetic inertia. The potential influence of habitat preference (lotic versus lentic) on body size clines was also considered. 3. Within‐lineage analyses showed negative relationships (i.e. converse Bergmann's rule), but only in two lineages (specifically in those that included both lotic and lentic species). By contrast, no relationship was found between body size and latitude for the whole dataset. 4. These results suggest that there may be no universal interspecific trends in latitudinal variation of body size in aquatic insects, even among closely related groups, and show the need to account for phylogenetic inertia. Furthermore, habitat preferences should be considered when exploring latitudinal clines in body size in aquatic taxa at the interspecific level.     

Habitat type predicts genetic population differentiation in freshwater invertebrates

A basic challenge in evolutionary biology is to establish links between ecology and evolution of species. One important link is the habitat template. It has been hypothesized, that the spatial and temporal settings of a habitat strongly influence the evolution of species dispersal propensity. Here, we evaluate the importance of the habitat type on genetic population differentiation of species using freshwater habitats as a model system. Freshwater habitats are either lentic (standing) or lotic (running). On average, lotic habitats are more stable and predictable over space and time than lentic habitats. Therefore, lentic habitats should favour the evolution of higher dispersal propensity which ensures population survival of lentic species. To test for such a relationship, we used extensive data on species' genetic population differentiation of lentic and lotic freshwater invertebrates retrieved from published allozyme studies. Overall, we analysed more than 150 species from all over the world. Controlling for several experimental, biological and geographical confounding effects, we always found that lentic invertebrates exhibit, on average, lower genetic population differentiation than lotic species. This pattern was consistent across insects, crustaceans and molluscs. Our results imply fundamental differences in genetic population differentiation among species adapted to either lentic or lotic habitats. We propose that such differences should occur in a number of other habitat types that differ in spatio-temporal stability. Furthermore, our results highlight the important role of lotic habitats as reservoirs for evolutionary processes and the potential for rapid speciation.     

Repeated evolution of terrestrial lineages in a continental lizard radiation

The “early‐burst” model of adaptive radiation predicts an early increase in phenotypic disparity concurrent with lineage diversification. Although most studies report a lack of this coupled pattern, the underlying processes are not identified. The continental radiation of Hemidactylus geckos from Peninsular India includes morphologically diverse species that occupy various microhabitats. This radiation began diversifying ~36 Mya with an early increase in lineage diversification. Here, we test the “early‐burst” hypothesis by investigating the presence of ecomorphs and examining the pattern of morphological diversification in a phylogenetic framework. Two ecomorphs—terrestrial and scansorial species—that vary significantly in body size and toepad size were identified. Unlike the prediction of the “early‐burst” model, we find that disparity in toepad morphology accumulated more recently ~14 Mya and fit the Ornstein‐Ulhenbeck model. Ancestral state reconstruction of the two ecomorphs demonstrates that terrestrial lineages evolved independently at least five times from scansorial ancestors, with the earliest diversification in terrestrial lineages 19–12 Mya. Our study demonstrates a delayed increase in morphological disparity as a result of the evolution of terrestrial ecomorphs. The diversification of terrestrial lineages is concurrent with the establishment of open habitat and grasslands in Peninsular India, suggesting that the appearance of this novel resource led to the adaptive diversification.     

Habitat type mediates equilibrium with climatic conditions in the distribution of Iberian diving beetles

Aim The contrasting habitat permanence over geological time‐scales of lotic and lentic habitats may impose different constraints on the dispersal ability of their macroinvertebrate populations, and ultimately on the degree of equilibrium with current climate. We aim to test for differences between species typical of either habitat type in their potential versus realized distributions as a surrogate measure of degree of climate equilibrium, both in refuges and more recently deglaciated areas. Location Western Europe. Methods We focus on 99 Iberian diving beetles (family Dytiscidae). A multidimensional envelope procedure was used to estimate their potential distributions, which were projected for different spatial scales. At the continental scale we calculated the percentage of countries with climatically suitable conditions for each species over those actually occupied (range filling). At the regional scale, we estimated realized distributions using: (1) convex hull polygons for Sweden and the Iberian Peninsula; and (2) generalized linear models for the Iberian Peninsula. Results In the Iberian Peninsula, differences in the degree of equilibrium with climatic conditions between lotic and lentic species were few, if any. However, at the continental scale we found significant differences, with lentic species closer to equilibrium than lotic species. In the recently deglaciated area (Sweden) the subset of species with ranges wide enough to encompass Iberia and Scandinavia were mostly lentic, and all were closer to climatic equilibrium without significant differences between habitat types. Main conclusions Our results show that, at continental scales, climate equilibrium is not concordant between the habitat types across western Europe. We hypothesize that: (1) the differences between refuge areas in dispersal ability are erased probably due to long‐term climatic stability, allowing enough time to reach equilibrium, and (2) the species with wide geographical ranges able to recolonize recently deglaciated areas should have the highest dispersal abilities, and are closer to climatic equilibrium.     

