RESUMEN
Within the theoretical framework of the correlation pleiades, floral phenotypic integration has been proposed as a consequence of selection mediated by pollinators acting on floral characters. Here, we analyzed that assumption by studying the floral biology and pollination of the late-winter species Narcissus bulbocodium L. We found that the flowers of N. bulbocodium are pollinator-dependent (mainly on Bombus terrestris) in terms of achieving optimal levels of seed production (xenogamy mean seed-to-ovule ratio 64%). Flowers are phenotypically integrated, and only the inclusion of the stigma within the corona seems to have a positive and significant influence on the deposition of the pollen. It has been hypothesized that by including the stigma within the corolla, the flower has some control over the contact between stigma and pollinators that could lead to an "ordered herkogamy" as a way to promote outcross and avoid self-interference. Therefore, herkogamy was also studied, and while most previous studies have assessed the evolutionary significance of herkogamy by considering its relationship with outcrossing rates, we approach this phenomenon from a novel direction assessing the relationship between a proxy for herkogamy and the precision of the pollination process. Our results seem to support the existence of an optimal herkogamy distance that could maintain maximum levels of both pollen export and (cross) pollen capture. On the basis of the broad variability of herkogamy that we have found in N. bulbocodium and other data in the literature, we discuss the universality of the adaptive origin of herkogamy.
RESUMEN
BACKGROUND AND AIMS: Oxalis pes-caprae is a widespread invasive weed in regions with a Mediterranean climate. In its native habitat (southern Africa) this species has been reported as heterostylous with trimorphic flowers and a self- and morph-incompatible reproductive system. In most of the areas invaded, only a pentaploid short-styled morphotype that reproduces mainly asexually by bulbils is reported, but this has only been confirmed empirically. This study aims to analyse the floral morph proportions in a wide distribution area, test the sexual female success, and explain the causes of low sexual reproduction of this species in the western area of the Mediterranean Basin. METHODS: Fifty-five populations of O. pes-caprae were sampled in the Iberian Peninsula and Morocco to evaluate the floral morph ratio and individual fruit set. In plants from a dimorphic population, hand-pollination experiments were performed to evaluate the effect of the pollen source on pollen tube growth through the style. The ploidy level and genome size of individuals of each floral morph were analysed using flow cytometry. KEY RESULTS: From the populations studied 89.1 % were monomorphic, with most of them containing the short-styled (SS) floral morph, and 10.9 % were dimorphic containing long-styled (LS) and SS morphs. In some of these, isoplethy was verified but no fruit production was observed in any population. A sterile form was also recorded in several populations. Hand-pollination experiments revealed that pollen grains germinated over recipient stigmas. In intermorph crossings, pollen tubes were able to develop and fruit initiation was observed in some cases, while in intramorph pollinations, pollen tube development was sporadic and no fruit initiation was observed. All individuals within each floral form presented the same DNA ploidy level: SS plants were pentaploid and LS and the sterile form were tetraploid. CONCLUSIONS: The low or null sexual reproduction success of this species in the area of invasion studied seems related with the high frequency of monomorphic populations, the unequal proportion of floral morphs in dimorphic populations and the presence of different ploidy levels between SS and LS morphs. The discovery of the occurrence of an LS floral morph and a sterile form, whose invading capacity in these areas is as yet unknown, will be valuable information for management programmes.