Homologies of the flower and inflorescence in the early-divergent grass anomochloa (poaceae)

Premise of the study: The grass subfamily Anomochlooideae is phylogenetically significant as the sister group to all other grasses. Thus, comparison of their structure with that of other grasses could provide clues to the evolutionary origin of these characters. Methods: We describe the structure, embryology, and development of the flower and partial inflorescence of the monotypic Brazilian grass Anomochloa marantoidea. We compare these features with those of other early-divergent grasses such as Pharus and Streptochaeta and closely related Poales such as Ecdeiocolea. Key results: Anomochloa possesses several features that are characteristic of Poaceae, notably a scutellum, a solid style, reduced stamen number, and an ovary with a single ovule that develops into a single indehiscent fruit. Interpretation of floral patterning in Anomochloa is problematic because the ramification pattern of the florets places the bracts and axes in unusual positions relative to the primary inflorescence axis. Our study indicates that there is a single abaxial carpel in Anomochloa, probably due to a cryptic type of pseudomonomery in Anomochloa that resembles the pseudomonomery of other grasses. On the other hand, the Anomochloa flower differs from the typical grass flower in lacking lodicules and possessing four stamens, in contrast with the tristaminate condition that characterizes many other grasses. Conclusions: Using the median part of the innermost bract as a locator, we tentatively homologize the inner bract of the Anomochloa partial inflorescence with the palea of other grasses. In this interpretation, the pattern of monosymmetry due to stamen suppression differs from that of Ecdeiocolea. © 2012 Botanical Society of America.

Microsporogenesis is simultaneous in the early-divergent grass Streptochaeta, but successive in the closest grass relative, Ecdeiocolea

Simultaneous microsporogenesis is described for the first time in a grass, Streptochaeta spicata Schrad., a tropical Brazilian species that belongs in the early-divergent subfamily Anomochlooideae. Microsporogenesis is successive in all other Poaceae examined so far, and most other members of the order Poales, to which grasses belong. The only other reports of simultaneous microsporogenesis in Poales are in Rapateaceae and some members of the cyperid clade (Juncaceae, Cyperaceae, Prionium and Thurnia). Among the graminids, Ecdeiocolea (the putative closest relative to Poaceae) is successive, as are Joinvillea, Flagellaria and all other Poaceae, indicating that the simultaneous condition is autapomorphic in Streptochaeta, though Anomochloa has yet to be examined. Anther wall development in Streptochaeta is of the reduced type, as also in another early-divergent grass Pharus, though most other Poales, including most grasses, have the monocot type. In Streptochaeta, as in Pharus, the endothecium lacks thickenings, unlike other grasses that have a persistent endothecium with thickenings. The centrifixed anthers and nonplumose stigmas of Streptochaeta suggest entomophily.

Floral development and embryology in the early-divergent grass Pharus

A detailed study of floral ontogeny, anatomy, and embryology in two (of six) species of Pharus is presented as part of a series of comparative investigations on early-divergent grasses. Pharus is a taxonomically isolated genus belonging to the earliest-diverging grass lineage with a true grass spikelet. It is unusual in possessing remarkably dimorphic florets: male florets possess two lodicules, six stamens, and a pistillode, whereas female florets lack lodicules entirely but possess six staminodes and a tricarpellary ovary with three stigmas. The rudimentary lodicules in male florets are initiated after the stamen whorls. There are most commonly six androecial organs, but in some florets, a five-staminate condition was observed, resulting from suppression of the abaxial stamen from the inner whorl, or even a four-staminate condition resulting from subsequent fusion of the two adaxial outer stamens (i.e., elements of both whorls). Thus, the pattern of floral zygomorphy in Pharus differs from that of many other grasses. Centrifixed anther attachment is reported for the first time in Pharus, resembling the condition in another early-divergent grass, Anomochloa, though anthers are introrse in Anomochloa compared with latrorse in Pharus. Anther wall development is of the reduced type in Pharus, in contrast to most other monocots. Microsporogenesis is of the successive type, as in many other monocots. The ovary develops from three distinct primordia and is unilocular with a single ovule and a pronounced ovary beak that is highly characteristic of Pharus. There is a hollow style, in contrast to the solid styles that are common in many other grasses. The embryo is highly differentiated, as in other grasses, with a distinct epiblast and a small cleft between the scutellum and the coleorhiza.