Plasticity in sex allocation in the plant Mercurialis annua is greater for hermaphrodites sampled from dimorphic than from monomorphic populations.

Plants are notoriously variable in gender, ranging in sex allocation from purely male through hermaphrodite to purely female. This variation can have both a genetic and an adaptive plastic component. In gynodioecious species, where females co-occur with hermaphrodites, hermaphrodites tend to shift their allocation towards greater maleness when growing under low-resource conditions, either as a result of hermaphrodites shifting away from an expensive female function, or because of enhanced siring advantages in the presence of females. Similarly, in the androdioecious plant Mercurialis annua, where hermaphrodites co-exist with males, hermaphrodites also tend to enhance their relative male allocation under low-resource conditions. Here, we ask whether this response differs between hermaphrodites that have been evolving in the presence of males, in a situation analogous to that supposed for gynodioecious populations, vs. those that have been evolving in their absence. We grew hermaphrodites of M. annua from populations in which males were either present or absent under different levels of nutrient availability and compared their reaction norms. We found that, overall, hermaphrodites from populations with males tended to be more female than those from populations lacking males. Importantly, hermaphrodites' investment in pollen and seed production was more plastic when they came from populations with males than without them, reducing their pollen production at low resource availability and increasing their seed production at high resource availability. These results are consistent with the hypothesis that plasticity in sex allocation is enhanced in hermaphrodites that have likely been exposed to variation in mating opportunities due to fluctuations in the frequency of co-occurring males.

Phenotypic plasticity rather than ecotypic differentiation explains the broad realized niche of a Neotropical orchid species.

Local adaptation is common in plant species, and knowing whether a population is locally adapted has fundamental and applied relevance. However, local adaptation in tropical plants remains largely less studied, and covering this gap is not simple since reciprocal transplantation - the gold standard for detecting local adaptation - is not feasible for most species. Here, we combined genetic, climatic and phenotypic data to investigate ecotypic differentiation, an important aspect of local adaptation, in coastal and inland populations of the orchid Epidendrum fulgens Brongn., a long-lived tropical plant for which reciprocal transplantation would not be feasible. We used nine microsatellite markers to estimate genetic divergence between inland and coastal populations. Moreover, occurrence data and climate data were used to test for differences in the realized niche of those populations. Finally, we assessed saturated water content, leaf specific area, height, and stomatal density in common garden and in situ to investigate the effects of ecotypic differentiation and plasticity on the phenotype. Coastal and inland groups' niches do not overlap, the former occupying a wetter and warmer area. However, this differentiation does not seem to be driven by ecotypic differentiation since there was no positive correlation between genetic structure and climate dissimilarity. Moreover, specific leaf area and leaf saturated water content, which are important phenotypic traits related to soil fertility and drought stress, were rather plastic. We conclude that ecotypic differentiation is absent, since phenotypic plasticity is an important mechanism explaining the niche broadness of this species.

Species-specific functional trait responses in two species coexisting along a shore-to-inland dune gradient.

Coastal dunes are characterised by strong gradients of abiotic stress, typically increasing in severity from inland areas towards the shoreline. Thus, dune gradients represent unique opportunities to study intraspecific responses to environmental changes and to investigate which factors drive community change. This study aims to examine functional trait variation in two coexisting species in response to environmental changes along a dune gradient in NW Spain. Trait convergence was also investigated and compared between both ends of the gradient. We measured functional leaf traits related to plant efficiency in the use of light, water and nutrients, also possible stressors (salt content and pH) and availability of limiting resources (water and nutrients) in the soil. Most soil variables showed changes following a non-directional gradient. Differences in soil variables were site specific and depended on growth of the study species. Structural and functional traits depended on species and/or plant position on the gradient, except for effective quantum yield of PSII and leaf δ15N. The pattern of variation was mostly directional for reflectance indices related to leaf physiology. Multivariate analyses showed significant interspecific differences in the set of traits they exhibited along positions in the gradient. Species also differed in the combination of traits selected under given environmental conditions. Coexisting species display a specific set of traits that reflects different strategies to environmental stress. Our study highlights the overly simplistic nature of some previous studies that assume dune gradients are monotonically directional, without considering that these gradients may be differentially modified by species activity.

Rapid morphological change in UK populations of Impatiens glandulifera.

The highly invasive Impatiens glandulifera (Himalayan balsam) is one of the most prolific and widespread invasive plants in the British Isles. Introduced in the early nineteenth century, it has now been reported in almost every vice county across the UK and is a fierce competitor that has adverse effects on the local community structure. Despite the negative impacts that invaders like I. glandulifera have on local communities, there have been very few studies which address the morphological changes that invasive plant populations have undergone since their initial introduction. This is the first study of its kind to investigate the morphological changes that have occurred in I. glandulifera. 315 herbarium specimens dating from 1865 to 2017 were used to measure changes in morphological traits such as leaf size, flower length and stomatal characteristics. We found that since 1865, there has been a significant reduction in overall leaf size, a significant reduction in stomatal density and a significant increase in the overall flower length. These results highlight the importance of monitoring the evolutionary change in prolific alien species over the course of their invasion, providing useful insights into changes in competitive ability which may prove useful in managing dispersal and providing options for potential management.

