Geographic variation and plasticity to water and nutrients in Pelargonium australe

Here, patterns of phenotypic plasticity and trait integration of leaf characteristics in six geographically discrete populations of the perennial herb Pelargonium australe were compared. It was hypothesized that populations would show local adaptation in trait means, but similar patterns of plasticity and trait integration. Further, it was questioned whether phenotypic plasticity was positively correlated with environmental heterogeneity and whether plasticity for water-use traits in particular was adaptive. Seedlings were grown in a glasshouse at six combinations of water and nutrient availability. Leaf anatomical, morphological and gas exchange traits were measured. High amounts of plasticity in leaf traits were found in response to changes in growth conditions and there was evidence of local adaptation among the populations. While there were significant correlations between plasticity and environmental heterogeneity, not all were positive. Notably, patterns of plasticity and trait integration varied significantly among populations. Despite that variation, some of the observed plasticity was adaptive: fitness was correlated with conservative water use when water was limiting. Pelargonium arrived in Australia approximately 5 million yr ago. It is concluded here that high amounts of plasticity, in some cases adaptive, and weak integration among traits may be key to the spread and success of this species.     

Low genetic diversity contrasts with high phenotypic variability in heptaploid Spartina densiflora populations invading the Pacific coast of North America

Species can respond to environmental pressures through genetic and epigenetic changes and through phenotypic plasticity, but few studies have evaluated the relationships between genetic differentiation and phenotypic plasticity of plant species along changing environmental conditions throughout wide latitudinal ranges. We studied inter‐ and intrapopulation genetic diversity (using simple sequence repeats and chloroplast DNA sequencing) and inter‐ and intrapopulation phenotypic variability of 33 plant traits (using field and common‐garden measurements) for five populations of the invasive cordgrass Spartina densiflora Brongn. along the Pacific coast of North America from San Francisco Bay to Vancouver Island. Studied populations showed very low genetic diversity, high levels of phenotypic variability when growing in contrasted environments and high intrapopulation phenotypic variability for many plant traits. This intrapopulation phenotypic variability was especially high, irrespective of environmental conditions, for those traits showing also high phenotypic plasticity. Within‐population variation represented 84% of the total genetic variation coinciding with certain individual plants keeping consistent responses for three plant traits (chlorophyll b and carotenoid contents, and dead shoot biomass) in the field and in common‐garden conditions. These populations have most likely undergone genetic bottleneck since their introduction from South America; multiple introductions are unknown but possible as the population from Vancouver Island was the most recent and one of the most genetically diverse. S. densiflora appears as a species that would not be very affected itself by climate change and sea‐level rise as it can disperse, establish, and acclimate to contrasted environments along wide latitudinal ranges.     

Phenotypic variation in invasive and biocontrol populations of the harlequin ladybird, Harmonia axyridis

Despite numerous releases for biological control purposes during more than 20 years in Europe, Harmonia axyridis failed to become established until the beginning of the 21st century. Its status as invasive alien species is now widely recognised. Theory suggests that invasive populations should evolve toward greater phenotypic plasticity because they encounter differing environments during the invasion process. On the contrary, populations used for biological control have been maintained under artificial rearing conditions for many generations; they are hence expected to become specialised on a narrow range of environments and show lower phenotypic plasticity. Here we compared phenotypic traits and the extent of adaptive phenotypic plasticity in two invasive populations and two populations commercialized for biological control by (i) measuring six phenotypic traits related to fitness (eggs hatching rate, larval survival rate, development time, sex ratio, fecundity over 6 weeks and survival time of starving adults) at three temperatures (18, 24 and 30°C), (ii) recording the survival rate and quiescence aggregation behaviour when exposed to low temperatures (5, 10 and 15°C), and (iii) studying the cannibalistic behaviour of populations in the absence of food. Invasive and biocontrol populations displayed significantly different responses to temperature variation for a composite fitness index computed from the traits measured at 18, 24 and 30°C, but not for any of those traits considered independently. The plasticity measured on the same fitness index was higher in the two invasive populations, but this difference was not statistically significant. On the other hand, invasive populations displayed significantly higher survival and higher phenotypic plasticity when entering into quiescence at low temperatures. In addition, one invasive population displayed a singular cannibalistic behaviour. Our results hence only partly support the expectation of increased adaptive phenotypic plasticity of European invasive populations of H. axyridis, and stress the importance of the choice of the environmental parameters to be manipulated for assessing phenotypic plasticity variation among populations.     

