Fruit wings accelerate germination in Anacyclus clavatus.

PREMISE: The lateral membranous expansions of fruits, commonly referred to as wings, have long been theorized to serve only dispersal functions. Alternatively, because winged fruits typically have earlier seed germination than unwinged fruits, we hypothesized that wings could increase the contact surface with water, ultimately triggering earlier germination. METHODS: We investigated this alternative hypothesis by exploring the potential role of fruit wings on germination in the heterocarpic species Anacyclus clavatus (Desf.) Pers. (Asteraceae), which produces both winged and unwinged fruits. First, we measured the speed and degree of water absorption in winged and unwinged fruits. Second, we investigated the effects of wings on germination performance, by either reducing wing size or by preventing water absorption by sealing wings with wax. Next, we tested the influence of water availability on the germination performance of winged and unwinged fruits by reducing the water potential. RESULTS: Winged fruits absorbed more water at a faster rate than unwinged fruits. The sealing of wings delayed germination, whereas germination time was not significantly altered by wing cutting. The restriction of water availability by decreasing water potential significantly delayed seed germination of unwinged fruits, whereas winged fruits remained unaffected. CONCLUSIONS: Altogether, our results support the effect of wings on germination and cast doubt on the unique role of wings in dispersal. Whether or not wings contribute to dispersal, we propose that they also improve seed germination and seedling establishment by facilitating water absorption after the release from their mother plants.

Unravelling plant diversification: Intraspecific genetic differentiation in hybridizing Anacyclus species in the western Mediterranean Basin.

PREMISE: The interfertile species Anacyclus clavatus, A. homogamos, and A. valentinus represent a plant complex coexisting in large anthropic areas of the western Mediterranean Basin with phenotypically mixed populations exhibiting a great floral variation. The goal of this study was to estimate the genetic identity of each species, to infer the role of hybridization in the observed phenotypic diversity, and to explore the effect of climate on the geographic distribution of species and genetic clusters. METHODS: We used eight nuclear microsatellites to genotype 585 individuals from 31 populations of three Anacyclus species for population genetic analyses by using clustering algorithms based on Bayesian models and ordination methods. In addition, we used ecological niche models and niche overlap analyses for both the species and genetic clusters. We used an expanded data set, including 721 individuals from 129 populations for ecological niche models of the genetic clusters. RESULTS: We found a clear correspondence between species and genetic clusters, except for A. clavatus that included up to three genetic clusters. We detected individuals with admixed genetic ancestry in A. clavatus and in mixed populations. Ecological niche models predicted similar distributions for species and genetic clusters. For the two specific genetic clusters of A. clavatus, ecological niche models predicted remarkably different areas. CONCLUSIONS: Gene flow between Anacyclus species likely explains phenotypic diversity in contact areas. In addition, we suggest that introgression could be involved in the origin of one of the two A. clavatus genetic clusters, which also showed ecological differentiation.

Within-individual phenotypic plasticity in flowers fosters pollination niche shift.

Phenotypic plasticity, the ability of a genotype of producing different phenotypes when exposed to different environments, may impact ecological interactions. We study here how within-individual plasticity in Moricandia arvensis flowers modifies its pollination niche. During spring, this plant produces large, cross-shaped, UV-reflecting lilac flowers attracting mostly long-tongued large bees. However, unlike most co-occurring species, M. arvensis keeps flowering during the hot, dry summer due to its plasticity in key vegetative traits. Changes in temperature and photoperiod in summer trigger changes in gene expression and the production of small, rounded, UV-absorbing white flowers that attract a different assemblage of generalist pollinators. This shift in pollination niche potentially allows successful reproduction in harsh conditions, facilitating M. arvensis to face anthropogenic perturbations and climate change.

The Mendelian inheritance of gynomonoecy: insights from Anacyclus hybridizing species.

