Colocasiomyia, a moderate‐sized genus in the subfamily Drosophilinae, comprises seventy (twenty‐six described and forty‐four undescribed) species. Several Colocasiomyia species have evolved intimate mutualisms with specific host plants, especially of the family Araceae: the flies depend throughout the entire life cycle, oviposition, larval growth, pupation, and adult feeding and mating, on inflorescences of their host plants, and in turn act as species‐specific pollinators for their host plants. To understand the evolution of this mutualism between Colocasiomyia flies and their host plants, the phylogenetic relationships of this genus and some possibly related taxa are inferred from a cladistic analysis based on sixty‐two characters of adult morphology. We conclude that Colocasiomyia is polyphyletic, with the C. arenga species group clearly separate. Colocasiomyia without the arenga group (Colocasiomyia proper) is sister to all other studied drosophilines, whereas the arenga group is relatively derived within the Drosophilinae. Within Colocasiomyia proper, four clades are recognized, three of which correspond to previously proposed species groups: the cristata, toshiokai and baechlii groups. The other clade, C. sp.1 aff. nepalensis+C. sp.2 aff. nepalensis, is defined as a new species group. Relationships amongst the four clades and three independent species (C. micheliae, C. gigantea and C. sp.K1) remain almost unresolved, except for a sister group relationship between the toshiokai and baechlii groups. The classification of species groups in Colocasiomyia is revised by erecting two new species groups (crassipes and zeylanica groups) in addition to the three known (baechlii, cristata and toshiokai) groups. Revision of the arenga group, which should be removed from Colocasiomyia, is left for future studies. The evolution of host plant selection in Colocasiomyia is discussed by mapping host plant taxa (families, subfamilies and tribes) on the phylogenetic tree deduced from the cladistic analysis. Cohabitation in the same host inflorescence by a pair of species with microallopatric niche separation on the spadix is hypothesized to have evolved independently at least more than twice in Colocasiomyia.
We evaluated the role of floral scents in the reproductive success of Alocasia odora C. Koch (Araceae). Alocasia odora is pollinated by its specific pollinators, Colocasiomyia alocasiae (Okada) and C. xenalocasiae (Okada) (Diptera: Drosophilidae). These flies use the spadix of A. odora as breeding sites. The appendix, which is at an upper part of the spadix and is the most attractive region, attracted these pollinators by emitting volatiles, although the male zone of the inflorescence was also attractive. The number of flies attracted was positively correlated with appendix size. During the pistillate phase of the protogynous spadix, attracted flies aggregated in the lower part (female zone) to mate, lay eggs, and perhaps obtain nutrients. The flies moved to the upper part (male zone) of the spadix by the tightening of the constriction separating the upper and lower parts, and then the staminate phase started. This movement of the flies on the spadix promotes outcrossing of A. odora. Removal of the appendix or the whole upper part of the spadix resulted in much reduced fruit set, suggesting that the absence of the scent-producing region leads to insufficient pollination because of reduced pollinator attraction.
Abstract.
Two closely related species of Colocasiomyia alocasiae (Okada) and C.xenalocasiae (Okada) (Diptera, Drosophilidae) breed in inflorescences of Alocasia odora C. Koch (Araceae), a hermaphroditic understorey clonal herb.
The two drosophilid species form a synhospitalic pair in Okinawa with alocasiae breeding in the upper half of the inflorescence and xenalocasiae breeding in the lower half.
C.alocasiae also has the following combination of life history traits: small body size, many eggs, and early reproductive maturity. In contrast, xenalocasiae can be described as having larger body size, fewer eggs, and delayed reproductive maturity.
Resource partitioning between the two species on the same host may be affected by these life history traits which are associated with their larval habitats.
Two aroid congeners, Alocasia odora and Alocasia cucullata , grow on Okinawa Island, Japan. Two floricolous species of Colocasiomyia alocasiae and Colocasiomyia xenalocasiae (Diptera: Drosophilidae), previously known to be specific pollinators for A. odora , were found pollinating A. cucullata . We collected the floral volatiles of A. odora and A. cucullata and compared them using gas chromatography and mass spectrometry because floral volatiles act as attractive signals for these pollinators. The volatile compositions detected were similar and dominated by methyl salicylate, 4,8-dimethyl-1,3,7-nonatriene, b b b b -caryophyllene, bicyclogermacrene, methyl benzoate and a a a a -humulene, which means it is likely that the flies cannot discriminate between the two plant species when they search for hosts. Interspecific hybridization did not occur when A. odora was hand pollinated with A. cucullata pollen, suggesting that the movement of flies from one host species to another will result in wastage of pollen. Adults of both Colocasiomyia flies emerged from A. cucullata inflorescences collected in the field, suggesting that their larvae can also develop on A. cucullata . We did not find any difference in larval performance of C. alocasiae on A. cucullata or A. odora . C. alocasiae does not appear to suffer any ill effects when using A. cucullata as a host plant. The partnership between A. cucullata and the two Colocasiomyia flies may be an example of exaptation, although other possibilities cannot be excluded.
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