Summary1. Flower colour polymorphism is traditionally attributed to pollinator selection although other factors, such as indirect selection on correlated traits, can play an important role. 2. Lysimachia arvensis is a widespread annual species with two colour morphs differing in anthocyanin composition. We explored the hypothesis that colour polymorphism is maintained by selection related to environmental heterogeneity. Morph frequencies and environmental traits were recorded in 51 populations along a wide geographical range. To explore the existence of morph-by-environment interactions, we conducted an experimental study comparing the two morphs under treatments differing in water and light availability. 3. A geographical pattern was found with a negative association between blue frequencies and latitude. The proportion of the blue morph increased with temperature and sunshine hours, but decreased with precipitation. Flowering onset and flower size differed between morphs and scarcely varied across treatments. In contrast, several fitness components such as germination, seedling survival, seedling mass and flower production showed important morph-by-environment interactions. The blue morph showed higher overall male and female fitness in all the treatment combinations excepting in sun-wet conditions where the red morph had higher fitness. 4. Synthesis. Our results indicate that the mechanism of selection on flower colour seems to be related to differences in fitness of both morphs due to abiotic factors. These differences could explain the geographical distribution of flower colour morphs and the maintenance of the colour polymorphism. The marked difference in flowering time between morphs leaves open the potential for assortative mating and speciation in Lysimachia arvensis.

Evolutionary transitions between hermaphroditism and dioecy have occurred numerous times in the land plants. We briefly review the factors thought to be responsible for these transitions, and we provide a synthesis of what has been learned from recent studies of the annual herb Mercurialis annua, in which dioecy (males and females), monoecy (functional hermaphrodites), and androdioecy (males and hermaphrodites) occur in different parts of its geographic range. Previous research on M. annua has revealed the importance of genome duplication and hybridization in the origin of much of the observed variation. Here we show, however, that spatial transitions in the sexual system also occur within the same ploidy level. In particular, we present an analysis, using flow cytometry data, of ploidy variation across a previously unstudied transition between hermaphroditism and androdioecy, in which we find that the sexual-system transition is uncoupled from the shift in ploidy levels. We review recent research that shows that such transitions between sexual systems in M. annua are consistent with differential selection at the regional level for reproductive assurance during colonization. We also present new experimental data that highlight both the importance of the resource status of plants and that of their local mating context in regulating gender strategies and sex ratios. The studies reviewed and the new results presented emphasize the role that shifts in the ecological and genetic context of plant populations may play in causing transitions between sexual systems.

The pollination of Aristolochia involves the temporary confinement of visitors inside the flower. A literature review has shown that some species are visited by one or a few dipteran families, while others are visited by a wider variety of dipterans, but only some of these are effective pollinators. We observed flowering phenology and temporal patterns of pollinator attendance in diverse populations of Aristolochia baetica and A. paucinervis, two species that grow in SW Spain, frequently in mixed populations. The two species had overlapping floral phenologies, extended flowering periods and long-lived flowers. A. baetica attracted a higher number of visitors than A. paucinervis. Drosophilids and, to a lesser extent, phorids, were the main pollinators of A. baetica, whereas in A. paucinervis, phorids were the only pollinators. Attendance to A. paucinervis flowers by phorids in mixed populations was markedly lower than in pure populations. This effect was more evident in years with lower pollinator density. Our results suggest that A. baetica and A. paucinervis may compete for pollinators in mixed populations.

Pollen limitation, resource limitation, fruit abortion, and predation have all been proposed as factors explaining low fruit set in hermaphroditic plants. We conducted a 5-year study combining field observations and pollination experiments to determine the causes of the low fruit set in Aristolochia paucinervis, a Mediterranean species with a specialized pollination system in two populations in SW Spain. Fruit initiation was markedly low, and between 28.6 and 75.0% of the flowering stems did not initiate any fruit. In most flowers, the number of germinated pollen grains was less than the number of ovules, and supplemental pollination significantly increased fruiting, indicating deficient pollination. In A. paucinervis, autonomous self-pollination seems to be a decisive factor in fruit production because the number of germinated pollen and the fruit set from flowers bagged before anthesis were similar to those in free-pollinated flowers. Only in 2005 did flowers that were successfully pollinated outnumber ripened fruits, suggesting that other factors limit fruiting. We found a significant positive correlation between tuber mass and fruit set. Deficient pollination and lack of resources could explain the low fruit set, but the relative consequences seem to vary spatially and temporarily.