PREMISE: Gynomonoecy is an infrequent sexual system in angiosperms, although widely represented within the Asteraceae family. Currently, the hypothesis of two nuclear loci controling gynomonoecy is the most accepted. However, the genic interactions are still uncertain. Anacyclus clavatus, A. homogamos, and A. valentinus differ in their sexual system and floral traits. Here, we investigate the inheritance of gynomonoecy in this model system to understand its prevalence in the family. METHODS: We selected six natural populations (two per species) for intra- and interspecific experimental crosses, and generated a total of 1123 individuals from the F1 generation, F2 , and backcrosses for sexual system characterization. The frequency of gynomonoecy observed for each cross was tested to fit different possible hypotheses of genic interaction. Additionally, the breeding system and the degree of reproductive isolation between these species were assessed. RESULTS: Complementary epistasis, in which two dominant alleles are required for trait expression, explained the frequencies of gynomonoecy observed across all generations. The heterozygosity inferred in Anacyclus valentinus, as well as its lower and variable seed set, is congruent with its hybrid origin. CONCLUSIONS: In our model system gynomonoecy is controlled by complementary epistasis of two genes. A common origin of this sexual system in Asteraceae, in which genic duplications, mutations, and hybridization between lineages played a key role, is hypothesized whereas independent evolutionary pathways and possibly diverse underlying genetic factors are suggested for gynomonoecy expression in other angiosperm families.

Phylogenetic, functional, and taxonomic richness have both positive and negative effects on ecosystem multifunctionality.

Biodiversity encompasses multiple attributes such as the richness and abundance of species (taxonomic diversity), the presence of different evolutionary lineages (phylogenetic diversity), and the variety of growth forms and resource use strategies (functional diversity). These biodiversity attributes do not necessarily relate to each other and may have contrasting effects on ecosystem functioning. However, how they simultaneously influence the provision of multiple ecosystem functions related to carbon, nitrogen, and phosphorus cycling (multifunctionality) remains unknown. We evaluated the effects of the taxonomic, phylogenetic, and functional attributes of dominant (mass ratio effects) and subordinate (richness effect) plant species on the multifunctionality of 123 drylands from six continents. Our results highlight the importance of the phylogenetic and functional attributes of subordinate species as key drivers of multifunctionality. In addition to a higher taxonomic richness, we found that simultaneously increasing the richness of early diverging lineages and the functional redundancy between species increased multifunctionality. In contrast, the richness of most recent evolutionary lineages and the functional and phylogenetic attributes of dominant plant species (mass ratio effects) were weakly correlated with multifunctionality. However, they were important drivers of individual nutrient cycles. By identifying which biodiversity attributes contribute the most to multifunctionality, our results can guide restoration efforts aiming to maximize either multifunctionality or particular nutrient cycles, a critical step to combat dryland desertification worldwide.

Evolutionary history of the buildup and breakdown of the heterostylous syndrome in Plumbaginaceae.

The evolutionary pathways leading to the heterostylous syndrome are not well understood, and models concerning the origins of distyly differ in the order in which reciprocal herkogamy and self-incompatibility evolve. We investigated the evolution and breakdown of distyly in Plumbaginaceae, a family with considerable diversity of floral traits and reproductive systems. Using Bayesian Markov chain Monte Carlo analyses and stochastic character mapping, we examined the evolutionary assembly and breakdown of the heterostylous syndrome based on a well-resolved phylogeny of 121 species of Plumbaginaceae and six outgroup taxa using five nuclear and plastid gene regions. We used the distribution of reproductive traits and reconstructed ancestral characters across phylogenies to evaluate competing models for the evolution of distyly. The most likely common ancestor of Plumbaginaceae was self-incompatible and monomorphic for sex-organ arrangement and pollen-stigma characters. Character state reconstructions indicated that reciprocal herkogamy evolved at least three times and that shifts to selfing and apomixis occurred on multiple occasions. Our results provide comparative support for the early ideas of H. G. Baker on evolutionary pathways in Plumbaginaceae, and the more recent selfing avoidance model by D. & B. Charlesworth in which distyly evolves from self-incompatible ancestors.

Fitness benefits and costs of floral advertising traits: insights from rayed and rayless phenotypes of Anacyclus (Asteraceae).