Evidence for plasticity,but not local adaptation,in invasive Japanese knotweed (<Emphasis Type="Italic">Reynoutria japonica</Emphasis>) in North America

While local adaptation and phenotypic plasticity are commonly observed in species occupying heterogeneous environments, these phenomena are less well understood in invasive species. However, plant invasions offer the opportunity to study these dynamics as they occur in species colonizing new habitats. In this study, we examined local adaptation and phenotypic plasticity in an invasive plant, Reynoutria japonica, across a broad latitudinal range within North America. We performed full-factorial reciprocal transplants using plants from three sites and examined fitness responses in both sexual and clonal reproductive measures, as well as vegetative responses in height, basal stem diameter, and biomass. For all vegetative traits, there was a significant effect of source population, indicating genetic differentiation among populations. There was also a significant effect of transplant site, suggesting phenotypic plasticity. However, there was no evidence of local adaptation at the North American meta-population level for either measure of sexual or clonal fitness. All three comparisons for sexual fitness failed to show any differences between source populations, indicating a lack of local adaptation. For clonal fitness, two of the three comparisons showed local maladaptation, and only one showed greater fitness at the home compared to foreign sites, but this population had greater fitness at all sites, indicating greater fitness overall for this population rather than local adaptation. The fact that we did not detect consistent patterns of local adaptation in these populations across a broad geographic range is somewhat surprising given that local adaptation appears common in many species, including invasives, and that the populations have been established for over a century. However, the lack of local adaptation observed in this species may indicate that phenotypic plasticity within the species is sufficient to allow the persistence of R. japonica in a variety of environments across its invaded range.     

Rapid local adaptation in both sexual and asexual invasive populations of monkeyflowers (Mimulus spp.)

Background and AimsTraditionally, local adaptation has been seen as the outcome of a long evolutionary history, particularly with regard to sexual lineages. By contrast, phenotypic plasticity has been thought to be most important during the initial stages of population establishment and in asexual species. We evaluated the roles of adaptive evolution and phenotypic plasticity in the invasive success of two closely related species of invasive monkeyflowers (Mimulus) in the UK that have contrasting reproductive strategies: M. guttatus combines sexual (seeds) and asexual (clonal growth) reproduction while M. × robertsii is entirely asexual.MethodsWe compared the clonality (number of stolons), floral and vegetative phenotype, and phenotypic plasticity of native (M. guttatus) and invasive (M. guttatus and M. × robertsii) populations grown in controlled environment chambers under the environmental conditions at each latitudinal extreme of the UK. The goal was to discern the roles of temperature and photoperiod on the expression of phenotypic traits. Next, we tested the existence of local adaptation in the two species within the invasive range with a reciprocal transplant experiment at two field sites in the latitudinal extremes of the UK, and analysed which phenotypic traits underlie potential local fitness advantages in each species.Key ResultsPopulations of M. guttatus in the UK showed local adaptation through sexual function (fruit production), while M. × robertsii showed local adaptation via asexual function (stolon production). Phenotypic selection analyses revealed that different traits are associated with fitness in each species. Invasive and native populations of M. guttatus had similar phenotypic plasticity and clonality. M. × robertsii presents greater plasticity and clonality than native M. guttatus, but most populations have restricted clonality under the warm conditions of the south of the UK.ConclusionsThis study provides experimental evidence of local adaptation in a strictly asexual invasive species with high clonality and phenotypic plasticity. This indicates that even asexual taxa can rapidly (<200 years) adapt to novel environmental conditions in which alternative strategies may not ensure the persistence of populations.     

Plant performance and morpho-functional differentiation in response to edaphic variation in Iberian columbines: cues for range distribution?

