Labellar Structure of the Maxillaria splendens Alliance (Orchidaceae: Maxillariinae) Indicates Floral Polyphenols as a Reward for Stingless Bees.

Several studies have reported stingless Meliponini bees gathering hairs from the labella of Maxillaria spp., including M. ochroleuca, a member of the M. splendens alliance. Such hairs usually contain food materials and are thought to have nutritional value. The papillose labella of representatives of the Maxillaria splendens alliance, however, bear scattered, simple 1-5-celled uniseriate trichomes (hairs) that lack food materials. By contrast, here, as well as polyphenolic compounds, typical labellar papillae usually contain small quantities of starch, protein, and minute droplets of lipid, the last probably involved in the production of fragrance. Towards the labellum apex occur elevated groups of papillae that lack food materials, but contain volatile compounds, probably fragrance precursors. In the past, the terms 'trichomes' or 'hairs' and 'papillae' have been used interchangeably, causing some confusion. Since the trichomes, however, unlike the papillae, are easily detachable and can fragment, it is most likely they, not the papillae, that have previously been observed being collected by bees, but their poor food content indicates that they do not function as food-hairs. Even so, our field observations of M. ochroleuca reveal that stingless bees scrape polyphenol-rich labellar tissue and possibly use this material to produce a resinous, complex, heterogeneous substance commonly referred to as 'bee glue', used for nest construction and repair.

Evidence of floral rewards in Brasiliorchis supports the convergent evolution of food-hairs in Maxillariinae.

PREMISE: Angiosperms distributed over a large geographical area may display considerable phenotypic variation that can be recognized at morphological and micromorphological levels. Here, we investigate the pollination biology and the presence of floral rewards in Brazilian populations of the widely distributed orchid, Brasiliorchis picta. Based on the new data presented here this study investigates the evolution of floral rewards in Maxillariinae, and tests for the occurrence of convergent evolution of food-hairs in this subtribe. METHODS: Micromorphological and histochemical analyses of the labellar tissues were conducted, together with chemical analysis of fragrance and experiments involving the use of chemical baits. The evolution of floral rewards in Maxillariinae were addressed. RESULTS: Microscopy revealed that B. picta offers food-hairs as a reward. Fragrance is produced by abaxially located labellar epidermal papillae. The main compound present in our samples (2-phenylethanol) also occurs in the aggregation pheromone produced by the mandible glands of pollinators, Meliponini bees. Our analyses indicate a high diversity of flower rewards and pollinators displayed by members of Maxillariinae, and support that edible trichomes evolved independently five times in the subtribe. CONCLUSIONS: The high diversity of floral rewards and pollinators displayed by members of Maxillariinae suggests that different pollinator pressures are involved in the evolution of this neotropical subtribe. In addition, the offering of food-hairs, which are generally infrequently encountered in Orchidaceae, arose by convergent evolution in Maxillariinae.

Nectar-Secreting and Nectarless Epidendrum: Structure of the Inner Floral Spur.

Epidendrum, the largest genus of Neotropical orchids, contains both nectar-secreting and nectarless species. Here, we compare the fine structure of the inner floral spur, termed the cuniculus, in nectariferous (E. difforme, E. nocturnum,E. porpax, E. rigidum, E. vesicatum) and seemingly nectarless (E. capricornu, E. ciliare, E. criniferum, E. pseudepidendrum, E. radicans, E. xanthoianthinum) species. This is the first time for such a detailed investigation of cuniculus structure to be undertaken for Epidendrum. Our aim was to characterize features indicative of secretory activity and to ascertain whether flowers presumed to be nectarless produce alternative pollinator food-rewards. The cuniculus is formed by fusion of the basal part of the labellum and column and extends alongside the ovary and transmitting tract. Our study indicates that all investigated species produce nectar or nectar-like secretion to varying degrees, and no alternative pollinator food-rewards were observed. Even though macroscopic investigation of presumed rewardless species failed to reveal the presence of secretion within the cuniculus, close observations of the cells lining the cuniculus by LM, SEM, and TEM revealed the presence of cuticular blisters and surface material. Moreover, the similarity of both the thick tangential cell walls (with the exception of E. vesicatum) and organelle complement of cuniculus epidermal cells in both copiously nectariferous species and those producing only small quantities of surface secretion confirmed the presence of secretory activity in species generally regarded to be rewardless. The secretory character was particularly obvious in the cells of the cuniculus of E. nocturnum, but also in E. ciliare, E. radicans and E. xanthoianthinum, since electron-dense cytoplasm and mitochondria, ER and secretory vesicles were abundant. Furthermore, cell wall protuberances occurred in E. nocturnum, which was indicative of intense transmembrane transport. This investigation highlights the need to examine more closely whether Epidendrum spp. considered to lack food-rewards based solely on macroscopic examination really are rewardless and deceptive.

Butterfly pollination in Pteroglossa (Orchidaceae, Orchidoideae): a comparative study on the reproductive biology of two species of a Neotropical genus of Spiranthinae.

Spiranthinae orchids are known for being self-compatible and offering nectar as a reward. Although data on their pollinators are scarce, members of this tribe are mostly pollinated by bees, hummingbirds and moths. Some of them even reproduce through facultative self-pollination. Nothing is known about the pollinators and reproduction system in Pteroglossa. Based on records on flowering phenology, floral morphology, reward production, pollinators and breeding system, this paper aims to study the reproductive biology of two Pteroglossa spp. Both species offer nectar as a resource and are pollinated exclusively by diurnal Lepidoptera at the studied areas. Nectar is produced by two glandular nectaries, and is stored in a spur. Pollinaria possess a ventrally adhesive viscidium that is deposited on the basal portion of butterfly proboscides. Both species are self-compatible but pollinator-dependent. The reproductive success is low when compared to other Spiranthinae. Although no evident mechanical barrier to avoid self-pollination or geitonogamy was identified, the erratic behavior of the butterflies, with their infrequent visits to only one flower per inflorescence, contributes to an increased fruit set produced through cross-pollination. The presence of ventrally adhesive viscidia in Spiranthinae is responsible for greater pollinator diversity when compared to bee-pollinated Goodyerinae with dorsally adhesive viscidia, adapted to attach to bee mouthparts.