PREMISE OF THE STUDY: Ray flowers commonly observed in daisies' flowering heads are a well-known example of advertising structures for enhancing pollinator attraction. Despite this, ray loss has occurred in multiple lineages, which still rely on pollinators, suggesting that rayless phenotypes could also be adaptive for animal-pollination. Here, we investigate the benefits and costs of these specialized floral advertising structures by comparing rayed and rayless phenotypes in two hybridizing closely related species. METHODS: We assessed the advantages and costs of ray production in terms of floral visitor's attraction, pollen limitation, and female reproductive success using the broad natural variation on ray size and number at the contact zone of A. clavatus (rayed) and A. valentinus (rayless). In addition, we experimentally explored the effect of rays under controlled neighborhoods and the effect of ray removal on fruit production. KEY RESULTS: In sympatry, rayed phenotypes attracted significantly more visitors than rayless plants, in which seed production was pollen limited. However, rayed phenotypes did not show higher fruit set or seed production than rayless phenotypes. Fruit set and seed production benefited from denser neighborhood displays and larger individual floral displays, respectively. The removal of ray florets did not appear to enable resource reallocation to fruit production. CONCLUSIONS: Rayless heads compensated their lower visitation rate by means of a higher number of flowers per head achieving similar fecundity levels to rayed plants. The larger size of rayless heads might thus indicate an inflorescence-level trade-off between attraction and fertility.

Characterization of microsatellite markers for Moricandia moricandioides (Brassicaceae) and related species.

PREMISE OF THE STUDY: Polymorphic microsatellite markers were developed to study population structure and mating patterns of the monocarpic herb Moricandia moricandioides (Brassicaceae). METHODS AND RESULTS: Illumina MiSeq sequencing was used to develop a panel of 15 polymorphic microsatellite markers that were tested across 77 individuals from three populations on the Iberian Peninsula. All markers were polymorphic in at least two studied populations, and the number of alleles ranged from one to 11 per locus. The levels of observed and expected heterozygosity ranged from 0.000 to 1.000 and from 0.153 to 0.865, respectively. Nine and 11 loci were successfully amplified in the congeneric species M. arvensis and M. foetida, respectively. CONCLUSIONS: The 15 microsatellite markers will be useful for population genetic studies of the genus Moricandia. These markers will serve as a useful tool for exploring population structure and mating patterns of M. moricandioides.

Kin discrimination allows plants to modify investment towards pollinator attraction.

Pollinators tend to be preferentially attracted to large floral displays that may comprise more than one plant in a patch. Attracting pollinators thus not only benefits individuals investing in advertising, but also other plants in a patch through a 'magnet' effect. Accordingly, there could be an indirect fitness advantage to greater investment in costly floral displays by plants in kin-structured groups than when in groups of unrelated individuals. Here, we seek evidence for this strategy by manipulating relatedness in groups of the plant Moricandia moricandioides, an insect-pollinated herb that typically grows in patches. As predicted, individuals growing with kin, particularly at high density, produced larger floral displays than those growing with non-kin. Investment in attracting pollinators was thus moulded by the presence and relatedness of neighbours, exemplifying the importance of kin recognition in the evolution of plant reproductive strategies.

The Evolution of Haploid Chromosome Numbers in the Sunflower Family.

Chromosome number changes during the evolution of angiosperms are likely to have played a major role in speciation. Their study is of utmost importance, especially now, as a probabilistic model is available to study chromosome evolution within a phylogenetic framework. In the present study, likelihood models of chromosome number evolution were fitted to the largest family of flowering plants, the Asteraceae. Specifically, a phylogenetic supertree of this family was used to reconstruct the ancestral chromosome number and infer genomic events. Our approach inferred that the ancestral chromosome number of the family is n = 9. Also, according to the model that best explained our data, the evolution of haploid chromosome numbers in Asteraceae was a very dynamic process, with genome duplications and descending dysploidy being the most frequent genomic events in the evolution of this family. This model inferred more than one hundred whole genome duplication events; however, it did not find evidence for a paleopolyploidization at the base of this family, which has previously been hypothesized on the basis of sequence data from a limited number of species. The obtained results and potential causes of these discrepancies are discussed.

