RESUMO
The wild carrot (Daucus carota) is famous for its dark flowers in the umbel centre. Several studies have been conducted to figure out their functional significance, but the evolution of the dark centre remains an enigma. In the present paper, we consider all known apioid species with dark-centred umbels to get a deeper understanding of their biology and evolution. Based on herbaria studies, literature and field work, we reconstructed the distribution area of 10 species (7 genera, 6 clades) of Apiaceae-Apioideae. To recognize homology of the dark structures, developmental studies were conducted in Artedia squamata and Echiophora trichophylla Field studies included architecture, flower morph distribution (andromonoecy) and flowering sequence within the plants, abundancy and behaviour of umbel visitors and preliminary manipulation experiments (removal/adding of dark structures). The dark structures are not homologous to each other. In the Daucus alliance, central flowers or umbellets are conspicuous, whereas in other species dark brush-like (A. squamata) or club-shaped structures (Dicyclophora persica, Echinophora trichophylla, Tordylium aegyptiacum, T. cappadocicum) develop from a naked receptacle. Species are andromonoecious, have a modular architecture and flower in multicyclic protandrous sequence. Among the many umbel visitors, beetles were the most abundant group. Only visitors found on umbels in both flowering phases were recognized as possible pollinators. Manipulation experiments indicated that the dark structures influence the behaviour of some, but not all umbel visitors. In Echinophora trichophylla, a massive gall infection was observed. It is evident that the dark structures evolved several times in parallel. The brush- and club-shaped structures are interpreted as the results of mutations affecting umbel development. Dark umbel centres are most likely stabilized by selection due to their general adaptive function. Their appearance in an area known as a hotspot of beetle pollination gives rise to the assumption that they may act as beetle marks.
RESUMO
BACKGROUND AND AIMS: Andromonoecy, the presence of hermaphrodite and male flowers in the same individual, is genetically fixed or induced, e.g. by fruit set. Little is known about the forces triggering andromonoecy in the Apiaceae. In the present study, a natural population of the protandrous Chaerophyllum bulbosum was investigated to elucidate architectural constraints and effects of resource reallocation. METHODS: Three sets of plants (each n = 15) were treated by hand pollination, pollinator exclusion and removal of low-order inflorescences. Fifteen untreated plants were left as controls. KEY RESULTS: Untreated plants produce umbels up to the third branch order, with increasing proportions of male flowers from 15 % (terminal umbel) to 100 % (third-order umbels). Fruit set correspondingly decreases from 70% (terminal umbel) to <10 % (second-order umbels). Insignificant differences from hand-pollinated plants do not reveal any sign of pollinator limitation at the study site. Bagged individuals show the same increase in male flowers with age as untreated plants, indicating that the presence of andromonoecy is not induced by fruit set. After umbel removal, individuals tend to present a higher number of hermaphrodite flowers and fruits in the umbels of second and third order. Three plants (25 %) produced an additional branch order composed of 100 % male umbels. CONCLUSIONS: Inherited andromonoecy and the plastic response to environmental conditions are interpreted as a self-regulating system saving investment costs and optimizing fruit set at the same time.
