Habitat-specific pigmentation in a freshwater isopod: adaptive evolution over a small spatiotemporal scale

Pigmentation in the freshwater isopod Asellus aquaticus (Crustacea) differed between habitats in two Swedish lakes. In both lakes, isopods had lighter pigmentation in stands of submerged vegetation, consisting of stoneworts (Chara spp.), than in nearby stands of reed (Phragmites australis). Experimental crossings of light and dark isopods in a common environment showed that pigmentation had a genetic basis and that genetic variance was additive. Environmental effects of diet or chromatophore adjustment to the background had minor influence on pigmentation, as shown by laboratory rearing of isopods on stonewort or reed substrates, as well as analyses of stable isotope ratios for isopods collected in the field. In both study lakes, the average phenotype became lighter with time (across generations) in recently established stonewort stands. Taken together, these results indicate that altered phenotype pigmentation result from evolutionary responses to local differences in natural selection. Based on the assumption of two generations per year, the evolutionary rate of change in pigmentation was 0.08 standard deviations per generation (haldanes) over 20 generations in one lake and 0.22 haldanes over two generations in the other lake. This genetic change occurred during an episode of population growth in a novel habitat, a situation known to promote adaptive evolution. In addition, stonewort stands constitute large and persistent patches, characteristics that tend to preserve local adaptations produced by natural selection. Results from studies on selective forces behind the adaptive divergence suggest that selective predation from visually oriented predators is a possible selective agent. We found no indications of phenotype-specific movements between habitats. Mating within stonewort stands was random with respect to pigmentation, but on a whole-lake scale it is likely that mating is assortative, as a result of local differences in phenotype distribution.     

Evolution and stability of the G-matrix during the colonization of a novel environment

Populations that undergo a process of rapid evolution present excellent opportunities to investigate the mechanisms driving or restraining adaptive divergence. The genetic variance-covariance matrix (G) is often considered to constrain adaptation but little is known about its potential to evolve during phenotypic divergence. We compared the G-matrices of ancestral and recently established ecotype populations of an aquatic isopod (Asellus aquaticus) that have diverged in parallel in two south Swedish lakes. Phenotypic changes after colonization involved a reduction in overall size, lost pigmentation and changes in shape. Comparisons between G-matrices reveal close similarity within the same ecotype from different lakes but some degree of differentiation among ecotypes. Phenotypic divergence has apparently not been much influenced by the orientation of G. Additive genetic variation in the newly colonized habitats has also decreased substantially. This suggests that a process of adaptation from standing genetic variation has occurred and has probably facilitated phenotypic divergence.     

Is size‐assortative mating important for rapid pigment differentiation in a freshwater isopod?

Identifying mechanisms behind assortative mating is central to the understanding of ecological divergence and speciation. Recent studies show that populations of the freshwater isopod Asellus aquaticus can rapidly become locally differentiated when submerged Chara vegetation expands in lakes. In the novel Chara habitat, isopods have become lighter pigmented and smaller than in ancestral reed stands. In this study, we used a laboratory multiple-choice experiment to investigate assortative mating as a possible prezygotic reproductive barrier between Chara and reed isopods. Mating was assortative when Chara isopods were experimentally mixed with isopods from an adjacent reed site with large-size individuals, suggesting a partial prezygotic reproductive barrier. No deviation from random mating could, however, be detected when Chara isopods were mixed with smaller sized isopods from another reed site. In both experiments, assortative mating was apparently based on size, as Chara isopods were larger and reed isopods smaller in mixed pairs than in assortative pairs. Pigmentation did not have any clear influence on mating. We suggest that divergence in pigmentation evolved through natural selection in conjunction with size-assortative mating indirectly causing assortative mating between Chara and reed isopods. Size-assortative mating is likely a by-product of natural selection, but its importance may hypothetically be transient, if selection erodes the correlation between pigmentation and size over time.     