The role of pollinators in the evolution of corolla shape variation, disparity and integration in a highly diversified plant family with a conserved floral bauplan.

BACKGROUND AND AIMS: Brassicaceae is one of the most diversified families in the angiosperms. However, most species from this family exhibit a very similar floral bauplan. In this study, we explore the Brassicaceae floral morphospace, examining how corolla shape variation (an estimation of developmental robustness), integration and disparity vary among phylogenetically related species. Our aim is to check whether these floral attributes have evolved in this family despite its apparent morphological conservation, and to test the role of pollinators in driving this evolution. METHODS: Using geometric morphometric tools, we calculated the phenotypic variation, disparity and integration of the corolla shape of 111 Brassicaceae taxa. We subsequently inferred the phylogenetic relationships of these taxa and explored the evolutionary lability of corolla shape. Finally, we sampled the pollinator assemblages of every taxon included in this study, and determined their pollination niches using a modularity algorithm. We explore the relationship between pollination niche and the attributes of corolla shape. KEY RESULTS: Phylogenetic signal was weak for all corolla shape attributes. All taxa had generalized pollination systems. Nevertheless, they belong to different pollination niches. There were significant differences in corolla shape among pollination niches even after controlling for the phylogenetic relationship of the plant taxa. Corolla shape variation and disparity was significantly higher in those taxa visited mostly by nocturnal moths, indicating that this pollination niche is associated with a lack of developmental robustness. Corolla integration was higher in those taxa visited mostly by hovering long-tongued flies and long-tongued large bees. CONCLUSIONS: Corolla variation, integration and disparity were evolutionarily labile and evolved very recently in the evolutionary history of the Brassicaceae. These floral attributes were strongly related to the pollination niche. Even in a plant clade having a very generalized pollination system and exhibiting a conserved floral bauplan, pollinators can drive the evolution of important developmental attributes of corolla shape.

Pre-dispersal predation effect on seed packaging strategies and seed viability.

An increased understanding of intraspecific seed packaging (i.e. seed size/number strategy) variation across different environments may improve current knowledge of the ecological forces that drive seed evolution in plants. In particular, pre-dispersal seed predation may influence seed packaging strategies, triggering a reduction of the resources allocated to undamaged seeds within the preyed fruits. Assessing plant reactions to pre-dispersal seed predation is crucial to a better understanding of predation effects, but the response of plants to arthropod attacks remains unexplored. We have assessed the effect of cone predation on the size and viability of undamaged seeds in populations of Juniperus thurifera with contrasting seed packaging strategies, namely, North African populations with single-large-seeded cones and South European populations with multi-small-seeded cones. Our results show that the incidence of predation was lower on the single-large-seeded African cones than on the multi-small-seeded European ones. Seeds from non-preyed cones were also larger and had a higher germination success than uneaten seeds from preyed cones, but only in populations with multi-seeded cones and in cones attacked by Trisetacus sp., suggesting a differential plastic response to predation. It is possible that pre-dispersal seed predation has been a strong selective pressure in European populations with high cone predation rates, being a process which maintains multi-small-seeded cones and empty seeds as a strategy to save some seeds from predation. Conversely, pre-dispersal predation might not have a strong effect in the African populations with single-large-seeded cones characterized by seed germination and filling rates higher than those in the European populations. Our results indicate that differences in pre-dispersal seed predators and predation levels may affect both selection on and intraspecific variation in seed packaging.

PHENIX: An R package to estimate a size-controlled phenotypic integration index.

