Androdioecy is a sexual system in which males co-occur with hermaphrodites, which have both male and female function. Stable androdioecy is rare in nature, and theory suggests that it requires that males sire more than twice as many offspring as hermaphrodites. In several members of the olive family (Oleaceae), androdioecy occurs with higher frequencies of males than predicted by theory. In Phillyrea angustifolia L., we found that high male frequencies can be maintained in natural populations because hermaphrodites belong exclusively to one of two self-incompatibility groups, and thus, each can fertilize only half of all pollen recipients. In contrast, males can pollinate all hermaphrodites. Thus, in this species, the reproductive disadvantage that males face due to the loss of female function is offset by the fact that all males are fully compatible with all pollen recipients.

Tlic reproductive capacities of the females (male-steriles or mS) of l h y m u~ uuigarii L. ha\(. Iwcti i t ivestigated, in terms of size and weight of plants, seed production, germination ability and pollination. The data, collected during nine years under controlled conditions and in natul-al situations are discussed in comparison with the results known in the liter-aturr. It is suggestcd that the large diversity of characteristics promoting sexual polymorphistn could justifv the high and variable rates of tnS met in natural conditions in T. vulgaris.

Nucleocytoplasmic determination of male sterility in Thymus vulgaris L. has been assumed in all papers attempting to explain the remarkably high frequencies of male steriles found in natural populations of this species. This paper provides strong evidence that both nuclear and cytoplasmic genes are involved in the determination of male sterility of this species, giving a complex inheritance. Interpopulation and intrapopulation crosses have shown that the ratio of females versus hermaphrodites among offsprings varied widely from 1∶0 to 1∶1. Furthermore, interpopulation crosses consistently yielded a higher frequency of females than intrapopulation crosses. Nucleocytoplasmic inheritance was demonstrated by an absence of male fertiles in backcrosses and asymmetrical segregation in reciprocal crosses. Molecular analysis of the mitochondrial DNA of some of the parents used in crosses suggested the involvement of different cytoplasms in the inheritance of male steriliy.

SummaryA rare homomorphic diallelic self-incompatibility (DSI) system discovered in Phillyrea angustifolia (family Oleaceae, subtribe Oleinae) can promote the transition from hermaphroditism to androdioecy. If widespread and stable in Oleaceae, DSI may explain the exceptionally high rate of androdioecious species reported in this plant family. Here, we set out to determine whether DSI occurs in another Oleaceae lineage.We tested for DSI in subtribe Fraxininae, a lineage that diverged from subtribe Oleinae c. 40 million yr ago. We explored the compatibility relationships in Fraxinus ornus using 81 hermaphrodites and 25 males from one natural stand and two naturalized populations using intra-and interspecific stigma tests performed on F. ornus and P. angustifolia testers.We uncovered a DSI system with hermaphrodites belonging to one of two selfincompatibility (SI) groups and males compatible with both groups, making for a truly androdioecious reproductive system. The two human-founded populations contained only one of the two SI groups.Our results provide evidence for the evolutionary persistence of DSI. We discuss how its stability over time may have affected transitions to other sexual systems, such as dioecy.

Androdioecy (the coexistence of two genders, cosexuals and males, in a single population) is a rare breeding system. In terms of functional gamete production, androdioecy has been reported in a small number of windpollinated and insect-pollinated species. In this study we examine the floral biology, stability of gender, and fruit and seed production after self-pollination and outcrossing pollination in a potentially androdioecious tree, Fraxinus ornus, in southern France. Individual trees have either hermaphrodite flowers or male flowers, the latter lacking a well-developed gynoecium. The two genders produce morphologically similar pollen and were present in 1 : 1 or slightly male-biased ratios. Only hermaphrodites set fruit during 2-4 yr of observation at four different sites in southern France. Experimental pollinations of eight trees in two different sites showed that hermaphrodites produce viable pollen in dehiscent anthers and viable seeds. Thus, in terms of functional gamete production, F. ornus is a new case of functional androdioecy. Seven hermaphrodites were self-compatible, and such self-compatibility of hermaphrodites might improve the colonizing ability of F. ornus in a region where this species is actively expanding its range.

LEPART, J. & DOMMEE B., 1992. Is Phillyrea angustifolia L. (Oleaceae) an androdioecious species? Observations during two or three years on three natural populations of Phillyrea angustifolia growing in contrasting environments in southern France show that two distinct phenotypes occur in approximately equal proportions: hermaphrodites, which produce pollen and seeds, and males, which produce pollen only. The populations studied are thus morphologically androdioecious. Biometric investigation of the two morphs shows that they are clearly distinguished by (1) the shape of the stigma and (2) the ovary which is well developed in hermaphrodites and aborted in males. Neither size of anthers and corolla, nor pollen germination, differ between the two sexes. In the Camargue, where P. angustifolia is presently an invader, controlled crosses between hermaphrodites (selfing and inter‐individual) are partially successful. In contrast, such crosses are sterile in the well‐established population of the La Gardiole. Hence the Camargue population appears to be functionally androdioecious and La Gardiole functionally dioecious. This breeding system is particularly puzzling, since the occurrence of anemogamy and possible selfing do not fit classical explanations of androdioecy. The ability of hermaphrodites to be functionally male and female could be particularly adaptive in founding populations. Since functional hermaphroditism occurs in expanding populations of the Camargue population, we consider that the androdioecy of P. angustifolia could be a kind of leaky dioecism.

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The purpose of this study was to describe morph-specific patterns of stigma-anther separation and style curling in the distylous Jasminum fruticans. We also examined whether variation in floral traits is correlated with variation in seed production. Stigma-anther separation is more variable in short-styled plants than in long-styled plants. In all populations studied, some plants bear flowers that lack any stigma-anther separation. These plants have the pollen characteristics (size and number) and compatibility relations of short-styled plants. Comparison with other distylous species illustrates that the variability of stigma-anther separation in short-styled plants is a novel finding for a distylous species. Long-styled plants have greater stigma-anther separation than short-styled plants, styles are often curled and protrude from the corolla, and anthers are placed well within the corolla tube. The frequency of long-styled plants with curled styles and the mean degree of style curling were significantly correlated with style length. Short-styled plants have larger corollas than long-styled plants and never have curled styles. Morph ratios are always 50 : 50 in natural populations. Mean values of each floral trait in the two morphs were significantly correlated among populations. There were no consistent differences in fecundity of the two morphs nor any correlation between floral traits and seed set for each morph in natural populations. We discuss the potential causes and significance of the two morph-specific patterns we describe ; reduced stigma-anther separation in short-styled plants and the presence of curled styles in long-styled plants.