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Revealing the environmental pressures determining the frequency of females amongst populations of sexually dimorphic plants is a key research question. Analyses of sex ratio variation have been mainly done in dioecious plants, which misses key plant sexual systems that might represent intermediate stages in the evolution of dioecy from hermaphroditism. • We investigated female frequency across populations of sexually dimorphic plant species in relation to environmental stressors (temperature, precipitation), totaling 342 species, 2011 populations, representing 40 orders and three different sexual systems (dioecy, gynodioecy and subdioecy). We also included the biome where the population was located to test how female frequency may vary more broadly with climate conditions. • After correcting for phylogeny, our results for gynodioecious systems showed a positive relationship between female frequency and increased environmental stress, with the main effects being temperature-related. Subdioecious systems also showed strong positive relationships with temperature, and positive and negative relationships related to precipitation, while no significant effects on sex ratio in dioecious plants were detected. • Combined, we show that female frequencies in an intermediate sexual system on the pathway from hermaphroditism to dioecy respond strongly to environmental stressors and have different selective agents driving female frequency. [ABSTRACT FROM AUTHOR]

Abstract: Most dioecious plants are perennial and subject to trade‐offs between sexual reproduction and vegetative performance. However, these broader life‐history trade‐offs have not usually been incorporated into theoretical analyses of the evolution of separate sexes. One such analysis has indicated that hermaphroditism is favoured over unisexuality when female and male sex functions involve the allocation of nonoverlapping types of resources to each sex function (e.g. allocations of carbon to female function vs. allocations of nitrogen to male function). However, some dioecious plants appear to conform to this pattern of resource allocation, with different resource types allocated to female vs. male sex functions. Using an evolutionarily stable strategy approach, we show that life‐history trade‐offs between sexual reproduction and vegetative performance enable the evolution of unisexual phenotypes even when there are no direct resource‐based trade‐offs between female and male sex functions. This result might help explain the preponderance of perennial life histories among dioecious plants and why many dioecious plants with annual life histories have indeterminate growth with ongoing trade‐offs between sexual reproduction and vegetative growth. [ABSTRACT FROM AUTHOR]

Gene flow in plant populations is heavily affected by species sexual systems. In order to study the effect of sexual systems on genetic structure, we examined plastid and nuclear DNA of 12 dioecious (males and females) and 18 trioecious (males, females and hermaphrodites) populations of Salix myrsinifolia-a boreal shrub with slow range expansion. Populations were located along latitudinal gradients across submarginal and marginal parts of the range. Individuals of each sex morph were all hexaploid. We identified 10 chloroplast DNA haplotypes and scored 205 polymorphic bands with amplified fragment length polymorphism. We found dioecious populations that differed from trioecious populations via the presence of four unique haplotypes and significant difference in Nei's gene diversity index (0.119 vs. 0.116) and down-weighed marker value (1.17 vs. 1.02). The latter parameter, together with haplotype and nucleotide diversity, significantly decreased with latitude similar to the expansion front. Also, we found that 89% of hermaphrodite individuals belong to one distinct in tree parsimony network haplotype. This frequency significantly decreased with latitude towards the expansion front. We suspect that the presence of hermaphrodites in trioecious populations may represent a trade-off between the possibility of producing progeny by single hermaphrodites and genetic variability loss through autogamy. S. myrsinifolia benefits from trioecious sexual systems under colonization events. This phenomenon is no longer a gain closer to the core of the species range. [ABSTRACT FROM AUTHOR]

Most Salix species are characterized by a complex sex structure with populations that contain different ratios of male, female (dioecious populations) and even hermaphroditic individuals (trioecious populations). However, our knowledge of the temporal and spatial variability of such populations is rudimentary. I studied how sexual structure in trioecious populations of Salix myrsinifolia changes over time (from 1995 to 2012) and space in 32 populations across NE Poland and Lithuania. Additionally, the sex of marked individuals was monitored in a population over eight seasons. A number of land cover variables, latitude and population size were investigated to determine what possible mechanisms/factors affect the occurrence of hermaphroditism in this species. Sex structure was found to differ between the northern and southern parts of the study area (primary and secondary ranges), with trioecious populations occurring significantly more often in the secondary part of the range. Females dominated in northern dioecious populations, while sex ratios were balanced in the southern part of the range and in trioecious populations. The mean frequency of hermaphrodites reached 14 % and was decreasing toward the north. The sex structure remained stable over the study period, with an increasing frequency of hermaphrodites (by nearly 6 %). A detailed study of marked individuals revealed that sex is labile in this species. Over 12 % of individuals changed sex from one to three times. Most changes (67 %) were to hermaphroditism, which frequently occurred (34 %) just before death or when entering the post-reproductive period. [ABSTRACT FROM AUTHOR]