PREMISE OF THE STUDY: Organisms usually show intercorrelations between all or some of their components leading to phenotypic integration, which may have deep consequences on the evolution of phenotypes. One of the main difficulties with phenotypic integration studies is how to correct the integration measures for size. This has been considered a challenging task. In this paper, we introduce an R package (PHENIX: PHENotypic Integration indeX), in which we provide functions to estimate a size-controlled phenotypic integration index, a bootstrapping method to calculate confidence intervals, and a randomization method to simulate null distributions and test the statistical significance of the integration. METHODS AND RESULTS: PHENIX is an open source package written in R. As usual for R packages, the manual and sample data are available at: http://cran.r-project.org/web/packages/PHENIX/index.html. Functions included in this package easily estimate phenotypic integration by controlling a third variable (e.g., the size of the studied organ). CONCLUSIONS: PHENIX helps to estimate and test the statistical significance of the magnitude of integration using one of the most-used methodological approaches, while taking size into account.

The effects of achene type and germination time on plant performance in the heterocarpic Anacyclus clavatus (Asteraceae).

UNLABELLED: • PREMISE OF THE STUDY: In heterocarpy, fruits with different morphologies have been associated with alternative strategies of dispersal, germination, dormancy, and seedling competitive ability. In heterocarpic species, it is common to find fruits with competitive or dispersal syndromes. The competitive advantage of nondispersing fruits has been frequently attributed to their larger size, but recent studies have suggested that this could also be mediated by germination time. The main objective of our study was to investigate which factor, fruit type or germination time, most affects plant performance and, consequently, competitive ability, using the heterocarpic species Anacyclus clavatus• METHODS: To explore the effects of achene type and germination time on plant performance, we followed an innovative experimental approach including two experiments: one allowing for differences in germination time, and the other evaluating the effect of achene type alone by synchronizing germination time.• KEY RESULTS: A significant effect of germination time on several postdispersal life-history traits was observed: Achenes that germinated earlier produced plants with higher biomass and reproductive effort. When germination time was controlled, no significant differences were observed in any of the traits.• CONCLUSIONS: The competitive advantage of achenes with different morphologies was mainly mediated by germination time and not by differences in size or other intrinsic traits. The consequences of these results are discussed in light of the dispersal-competition trade-off. Our experimental approach (i.e., the synchronization of germination time) revealed the importance of manipulative experiments for testing the effects of germination time on plant survival and performance.

Functional traits determine plant co-occurrence more than environment or evolutionary relatedness in global drylands.

Plant-plant interactions are driven by environmental conditions, evolutionary relationships (ER) and the functional traits of the plants involved. However, studies addressing the relative importance of these drivers are rare, but crucial to improve our predictions of the effects of plant-plant interactions on plant communities and of how they respond to differing environmental conditions. To analyze the relative importance of -and interrelationships among- these factors as drivers of plant-plant interactions, we analyzed perennial plant co-occurrence at 106 dryland plant communities established across rainfall gradients in nine countries. We used structural equation modeling to disentangle the relationships between environmental conditions (aridity and soil fertility), functional traits extracted from the literature, and ER, and to assess their relative importance as drivers of the 929 pairwise plant-plant co-occurrence levels measured. Functional traits, specifically facilitated plants' height and nurse growth form, were of primary importance, and modulated the effect of the environment and ER on plant-plant interactions. Environmental conditions and ER were important mainly for those interactions involving woody and graminoid nurses, respectively. The relative importance of different plant-plant interaction drivers (ER, functional traits, and the environment) varied depending on the region considered, illustrating the difficulty of predicting the outcome of plant-plant interactions at broader spatial scales. In our global-scale study on drylands, plant-plant interactions were more strongly related to functional traits of the species involved than to the environmental variables considered. Thus, moving to a trait-based facilitation/competition approach help to predict that: 1) positive plant-plant interactions are more likely to occur for taller facilitated species in drylands, and 2) plant-plant interactions within woody-dominated ecosystems might be more sensitive to changing environmental conditions than those within grasslands. By providing insights on which species are likely to better perform beneath a given neighbour, our results will also help to succeed in restoration practices involving the use of nurse plants.

Floral development and evolution of capitulum structure in Anacyclus (Anthemideae, Asteraceae).