Dispersal is linked to habitat use in 59 species of water beetles (Coleoptera: Adephaga) on Madagascar

Lentic habitats (standing water, such as ponds and lakes) differ from lotic habitats (running water; streams and rivers) in their spatiotemporal persistence, with lentic habitats being more ephemeral in evolutionary time. This habitat instability is thought to select for dispersal, and several phylogenetic and macroecological studies have suggested that high rates of dispersal are more characteristic of lentic than lotic species. We tested this hypothesis using a comparative population genetic and phylogeographic approach based on mitochondrial DNA for 59 aquatic beetle species, sampled across Madagascar. Species were classified as lotic (n = 25), lentic (n = 25), or lotolentic (associated with both running and standing water; n = 9). Hierarchical population genetic structure (AMOVA), nucleotide diversity (π), and geographic structure were compared among habitat types. Lotic species had significantly greater population structure (Ф_ST = 0.55, hierarchical AMOVA) than lentic (Ф_ST = 0.13) and lotolentic (Ф_ST = 0.19) species using phylogenetic generalized least squares (PGLS) to correct for phylogeny. Body size was independent of habitat preference, and did not explain any of the intraspecific variation. A greater proportion of lotic species were endemic to Madagascar and lotic species had more pronounced geographic structure in their haplotype networks. The results indicate that dispersal is consistently lower among lotic species, independent of phylogenetic relatedness. This has macroevolutionary and biogeographical consequences for the freshwater fauna of this tropical biodiversity hotspot where remaining natural habitats are becoming increasingly isolated from one another.     

Habitat stability affects dispersal and the ability to track climate change

Habitat persistence should influence dispersal ability, selecting for stronger dispersal in habitats of lower temporal stability. As standing (lentic) freshwater habitats are on average less persistent over time than running (lotic) habitats, lentic species should show higher dispersal abilities than lotic species. Assuming that climate is an important determinant of species distributions, we hypothesize that lentic species should have distributions that are closer to equilibrium with current climate, and should more rapidly track climatic changes. We tested these hypotheses using datasets from 1988 and 2006 containing all European dragon- and damselfly species. Bioclimatic envelope models showed that lentic species were closer to climatic equilibrium than lotic species. Furthermore, the models over-predicted lotic species ranges more strongly than lentic species ranges, indicating that lentic species track climatic changes more rapidly than lotic species. These results are consistent with the proposed hypothesis that habitat persistence affects the evolution of dispersal.     

Lentic odonates have larger and more northern ranges than lotic species

Aim We analysed latitudinal range, centres of distribution and northern range boundaries of dragonflies and damselflies occurring in Europe and North America with respect to larval habitat (standing water = lentic and running water = lotic). As lentic water bodies are thought to be less predictable in space and time than lotic habitats, species adapted to standing waters depend on effective dispersal ability for long‐term survival. If species occurring in lentic habitats have a higher propensity for dispersal, then larger range sizes in lentic than in lotic species, as well as an increase in the proportion of lentic species with latitude, would be expected. Location Europe, North America. Methods Distributional and habitat data were collected from published sources for all odonates of Europe and North America. Species were assigned to lentic and lotic habitats according to the habitat of the larvae. From distribution maps we estimated the latitudinal range, centre of distribution and northern range boundary of each species. Differences in these distribution variables between lentic and lotic species were evaluated using anova . We related the proportion of lentic species by latitudinal interval in Europe, and by political unit (state, province) in North America, to area, altitudinal range, longitude (only for North America) and latitude by means of generalized linear models. Results Lentic damselflies and dragonflies had larger latitudinal spans, and more northern distribution centres and range boundaries, than lotic species. The proportion of lentic species increased with latitude. These findings were consistent between continents. Main conclusions Our results support previous findings that distribution patterns of freshwater species depend on habitat preference. Evolution of dispersal propensity according to habitat characteristics is the most likely explanation. However, at present, alternative explanations, such as an increase in lentic habitats with latitude, cannot be ruled out.     

Stream flow alone does not predict population structure of diving beetles across complex tropical landscapes

Recent theoretical advances have hypothesized a central role of habitat persistence on population genetic structure and resulting biodiversity patterns of freshwater organisms. Here, we address the hypothesis that lotic species, or lineages adapted to comparably geologically stable running water habitats (streams and their marginal habitats), have high levels of endemicity and phylogeographic structure due to the persistent nature of their habitat. We use a nextRAD DNA sequencing approach to investigate the population structure and phylogeography of a putatively widespread New Guinean species of diving beetle, Philaccolilus ameliae (Dytiscidae). We find that P. ameliae is a complex of morphologically cryptic, but geographically and genetically well‐differentiated clades. The pattern of population connectivity is consistent with theoretical predictions associated with stable lotic habitats. However, in two clades, we find a more complex pattern of low population differentiation, revealing dispersal across rugged mountains and watersheds of New Guinea up to 430 km apart. These results, while surprising, were also consistent with the original formulation of the habitat template concept by Southwood, involving lineage‐idiosyncratic evolution in response to abiotic factors. In our system, low population differentiation might reflect a young species in a phase of range expansion utilizing vast available habitat. We suggest that predictions of life history variation resulting from the dichotomy between lotic and lentic organisms require more attention to habitat characterization and microhabitat choice. Our results also underpin the necessity to study fine‐scale processes but at a larger geographical scale, as compared to solely documenting macroecological patterns, to understand ecological drivers of regional biodiversity. Comprehensive sampling especially of tropical lineages in complex and threatened environments such as New Guinea remains a critical challenge.     