Pterin-based pigmentation in animals

Pterins are one of the major sources of bright coloration in animals. They are produced endogenously, participate in vital physiological processes and serve a variety of signalling functions. Despite their ubiquity in nature, pterin-based pigmentation has received little attention when compared to other major pigment classes. Here, we summarize major aspects relating to pterin pigmentation in animals, from its long history of research to recent genomic studies on the molecular mechanisms underlying its evolution. We argue that pterins have intermediate characteristics (endogenously produced, typically bright) between two well-studied pigment types, melanins (endogenously produced, typically cryptic) and carotenoids (dietary uptake, typically bright), providing unique opportunities to address general questions about the biology of coloration, from the mechanisms that determine how different types of pigmentation evolve to discussions on honest signalling hypotheses. Crucial gaps persist in our knowledge on the molecular basis underlying the production and deposition of pterins. We thus highlight the need for functional studies on systems amenable for laboratory manipulation, but also on systems that exhibit natural variation in pterin pigmentation. The wealth of potential model species, coupled with recent technological and analytical advances, make this a promising time to advance research on pterin-based pigmentation in animals.     

Rapid adaptive divergence between ecotypes of an aquatic isopod inferred from FST–QST analysis

Divergent natural selection is often thought to be the principal factor driving phenotypic differentiation between populations. We studied two ecotypes of the aquatic isopod Asellus aquaticus which have diverged in parallel in several Swedish lakes. In these lakes, isopods from reed belts along the shores colonized new stonewort stands in the centre of the lakes and rapid phenotypic changes in size and pigmentation followed after colonization. We investigated if selection was likely to be responsible for these observed phenotypic changes using indirect inferences of selection (F_ST–Q_ST analysis). Average Q_ST for seven quantitative traits were higher than the average F_ST between ecotypes for putatively neutral markers (AFLPs). This suggests that divergent natural selection has played an important role during this rapid diversification. In contrast, the average Q_ST between the different reed ecotype populations was not significantly different from the mean F_ST. Genetic drift could therefore not be excluded as an explanation for the minor differences between allopatric populations inhabiting the same source habitat. We complemented this traditional F_ST–Q_ST approach by comparing the F_ST distributions across all loci (n = 67–71) with the Q_ST for each of the seven traits. This analysis revealed that pigmentation traits had diverged to a greater extent and at higher evolutionary rates than size‐related morphological traits. In conclusion, this extended and detailed type of F_ST–Q_ST analysis provides a powerful method to infer adaptive phenotypic divergence between populations. However, indirect inferences about the operation of divergent selection should be analyzed on a per‐trait basis and complemented with detailed ecological information.     

Contemporary Parallel Diversification,Antipredator Adaptations and Phenotypic Integration in an Aquatic Isopod

It is increasingly being recognized that predation can be a strong diversifying agent promoting ecological divergence. Adaptations against different predatory regimes can emerge over short periods of time and include many different traits. We studied antipredator adaptations in two ecotypes of an isopod (Asellus aquaticus) that have, diverged in parallel in two Swedish lakes over the last two decades. We quantified differences in escape speed, morphology and behavior for isopods from different ecotypes present in these lakes. Isopods from the source habitat (reed) coexist with mainly invertebrate predators. They are more stream-profiled and have higher escape speeds than isopods in the newly colonized stonewort habitat, which has higher density of fish predators. Stonewort isopods also show more cautious behaviors and had higher levels of phenotypic integration between coloration and morphological traits than the reed isopods. Colonization of a novel habitat with a different predation regime has thus strengthened the correlations between pigmentation and morphology and weakened escape performance. The strong signature of parallelism for these phenotypic traits indicates that divergence is likely to be adaptive and is likely to have been driven by differences in predatory regimes. Furthermore, our results indicate that physical performance, behavior and morphology can change rapidly and in concert as new habitats are colonized.     

Evidence for coloration plasticity in the yellow‐bellied toad,Bombina variegata

Phenotypic adaptation in terms of background color matching to the local habitat is an important mechanism for survival in prey species. Thus, intraspecific variation in cryptic coloration is expected among localities with dissimilar habitat features (e.g., soil, vegetation). Yellow‐bellied toads (Bombina variegata) display a dark dorsal coloration that varies between populations, assumed to convey crypsis. In this study, we explored I) geographic variation in dorsal coloration and II) coloration plasticity in B. variegata from three localities differing in substrate coloration. Using avian visual modeling, we found that the brightness contrasts of the cryptic dorsa were significantly lower on the local substrates than substrates of other localities. In experiments, individuals from one population were able to quickly change the dorsal coloration to match a lighter substrate. We conclude that the environment mediates an adaptation in cryptic dorsal coloration. We suggest further studies to test the mechanisms by which the color change occurs and explore the adaptive potential of coloration plasticity on substrates of varying brightness in B. variegata and other species.     