BACKGROUND AND AIMS: Most of the diversity in the pseudanthia of Asteraceae is based on the differential symmetry and sexuality of its flowers. In Anacyclus, where there are (1) homogamous capitula, with bisexual, mainly actinomorphic and pentamerous flowers; and (2) heterogamous capitula, with peripheral zygomorphic, trimerous and long-/short-rayed female flowers, the floral ontogeny was investigated to infer their origin. METHODS: Floral morphology and ontogeny were studied using scanning electron microscope and light microscope techniques. KEY RESULTS: Disc flowers, subtended by paleae, initiate acropetally. Perianth and androecium initiation is unidirectional/simultaneous. Late zygomorphy occurs by enlargement of the adaxial perianth lobes. In contrast, ray flowers, subtended by involucral bracts, initiate after the proximal disc buds, breaking the inflorescence acropetal pattern. Early zygomorphy is manifested through the fusion of the lateral and abaxial perianth lobes and the arrest of the adaxials. We report atypical phenotypes with peripheral 'trumpet' flowers from natural populations. The peripheral 'trumpet' buds initiate after disc flowers, but maintain an actinomorphic perianth. All phenotypes are compared and interpreted in the context of alternative scenarios for the origin of the capitulum and the perianth identity. CONCLUSIONS: Homogamous inflorescences display a uniform floral morphology and development, whereas the peripheral buds in heterogamous capitula display remarkable plasticity. Disc and ray flowers follow different floral developmental pathways. Peripheral zygomorphic flowers initiate after the proximal actinomorphic disc flowers, behaving as lateral independent units of the pseudanthial disc from inception. The perianth and the androecium are the most variable whorls across the different types of flowers, but their changes are not correlated. Lack of homology between hypanthial appendages and a calyx, and the perianth double-sided structure are discussed for Anacyclus together with potential causes of its ray flower plasticity.

Evolutionary relationships can be more important than abiotic conditions in predicting the outcome of plant-plant interactions.

Positive and negative plant-plant interactions are major processes shaping plant communities. They are affected by environmental conditions and evolutionary relationships among the interacting plants. However, the generality of these factors as drivers of pairwise plant interactions and their combined effects remain virtually unknown. We conducted an observational study to assess how environmental conditions (altitude, temperature, irradiance and rainfall), the dispersal mechanism of beneficiary species and evolutionary relationships affected the co-occurrence of pairwise interactions in 11 Stipa tenacissima steppes located along an environmental gradient in Spain. We studied 197 pairwise plant-plant interactions involving the two major nurse plants (the resprouting shrub Quercus coccifera and the tussock grass S. tenacissima) found in these communities. The relative importance of the studied factors varied with the nurse species considered. None of the factors studied were good predictors of the co-ocurrence between S. tenacissima and its neighbours. However, both the dispersal mechanism of the beneficiary species and the phylogenetic distance between interacting species were crucial factors affecting the co-occurrence between Q. coccifera and its neighbours, while climatic conditions (irradiance) played a secondary role. Values of phylogenetic distance between 207-272.8 Myr led to competition, while values outside this range or fleshy-fruitness in the beneficiary species led to positive interactions. The low importance of environmental conditions as a general driver of pairwise interactions was caused by the species-specific response to changes in either rainfall or radiation. This result suggests that factors other than climatic conditions must be included in theoretical models aimed to generally predict the outcome of plant-plant interactions. Our study helps to improve current theory on plant-plant interactions and to understand how these interactions can respond to expected modifications in species composition and climate associated to ongoing global environmental change.

Environmental conditions and biotic interactions acting together promote phylogenetic randomness in semi-arid plant communities: new methods help to avoid misleading conclusions.