Aims Experimental studies of the response to environmental variation of closely related taxa are needed to understand the mechanisms underlying phenotypic divergence, habitat segregation and range span within a radiating genus. We explored the magnitude of phenotypic differentiation and adaptive plasticity in relation to edaphic variation and its significance for habitat segregation and range span in Iberian columbines.Methods We performed a common garden experiment varying soil type (calcareous vs. siliceous) and depth (rocky-shallow vs. deep) with two pairs of widespread and narrowly distributed subspecies of Iberian columbines: Aquilegia vulgaris (subspp. vulgaris and nevadensis) and Aquilegia pyrenaica (subspp. pyrenaica and cazorlensis). We compared tolerance to edaphic variation, trait differentiation and adaptive plasticity of 10 morpho-functional traits between species and distribution ranges. Additionally, we obtained estimates of the competitive environment faced by each taxon in two to four populations per taxa.Important findings Results partially support hypotheses of higher competitive ability and tolerance to edaphic variation in widespread than in narrow endemic taxa. At the species level, the widely distributed taxon was the most tolerant to edaphic variation. Within species, no consistent pattern emerged since the population from the most widespread subspecies was the most tolerant in A. vulgaris but not in A. pyrenaica. Columbines were differentiated in many traits at species and range level. However, the pattern of differentiation does not fully support the hypothesis of higher specialization and stress tolerance in narrow endemics. Although plasticity was generally low, the results support the hypothesis of adaptive plasticity in widespread but not in restricted taxa at least at species level. Ecological differences (adaptive plasticity and competitive ability among others) may have contributed to phenotypic divergence and edaphic niche segregation, as well as to differences in range span among columbines.     

Environmental Heterogeneity and Population Differentiation in Plasticity to Drought in <Emphasis Type="Italic">Convolvulus Chilensis</Emphasis> (Convolvulaceae)

Plant populations may show differentiation in phenotypic plasticity, and theory predicts that greater levels of environmental heterogeneity should select for higher magnitudes of phenotypic plasticity. We evaluated phenotypic responses to reduced soil moisture in plants of Convolvulus chilensis grown in a greenhouse from seeds collected in three natural populations that differ in environmental heterogeneity (precipitation regime). Among several morphological and ecophysiological traits evaluated, only four traits showed differentiation among populations in plasticity to soil moisture: leaf area, leaf shape, leaf area ratio (LAR), and foliar trichome density. In all of these traits plasticity to drought was greatest in plants from the population with the highest interannual variation in precipitation. We further tested the adaptive nature of these plastic responses by evaluating the relationship between phenotypic traits and total biomass, as a proxy for plant fitness, in the low water environment. Foliar trichome density appears to be the only trait that shows adaptive patterns of plasticity to drought. Plants from populations showing plasticity had higher trichome density when growing in soils with reduced moisture, and foliar trichome density was positively associated with total biomass. Co-ordinating editor: F. Stuefer     

Evolution of chinook salmon (Oncorhynchus tshawytscha) populations in New Zealand: pattern,rate, and process

Chinook salmon, Oncorhynchus tshawytscha, from the Sacramento River, California, USA were introduced to New Zealand between 1901 and 1907, and colonized most of their present-day range within about 10 years. The New Zealand populations now vary in phenotypic traits typically used to differentiate salmon populations within their natural range: growth in freshwater and at sea, age at maturity, dates of return to fresh water and reproduction, morphology, and reproductive allocation. This paper reviews a large research program designed to determine the relative contributions of phenotypic plasticity and genetic adaptation to this variation, in an effort to understand the processes underlying the natural evolution of new populations. We found strong evidence of trait divergence between populations within at most 30 generations, particularly in freshwater growth rate, date of return, and reproductive output, with plausible adaptive bases for these differences. Importantly, we also demonstrated not only a genetic basis for post-release survival but higher survival, and hence fitness, of a population released from its established site compared to another population released from the same site. We conclude that divergence of salmon in different rivers probably resulted initially from phenotypic plasticity (e.g., habitat-specific growth rates, and effects of upriver migration on ovarian investment). Philopatry (homing to natal streams) combined with rapid evolution of distinct breeding periods to restrict gene flow, facilitating divergence in other traits. We also suggest that in addition to genetic divergence resulting from random founder effects, divergence may also arise during the very early stages of colonization when the original colonists are a non-random, pre-adapted subset of the source population. This favored founders effect immediately improves the fitness of the new population. Overall, this research reveals the complex interplay of environmental and genetic controls over behavior, physiology and life history that characterize the early stages of population differentiation, a process that has taken place repeatedly during the history of salmon populations.     

