Müllerian mimicry of a quantitative trait despite contrasting levels of genomic divergence and selection

Hybrid zones, where distinct populations meet and interbreed, give insight into how differences between populations are maintained despite gene flow. Studying clines in genetic loci and adaptive traits across hybrid zones is a powerful method for understanding how selection drives differentiation within a single species, but can also be used to compare parallel divergence in different species responding to a common selective pressure. Here, we study parallel divergence of wing colouration in the butterflies Heliconius erato and H. melpomene, which are distantly related Müllerian mimics which show parallel geographic variation in both discrete variation in pigmentation, and quantitative variation in structural colour. Using geographic cline analysis, we show that clines in these traits are positioned in roughly the same geographic region for both species, which is consistent with direct selection for mimicry. However, the width of the clines varies markedly between species. This difference is explained in part by variation in the strength of selection acting on colour traits within each species, but may also be influenced by differences in the dispersal rate and total strength of selection against hybrids between the species. Genotyping‐by‐sequencing also revealed weaker population structure in H. melpomene, suggesting the hybrid zones may have evolved differently in each species, which may also contribute to the patterns of phenotypic divergence in this system. Overall, we conclude that multiple factors are needed to explain patterns of clinal variation within and between these species, although mimicry has probably played a central role.     

THE QUANTITATIVE AND MOLECULAR GENETIC ARCHITECTURE OF A SUBDIVIDED SPECIES

In an effort to elucidate the evolutionary mechanisms that determine the genetic architecture of a species, we have analyzed 17 populations of the microcrustacean Daphnia pulex for levels of genetic variation at the level of life-history characters and molecular markers in the nuclear and mitochondrial genomes. This species is highly subdivided, with approximately 30% of the variation for nuclear molecular markers and 50% of the variation for mitochondrial markers being distributed among populations. The average level of genetic subdivision for quantitative traits is essentially the same as that for nuclear markers, which superficially suggests that the life-history characters are diverging at the neutral rate. However, the existence of a strong correlation between the levels of population subdivision and broadsense heritabilities of individual traits argues against this interpretation, suggesting instead that the among-population divergence of some quantitative traits (most notably body size) is being driven by local adaptation to different environments. The fact that the mean phenotypes of the individual populations are also strongly correlated with local levels of homozygosity indicates that variation in local inbreeding plays a role in population differentiation. Rather than being a passive consequence of local founder effects, levels of homozygosity may be selected for directly for their effects on the phenotype (adaptive inbreeding depression). There is no relationship between the levels of variation within populations for molecular markers and quantitative characters, and this is explained by the fact that the average standing genetic variation for life-history characters in this species is equivalent to only 33 generations of variation generated by mutation.     

Right here,right now: Populations of Actinotus helianthi differ in their early performance traits and interactions

Populations across the geographical distribution of a species are shaped by different local environments to produce distinctive patterns of variation in plant traits. Among‐population variation is, therefore, important for understanding potential shifts in distributions under changing environments, but is often not included in studies. In particular, critical data on the suitability of local environments for plant traits expressed at different life stages are lacking. To address this we performed two experiments to disentangle the influence of the local environment on multiple plant traits for populations of Actinotus helianthi from across its latitudinal range. A common environment experiment was used to compare early plant traits of germination, early seedling growth and survival for 17 populations of A. helianthi. To examine how biotic interactions vary across populations, we evaluated whether plant traits, including height and number of pseudanthia, influence visitor diversity and abundance, and if insect visitor abundance or diversity was associated with seed set success. We found that populations varied in germination success between 0.2 ± 0.1% and 64.2 ± 2.3%. Seedling growth and early survival varied among populations by as much as a factor of two and 44 respectively. We recorded variation in plant traits across hierarchical spatial scales from the maternal plant to biogeographical regions. The abundance and diversity of insect visitors also varied among populations and seed set was found to be site specific. There was a trend for populations with taller plants and larger floral display sizes to be more frequently visited by pollinators. We also identified a positive linear relationship between the number of visits by flies and seed set success. These results suggest that the local environment has a strong role in directly and indirectly influencing variation in plant traits within populations of A. helianthi, and potentially other perennial species.     