Phenotypic similarity and differentiation among sympatric cryptic species in a freshwater amphipod species complex

1. Cryptic species diversity poses evolutionary questions about its origin and maintenance, and ecological questions about the coexistence of seemingly identical species. 2. We examine patterns of morphological and life history differentiation in three sympatric cryptic species of freshwater amphipods within the Hyalella azteca species complex. These species are separated by extensive molecular evolution, but appear similar in phenotype. Species were collected from the littoral zone of a small kettle lake in Michigan, U.S.A., and identified to species by molecular genetic methods. 3. Two of the species were similar, differing only in female size, whereas the third was larger in body size and had larger clutches of smaller eggs than the other two. There were differences between the species also in pleon spine length and antennal segment number. 4. An analysis of allozyme variation among the cryptic species in three lakes suggests that the species are reproductively isolated within lakes. 5. We suggest that phenotypic similarity of these species is maintained by size‐selective predation by fish. The small, but statistically significant, differences in body size may form the axis for a tradeoff between resource exploitation and predator avoidance, a condition that can foster coexistence of phenotypically similar species.     

Parallelism and historical contingency during rapid ecotype divergence in an isopod

Recent studies on parallel evolution have focused on the relative role of selection and historical contingency during adaptive divergence. Here, we study geographically separate and genetically independent lake populations of a freshwater isopod (Asellus aquaticus) in southern Sweden. In two of these lakes, a novel habitat was rapidly colonized by isopods from a source habitat. Rapid phenotypic changes in pigmentation, size and sexual behaviour have occurred, presumably in response to different predatory regimes. We partitioned the phenotypic variation arising from habitat ('selection': 81–94%), lake ('history': 0.1–6%) and lake × habitat interaction ('unique diversification': 0.4–13%) for several traits. There was a limited role for historical contingency but a strong signature of selection. We also found higher phenotypic variation in the source populations. Phenotype sorting during colonization and strong divergent selection might have contributed to these rapid changes. Consequently, phenotypic divergence was only weakly influenced by historical contingency.     

LIFE-HISTORY EVOLUTION IN GUPPIES (POECILIA RETICULATA) 6. DIFFERENTIAL MORTALITY AS A MECHANISM FOR NATURAL SELECTION

We have previously reported a correlation between the life-history patterns of guppies and the types of predators with which they coexist. Guppies from localities with an abundance of large predators (high predation localities) mature at an earlier age and devote more resources to reproduction than those found in localities with only a single, small species of predator (low predation localities). We also found that when guppies were introduced from a high to low predation locality, the guppy life history evolved to resemble what was normally found in this low predation locality. The presumed mechanism of natural selection is differences among localities in age/size-specific mortality (the age/size-specific mortality hypothesis); in high predation localities we assumed that guppies experienced high adult mortality rates while in the low predation localities we assumed that guppies experienced high juvenile mortality rates. These assumptions were based on stomach content analyses of wild-caught predators and on laboratory experiments. Here, we evaluate these assumptions by directly estimating the mortality rates of guppies in natural populations. We found that guppies from high predation localities experience significantly higher mortality rates than their counterparts from low predation localities, but that these higher mortality rates are uniformly distributed across all size classes, rather than being concentrated in the larger size classes. This result appears to contradict the predictions of the age/size-specific predation hypothesis. However, we argue, using additional data on growth rates and the probabilities of survival to maturity in each type of locality, that the age-specific mortality hypothesis remains plausible. This is because the probability of survival to first reproduction is very similar in each type of locality, but the guppies from high predation localities have a much lower probability of survival per unit time after maturity. We also argue for the plausibility of two other mechanisms of natural selection. These results thus reveal mortality patterns that provide a potential cause of natural selection, but expand, rather than narrow, the number of possible mechanisms responsible for life-history evolution in guppies.     

