Jacques Fournel

Independent recruitment of saprotrophic fungi as mycorrhizal partners by tropical achlorophyllous orchids

Mycoheterotrophic orchids have adapted to shaded forest understory by shifting to achlorophylly and receiving carbon from their mycorrhizal fungi. In temperate forests, they associate in a highly specific way with fungi forming ectomycorrhizas on nearby trees, and exploiting tree photosynthates. However, many rainforests lack ectomycorrhizal fungi, and there is evidence that some tropical Asiatic species associate with saprotrophic fungi. To investigate this in different geographic and phylogenetic contexts, we identified the mycorrhizal fungi supporting two tropical mycoheterotrophic orchids from Mascarene (Indian Ocean) and Caribbean islands. We tested their possible carbon sources by measuring natural nitrogen ((15)N) and carbon ((13)C) abundances. Saprotrophic basidiomycetes were found: Gastrodia similis associates with a wood-decaying Resinicium (Hymenochaetales); Wullschlaegelia aphylla associates with both litter-decaying Gymnopus and Mycena species, whose rhizomorphs link orchid roots to leaf litter. The (15)N and (13)C abundances make plausible food chains from dead wood to G. similis and from dead leaves to W. aphylla. We propose that temperature and moisture in rainforests, but not in most temperate forests, may favour sufficient saprotrophic activity to support development of mycoheterotrophs. By enlarging the spectrum of mycorrhizal fungi and the level of specificity in mycoheterotrophic orchids, this study provides new insights on orchid and mycorrhizal biology in the tropics.

Sapromyiophily in the native orchid, Bulbophyllum variegatum , on Réunion (Mascarene Archipelago, Indian Ocean)

Orchid species are well known for their highly specialized pollinator interactions. To better understand the reproductive biology of the tropical epiphytic orchid Bulbophyllum variegatum on Reunion, we investigated the floral morphology, breeding system, pollinator diversity, floral scent profile and fruiting success of about 30 individuals in three natural populations during two consecutive flowering seasons. Controlled hand-pollination experiments in two populations showed that the species is self-compatible, but requires pollinator service to achieve reproduction. Videotape pollinator observations were conducted during two flowering seasons for 56 h and revealed that B. variegatum is pollinated by a single species of fly from the Platystomatidae. This fly seems to be attracted by the unpleasant scent produced by the flowers, and does not receive any reward after achieving pollination. In addition, no egg-laying behaviour was observed. Bulbophyllum variegatum thus exhibits a typical sapromyiophilous pollination syndrome which constitutes the first proven case of sapromyiophily within the genus Bulbophyllum on the Mascarene Archipelago. Hand pollinations further showed that fruit set was not significantly higher for flowers that received outcross pollen than for those that were self-crossed (53% and 44% respectively). Fruit sets under natural conditions were significantly different among populations, ranging from 0.5% to 24.3%. This low fruit production is likely due to infrequent pollinator visits, particularly in disturbed forests where the pollinator has never been observed.

Orchid-bird interactions: a case study from Angraecum (Vandeae, Angraecinae) and Zosterops (white-eyes, Zosteropidae) on Reunion Island

Although angraecoid orchids are famous for their spectacular flowers, pollinated by hawkmoths in Madagascar, two species of Angraecum endemic to the Mascarenes (sect. Hadrangis ) have atypical unscented and short-spurred flowers that are pollinated by birds. We focused on the pollination and reproductive success of Angraecum bracteosum Balf.f. & S.Moore, a member of sect. Hadrangis . We investigated the breeding system by performing a set of hand-pollination ex- periments and by observing natural pollinators using video-captures. Pollinator efficiency (pollen removal and deposition) and reproductive success (fruit set) were quantified once a week in two natural island populations during three consecutive flowering seasons (2003–2005). The orchid is self-compatible but requires pollinator services to achieve reproduction. The two white-eye song-bird species (Zosteropidae) that occur on Reunion, Zosterops borbonicus Boddaert and Zosterops oliva- Ceus L., were recorded as pollinators. However, while pollination events were rare and occasional following flower visits by Z. borbonicus , foraging of Z. oliveaceus resulted in a high frequency of successful pollinations. White-eyes typically perched on inflorescences, and probed most of the ‘‘fresh-looking’’ flowers for nectar. This study documents a probable case of bird-pollination specialization within Angraecum section Hadrangis , and sheds light on the role played by pollina- tors during the radiation of section Hadrangis in the Mascarene Archipelago.

Mouthparts and nectar feeding of the flower visiting cricket Glomeremus orchidophilus (Gryllacrididae).

Glomeremus orchidophilus (Gryllacrididae) is a flower visiting cricket on the tropical island La Réunion. This species is the only Orthoptera shown to be a pollinator of a plant. We studied its nectar feeding behavior and mouthpart morphology in detail. Since G. orchidophilus possesses biting-and-chewing mouthparts, our objective was to find behavioral and/or structural specializations for nectar-feeding. The comparative analysis of feeding behavior revealed that fluid is taken up without movements of the mouthparts in Glomeremus. A comparative morphological examination of two Glomeremus species, together with several representatives of other Gryllacrididae and other Ensifera taxa revealed subtle adaptations to fluid feeding in Glomeremus. All representatives of Gryllacrididae were found to possess a distinct patch of microtrichia at the tip of their galeae. However, in Glomeremus a channel is formed between the distal components of the maxillae and the mandibles on each side of the body. Micro-CT and SEM examination revealed a longitudinal groove that extends over the galea beginning at the patch of microtrichia in the studied Glomeremus species. We hypothesize that the microtrichia take up fluid by capillarity and the action of the cibarium and pharyngeal pumps transports fluid along the channels between the maxillae and mandibles into the preoral cavity. These mouthpart features allow nectar uptake from flowers that is unique in Orthoptera.