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BACKGROUND AND AIMS: Androdioecy, the co-occurrence of males and hermaphrodites, is a rare reproductive system. Males can be maintained if they benefit from a higher male fitness than hermaphrodites, referred to as male advantage. Male advantage can emerge from increased fertility owing to resource reallocation. However, empirical studies usually compare sexual phenotypes over a single flowering season, thus ignoring potential cumulative effects over successive seasons in perennials. In this study, we quantify various components of male fertility advantage, both within and between seasons, in the long-lived perennial shrub Phillyrea angustifolia (Oleaceae). Although, owing to a peculiar diallelic self-incompatibility system and female sterility mutation strictly associated with a breakdown of incompatibility, males do not need fertility advantage to persist in this species, this advantage remains an important determinant of their equilibrium frequency. METHODS: A survey of >1000 full-sib plants allowed us to compare males and hermaphrodites for several components of male fertility. Individuals were characterized for proxies of pollen production and vegetative growth. By analysing maternal progeny, we compared the siring success of males and hermaphrodites. Finally, using a multistate capture-recapture model we assessed, for each sexual morph, how the intensity of flowering in one year impacts next-year growth and reproduction. KEY RESULTS: Males benefitted from a greater vegetative growth and flowering intensity. Within one season, males sired twice as many seeds as equidistant, compatible hermaphroditic competitors. In addition, males more often maintained intense flowering over successive years. Finally, investment in male reproductive function appeared to differ between the two incompatibility groups of hermaphrodites. CONCLUSION: Males, by sparing the cost of female reproduction, have a higher flowering frequency and vegetative growth, both of which contribute to male advantage over an individual lifetime. This suggests that studies analysing sexual phenotypes during only single reproductive periods are likely to provide inadequate estimates of male advantage in perennials.

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Androdioecy, the occurrence of males and hermaphrodites in a single population, is a rare breeding system because the conditions for maintenance of males are restrictive. In the androdioecious shrub Phillyrea angustifolia, high male frequencies are observed in some populations. The species has a sporophytic self-incompatibility (SI) system with two self-incompatibility groups, which ensures that two groups of hermaphrodites can each mate only with the other group, whereas males can fertilize hermaphrodites of both groups. Here, we analyse a population genetic model to investigate the dynamics of such an androdioecious species, assuming that self-incompatibility and sex phenotypes are determined by a single locus. Our model confirms a previous prediction that a slight reproductive advantage of males relative to hermaphrodites allows the maintenance of males at high equilibrium frequencies. The model predicts different equilibria between hermaphrodites of the two SI groups and males, depending on the male advantage, the initial composition of the population and the population size, whose effect is studied through stochastic simulations. Although the model can generate high male frequencies, observed frequencies are considerably higher than the model predicts. We finally discuss how this model may help explain the large male frequency variation observed in other androdioecious species of Oleaceae: some species show only androdioecious populations, as P. angustifolia, whereas others show populations either completely hermaphrodite or androdioecious.

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Inbreeding depression is a major selective factor acting to maintain outcrossing in hermaphroditic plants. Recently it has been shown that environmental conditions may greatly affect the levels of inbreeding depression. In this study, the effects of intraspecific competition, from either crossed or inbred progeny, and plant density on the expression of inbreeding depression were estimated for the allogamous colonizing plant Crepis sancta (Asteraceae). The population used in this experiment showed a very high outcrossing rate (t = 0.99). Inbreeding depression was measured for germination, survival to reproduction, number of capitula, and a multiplicative fitness estimate. We found that inbreeding depression on survival, the number of capitula, and total fitness was the greatest when inbred plants compete with outbred plants. The effect of density on inbreeding depression was less evident. The major implication of our study is that the relative fitness of selfed progeny is strongly influenced by the type of competitors (outbred or inbred). These results support the hypothesis that inbreeding depression varies according to the density and frequency of outbred plants and suggest that it could be considered as a density- and frequency-dependent phenomenon.

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Androdioecy is a rare breeding system in which low male frequency is expected in populations because males require a strong increase in their fertility to be maintained by selection. Phillyrea angustifolia L. has previously been reported as possibly functionally androdioecious. However, 1&rcolon;1 sex ratios have been reported and suggest functional dioecy. In this article, we compared both pollen tube growth and siring success of male and hermaphrodite pollen in two single-donor pollination experiments. We verified at both pre- and postzygotic levels that hermaphrodites produce functional pollen. Self-incompatibility was also clearly established. However, pollen from hermaphrodites was less efficient than male pollen. The probability of a pollen tube growing through the style was higher for male than for hermaphrodite pollen donors, and males sired twice as many fruits as hermaphrodites. The twofold male advantage in relative fecundity was mainly because of lower pollen fertility of hermaphrodites and possible cross-incompatibility among hermaphrodites.