Is seasonal variation in leaf traits adaptive for the annual plant Dicerandra linearifolia?

Empirical studies of phenotypic plasticity have often relied on the plausibility that a plastic response to the environment would increase fitness in order to diagnose the response as adaptive. I conducted a test of the hypothesis that seasonal variation in leaf traits is an adaptive response to seasonal variation in environmental conditions faced by the annual plant Dicerandralinearifolia. This species exhibits variation in leaf morphology and anatomy in response to temperature that is consistent with the expectations for adaptive plasticity. I examined variation in the size, thickness and density of stomata of leaves that develop in summer and winter and used analysis of phenotypic selection during winter and summer seasons to test the hypothesis that seasonal variation in these traits is adaptive. Regression analyses of estimated dry mass (as a proxy for fitness) on leaf traits revealed no evidence supporting the adaptive hypothesis. Selection favoured individuals with large and thick leaves in both winter and summer, and density of stomata had little or no effect on estimated relative fitness in any season. Correspondence between seasonal variation in leaf thickness and density of stomata and expectations for adaptive plasticity appears to be purely fortuitous. Seasonal variation in leaf traits may persist simply because there is no selection against individuals in which these traits vary. My results underscore the importance of definitive tests of the hypothesis of adaptation to distinguish adaptive plasticity from neutral or nonadaptive phenotypic plasticity.     

Patterns of genetic diversity vary among shoot and root functional traits in Norway spruce Picea abies along a latitudinal gradient

Roots constitute a major segment of plant biomass, and variation in belowground traits in situ correlates with environmental gradients at large spatial scales. Local adaptation of populations maintains intraspecific genetic variation in various shoot traits, but the contribution of genetic factors to adaptation to soil heterogeneity remains poorly known. I established a common-garden experiment with three Norway spruce Picea abies populations sampled between 60° and 67° N in Finland, each represented by 13 or 15 maternal families, to determine whether belowground traits are as genetically differentiated among populations as those in the shoot along a collective latitudinal gradient of temperature and soil heterogeneity. Two growing season simulations enabled testing for among-population differences in phenotypic plasticity. I phenotyped 777 first-year seedlings from shoot to root to capture functional traits that may influence survival in the wild: autumn phenology, shoot growth, root system size, root architecture, root morphology and growth allocation. All traits exhibited within-population genetic diversity, but among-population differentiation ranged from strong in shoot traits to nonexistent in root system architecture and morphology that are scaled to root system size. However, latitudinal trends characterised root-to-shoot ratio and root tip-to-shoot ratio that account for among-population differences in aboveground growth. Overall trait variability was multidimensional with variable among- versus within-population trends: for example, phenology and shoot growth covaried across populations, but their association within individual populations was variable. Shoot growth correlated positively with root system size, but not with root architecture or morphology. Finally, the two higher-latitude populations exhibited greater phenotypic plasticity in shoot traits and growth allocation. The results demonstrate varying patterns of genetic variation in functional traits of Norway spruce in the boreal zone, suggesting simultaneous adaptation to multiple environmental factors. Functional traits that exhibit phenotypic plasticity, genetic diversity and little covariation will promote long-term survival of populations in fluctuating environments.     

Translating environmental gradients into discontinuous reaction norms via hormone signalling in a polyphenic butterfly

Polyphenisms—the expression of discrete phenotypic morphs in response to environmental variation—are examples of phenotypic plasticity that may potentially be adaptive in the face of predictable environmental heterogeneity. In the butterfly Bicyclus anynana, we examine the hormonal regulation of phenotypic plasticity that involves divergent developmental trajectories into distinct adult morphs for a suite of traits as an adaptation to contrasting seasonal environments. This polyphenism is induced by temperature during development and mediated by ecdysteroid hormones. We reared larvae at separate temperatures spanning the natural range of seasonal environments and measured reaction norms for ecdysteroids, juvenile hormones (JHs) and adult fitness traits. Timing of peak ecdysteroid, but not JH titres, showed a binary response to the linear temperature gradient. Several adult traits (e.g. relative abdomen mass) responded in a similar, dimorphic manner, while others (e.g. wing pattern) showed a linear response. This study demonstrates that hormone dynamics can translate a linear environmental gradient into a discrete signal and, thus, that polyphenic differences between adult morphs can already be programmed at the stage of hormone signalling during development. The range of phenotypic responses observed within the suite of traits indicates both shared regulation and independent, trait-specific sensitivity to the hormone signal.     

