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.     

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.     

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.     

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.     

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.     

Habitat type as a determinant of species range sizes: the example of lotic–lentic differences in aquatic Goleoptera

Species differ in the size of their geographical ranges, but it is unclear how this is affected by the intrinsic properties of various habitat types. Using data on range sizes for 490 species of aquatic Coleoptera from the Iberian Peninsula we show that running-water (lotic) species have much smaller distributional ranges than those occurring in standing water (lentic). This robust association of habitat type and range size has independently arisen in at least four monophyletic coleopteran lineages, in Hydradephaga, Hydrophiloidea, Hydraenidae and Byrrhoidea, and several more times within these main groups. We propose that this pattern is due to different evolutionary dynamics of both habitat types: stagnant water bodies are more likely to completely disappear, requiring frequent migration of resident populations. Rivers and streams, on the contrary, have more temporal and spatial continuity, and therefore permit the long-term persistence of local populations. In less permanent habitats species will require a greater geographical mobility, which indirectly results in a larger size range. The less dispersive populations of running water should also have reduced gene flow, increasing the probability of allopatric speciation, and thus reducing the average range of more widespread ancestral species. These differences in population parameters, and the frequency of transitions between the two habitat types, may have strong macroevolutionary consequences, in particular regarding speciation rates and possible morphological specializations.     

Comparative phylogeography of tenebrionid beetles in the Aegean archipelago: the effect of dispersal ability and habitat preference

Comparative phylogeographical studies in island archipelagos can reveal lineage-specific differential responses to the geological and climatic history. We analysed patterns of genetic diversity in six codistributed lineages of darkling beetles (Tenebrionidae) in the central Aegean archipelago which differ in wing development and habitat preferences. A total of 600 specimens from 30 islands and eight adjacent mainland regions were sequenced for mitochondrial cytochrome oxidase I and nuclear Muscular protein 20. Individual gene genealogies were assessed for the presence of groups that obey an independent coalescent process using a mixed Yule coalescent model. The six focal taxa differed greatly in the number of coalescent groups and depth of lineage subdivision, which was closely mirrored by the degree of geographical structuring. The most severe subdivision at both mitochondrial DNA and nuclear DNA level was found in flightless lineages associated with presumed stable compact-soil habitats (phrygana, maquis), in contrast to sand-obligate lineages inhabiting ephemeral coastal areas that displayed greater homogeneity across the archipelago. A winged lineage, although associated with stable habitats, showed no significant phylogenetic or geographical structuring. Patterns of nucleotide diversity and local genetic differentiation, as measured using Φ_ST and hierarchical amova , were consistent with high levels of ongoing gene flow in the winged taxon; frequent local extinction and island recolonisation for flightless sand-obligate taxa; and very low gene flow and geographical structure largely defined by the palaeogeographical history of the region in flightless compact-soil taxa. These results show that differences in dispersal rate, mediated by habitat persistence, greatly influence the levels of phylogeographical subdivision in lineages that are otherwise subjected to the same geological events and palaeoclimatic changes.     

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.     

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.     

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。