Intraspecific trait variation and trade‐offs within and across populations of a toxic dinoflagellate

Intraspecific trait diversity can promote the success of a species, as complementarity of functional traits within populations may enhance its competitive success and facilitates resilience to changing environmental conditions. Here, we experimentally determined the variation and relationships between traits in 15 strains of the toxic dinoflagellate Alexandrium ostenfeldii derived from two populations. Measured traits included growth rate, cell size, elemental composition, nitrogen uptake kinetics, toxin production and allelochemical potency. Our results demonstrate substantial variation in all analysed traits both within and across populations, particularly in nitrogen affinity, which was even comparable to interspecific variation across phytoplankton species. We found distinct trade‐offs between maximum nitrogen uptake rate and affinity, and between defensive and competitive traits. Furthermore, we identified differences in trait variation between the genetically similar populations. The observed high trait variation may facilitate development and resilience of harmful algal blooms under dynamic environmental conditions.     

Heritability and evolvability of fitness and nonfitness traits: Lessons from livestock

Data from natural populations have suggested a disconnection between trait heritability (variance standardized additive genetic variance, V_A) and evolvability (mean standardized V_A) and emphasized the importance of environmental variation as a determinant of trait heritability but not evolvability. However, these inferences are based on heterogeneous and often small datasets across species from different environments. We surveyed the relationship between evolvability and heritability in >100 traits in farmed cattle, taking advantage of large sample sizes and consistent genetic approaches. Heritability and evolvability estimates were positively correlated (r = 0.37/0.54 on untransformed/log scales) reflecting a substantial impact of V_A on both measures. Furthermore, heritabilities and residual variances were uncorrelated. The differences between this and previously described patterns may reflect lower environmental variation experienced in farmed systems, but also low and heterogeneous quality of data from natural populations. Similar to studies on wild populations, heritabilities for life‐history and behavioral traits were lower than for other traits. Traits having extremely low heritabilities and evolvabilities (17% of the studied traits) were almost exclusively life‐history or behavioral traits, suggesting that evolutionary constraints stemming from lack of genetic variability are likely to be most common for classical “fitness” (cf. life‐history) rather than for “nonfitness” (cf. morphological) traits.     

Latitudinal clines in the grasshopper Dichroplus elongatus: Coevolution of the A genome and B chromosomes?

Argentine populations of Dichroplus elongatus (Orthoptera: Acrididae) are polymorphic for B chromosomes. Previous studies showed that B chromosomes affect body size and some fitness components in Northwestern populations. We studied phenotype and B′s variation patterns along a latitudinal cline as well as the relationship between karyotype and body size related traits in 17 populations from East. Body size related traits showed a ‘saw tooth’ pattern of variation being small at low and high latitudes and large at intermediate latitudes in most of the analysed populations. Analyses of variance and principal components demonstrated that in most analysed populations B carrier males are associated with a decrease in body size related traits with respect to individuals with standard karyotype. Accordingly with the relationship between karyotype and body size, an opposite pattern of latitudinal variation in the frequencies of B′s with respect to body size variation was observed in this area. i.e. smaller individuals tend to have a higher frequency of B chromosomes. The comparison of the differentiation of both karyotype and body size traits with molecular neutral markers demonstrated the relative importance of selection moulding chromosome and phenotype variation. The observed pattern of phenotypic variation is likely to be the result of local adaptation to season length along the latitudinal gradient. The observed contrary pattern of B′s clinal variation may reflect the population ability to maintain this chromosome in relation to the local adaptation. The available evidence indicates that the distribution of B chromosome frequency was shaped by selective factors.     