To cite this version:M. Arista, R. Berjano, J. Viruel, M. A. Ortiz, M. Talavera, et al.. Uncertain pollination environment promotes the evolution of a stable mixed reproductive system in the self-incompatible Hypochaeris salzmanniana (Asteraceae). Annals of Botany, Oxford University Press (OUP), 2017, 120 (3), pp. 447-456. 10.1093/aob/mcx059. hal-01681643 Uncertain pollination environment promotes the evolution of a stable mixed reproductive system in the self-incompatible Hypochaeris salzmanniana (Asteraceae) Background and aims The transition from outcrossing to selfing is a repeated pattern in angiosperm diversification and according to general theory this transition should occur quickly and mixed reproductive systems should be infrequent. However, a large proportion of flowering plants have mixed reproductive systems, even showing inbreeding depression. Recently, several theoretical studies have shown that mixed mating systems can be stable, but empirical studies supporting these assumptions are still scarce.Methods Hypochaeris salzmanniana, an annual species with populations differing in their self-incompatibility expression, was used as a study case to assess the stability of its mixed reproductive system. Here a descriptive study of the pollination environment was combined with measurements of the stability of the self-incompatibility system, outcrossing rate, reproductive assurance and inbreeding depression in four populations for two consecutive years.Key Results The reproductive system of populations exhibited a geographical pattern: the proportion of plants decreased from west to east. Pollinator environment also varied geographically, being less favourable from west to east. The self-incompatibility expression of some populations changed markedly in only one year. After selfing, progeny was mainly self-compatible, while after outcrossing both self-incompatible and self-compatible plants were produced. In general, both reproductive assurance and high inbreeding depression were found in all populations and years. The lowest values of inbreeding depression were found in 2014 in the easternmost populations, which experienced a marked increase in self-compatibility in 2015.Conclusions The mixed reproductive system of H. salzmanniana seems to be an evolutionarily stable strategy, with selfing conferring reproductive assurance when pollinator attendance is low, but strongly limited by inbreeding depression. The fact that the highest frequencies of self-compatible plants appeared in the environments most unfavourable to pollination suggests that these plants are selected in these sites, although high rates of inbreeding depression should impede the complete loss of self-incompatibility. In H. salzmanniana, year-to-year changes in the frequency of self-incompatible individuals are directly derived from the balance between reproductive assurance and inbreeding depression.

Summary• Speciation via race formation is an important evolutionary process in parasites, producing changes that favour their development on particular host species. Here, the holoparasitic plant Cytinus, which has diverse host species in the family Cistaceae, has been used to study the occurrence of such races.• Amplified fragment length polymorphism (AFLP) analyses were performed on 174 individuals of 22 populations parasitizing 10 Cistaceae species in the Western Mediterranean basin.• Neighbour-joining, multivariate ordination analyses, and individual-based Bayesian analyses, clustered Cytinus populations into five well-characterized genetic races that, overall, agreed with the taxonomic sections of their hosts. In the AMOVA, among-races differences accounted for almost 50% of the genetic variation. The isolation-by-distance model was not supported by a Mantel test among Cytinus populations (r = 0.012; P = 0.456). All races showed low within-population genetic diversity, probably as a result of restricted pollen flow aggravated by flowering asynchrony, restricted seed dispersion, or stochastic processes.• The genetic differentiation among the five races of Cytinus is congruent with the view that these races are well-characterized lineages that have evolved independently as a result of selective pressures imposed by their hosts. This pattern, with genetically distinctive groups associated with the infrageneric sections of the host species, has not been reported previously for parasitic angiosperms.

Determining the sources of floral variation is crucial to the understanding of floral evolution. Architectural effects and phenotypic plasticity in development can play an important role in intraplant floral variation, giving rise to gender dimorphism or sexual specialization. Amphicarpic plants have another source of floral variation that could also be influenced by positional effects. We studied the effects of resource availability and architecture in intraplant floral variability in two ecotypes of the amphicarpic Emex spinosa. Male flowers were smaller than females, irrespective of position or resource availability. Emex spinosa shows gender dimorphism not influenced by positional effects. Flower size differences among positions were mainly due to architecture, because the effects of resources on flower size were minimal. Architectural effects caused a decrease in female flower size from ground to apical positions but an increase in most male traits that resulted in sexual specialization at the end of flowering. In general, the ecotypes were similarly affected by resources or architecture. Differences between subterranean and aerial female flowers seem also to be a consequence of architecture. Our results contribute to the evidence that resource limitation is an overestimated effect and that architectural effects must be considered in studies of floral or fruit variation.

This study compares how Lantana camara, an invasive species, and L. peduncularis, an autochthonous one, cope with drought in Galapagos. Soil surface temperature was the abiotic environmental parameter that best explained variations in photosynthetic stress. Higher soil surface temperatures were recorded in the lowlands and in rain-shadow areas, which were also the driest areas. L. peduncularis, with a shallow root system, behaved as a drought-tolerant species, showing lower relative growth rates, which decreased with leaf water content and higher photosynthetic stress levels in the lowlands and in a northwest rain-shadow area in comparison with higher and wetter locations. Its basal and maximal fluorescences decreased at lower altitudes, reflecting the recorded drops in chlorophyll concentration. In contrast, L. camara with a deep root system behaved as a drought-avoiding species, showing leaf and relative water contents higher than 55% and avoiding permanent damage to its photosynthetic apparatus even in the driest area where it showed very low chlorophyll content. Its relative growth rate decreased more in dry areas in comparison to wetter zones than did that of L. peduncularis, even though it had greater water content. Furthermore, L. camara showed higher water contents, growth rate, and lower photosynthetic stress levels than L. peduncularis in the arid lowlands. Thus, L. peduncularis maintained lower maximum quantum efficiency of photosystem II photochemistry (F v /F m ) than L. camara even at sunrise, due to higher basal fluorescence values with similar maximal fluorescence, which indicated Ó Springer Science+Business Media B.V. 2006 permanent damage to PSII reaction centres. Our results help to explain the success and limitations of L. camara in the invasion of arid and sub-arid environments.