STOCHASTIC LOSS OF STYLE MORPHS FROM POPULATIONS OF TRISTYLOUS LYTHRUM SALICARIA AND DECODON VERTICILLATUS (LYTHRACEAE)

Despite the theoretical importance of stochastic processes in evolution there have been few empirical studies of the interaction between genetic drift and selection on the maintenance of polymorphisms in plant populations. We used computer models to investigate the interaction between drift and frequency-dependent selection in affecting style morph frequencies in populations of tristylous species. Drift produces a distinct pattern of morph frequency variation involving: 1) the loss of the S morph and, to a lesser extent, the M morph; 2) no consistent bias in frequencies within populations; 3) a restricted pattern of variation involving a deficiency of one morph and equal excesses of the other two. Morph frequencies were surveyed in 137 populations of Lythrum salicaria from both its native range in Europe (N = 35) and recent adventive range in Ontario (N = 102), and 133 populations of Decodon verticillatus from four regions in eastern North America with different glacial histories to assess these theoretical predictions. There was a negative relationship between morph loss and population size in both species; the relationship was weaker in D. verticillatus than in L. salicaria. Morph loss was more frequent in the adventive than native range of L. salicaria, and in populations of D. verticillatus from glaciated northern regions compared with the unglaciated southern portion of its range. Simulations incorporating variation in life history, regeneration strategy and mating patterns revealed that the degree of morph loss was strongly influenced by year to year survival, clonal propagation, self-fertilization and departures from disassortative mating. Comparing the pattern of morph frequency variation between species supported these predictions. Morph loss was lower in self-incompatible L. salicaria (0% in Europe; 23% in Ontario), which reproduces through seed compared to self-compatible, clonal D. verticillatus (52%). A stochastic model provides the most parsimonious explanation for observed patterns of morph frequency variation in both species.     

The maintenance of genetic diversity under host–parasite coevolution in finite,structured populations

As a corollary to the Red Queen hypothesis, host–parasite coevolution has been hypothesized to maintain genetic variation in both species. Recent theoretical work, however, suggests that reciprocal natural selection alone is insufficient to maintain variation at individual loci. As highlighted by our brief review of the theoretical literature, models of host–parasite coevolution often vary along multiple axes (e.g. inclusion of ecological feedbacks or abiotic selection mosaics), complicating a comprehensive understanding of the effects of interacting evolutionary processes on diversity. Here we develop a series of comparable models to explore the effect of interactions between spatial structures and antagonistic coevolution on genetic diversity. Using a matching alleles model in finite populations connected by migration, we find that, in contrast to panmictic populations, coevolution in a spatially structured environment can maintain genetic variation relative to neutral expectations with migration alone. These results demonstrate that geographic structure is essential for understanding the effect of coevolution on biological diversity.     

GENETIC CONSTRAINTS ON THE INDEPENDENT EVOLUTION OF MALE AND FEMALE REPRODUCTIVE CHARACTERS IN THE TRISTYLOUS PLANT LYTHRUM SALICARIA

Here we test whether the potential exists for the independent evolution of allocation to male, female, and attractive functions within a flower. We employed half-sib and parent-offspring regression methods in the tristylous plant Lythrum salicaria to determine whether there is additive genetic variation for characters important to male and female reproductive success and whether genetic correlations could constrain the independent evolution of male and female function. Although significance levels were not consistent among morph types or between populations, there were significant narrow-sense heritabilities for several traits including stamen mass, pistil mass, perianth mass, petal length, and calyx length. Traits that might be under strong stabilizing selection to promote specific pollen transfer, such as stamen and style lengths, had little heritable variation. In the majority of cases in which heritable variation was present, there were positive genetic correlations among floral traits. A strong positive genetic correlation appeared between stamen and pistil mass in the short-styled morph from one of the populations studied. This suggests that selection might not be able to act independently on biomass allocation to male and female flower parts. No evidence of negative genetic correlations appeared that would suggest trade-offs and that could augment a selection response towards sexual specialization. The observed positive correlations could be explained if we consider the “functional architecture” that underlies the covariance structure. If there is more covariance generated by pleiotropic loci controlling overall flower size than at loci controlling male versus female allocation, it could result in the observed positive covariance. At the phenotypic level, we did find significant negative partial correlations between male and female traits when flower size was controlled, but these trade-offs were among rather than within morphs.     

Female reproductive success and the evolution of mating-type frequencies in tristylous populations

In tristylous populations, mating-type frequencies are governed by negative frequency-dependent selection typically resulting in equal morph ratios at equilibrium. However, Narcissus triandrus generally exhibits long-styled (L)-biased populations with a deficiency of the mid-styled (M)-morph. Here we used a pollen-transfer model and measurements of female fertility in natural populations to investigate whether these uneven morph ratios were associated with variation in female reproductive success. Our theoretical analysis demonstrated that morph ratio bias can result from maternal fitness differences among the morphs, and that these effects were magnified by asymmetrical mating. In nine out of 15 populations of N. triandrus, seed set differed significantly among the morphs, but pollen limitation occurred in only two of 11 populations investigated. Average seed set of the M-morph was positively associated with its frequency in populations. Flower size was negatively correlated with the seed set of the M-morph. Our results suggest that interactions between mating patterns and female fertility are responsible for variation in morph frequencies and loss of the M-morph from tristylous populations of N. triandrus.     