Comparative studies of the population genetic structure of the Brachionus calyciflorus species complex from four inland lakes in Wuhu,China

To study how the population genetic structure in zooplankton respond to environmental conditions, using comparative limnology, the genetic diversity and genetic differentiation of the B. calyciflorus complex collected from four inland lakes in Wuhu City, China were investigated based on the 16S rRNA gene and nuDNA ITS sequences. The results displayed a high genetic diversity, and the nucleotide diversity of the 16S rRNA gene was higher than that of the ITS sequence. The phylogenetic analyses grouped the four populations into two cryptic species (Bc-JT and Bc-FL) with strong support. The two cryptic species were found in lakes with different trophic levels, demonstrating significant ecological specialization. The origins of clone TW12 were not consistent in the phylogenetic trees between two genetic markers, which might be attributed to the effects of male-mediated gene flow on the phylogenetic relationships of rotifers. The nucleotide diversity of the cryptic species Bc-JT was higher than that of Bc-FL, indicating that eutrophication might decrease the genetic diversity of cryptic species. The total phosphorus concentration in water bodies might be the most important factor affecting the genetic diversity of species.     

A serological investigation of the diet of Helobdella stagnalis (Hirudinea: Glossiphoniidae) in British lakes

The food of Helobdella stagnalis living in unproductive and productive British lakes has been determined using the precipitin test. Antisera were produced against each of ten potential prey groups. Correction factors were applied to basic data to compensate for differences in leech size and seasonal changes in field temperatures. In both types of lake, Oligochaeta followed by Chironomidae were most heavily utilized, whilst Trichoptera, Cladocera and Tricladida were least favoured. More Mollusca were eaten in productive than unproductive lakes, reflecting availability. In productive lakes, Asellus and Amphipoda together constituted the same proportion (about 25%) of the overall diet as Ephemeroptera and Plecoptera in unproductive lakes. Diet did not vary much during the course of a year, though diversity was lower in winter. Feeding activity varied seasonally being lowest in winter. Prey category utilization by small, medium and large leeches did not differ significantly.     

Cuticular hydrocarbons as potential mediators of cryptic species divergence in a mutualistic ant association

Upon advances in sequencing techniques, more and more morphologically identical organisms are identified as cryptic species. Often, mutualistic interactions are proposed as drivers of diversification. Species of the neotropical parabiotic ant association between Crematogaster levior and Camponotus femoratus are known for highly diverse cuticular hydrocarbon (CHC) profiles, which in insects serve as desiccation barrier but also as communication cues. In the present study, we investigated the association of the ants’ CHC profiles with genotypes and morphological traits, and discovered cryptic species pairs in both genera. To assess putative niche differentiation between the cryptic species, we conducted an environmental association study that included various climate variables, canopy cover, and mutualistic plant species. Although mostly sympatric, the two Camponotus species seem to prefer different climate niches. However in the two Crematogaster species, we could not detect any differences in niche preference. The strong differentiation in the CHC profiles may thus suggest a possible role during speciation itself either by inducing assortative mating or by reinforcing sexual selection after the speciation event. We did not detect any further niche differences in the environmental parameters tested. Thus, it remains open how the cryptic species avoid competitive exclusion, with scope for further investigations.     

