The comparison of the molecular analysis with morphological and anatomical data presented here represents an important basis for a new formal classification for the Araceae and for the understanding of the evolution of this ancient family, a monocot group known in the fossil record from the early Cretaceous.
Field studies integrating pollination investigations with an assessment of floral scent composition and thermogenesis in tropical aroids are rather few. Thus, this study aimed to investigate the pollination biology of nine species belonging to Schismatoglottis Calyptrata Complex Clade. The flowering mechanism, visiting insect activities, reproductive system, thermogenesis and floral scent composition were examined. Anthesis for all species started at dawn and lasted 25-29 h. Colocasiomyia (Diptera, Drosophilidae) are considered the main pollinators for all the investigated species. Cycreon (Coleoptera, Hydrophilidae) are considered secondary pollinators as they are only present in seven of the nine host plants, despite the fact that they are the most effective pollen carrier, carrying up to 15 times more pollen grains than Colocasiomyia flies. However, the number of Colocasiomyia individuals was six times higher than Cycreon beetles. Chaloenus (Chrysomelidae, Galeuricinae) appeared to be an inadvertent pollinator. Atheta (Coleoptera, Staphylinidae) is considered a floral visitor in most investigated species of the Calyptrata Complex Clade in Sarawak, but a possible pollinator in S. muluensis. Chironomidae midges and pteromalid wasps are considered visitors in S. calyptrata. Thermogenesis in a biphasic pattern was observed in inflorescences of S. adducta, S. calyptrata, S. giamensis, S. pseudoniahensis and S. roh. The first peak occurred during pistillate anthesis; the second peak during staminate anthesis. Inflorescences of all investigated species of Calyptrata Complex Clade emitted four types of ester compound, with methyl ester-3-methyl-3-butenoic acid as a single major VOC (volatile organic compound). The appendix, pistillate zone, staminate zone and spathe emitted all these compounds. A mixed fly-beetle pollination system is considered an ancestral trait in the Calyptrata Complex Clade, persisting in Sarawak taxa, whereas the marked reduction of interpistillar staminodes in taxa from Peninsular Malaysia and especially, Ambon, Indonesia, is probably linked to a shift in these taxa to a fly-pollinated system.
Pollen characters in Araceae accord well with recent DNA-based phylogenies, and here we provide a new example of “compass needle” quality in Araceae on the basis of two closely related genera, Schismatoglottis and Apoballis. All investigated Schismatoglottis pollen is psilate (smooth pollen surface) with calcium crystals covering the pollen surface. By contrast, pollen of species transferred to recently resurrected Apoballis (Apoballis acuminatissima and A. mutata) is distinctively echinate (spiny). A unique layer covers the endexine of Schismatoglottis, and the whole pollen surface of Apoballis. Our findings strongly suggest that “Schismatoglottis” species with echinate pollen fall into the genus Apoballis. Moreover, all schismatoglottid taxa perform spathe movements during anthesis to control the movement of pollinators. The spathe movements of Apoballis acuminatissima clearly differ from those known in Schismatoglottis species, and indeed are so far unique for the entire family. This, together with differences in floral odour is strongly suggestive of differences in pollination ecology between the genera Schismatoglottis and Apoballis.
Homoplastic evolution of 'unique' morphological characteristics in the Schismatoglottideae -many previously used to define genera -prompted this study to compare morphology and function in connection with pollination biology for Aridarum nicolsonii, Phymatarum borneense and Schottarum sarikeense. Aridarum nicolsonii and P. borneense extrude pollen through a pair of horned thecae while S. sarikeense sheds pollen through a pair of pores on the thecae. Floral traits of spathe constriction, presence and movement of sterile structures on the spadix, the comparable role of horned thecae and thecae pores, the presence of stamen-associated calcium oxalate packages, and the timing of odour emission are discussed in the context of their roles in pollinator management. Pollinators for all investigated species were determined to be species of Colocasiomyia (Diptera: Drosophilidae).
Tribe Schismatoglottideae comprises one large genus, Schismatoglottis, and six small 'satellite' genera. A combined molecular phylogenetic analysis of matK, the 3′ portion of the trnK intron, and trnL‐F sequence data was carried out on 77 taxa representing all genera in the tribe, all informal groups in Schismatoglottis, together with sister tribe Cryptocoryneae, and outgroups from Araceae. Analyses of combined datasets with parsimony, maximum likelihood, and Bayesian methods revealed tribe Schismatoglottideae to be a polyphyletic assemblage. Neotropical Schismatoglottis is shown to be sister to the palaeotropical Schismatoglottideae + Cryptocoryneae. Schismatoglottis acuminatissima is a sister clade to the rest of the Schismatoglottideae. Palaeotropical Schismatoglottis is unsupported as a monophyletic genus. A new neotropical tribe of Araceae, Philonotieae S.Y. Wong & P.C. Boyce, sister to Cryptocoryneae + palaeotropical Schismatoglottideae, is proposed.
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