Embryology of Manekia naranjoana (Piperaceae) and its Implications for the Origin of the Sixteen-nucleate Female Gametophyte in Piperales

Piperaceae is unique among Piperales because it is the only tetrasporic group in the order and a great deal of diversity in the ontogenetic trajectories of the female gametophyte is found in its genera. The evolutionary developmental origin of the sixteen-nucleate female gametophyte remains unclear in the family until now. In Piperaceae, Manekia has been identified as sister to Zippelia, and this clade is sister to core Piperaceae (Piper, Peperomia). This research is the first attempt to understand the development of the female gametophyte of Manekia naranjoana in order to provide critical data on the origin of tetrasporic development in the family. Several aspects of the floral biology and phenological events taking place in the ovary, the flower and the inflorescence were explored. Manekia has a tetrasporic, sixteen nuclei female gametophyte, that is being produced from a single archesporial cell. The egg apparatus is located at the micropylar end of the female gametophyte. It is constituted of three cells, two synergids and an egg. The central cell nuclei consist of two nuclei, one from the micropylar end and the other one from the chalazal one. The eleven remaining nuclei are arranged toward the chalazal pole of the female gametophyte, and sometimes fuse. This description corresponds mostly to the Drusa type. But Penaea type is also occasionally reported for first time in this study for the genus. Manekia and Zippelia share a similar structure of the female gametophyte with a total of 16 nuclei, and two nuclei in a central cell suggesting a triploid endosperm. The transition from monosporic to tetraporic female gametophyte development can be explained through the theory of modular construction and several kind modifications in the ontogenetic trajectories. Heterochronic and heterotopic changes, additions, and deletions in the development of the female gametophytes reflect evolutionary histories of the particular taxa implicated. A great deal of plasticity in terms of lack of polarity and nuclear fusion of antipodals was found in the chalazal module of the female gametophyte of Manekia.