基于植物多样性特征的武汉市城市湖泊湿地植被分类保护和恢复

湿地植被多样性特征及其影响因素的调查分析是湿地植被保护与恢复策略制定的基础。借鉴生物多样性热点分析原理,在武汉市城市湖泊湿地植物多样性调查的基础上,研究了湖泊湿地的植被多样性特征,探讨了城市湖泊湿地植被分类保护与恢复对策。结果表明,武汉市湿地维管束植物的物种丰富度、植物多样性、优势度和均匀度指数在各湖泊间的变化趋势较为一致,但在空间变化幅度上存在一定差异。按照物种丰富度、多样性、优势度、均匀度、湿地植被群丛数目,以及典型湿地植物的物种所占比例、丰富度和优势度的差异,可将调查涉及的26个典型湖泊湿地分为原生植被湖泊、次生植被湖泊、人工植被湖泊和退化植被湖泊4类。原生植被湖泊应建立相对严格的湿地保护区,优先保护原有湿地植被。次生植被湖泊最多,城市发展区内的次生植被湖泊应建立30-100m的植被缓冲带,促进植被自然恢复和发育;而农业区的次生植被湖泊应引导和规范湖泊周围的农业生产模式,以减少人类活动干扰。人工植被湖泊应通过建立城市湿地公园,人工促进植被的近自然恢复。而退化植被湖泊则应尽快采用生态工程法促进湿地植被生境改善,并积极开展近自然湿地植被重建与恢复。     

The influence of prey size and abundance, and individual phenotype on prey choice by the three-spined stickleback, Gasterosteus aculeatm L.

Prey selection behaviour of three-spined sticklebacks, Gasterosteus aculeatus L., was studied in two experiments. Where possible, the experimental apparatus satisfied the assumptions of the simplest optimal diet model (the basic prey model); prey were presented sequentially, the fish could not search for and handle prey at the same time, and net energy gain, handling time and encounter rate were fixed. Experiment 1 presented fish with a range of Asellus sizes so that pursuit ( p ) and handling ( h ) time could be related to prey size. Published energy values of Asellus together with pursuit and handling times were used to calculate E /( p+h ) for Asellus measuring 3,4,5,6,7 and 9 mm. Pursuit times did not differ with prey size but handling times did. E /( p+h ) was very variable particularly at the larger prey sizes. Experiment 2 presented fish with two sequences of prey differing in the encounter rate with the most profitable prey sizes. Fish did not select the diet predicted by the basic prey model tending to always ignore the largest prey even when net energy gain would have been maximized by including them in the diet. Further analysis showed that the probability of a prey size being taken was a function of prey size, fish stomach fullness and encounter rate. It is concluded that the basic prey model is too simple to capture the behaviour of the fish. One of its main faults is that the changing state of the fish through the feeding bout is ignored.     

Autotrophic and Heterotrophic Picoplankton in Wetlands: Differences with Lake Patterns

This study describes the occurrence, importance and seasonal patterns of picoplankton in two wetlands (TDNP and La Safor), and compares them to a system of fifteen interconnected lakes (Ruidera). In TDNP we performed a six‐year monthly study in three sites of the wetland. Bacterial abundance increased throughout time and the autotrophic picoplankton (APP) range was wide (up to 33 × 10⁶ cells/ml). The annual averaged APP contribution to total picoplankton and phytoplankton biovolumes was 0.5–22% and 0.03–6% respectively. There were large differences among sites in terms of APP absolute and relative abundance and seasonal patterns. In La Safor, the APP relative contribution to picoplankton and phytoplankton biovolumes was 0–25% and 0–40%, respectively, while in the Ruidera lakes was 0–47% and 0–5%, respectively. In the three systems there was a significant correlation between bacterial abundance and chlorophyll a but the slopes of the linear regressions were different. No significant relationships were found of APP abundance and trophic status in the wetlands, but were noted in the lake system. There was no clear relationship of APP contribution to total phytoplankton biomass to the trophic gradient in wetlands. In the lakes, the higher contribution of APP was found in those with higher trophic levels.     

The role of selection and historical factors in driving population differentiation along an elevational gradient in an island bird

Adaptation to local environmental conditions and the range dynamics of populations can influence evolutionary divergence along environmental gradients. Thus, it is important to investigate patterns of both phenotypic and genetic variations among populations to reveal the respective roles of these two types of factors in driving population differentiation. Here, we test for evidence of phenotypic and genetic structure across populations of a passerine bird (Zosterops borbonicus) distributed along a steep elevational gradient on the island of Réunion. Using 11 microsatellite loci screened in 401 individuals from 18 localities distributed along the gradient, we found that genetic differentiation occurred at two spatial levels: (i) between two main population groups corresponding to highland and lowland areas, respectively, and (ii) within each of these two groups. In contrast, several morphological traits varied gradually along the gradient. Comparison of neutral genetic differentiation (F_ST) and phenotypic differentiation (P_ST) showed that P_ST largely exceeds F_ST at several morphological traits, which is consistent with a role for local adaptation in driving morphological divergence along the gradient. Overall, our results revealed an area of secondary contact midway up the gradient between two major, cryptic, population groups likely diverged in allopatry. Remarkably, local adaptation has shaped phenotypic differentiation irrespective of population history, resulting in different patterns of variation along the elevational gradient. Our findings underscore the importance of understanding both historical and selective factors when trying to explain variation along environmental gradients.     