Demographic responses to weather fluctuations are context dependent in a long‐lived amphibian

Weather fluctuations have been demonstrated to affect demographic traits in many species. In long‐lived organisms, their impact on adult survival might be buffered by the evolution of traits that reduce variation in interannual adult survival. For example, skipping breeding is an effective behavioral mechanism that may limit yearly variation in adult survival when harsh weather conditions occur; however, this in turn would likely lead to strong variation in recruitment. Yet, only a few studies to date have examined the impact of weather variation on survival, recruitment and breeding probability simultaneously in different populations of the same species. To fill this gap, we studied the impact of spring temperatures and spring rainfall on survival, on reproductive skipping behavior and on recruitment in five populations of a long‐lived amphibian, the yellow‐bellied toad (Bombina variegata). Based on capture–recapture data, our findings demonstrate that survival depends on interactions between age, population and weather variation. Varying weather conditions in the spring result in strong variation in the survival of immature toads, whereas they have little effect on adult toads. Breeding probability depends on both the individual's previous reproductive status and on the weather conditions during the current breeding season, leading to high interannual variation in recruitment. Crucially, we found that the impact of weather variation on demographic traits is largely context dependent and may thus differ sharply between populations. Our results suggest that studies predicting the impact of climate change on population dynamics should be taken with caution when the relationship between climate and demographic traits is established using only one population or few populations. We therefore highly recommend further research that includes surveys replicated in a substantial number of populations to account for context‐dependent variation in demographic processes.     

Conservation of genetic diversity in old‐growth forest communities of the southeastern United States

Question: How do studies of the distribution of genetic diversity of species with different life forms contribute to the development of conservation strategies? Location: Old‐growth forests of the southeastern United States. Methods: Reviews of the plant allozyme literature are used to identify differences in genetic diversity and structure among species with different life forms, distributions and breeding systems. The general results are illustrated by case studies of four plant species characteristic of two widespread old‐growth forest communities of the southeastern United States: the Pinus palustris – Aristida stricta (Longleaf pine – wiregrass) savanna of the Coastal Plain and the Quercus – Carya – Pinus (Oak‐hickory‐pine) forest of the Piedmont. Genetic variation patterns of single‐gene and quantitative traits are also reviewed. Results: Dominant forest trees, represented by Pinus palustris(longleaf pine) and Quercus rubra (Northern red oak), maintain most of their genetic diversity within their populations whereas a higher proportion of the genetic diversity of herbaceous understorey species such as Sarracenia leucophylla and Trillium reliquum is distributed among their populations. The herbaceous species also tend to have more population‐to‐population variation in genetic diversity. Higher genetic differentiation among populations is seen for quantitative traits than for allozyme traits, indicating that interpopulation variation in quantitative traits is influenced by natural selection. Conclusion: Developing effective conservation strategies for one or a few species may not prove adequate for species with other combinations of traits. Given suitable empirical studies, it should be possible to design efficient conservation programs that maintain natural levels of genetic diversity within species of conservation interest.     

Genetic architecture and phenotypic plasticity of thermally-regulated traits in an eruptive species, <Emphasis Type="Italic">Dendroctonus ponderosae</Emphasis>

Phenotypic plasticity in thermally-regulated traits enables close tracking of changing environmental conditions, and can thereby enhance the potential for rapid population increase, a hallmark of outbreak insect species. In a changing climate, exposure to conditions that exceed the capacity of existing phenotypic plasticity may occur. Combining information on genetic architecture and trait plasticity among populations that are distributed along a latitudinal cline can provide insight into how thermally-regulated traits evolve in divergent environments and the potential for adaptation. Dendroctonus ponderosae feed on Pinus species in diverse climatic regimes throughout western North America, and show eruptive population dynamics. We describe geographical patterns of plasticity in D. ponderosae development time and adult size by examining reaction norms of populations from multiple latitudes. The relative influence of additive and non-additive genetic effects on population differences in the two phenotypic traits at a single temperature is quantified using line-cross experiments and joint-scaling tests. We found significant genetic and phenotypic variation among D. ponderosae populations. Simple additive genetic variance was not the primary source of the observed variation, and dominance and epistasis contributed greatly to the genetic divergence of the two thermally-regulated traits. Hybrid breakdown was also observed in F₂ hybrid crosses between northern and southern populations, further indication of substantial genetic differences among clinal populations and potential reproductive isolation within D. ponderosae. Although it is unclear what maintains variation in the life-history traits, observed plasticity in thermally-regulated traits that are directly linked to rapid numerical change may contribute to the outbreak nature of D. ponderosae, particularly in a changing climate.     

