Reproductive biology and pollinators in two enantiostylous Qualea species (Vochysiaceae) in the Brazilian Cerrado.

Enantiostyly is a floral polymorphism in which two floral forms in the same species differ in deflection of the stigma to right or left position. In monomorphic enantiostylous plants, flowers of the two morphs occur within the same individual, usually in the same proportion. In self-compatible species the function of monomorphic enantiostyly is proposed to increase outcrossing rates and offer a reproductive advantage under pollination limitation. Enantiostylous species are usually self-compatible and show heteranthery, with poricide anthers and pollen as pollinator reward; however, there are families, such as Vochysiaceae, that have different characteristics. We analysed the reproductive system and pollination biology of Qualea parviflora and Q. multiflora, two enantiostylous species from the Brazilian Cerrado that have specific morphological and physiological traits. For this, we characterized flower traits, performed hand pollinations and studied floral visitors. We found no differences between morphs in the proportion of flowers, nectar produced or its concentration, pollen quantity and fruit set. Both species were self-incompatible and quite generalist regarding floral visitors. Enantiostyly in self-incompatible plants seems to confer a reproductive advantage by reducing self-interference resulting from stigma clogging. This novel result helps to expand our knowledge on this complex floral polymorphism and opens new avenues for future research on this topic.

Reciprocal style polymorphisms are not easily categorised: the case of heterostyly in Lithodora and Glandora (Boraginaceae).

Interest in reciprocal floral polymorphisms, such as heterostyly, has increased in recent decades because they can be used as suitable model systems to study mechanisms of outbreeding and disassortative mating in plants. Heterostylous plants are characterised by the presence of discrete morphs that differ in sex organ position and in some other ancillary traits. As regards sex organ deployment, different types of polymorphisms have been described, depending on number and type of discrete classes present in populations and degree of reciprocity between them. However, a clear-cut characterisation of stylar polymorphisms does not appear to be the best approach when there is great variability among populations because of continuous variation of some of traits examined. A recent study in Lithodora sensu lato (recently split into two separate genera, Lithodora and Glandora) showed a wide variation in sex organ position across species in the genus, which warrants precise population analysis of stylar polymorphism and its reciprocity. We provide a detailed morphometric analysis of flower sexual traits and include those considered to be ancillary characters. We report a wide variation in these traits in populations of Lithodora s.l. and highlight the subjectivity of the former characterisation of style polymorphism based on visual inspection. Ancillary traits appear repeatedly in Lithodora and Glandora, particularly in the latter. The appearance of these traits seems to be related to greater reciprocity between sexual whorls in Glandora, with the exception of G. prostrata. These results agree with evolutionary steps proposed in the build-up of heterostyly according to some evolutionary models. We also examined variation in polymorphisms in light of current models for evolution of heterostyly, and, more specifically, we sought to verify the prediction that flower traits as a whole (i.e., flower integration) respond to selective pressure to assure the exact location of pollen on the pollinator body. Most reciprocal populations and species, where between-morph pollen transfer is expected to be higher, would show greater integration. Our results confirm this hypothesis.