Isozyme characterization of genetic diversity in Minnesota populations of purple loosestrife,Lythrum salicaria (lythraceae)

Starch gel electrophoresis of plant proteins was used to identify purple loosestrife (Lythrum spp.) cultivars and weedy populations. Preliminary determinations were made as to what degree weedy loosestrife populations were related (or genetically similar) to populations of L. alatum, L. virgatum, and horticultural cultivars. Cluster analysis of the data indicated that native L. alatum was genetically different from all populations of purple loosestrife and cultivars examined. The L. salicaria and L. virgatum cultivars, as groups, were not genetically distinguishable from the weedy populations analyzed. Seven cultivars of L. salicaria origin analyzed as a group were not distinguishable from the eight cultivars of L. virgatum origin, indicating that separation by cultivar origin may not be feasible. While the two “groups” were not distinguishable, most individual cultivars could be distinguished from one another by isozyme phenotype. Genetic variation was high within populations of weedy purple loosestrife but low among populations, which is characteristic of polyploid, perennial plant species that are widely distributed. Geographic location did not consistently correlate with genetic similarity.     

The effect of color polymorphism on mortality in the aphidMacrosiphoniella yomogicola

We examined body color polymorphism in the aphidMacrosiphoniella yomogicola from July to September 1993. We classified body color into eight types: green 1, green 2, red 1, red 2, white, orange, yellow and mist. The frequencies of body color varied with time and among patches of the host plant, yomogi (Artemisia spp.). Color diversity within a shoot was calculated using the Shannon diversity index. Of five usable data sets, three showed negative relationships between color diversity and mortality. The regression coefficients for two of these relationships were significant. No significant relationship between mortality and the number of aphids was found. The color diversity was not significantly related to a particular body color found on a yomogi shoot. Color polymorphism may be maintained because selection may favor a high color diversity on the host plant shoot.     

Adaptation to local ultraviolet radiation conditions among neighbouring Daphnia populations

Understanding the historical processes that generated current patterns of phenotypic diversity in nature is particularly challenging in subdivided populations. Populations often exhibit heritable genetic differences that correlate with environmental variables, but the non-independence among neighbouring populations complicates statistical inference of adaptation. To understand the relative influence of adaptive and non-adaptive processes in generating phenotypes requires joint evaluation of genetic and phenotypic divergence in an integrated and statistically appropriate analysis. We investigated phenotypic divergence, population-genetic structure and potential fitness trade-offs in populations of Daphnia melanica inhabiting neighbouring subalpine ponds of widely differing transparency to ultraviolet radiation (UVR). Using a combination of experimental, population-genetic and statistical techniques, we separated the effects of shared population ancestry and environmental variables in predicting phenotypic divergence among populations. We found that native water transparency significantly predicted divergence in phenotypes among populations even after accounting for significant population structure. This result demonstrates that environmental factors such as UVR can at least partially account for phenotypic divergence. However, a lack of evidence for a hypothesized trade-off between UVR tolerance and growth rates in the absence of UVR prevents us from ruling out the possibility that non-adaptive processes are partially responsible for phenotypic differentiation in this system.     

Frequency-dependent variation in mimetic fidelity in an intraspecific mimicry system

Contemporary theory predicts that the degree of mimetic similarity of mimics towards their model should increase as the mimic/model ratio increases. Thus, when the mimic/model ratio is high, then the mimic has to resemble the model very closely to still gain protection from the signal receiver. To date, empirical evidence of this effect is limited to a single example where mimicry occurs between species. Here, for the first time, we test whether mimetic fidelity varies with mimic/model ratios in an intraspecific mimicry system, in which signal receivers are the same species as the mimics and models. To this end, we studied a polymorphic damselfly with a single male phenotype and two female morphs, in which one morph resembles the male phenotype while the other does not. Phenotypic similarity of males to both female morphs was quantified using morphometric data for multiple populations with varying mimic/model ratios repeated over a 3 year period. Our results demonstrate that male-like females were overall closer in size to males than the other female morph. Furthermore, the extent of morphological similarity between male-like females and males, measured as Mahalanobis distances, was frequency-dependent in the direction predicted. Hence, this study provides direct quantitative support for the prediction that the mimetic similarity of mimics to their models increases as the mimic/model ratio increases. We suggest that the phenomenon may be widespread in a range of mimicry systems.     

Comparison of quantitative and molecular genetic variation of native vs. invasive populations of purple loosestrife (Lythrum salicaria L., Lythraceae)

Study of adaptive evolutionary changes in populations of invasive species can be advanced through the joint application of quantitative and population genetic methods. Using purple loosestrife as a model system, we investigated the relative roles of natural selection, genetic drift and gene flow in the invasive process by contrasting phenotypical and neutral genetic differentiation among native European and invasive North American populations ( Q _ST −  F _ST analysis). Our results indicate that invasive and native populations harbour comparable levels of amplified fragment length polymorphism variation, a pattern consistent with multiple independent introductions from a diverse European gene pool. However, it was observed that the genetic variation reduced during subsequent invasion, perhaps by founder effects and genetic drift. Comparison of genetically based quantitative trait differentiation ( Q _ST) with its expectation under neutrality ( F _ST) revealed no evidence of disruptive selection ( Q _ST >  F _ST) or stabilizing selection ( Q _ST <  F _ST). One exception was found for only one trait (the number of stems) showing significant sign of stabilizing selection across all populations. This suggests that there are difficulties in distinguishing the effects of nonadaptive population processes and natural selection. Multiple introductions of purple loosestrife may have created a genetic mixture from diverse source populations and increased population genetic diversity, but its link to the adaptive differentiation of invasive North American populations needs further research.