Subdioecy is thought to occupy a transitional position in the gynodioecy-dioecy pathway, explaining one of the evolutionary routes from hermaphroditism to dioecy. Quantifying any female reproductive advantage of females versus hermaphrodites is fundamental to examining the spectrum between subdioecy and dioecy; however, this is challenging, as multiple interacting factors, such as pollen limitation and resource availability, affect plant reproduction. We compared the female reproductive success of females and hermaphrodites via a field experiment in which we hand-pollinated individuals of the subdioecious shrub Eurya japonica of similar size growing under similar light conditions. Effects of pollen limitation and seed quality were also evaluated through comparing the results of hand- and natural-pollination treatments and performing additional laboratory and greenhouse experiments. Overall, females had higher fruit set and produced heavier fruit and more seeds than hermaphrodites, and these results were more pronounced for hand-pollinated than for natural-pollinated plants of both sexes. We also found that seeds naturally produced by females had a higher mean germination rate. These results indicate that females had a pronounced advantage in female reproductive success under conditions of no pollen limitation. The sexual difference in the degree of pollen limitation suggests a pollinator-mediated interaction, whereas the higher female reproductive success of females even under natural conditions implies that E. japonica is a good model species for elucidating the later stages of the gynodioecy-dioecy pathway. [ABSTRACT FROM AUTHOR]

In subdioecious populations, functional female, male and hermaphrodite individuals coexist. Subdioecy may be a transitional state towards dioecy or a breakdown of dioecy, although lability in sex expression may maintain subdioecy as a stable condition. To better understand the ecological aspects involved in sex ratio dynamics and breeding system evolution, we studied the pollination and female fitness components of female and hermaphrodite individuals of the subdioecious shrub Fuchsia microphylla. In two natural populations at the Trans-Mexican Volcanic Belt we estimated female frequency and several reproductive components of female and hermaphrodite plants under natural pollination and experimental pollination treatments. Average female frequency was 42%, and on average, 42.5% of hermaphrodites produced fruits. Female plants showed a 17-fold female fertility advantage over hermaphrodites through increased fruit production, as the number of seeds and germination rates did not differ between morphs. Hermaphrodite flowers were larger, with similar nectar production and concentration to female flowers, and pollinators did not show consistent morph preferences. Some hermaphrodites produced fruits autonomously, and female flowers excluded from pollinators produced fruits putatively by apomixis. Fruit production in hermaphrodites, but not in females, was related to height, suggesting increased investment of hermaphrodites in the female function at higher resource status. For sex ratios to be at equilibrium, the female fertility advantage should be reduced about eightfold. However, it may be that hermaphrodites are maintained by producing fruits at no cost to the male function at higher resource status, as the gender plasticity hypothesis proposes. [ABSTRACT FROM AUTHOR]

Bisexualism (and its derivatives) is a rare exception in species which already acquired the dioecious breeding system. Although it occurs in some tree species, it is still rather uncommon, except for willows, in which bisexualism occurs in at least 18 species. In most willows such unusual individuals are subdioecious, hermaphroditic or monoecious, or produce intersexes (staminate flowers transform into pistillate flowers or the other way round). The frequency of non-dioecy can vary from a single individual to whole populations, but is in principle rare. Its possible causes are both environmental and genetic. In Salix sex lability is known to be affected by parasites and anthropogenic habitat change. In interspecific willow crosses there are numerous exceptions from dioecy that are mostly caused by polyploidization. Since sex determination in willows is genetic, environmental factors are expected to determine exclusively phenotypic sex traits. [ABSTRACT FROM AUTHOR]