The lentic and lotic characteristics of habitats determine the distribution of benthic macroinvertebrates in Mediterranean rivers

1. The importance of flow‐related factors to benthic organisms, as well as the role of habitat conditions in shaping aquatic communities during low‐flow periods, have been recognised. Despite this, the preferences of macroinvertebrates to the ratio of lentic to lotic habitats at the reach scale have not been accurately quantified in most instances.
2. Aquatic invertebrates and habitat features in a range of temporary rivers in Sardinia were investigated. The investigation focused on the flow‐related characteristics that contribute to defining the lentic–lotic condition of the river reaches. The relation of habitat features to benthic taxa distributions was assessed using multidimensional scaling. The main aim of the paper was to quantify the responses of taxa to the different lentic and lotic habitat conditions by applying hierarchical logistic regressions. Finally, taxon optima were aligned along the lentic–lotic gradient and the responses of different taxonomic groups compared.
3. Unbroken waves and imperceptible flow were correlated with benthic taxa variability, suggesting local hydraulics and turbulence have a major role in regulating community composition. The overall lentic–lotic character of the river reaches was also clearly related to the benthic taxa distribution. More than 80% of taxa were significantly related to the lentic–lotic gradient, and an asymmetrical response curve was the predominant model.
4. Benthic groups showed taxon optima clustered in different ranges of the lentic–lotic gradient. Odonata, Coleoptera, Hemiptera, and Mollusca preferred clearly lentic conditions. Diptera mainly ranged on the lotic side of the gradient, while Trichoptera were relatively uniformly distributed across the gradient. Ephemeroptera taxa clustered in intermediate lentic–lotic conditions, with two species preferring extremely lentic habitats. In general, optima converged at intermediate and extremely lentic conditions, presumably due, respectively, to the coexistence of different lentic and lotic features and to the highly diverse environmental characteristics under extremely lentic situations.
5. These results support the conclusion that dissimilar ecological factors act on benthic taxa along the lentic–lotic range and species favouring different lentic–lotic conditions are subjected to pressures of different nature. This should not be ignored when defining species preferences and studying community structure or relationships between species in Mediterranean rivers, which cyclically vary their habitat composition. In addition, the uneven distribution of optima of different groups along the lentic–lotic gradient might affect macroinvertebrate metrics when assessing ecological status or establishing reference conditions under variable climatic conditions.

     

Getting in shape: habitat‐based morphological divergence for two sympatric fishes

Freshwater fishes often show large amounts of body shape variation across divergent habitats and, in most cases, the observed differences have been attributed to the environmental pressures of living in lentic or lotic habitats. Previous studies have suggested a distinct set characters and morphological features for species occupying each habitat under the steady–unsteady swimming performance model. We tested this model and assessed body shape variation using geometric morphometrics for two widespread fishes, Goodea atripinnis (Goodeidae) and Chirostoma jordani (Atherinopsidae), inhabiting lentic and lotic habitats across the Mesa Central of Mexico. These species were previously shown to display little genetic variation across their respective ranges. Our body shape analyses reveal morphometric differentiation along the same axes for both species in each habitat. Both possess a deeper body shape in lentic habitats and a more streamlined body in lotic habitats, although the degree of divergence between habitats was less for C. jordani. Differences in the position of the mouth differed between habitats as well, with both species possessing a more superior mouth in lentic habitats. These recovered patterns are generally consistent with the steady–unsteady swimming model and highlight the significance of environmental forces in driving parallel body shape differences of organisms in divergent habitats. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 114 , 152–162.     

Morphological variation between non‐native lake‐ and stream‐dwelling pumpkinseed Lepomis gibbosusin the Iberian Peninsula

The objective of this study was to test if morphological differences in pumpkinseed Lepomis gibbosus found in their native range (eastern North America) that are linked to feeding regime, competition with other species, hydrodynamic forces and habitat were also found among stream‐ and lake‐ or reservoir‐dwelling fish in Iberian systems. The species has been introduced into these systems, expanding its range, and is presumably well adapted to freshwater Iberian Peninsula ecosystems. The results show a consistent pattern for size of lateral fins, with L. gibbosus that inhabit streams in the Iberian Peninsula having longer lateral fins than those inhabiting reservoirs or lakes. Differences in fin placement, body depth and caudal peduncle dimensions do not differentiate populations of L. gibbosus from lentic and lotic water bodies and, therefore, are not consistent with functional expectations. Lepomis gibbosus from lotic and lentic habitats also do not show a consistent pattern of internal morphological differentiation, probably due to the lack of lotic–lentic differences in prey type. Overall, the univariate and multivariate analyses show that most of the external and internal morphological characters that vary among populations do not differentiate lotic from lentic Iberian populations. The lack of expected differences may be a consequence of the high seasonal flow variation in Mediterranean streams, and the resultant low‐ or no‐flow conditions during periods of summer drought.