Waterbird use of artificial wetlands in an Australian urban landscape

With the loss of natural wetlands, artificial wetlands are becoming increasingly important as habitat for waterbirds. We investigated the relationships between waterbirds and various biophysical parameters on artificial wetlands in an Australian urban valley. The densities (birds per hectare) of several species were correlated (mostly positively) with wetland area, and correlations were observed between certain species and other physical and water chemistry variables. Waterbird community structure, based on both abundance (birds per wetland) and density data, was most consistently positively correlated with the relative amount of wetland perimeter that was vegetated, surface area, distance to nearest wetland, public accessibility and shoreline irregularity. We also compared the relative use of the two types of urban wetlands, namely urban lakes and stormwater treatment wetlands, and found for both abundance and density that the number of individuals and species did not vary significantly between wetland types but that significant differences were observed for particular species and feeding guilds, with no species or guild being more abundant or found in greater density on an urban lake than a stormwater treatment wetland. Designing wetlands to provide a diversity of habitat will benefit most species.     

Geographic pattern of cranial differentiation in the Asian Midday Jird Meriones meridianus (Rodentia: Muridae: Gerbillinae) and its taxonomic implications

The existence of cryptic species in the midday jird (Meriones meridianus) has been suggested in literature, although based on little empirical data to support this hypothesis. In this study, a two‐dimensional landmark‐based geometric morphometric approach was used to investigate whether patterns in intraspecific variation in skull shape and size exist, using 110 skull specimens from more than 20 different localities along the distribution range of M. meridianus. This is the first study of morphological differences on such a big sample size and geographical range, and it tries to find whether skull shape variation in this species is best described as being clinal or rather reflecting cryptic diversity. The latter seems to be the case, as a dimorphic skull phenotype was found, reflecting a geographic disparity between the Middle East and the Far East specimens both in shape and in size. Distinct cranial differences were found in the overall cranial size and, also at the level of the inflation of the bulla, the elongation of the nasal, the length of the teeth row and the incisive foramen, as well as the distance in between the latter two. It thus seems that M. meridianus from Middle East is morphologically distinct from that of the Far East. Furthermore, our results also demonstrate that clinal variation could explain variation within Middle East populations, whereas a more heterogenous pattern is found for those of the Far East. The hypothesis that the observed phenotypic variation may reflect cryptic species is discussed, with the recommendation for a thorough taxonomical revision of the genus in the region.     

Cryptic barriers to dispersal within a lake allow genetic differentiation of Eurasian perch

Gene flow between coexisting or nearby populations normally prevents genetic divergence and local adaptation. Despite this, there are an increasing number of reports of sympatric sister taxa, indicating potential divergence and speciation in the face of gene flow. A large number of such reported cases involve lake-dwelling fish, which are expected to run into few physical barriers to dispersal within their aquatic habitat. However, such cases may not necessarily reflect sympatric speciation if cryptic dispersal barriers are common in lakes and other aquatic systems. In this study, we examined genetic differentiation in perch (Perca fluviatilis L.) from nine locations in a single, small lake (24 km(2)), using microsatellites. We detected significant genetic differentiation in all but two pairwise comparisons. These patterns were not consistent with divergence by distance or the existence of kin groups. Instead, they suggest that cryptic barriers to dispersal exist within the lake, allowing small-scale genetic divergence. Such an observation suggests that allopatric (or parapatric) divergence may be possible, even in small, apparently homogenous environments such as lakes. This has important consequences for how we currently view evidence from nature for sympatric speciation.