Adaptive phenotypic plasticity contributes to divergence between lake and river populations of an East African cichlid fish

Adaptive phenotypic plasticity and fixed genotypic differences have long been considered opposing strategies in adaptation. More recently, these mechanisms have been proposed to act complementarily and under certain conditions jointly facilitate evolution, speciation, and even adaptive radiations. Here, we investigate the relative contributions of adaptive phenotypic plasticity vs. local adaptation to fitness, using an emerging model system to study early phases of adaptive divergence, the generalist cichlid fish species Astatotilapia burtoni. We tested direct fitness consequences of morphological divergence between lake and river populations in nature by performing two transplant experiments in Lake Tanganyika. In the first experiment, we used wild‐caught juvenile lake and river individuals, while in the second experiment, we used F1 crosses between lake and river fish bred in a common garden setup. By tracking the survival and growth of translocated individuals in enclosures in the lake over several weeks, we revealed local adaptation evidenced by faster growth of the wild‐caught resident population in the first experiment. On the other hand, we did not find difference in growth between different types of F1 crosses in the second experiment, suggesting a substantial contribution of adaptive phenotypic plasticity to increased immigrant fitness. Our findings highlight the value of formally comparing fitness of wild‐caught and common garden‐reared individuals and emphasize the necessity of considering adaptive phenotypic plasticity in the study of adaptive divergence.     

Behavioral constraints on local adaptation and counter‐gradient variation: Implications for climate change

Resource allocation to growth, reproduction, and body maintenance varies within species along latitudinal gradients. Two hypotheses explaining this variation are local adaptation and counter‐gradient variation. The local adaptation hypothesis proposes that populations are adapted to local environmental conditions and are therefore less adapted to environmental conditions at other locations. The counter‐gradient variation hypothesis proposes that one population out performs others across an environmental gradient because its source location has greater selective pressure than other locations. Our study had two goals. First, we tested the local adaptation and counter‐gradient variation hypotheses by measuring effects of environmental temperature on phenotypic expression of reproductive traits in the burying beetle, Nicrophorus orbicollis Say, from three populations along a latitudinal gradient in a common garden experimental design. Second, we compared patterns of variation to evaluate whether traits covary or whether local adaptation of traits precludes adaptive responses by others. Across a latitudinal range, N. orbicollis exhibits variation in initiating reproduction and brood sizes. Consistent with local adaptation: (a) beetles were less likely to initiate breeding at extreme temperatures, especially when that temperature represents their source range; (b) once beetles initiate reproduction, source populations produce relatively larger broods at temperatures consistent with their local environment. Consistent with counter‐gradient variation, lower latitude populations were more successful at producing offspring at lower temperatures. We found no evidence for adaptive variation in other adult or offspring performance traits. This suite of traits does not appear to coevolve along the latitudinal gradient. Rather, response to selection to breed within a narrow temperature range may preclude selection on other traits. Our study highlights that N. orbicollis uses temperature as an environmental cue to determine whether to initiate reproduction, providing insight into how behavior is modified to avoid costly reproductive attempts. Furthermore, our results suggest a temperature constraint that shapes reproductive behavior.     

Phenotypic plasticity as a component of evolutionary change

Phenotypic plasticity has often been assumed to buffer the effects of natural selection and thus act as a constraint on evolutionary change. It has become increasingly clear, however, that phenotypic plasticity actually represents a fundamental component of evolutionary change. Where genetic variation for plasticity exists, a population with a different mean plasticity can evolve. Recent attention has been focused on the conditions necessary for the evolution of phenotypic plasticity, i.e. those under which a generalist strategy, as opposed to a range of genetically differentiated specialists, will be favoured. It is also now clear that genotypes that perform best in one environment usually perform less well than other genotypes in a different environment; hence, their greater response is not an adaptation to environmental variation. A response to environmental variation is only adaptive if it represents a mechanism by which relative fitness is maintained in the face of environmental variation. Adaptive plasticity may thus involve both physiological homeostasis and morphological response.     