Are populations of mayflies living in adjacent fish and fishless streams genetically differentiated?

1. Conspecific populations living in habitats with different risks of predation often show phenotypic variation in defensive traits. Traits of two species of mayflies (Baetidae: Baetis bicaudatus and Baetis sp. nov.) differ between populations living in fish and fishless streams in a high altitude drainage basin in western Colorado, U.S.A. We tested for genetic differentiation between mayfly populations in these two habitat types, assuming that lack of genetic differentiation would be consistent with the hypothesis that those traits are phenotypically plastic. 2. Previous work has shown that larvae of both species behave differently and undergo different developmental pathways in adjacent fish and fishless streams. These phenotypic differences in behaviour and development have been induced experimentally, suggesting that populations from fishless streams have the genetic capability to respond to fish. 3. During summer 2001 we collected Baetis larvae from several fish and fishless streams, and from fish and fishless sections of the same streams. We used allozymes and a fragment of the cytochrome oxidase subunit 1 mitochondrial gene to examine genetic variation of Baetis individuals within and among streams. 4. Results showed that genetic variation exists among populations of the same species of Baetis from different streams, but none of that variation was associated with the presence or absence of fish. These data confirm that populations of Baetis living in fish and fishless streams are not genetically distinct, and are consistent with the hypothesis that traits associated with environments of different risk are phenotypically plastic.     

Contrasting levels of genotype by environment interaction for life history and morphological traits in invasive populations of Zaprionus indianus (Diptera: Drosophilidae)

It has been demonstrated that phenotypic plasticity and genotype by environment interaction are important for coping with new and heterogeneous environments during invasions. Zaprionus indianus Gupta (Diptera: Drosophilidae) is an Afrotropical invasive fly species introduced to the South American continent in 1999. This species is generalist and polyphagous, since it develops and feeds in several different fruit species. These characteristics of Z. indianus suggest that phenotypic plasticity and genotype by environment interaction may be important in this species invasion process. In this sense, our aim was to investigate the role of genetic variation for phenotypic plasticity (genotype by environment interaction) in Z. indianus invasion of the South American continent. Specifically, we quantified quantitative genetic variation and genotype by environment interactions of morphological and life history traits in different developmental environments, that is, host fruits. This was done in different populations in the invasive range of Z. indianus in Argentina. Results showed that Z. indianus populations have considerable amounts of quantitative genetic variation. Also, genotype by environment interactions was detected for the different traits analyzed in response to the different developmental environments. Interestingly, the amounts and patterns of these parameters differed between populations. We interpreted these results as the existence of differences in evolutionary potential between populations that have an important role in the short‐ and long‐term success of the Z. indianus invasion process.     

Intraspecific morphological divergence in two Cichlid species from Benin

Selection on morphological traits can vary across the range of species, inducing a mosaic of phenotypes across populations. Intraspecific morphological divergence had been demonstrated for many fish groups inhabiting environments with varying abiotic or biotic selective pressures. Such intraspecific phenotypic variation can have a strong influence on the ecologies of species. In the current study, we examined patterns of intraspecific morphological divergence between two populations of Sarotherodon melanotheron and Coptodon guineensis in Lake Ahémé and Porto-Novo lagoon, Benin. Using multiple morphological traits, we demonstrated intraspecific morphological divergence between Lake Ahémé and Porto-Novo lagoon for these species. However, evidence for parallel divergence was found for these two species, implying a similar response to selective pressures might have been acting on labile traits. In addition, species specific morphological changes observed in the current study might be because of differing responses to similar selective forces or taxon-specific selective forces acting on labile traits. The intraspecific trait divergence demonstrated in the current study suggests several possible selective pressures acting on these populations, yet the cause of this divergence remains unknown and additional studies are required to test these inferences.     