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Growth, tolerance and zinc and cadmium hyperaccumulation of Thlaspi caerulescens populations from three metal contaminated soils and three normal soils were compared under controlled conditions. Individuals of six populations were cultivated on five soils with increasing concentrations of zinc (50-25000 µg g-1 ) and cadmium (1-170 µg g-1 ). There was no mortality of normal soil populations in the four metal-contaminated soils, but plant growth was reduced to half that of populations from metal-contaminated soils. However, in noncontaminated soil, the growth of individuals from normal soils was greater than that of individuals from metal-contaminated soils. Individuals from normal soils concentrated three times more zinc in the aboveground biomass than those from metal-contaminated soils, but the latter accumulated twice as much cadmium. We conclude that populations of T. caerulescens from both normal and metal-contaminated soils are interesting material for phytoextraction of zinc and cadmium, but to optimize the process of phytoextraction it is necessary to combine the extraction potentials of both type of populations.

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The effects of simulated herbivory (early or late defoliation and cutting of the flowering shoot) on the growth and reproduction of three species of monocarpic composite forbs (Crepis pulchra, Picris hieracioides and C. foetida) with different inflorescence architectures were studied in experimental plots. For the three species studied, early defoliation had no significant effect on subsequent growth. In contrast, late defoliation, occurring at the start of the season of drought, had a negative effect on growth and reproduction in the two Crepis species, particularly C. foetida, but had less effect on P. hieracioides. Sexual biomass was more clearly affected by late defoliation than was vegetative biomass, although the effects differed markedly among species possibly as a result of differences in phenology. Clipping the flowering shoot removed about 3 times less biomass than late defoliation and had little effect on vegetative biomass. It had much greater effects on the sexual biomass in P. hieracioides and C. pulchra, and resulted in the production of many shoots sprouting from the rosette, allowing the treated plants to regain a vegetative biomass close to that of control plants. Clipping did however lead to the production of shorter shoots and a reduction in the number of capitula formed. In C. foetida, much branching occurred even when the main shoot was not cut; the architecture of individual plants was therefore only slightly changed by clipping the apical bud and the sexual biomass of this species was not affected by ablation of the flowering shoot. Overcompensation was found in only two families of C. pulchra for vegetative biomass. No over-compensation was found for sexual biomass, despite an increase in the number of flowering shoots in C. pulchra and P. hieracioides following clipping. However situations close to compensation for the vegetative biomass in the three species and in P. hieracioides for the sexual biomass were recorded. The response of the three study species to simulated herbivory were related to their architecture and to the time of defoliation.

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The two dominant species of the Corsican mattoral,Arbutus unedo L. andErica arborea L., can produce abundant sprouts from the lignotuber not only immediately after fire but also more or less continuously in the absence of major disturbance. The lignotuber appears to be more important during the early stages of development; the result is an increase in the number of sprouts during the 25 years following the establishment of the individuals. Later the lignotuber seems to lose the ability to ensure the development of new basal sprouts. A hypothesis is that the presence of a lignotuber is related to the growth form.Arbutus unedo andErica arborea show behaviour intermediate between acrotony and basitony, as the shoots show acrotony, and continuous sprouting is characteristic of basitonic species. The fact that sprouting from the lignotuber is not necessarily a result of fire suggests that the relation between fire and vegetation in the Mediterranean region should be reconsidered.

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Numerical cladistics is applied to the phyletic analysis of the genus Leishmania Ross, 1903. 128 Old World stocks are characterized by means of electrophoretic techniques using 13 enzymes. The 68 electromorphs detected allow to group the stocks into 17 zymodemes. Data are treated with the Farris' method (1970). The resultant cladogram appears to be coherent. It recognizes classical systematics entities. It sets off two sister groups: major-gerbilli on one hand, donovani-tropica-aethiopica on the other. It confirms L. aethiopica and L. gerbilli as archaic groups and points out a certain divergence between L. infantum and L. donovani s. st.

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Using enzyme characters determined by starch gel electrophoresis, the authors have applied the method of Numerical Taxonomy to the genus Leishmania Ross, 1903. Eight isoenzymes (PGI, ME, PGM, GOT, G-6-PDH, 6-PGDH, MDH and IDH) of 146 Old World strains are examined. 35 electromorphs, corresponding to equivalent number of isoenzymes, are identified by this method, and then grouped into 14 zymodemes. These zymodemes were used as Operational Taxonomy Units (OTU) and pairs were compared, using Jaccard's index of similarity. A matrix of association was constructed using these indices and it forms the basis for the taxonomic scheme elaborated. The final relationships are exhibited in the agglomerative dendrogram, constructed using complete linkage. The separation into phylons is confirmed by correspondence analysis. It is concluded that the original lines fall into five groups corresponding to the complexes Leishmania donovani, Leishmania tropica, Leishmania major, Leishmania aethiopica and cf. Leishmania tarentolae. The phylons as recognised by Numerical Taxonomy, can be equated with the taxa of traditional systematics. The phyletic significance of the individualized phylons is then provided by a genetic approach. Thus, it is possible to compute the genetic distances of Nei with allozyme frequency values of 0,0.5 and 1. The new dendrogram is similar to the previous one: Leishmania major constitutes an homogeneous taxon, long isolated from the others. Leishmania donovani and Leishmania tropica have retained a nonnegligible amount of genetic similarity, attesting both a common origin and a more recent evolutionary divergence.

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