The role of environment and core‐margin effects on range‐wide phenotypic variation in a montane grasshopper

The integration of genetic information with ecological and phenotypic data constitutes an effective approach to gain insight into the mechanisms determining interpopulation variability and the evolutionary processes underlying local adaptation and incipient speciation. Here, we use the Pyrenean Morales grasshopper (Chorthippus saulcyi moralesi) as study system to (i) analyse the relative role of genetic drift and selection in range‐wide patterns of phenotypic differentiation and (ii) identify the potential selective agents (environment, elevation) responsible for variation. We also test the hypothesis that (iii) the development of dispersal‐related traits is associated with different parameters related to population persistence/turnover, including habitat suitability stability over the last 120 000 years, distance to the species distribution core and population genetic variability. Our results indicate that selection shaped phenotypic differentiation across all the studied morphological traits (body size, forewing length and shape). Subsequent analyses revealed that among‐population differentiation in forewing length was significantly explained by a temperature gradient, suggesting an adaptive response to thermoregulation or flight performance under contrasting temperature regimes. We found support for our hypothesis predicting a positive association between the distance to the species distribution core and the development of dispersal‐related morphology, which suggests an increased dispersal capability in populations located at range edges that, in turn, exhibit lower levels of genetic variability. Overall, our results indicate that range‐wide patterns of phenotypic variation are partially explained by adaptation in response to local environmental conditions and differences in habitat persistence between core and peripheral populations.     

Range margin populations show high climate adaptation lags in European trees

How populations of long‐living species respond to climate change depends on phenotypic plasticity and local adaptation processes. Marginal populations are expected to have lags in adaptation (i.e. differences between the climatic optimum that maximizes population fitness and the local climate) because they receive pre‐adapted alleles from core populations preventing them from reaching a local optimum in their climatically marginal habitat. Yet, whether adaptation lags in marginal populations are a common feature across phylogenetically and ecologically different species and how lags can change with climate change remain unexplored. To test for range‐wide patterns of phenotypic variation and adaptation lags of populations to climate, we (a) built model ensembles of tree height accounting for the climate of population origin and the climate of the site for 706 populations monitored in 97 common garden experiments covering the range of six European forest tree species; (b) estimated populations' adaptation lags as the differences between the climatic optimum that maximizes tree height and the climate of the origin of each population; (c) identified adaptation lag patterns for populations coming from the warm/dry and cold/wet margins and from the distribution core of each species range. We found that (a) phenotypic variation is driven by either temperature or precipitation; (b) adaptation lags are consistently higher in climatic margin populations (cold/warm, dry/wet) than in core populations; (c) predictions for future warmer climates suggest adaptation lags would decrease in cold margin populations, slightly increasing tree height, while adaptation lags would increase in core and warm margin populations, sharply decreasing tree height. Our results suggest that warm margin populations are the most vulnerable to climate change, but understanding how these populations can cope with future climates depend on whether other fitness‐related traits could show similar adaptation lag patterns.     

Phenotypic plasticity in gene expression contributes to divergence of locally adapted populations of Fundulus heteroclitus

We examine the interaction between phenotypic plasticity and evolutionary adaptation using muscle gene expression levels among populations of the fish Fundulus heteroclitus acclimated to three temperatures. Our analysis reveals shared patterns of phenotypic plasticity due to thermal acclimation as well as non‐neutral patterns of variation among populations adapted to different thermal environments. For the majority of significant differences in gene expression levels, phenotypic plasticity and adaptation operate on different suites of genes. The subset of genes that demonstrate both adaptive differences and phenotypic plasticity, however, exhibit countergradient variation of expression. Thus, expression differences among populations counteract environmental effects, reducing the phenotypic differentiation between populations. Finally, gene‐by‐environment interactions among genes with non‐neutral patterns of expression suggest that the penetrance of adaptive variation depends on the environmental conditions experienced by the individual.     