The impact of species-specific traits and phylogenetic relatedness on allozyme diversity inCarex sect.Phyllostachys (Cyperaceae)

Allozyme variation was examined inCarex sect.Phyllostachys (Cyperaceae) to study the effects of species-specific traits and phylogenetic relatedness on genetic structure. In contrast to the findings of similar studies, genetic variability in thePhyllostachys is poorly correlated with geographic range and putative differences in breeding systems (as inferred from morphology). This suggests that other patterns of evolution, colonization, and gene flow characterize the species found in this section. Fixation indices are negative for all populations suggesting that mechanisms such as disassortative mating and selection are maintaining heterozygous excess within populations. Closely related taxa often exhibit different genetic variability statistics. In some instances, however, clades (e.g.C. jamesii andC. juniperorum) display very similar levels of genetic variability despite marked differences in species-specific traits. Recent speciation coupled with the ability to maintain historical levels of variation within populations may be factors accounting for this phenomenon. Contrary to similar studies, species restricted to known glacial refugia have lower genetic diversity than those species that underwent mass migrations in response to deglaciation. Narrowly endemic species were found to partition their genetic diversity within, as opposed to between populations. The opposite trend was evident in wider ranging congeners.     

Genetic variation of two species with different life‐history traits in the endangered renosterveld of South Africa – a comparative analysis of Eriocephalus africanus and Hemimeris racemosa

We tested the effects of life‐history traits on genetic variation and conducted a comparative analysis of two plant species with differing life‐history traits co‐occurring in the highly endangered renosterveld of South Africa. We selected eighteen renosterveld remnants with varying degrees of size and isolation where populations of the herbaceous, annual and insect‐pollinated Hemimeris racemosa and the shrubby perennial and both wind‐ and insect‐pollinated Eriocephalus africanus occurred. We postulated a lower genetic variation within populations and increased genetic variation between populations in the annual than in the perennial species. Genetic variation was lower within populations of H. racemosa than within E. africanus, as is typical for annual compared to perennial species. Variation within populations was, however, not correlated with fragment size or distance in either of the two species and genetic variation between populations of the two species was comparable (Φ_ST = 0.10, 0.09).     

CONTRASTING PATTERNS OF HERITABLE GEOGRAPHIC VARIATION IN SHELL MORPHOLOGY AND GROWTH POTENTIAL IN THE MARINE GASTROPOD BEMBICIUM VITTATUM: EVIDENCE FROM FIELD EXPERIMENTS

Similar phenotypes do not always imply similar genotypes. In species distributed over a broad latitudinal range, geographical variation in morphological and life-history traits may reflect very different relations between genotypic and environmental effects on these traits. Patterns of selection among latitudinally separated sites may minimize phenotypic differences in life-history traits but promote phenotypic differences in form. Thus, for example, latitudinal variation in temperature often leads to genetically based metabolic differences that minimize differences in growth rate among populations at different latitudes (countergradient variation). However, variation in habitat experienced by the same populations may promote genetically based differences in shell form (cogradient variation). Few attempts have been made to assess simultaneously such mosaic effects of natural selection on the genetic basis of variation in both morphological and life-history traits among geographically separated populations. I quantified the extent to which widely separated populations of the rocky shore marine gastropod Bembicium vittatum exhibited genetic differences in shell shape, shell pattern, and growth rate. Bembicium vittatum occurs naturally at only three widely separated locations on the Western Australian coast. Individuals were transplanted from all three locations to a latitudinally intermediate site, where they were released in different pairwise combinations and allowed to reproduce. F₁ offspring from crosses between same- or different-source parents were identified using allozyme markers. When grown in a common environment, offspring from same-source parents exhibited similar differences in shell shape and pattern, but dramatic differences in growth rates, compared to native populations. Genetic variation therefore exists for all three traits. Growth rates in the common environment were positively correlated with latitude of the source population, confirming the existence of countergradient variation for growth associated with metabolic compensation. In addition, for both shell shape and growth rate, hybrids exhibited phenotypes roughly midway between the same-source parents, suggesting that genetic differences have a large additive component. In contrast, when one parent had pigmented spots, the offspring also had spots, suggesting a strong dominance component to the genetic basis of shell pattern. Genetic differences therefore yield different morphological phenotypes but similar life-history phenotypes, among latitudinally distant populations, and confirm a pattern of mosaic evolution in B. vittatum.     