Sexual dimorphism at different life stages: early life sexual differences in root growth in Silene latifolia.

Male and female dioecious plants often show sexual dimorphism, differing in morphological, physiological and life-history traits. Most previous studies have focused on differences between males and females during or after reproduction, paying little attention to the pre-reproductive stages of the individuals. Here we assessed the response of male and female individuals of the dioecious plant Silene latifolia to abiotic stress at different life stages, including pre-reproductive (i.e. seedlings and young plants) and reproductive individuals. We measured growth, resource allocation and discrimination against 13C under nutrient deficiency, water stress, as well as their interaction. We observed sexual dimorphism in root growth, with female seedlings having longer main roots than male plants. Pre-reproductive male and female plants also responded differently, in terms of root allocation, to nutrient and water availability. At reproduction, females grew more roots than males when water was not limiting. These differences could help explain the female-skewed sex ratios found in natural populations of S. latifolia. We found no evidence of sexual dimorphism in aboveground dry mass, although females had longer leaves than males at the seedling stage. We conclude that sexual dimorphism in S. latifolia may occur not as a consequence of reproduction, but well before it.

Differential niche modification by males and females of a dioecious herb: extending the Jack Sprat effect.

Males and females of dioecious plants often differ in morphological, physiological and life-history traits, probably as a result of their different requirements for reproduction. We found that the growth and reproductive effort of individuals of the dioecious herb Mercurialis annua depended on whether males or females had been growing in the soil previously. This suggests that males and females of M. annua differentially modify the soil in which they are growing. Our study indicates that sexual dimorphism in dioecious plants can give rise to increased environmental heterogeneity as a consequence of sex-specific niche modification.

Sex-specific physiological, allocation and growth responses to water availability in the subdioecious plant Honckenya peploides.

The gender of dimorphic plant species is often affected by ecophysiological variables. Differences have been interpreted as a response of the sexes to meet specific resource demands associated with reproduction. This study investigated whether sex-specific variations in ecophysiological traits in response to water availability determine the performance of each sex in different habitats, and therefore promote extreme spatial segregation of the sexes in the subdioecious plant, Honckenya peploides. Twenty-seven plants of each sex were individually potted in dune sand and assigned randomly to one of three water treatments. Well-watered plants were watered daily to field capacity, whereas plants in the moderate and high-water stress treatments received 40% and 20%, respectively, of the water given to well-watered plants. Photochemical efficiency, leaf spectral properties and components of relative growth rate (leaf area ratio and net assimilation rate) were measured. Photochemical efficiencies integrated over time were higher in male than in female plants. Water deficit decreased maximum quantum yield in female plants more rapidly than in male plants, but female plants (unlike male plants) had recovered to initial values by the end of the experiment. Maximum quantum yield in male plants was more affected by water stress than in female plants, indicating that male plants were more susceptible to photoinhibition. The two sexes did not differ in growth rate, but male plants invested a higher proportion of their biomass in leaves, had a higher leaf area per unit biomass and lower net assimilation rate relative to female plants. Female plants had a higher water content and succulence than male plants. Differences in stomatal density between the sexes depended on water availability. The results suggest that the two sexes of H. peploides have different strategies for coping with water stress. The study also provides evidence of sex differences in allocation traits. We conclude that between-sex differences in ecophysiological and allocation traits may contribute to explain habitat-related between-sex differences in performance and, therefore, the spatial segregation of the sexes.

Sexual dimorphism in intra- and interspecific competitive ability of the dioecious herb Mercurialis annua.

Males and females of dioecious plant species often show different responses to competition with individuals of the same or opposite gender, but almost no data are available on the outcome of competition with members of other species. Here, we show that male and female individuals of the wind-pollinated herb Mercurialis annua are sexually dimorphic in both their intraspecific and interspecific competitive abilities. In a controlled experiment, we found that both sexes of M. annua were negatively affected by interspecific competition, but the sensitivity of males and females depended on the identity of their competitor species, with females tending to suppress the aboveground growth of competitor species more than males. Further, we found that intrasexual and intersexual competition affected the aboveground growth of males but not that of females: only males showed a significant reduction in growth when growing with conspecific competitors (male or female). We discuss our results with reference to related studies that suggest that males and females of M. annua have different resource requirements for reproduction, which in turn affect their competitive abilities.

Unexpectedly high genetic variation in large unisexual clumps of the subdioecious plant Honckenya peploides (Caryophyllaceae).

Honckenya peploides is a subdioecious dune plant that reproduces both sexually and by clonal growth. In northwest Spain this species was found to exhibit an extreme spatial segregation of the sexes, and our objective was to investigate genetic variation in unisexual clumps. Genetic variation was studied in six unisexual clumps of H. peploides, three of them exclusively composed of males and three exclusively female. In total, 193 samples were analysed using isozyme analysis and 80 samples were analysed using two AFLP primer combinations. Both techniques revealed considerably high genetic diversity (average proportion of distinguishable genotypes: 0.22 for isozymes and 0.36 for AFLP; average Simpson's D: 0.65 for isozymes and 0.68 for AFLP). Our results show that, in spite of clonal growth, each unisexual clump consists of different genotypes. Genetic diversity within clumps is similar for both sexual morphs. Reasons for unisexuality of the clumps are discussed.