Plasticity in reproduction and growth among 52 range‐wide populations of a Mediterranean conifer: adaptive responses to environmental stress

A plastic response towards enhanced reproduction is expected in stressful environments, but it is assumed to trade off against vegetative growth and efficiency in the use of available resources deployed in reproduction [reproductive efficiency (RE)]. Evidence supporting this expectation is scarce for plants, particularly for long‐lived species. Forest trees such as Mediterranean pines provide ideal models to study the adaptive value of allocation to reproduction vs. vegetative growth given their among‐population differentiation for adaptive traits and their remarkable capacity to cope with dry and low‐fertility environments. We studied 52 range‐wide Pinus halepensis populations planted into two environmentally contrasting sites during their initial reproductive stage. We investigated the effect of site, population and their interaction on vegetative growth, threshold size for female reproduction, reproductive–vegetative size relationships and RE. We quantified correlations among traits and environmental variables to identify allocation trade‐offs and ecotypic trends. Genetic variation for plasticity was high for vegetative growth, whereas it was nonsignificant for reproduction. Size‐corrected reproduction was enhanced in the more stressful site supporting the expectation for adverse conditions to elicit plastic responses in reproductive allometry. However, RE was unrelated with early reproductive investment. Our results followed theoretical predictions and support that phenotypic plasticity for reproduction is adaptive under stressful environments. Considering expectations of increased drought in the Mediterranean, we hypothesize that phenotypic plasticity together with natural selection on reproductive traits will play a relevant role in the future adaptation of forest tree species.     

The fitness costs of developmental canalization and plasticity

Organisms are capable of an astonishing repertoire of phenotypic responses to the environment, and these often define important adaptive solutions to heterogeneous and unpredictable conditions. The terms ‘phenotypic plasticity’ and ‘canalization’ indicate whether environmental variation has a large or small effect on the phenotype. The evolution of canalization and plasticity is influenced by optimizing selection‐targeting traits within environments, but inherent fitness costs of plasticity may also be important. We present a meta‐analysis of 27 studies (of 16 species of plant and 7 animals) that have measured selection on the degree of plasticity independent of the characters expressed within environments. Costs of plasticity and canalization were equally frequent and usually mild; large costs were observed only in studies with low sample size. We tested the importance of several covariates, but only the degree of environmental stress was marginally positively related to the cost of plasticity. These findings suggest that costs of plasticity are often weak, and may influence phenotypic evolution only under stressful conditions.     

Testing the adaptive plasticity of Iris pumila leaf traits to natural light conditions using phenotypic selection analysis

A multivariate selection analysis has been used to test the adaptiveness of several Iris pumila leaf traits that display plasticity to natural light conditions. Siblings of a synthetic population comprising 31 families of two populations from contrasting light habitats were grown at an open dune site and in the understory of a Pinus nigra stand in order to score variation in phenotypic expression of six leaf traits: number of senescent leaves, number of live leaves, leaf length, leaf width, leaf angle, and specific leaf area. The ambient light conditions affected the values of all traits studied except for specific leaf area. In accordance to ecophysiological expectations for an adaptive response to light, both leaf length and width were significantly greater while the angle between sequential leaves was significantly smaller in the woodland understory than at the exposed dune site. The relationship between leaf traits and vegetative fitness (total leaf area) differed across light habitats as predicted by functional hypotheses. The standardized linear selection gradient (β′) for leaf length and width were positive in sign in both environments, but their magnitude for leaf length was higher in the shade than under full sunlight. Since plasticity of leaf length in the woodland shade has been recognized as adaptive, fitness cost of producing plastic change in leaf length was assessed. In both of the available methods used, the two-step and the multivariate regression procedures, a rather high negative association between the fitness value and the plasticity of leaf length was obtained, indicating a cost of plasticity. The selection gradient for leaf angle was weak and significant only in the woodland understory. Genetic correlations between trait expressions in contrasting light environments were negative in sign and low in magnitude, implying a significant genetic variation for plasticity in these leaf traits. Furthermore, leaf length and leaf width were found to be genetically positively coupled, which indicates that there is a potential for these two traits to evolve toward their optimal phenotypic values even faster than would be expected if they were genetically independent.