Associations between physical isolation and geographical variation within three species of Neotropical birds

We studied effects of physical isolation on geographical variation in mtDNA RFLP polymorphisms and a suite of morphological characters within three species of neotropical forest birds; the crimson-backed tanager Ramphocelus dimidiatus, the blue-gray tanager Thraupis episcopus, and the streaked saltator Saltator albicollis. Variation among populations within continuous habitat on the Isthmus of Panama was compared with that among island populations isolated for about 10000 years. Putative barriers to dispersal were influential, but apparent isolation effects varied by species, geographical scale, and whether molecular or morphological traits were being assessed. We found no geographical structuring among the contiguous, mainland sampling sites. Migration rates among the islands appeared sufficient to maintain homogeneity in mtDNA haplotype frequencies. In contrast, variation in external morphology among islands was significant within two of three species. For all species, we found significant variation in genetic and morphological traits between the island (collectively) and mainland populations. Interspecific variation in the effects of isolation was likely related to differential vagility. These data generally corroborate other studies reporting relatively great geographical structuring within tropical birds over short distances. Behaviourally based traits - low vagility and high ‘sensitivity’ to geographical barriers - may underlie extensive diversification within neotropical forest birds, but more extensive ecological and phylogeographic information are needed on a diverse sample of species.     

Geographic variation of flower traits in Narcissus papyraceus (Amaryllidaceae): do pollinators matter?

Aim The geographic clinal variation of traits in organisms can indicate the possible causes of phenotypic evolution. We studied the correlates of flower trait variation in populations of a style‐dimorphic plant, Narcissus papyraceus Ker‐Gawl., within a region of high biogeographical significance, the Strait of Gibraltar. This species shows a geographic gradient in the style‐morph ratio, suggested to be driven by pollinator shifts. We tested whether parallel geographic variation of perianth traits also exists, concomitant with vegetative trait variation or genetic similarity of plant populations. Location The Strait of Gibraltar region (SG hereafter, including both south‐western Iberian Peninsula and north‐western Morocco). Methods We used univariate and multivariate analyses of flower and vegetative traits in 23 populations. We applied Mantel tests and partial Mantel correlations on vegetative and flower traits and geographic locations of populations to test for spatial effects. We used Moran’s autocorrelation analyses to explore the spatial structure within the range, and performed the analyses with and without the Moroccan samples to test for the effects of the SG on spatial patterns. Amplified fragment length polymorphism data were used to estimate the genetic distance between populations and to ascertain its relationship with morphometric distance. Results There was high variation between and within populations in both flower and vegetative traits. Mantel correlations between geographic and morphometric distances were not significant, but the exclusion of Moroccan populations revealed some distance effect. Partial Mantel correlation did not detect a significant correlation between flower and vegetative morphometric distances after controlling for geographic distance. There were opposite trends in spatial autocorrelograms of flower and vegetative traits. The genetic distance between pairs of populations was directly correlated with geographic distance; however, flower morphometric and genetic distances were not significantly correlated. Main conclusions The SG had some influence on phenotypes, although the causes remain to be determined. The opposite trend of variation in flower and vegetative traits, and the lack of correlation between genetic distance and dissimilarity of flower phenotypes favour the hypothesis of pollinator‐mediated selection on flower morphology, although this may affect only particular traits and populations rather than overall phenotypes. Although stochastic population processes may have a small effect, other factors may account for the high flower variation within and between populations.     