QUESTIONS: Molecular phylogenies are being increasingly used to better understand the mechanisms structuring natural communities. The prevalent theory is that environmental factors and biotic interactions promote the phylogenetic clustering and overdispersion of plant communities, respectively. However, both environmental filtering and biotic interactions are very likely to interact in most natural communities, jointly affecting community phylogenetic structure. We asked the following question: How do environmental filters and biotic interactions jointly affect the phylogenetic structure of plant communities across environmental gradients? LOCATION: Eleven Stipa tenacissima L. grasslands located along an environmental gradient from central to south-east Spain, which covers the core of the distribution area of this vegetation type in Europe. METHODS: We jointly evaluated the effects of environmental conditions and plant-plant interactions on the phylogenetic structure -measured with the mean phylogenetic distance index- of the studied communities. As an indicator of environmental conditions, we used a PCA ordination including eight climatic variables. Different metrics were used to measure the following processes: i) competition/facilitation shifts at the entire community level (Species Combination index), and ii) the effect of the microclimatic amelioration provided by the two most important nurses on neighbours composition (similarity indices and comparison of the phylogenetic pattern between canopy patches and bare ground areas). RESULTS: Biotic interactions and, to a less extent, environmental conditions, affected the phylogenetic pattern of the studied communities. While positive plant-plant interactions (both at the community and the scale of individual nurses) increased phylogenetic overdispersion, higher rainfall increased phylogenetic clustering. The opposed effects of environmental conditions and biotic interactions could be the main cause of the overall random phylogenetic structure found in most of these communities. CONCLUSIONS: Our results illustrate, for the first time, how an overall random phylogenetic pattern may not be promoted only by the lack of influence of either environmental filtering or biotic interactions, but rather by their joint and opposing effects. They caution about making inferences about the underlying mechanisms shaping plant communities from the sole use of their phylogenetic pattern. We also provide a comprehensive set of easy-to-measure tools to avoid misleading conclusions when interpreting phylogenetic structure data obtained from observational studies.

Where do monomorphic sexual systems fit in the evolution of dioecy? Insights from the largest family of angiosperms.

A range of hypothesized evolutionary pathways has been proposed for describing the evolution of dioecy. However, the evolutionary links between other sexual systems not directly involved in dioecy evolution remain largely unexplored, and hence, a comprehensive picture of evolutionary transitions between sexual systems is still lacking. Here, we explored the diversity and evolution of sexual systems in Asteraceae, the largest family of flowering plants, where almost all sexual systems are present. We used a phylogenetic approach to build a model of evolutionary transitions between sexual systems. The best model involved nine transitions, including those from hermaphroditism to andromonoecy, gynomonoecy and gynodioecy, those from gynomonoecy to monoecy and trimonoecy, two transitions to dioecy -one through gynodioecy and the other through monoecy - and reversals from monoecy to gynomonoecy and from gynomonoecy to hermaphroditism. Our reconstruction of the evolution of sexual systems in Asteraceae provided, for the first time, a joint view of the evolutionary transitions between seven sexual systems, unveiling the evolutionary links between monomorphic sexual systems. A pathway from hermaphroditism to monoecy through gynomonoecy, instead of from andromonoecy, was highly supported, which was consistent with a gradient of floral gender specialization.

Fruit size decline from the margin to the center of capitula is the result of resource competition and architectural constraints.

Plants produce repeated structures, such as leaves, flowers, and fruits, which differ in size and shape. One example of this is fruit size, which is commonly observed to decrease from proximal to distal positions within an inflorescence. The resource limitation hypothesis proposes that because proximal fruits usually develop first, they have temporal priority on access to resources over distal fruits. The non-mutually exclusive architectural effects hypothesis suggests that these position effects in fruit size may also be due to inherent architectural variation along infructescence axes. We separated out the effects of resource competition and inflorescence architecture by removing the outer or the inner flowers within capitula of Tragopogon porrifolius. We also studied if fruit position influenced germination and seedling performance in order to assess fitness consequences of position effects. Inner fruits were significantly heavier when outer flowers were removed. However, outer fruits did not significantly increase when inner flowers were removed, suggesting later fruits were limited by the development of early fruits. Our findings also suggest that architectural constraints restricted the size of inner fruits in comparison with outer ones. We found that both resource competition and inflorescence architecture affected the fruit size of T. porrifolius, even though this species does not have linear, indeterminate inflorescences. We advance the hypothesis that, when such effects on fitness occur, resource competition-mediated position effects could turn, in evolutionary time, into architectural position effects.