The Role of Climatic Factors in the Expression of an Intrasexual Signal in the Paper Wasp Polistes dominulus

Within a species, variation in the use of sexual signals is observed between different populations. For intersexual traits, differences in the environmental conditions experienced by populations can play an important role in driving variation in male ornaments and female preferences. However, little is known about the factors maintaining variation in intrasexual traits used in competition. In this study, we investigate the role of climatic conditions in maintaining variation in the expression of an intrasexual signal in the paper wasp Polistes dominulus. The results of an experiment in which pupae were housed under different temperature and humidity conditions revealed a strong effect of temperature during pupal development on the expression of the signal. Furthermore, a comparison of survival and body weight between wasps reared at different temperatures indicates that signal expression exhibits phenotypic plasticity in response to developmental temperature. The effect of temperature on signal expression is consistent with patterns of signal expression observed across populations in the wild and suggests that climatic conditions may act to constrain signal expression in some populations but not in others, driving variation in signal use within P. dominulus. Environmental conditions may therefore be important in defining the scope for intrasexual signalling in animal populations and, in doing so, may play a role in maintaining variation in intrasexual traits in the face of sexual selection.     

Analyzing nested variation in the body form of Lepomid sunfishes

Phylogenetic hierarchies are often composed of younger diverging lineages nested within older diverging lineages. Comparing phenotypic variation among several hierarchical levels can be used to test hypotheses about selection, phenotypic evolution and speciation. Such hierarchical comparisons have only been performed in threespine stickleback, and so here we use a hierarchical pattern of divergences between near-shore littoral and off-shore pelagic habitats to test for selection on the evolution of body form in Lepomis sunfish in lakes. We compare variation in external body form between fish from littoral and pelagic habitats at three levels: among ecomorphs within individual lake populations (intrapopulation), among populations of the same species in different lakes (interpopulation), and between bluegill and pumpkinseed sunfish species (interspecifically). Using geometric morphometric methods, we first demonstrate that interpopulation variation in mean body form of pumpkinseed sunfish varies with the presence of pelagic habitat. We then incorporate these results with existing data in order to test the similarity of phenotypic divergence between littoral and pelagic habitats at different hierarchical levels. Parallel relationships between certain body form traits (head length, caudal length and pectoral length) and habitat occur at all three levels suggesting that selection persistently acts at all levels to diversify these traits and so may contribute to species formation. For other traits (caudal depth and pectoral altitude), divergence between habitats is inconsistent at different hierarchical levels. Thus, nested biological variation in Lepomid body form reflects a history of deterministic selection and historical contingency, and also identifies traits that likely have likely influenced fitness and so serve important functions.     

Geographic structure of genetic and phenotypic variation in the hybrid zone between Quercus affinis and Q. laurina in Mexico

Analyzing the structure of hybrid zones is important for inferring their origin, dynamics and evolutionary significance. We examined the geographic structure of phenotypic and genetic variation in the contact zone between two Mexican red oaks, Quercus affinis and Q. laurina. A total of 105 individuals from seven populations were sampled along a 600‐km latitudinal gradient representing the distribution area of the two species and their contact zone. Individuals were genotyped for nine nuclear and four chloroplast DNA microsatellite loci (ncSSR and cpSSR, respectively), and characterized for several leaf and acorn traits. The cpSSR data revealed extensive haplotype sharing among populations of the two species, while a Bayesian assignment analysis based on ncSSRs identified two main genetic groups, each corresponding to one of the species, and two populations in the contact zone showing evidence of admixture. The proportion of genetic ancestry in the populations was strongly associated with latitude and showed a pattern of variation with the shape of a narrow sigmoidal cline. The variation in three of the seven phenotypic traits was partially congruent with molecular variation, while the other traits did not conform to a geographic cline but instead were correlated with environmental variables. In conclusion, the hybrid zone between the two oak species has some of the characteristics of a tension zone, but heterogeneous variation across traits suggests differential introgression and the action of extrinsic selection.