Embryo sac, endosperm, and seed of Nemophila (Boraginaceae) relative to taxonomy, with a remark on embryogeny in Pholistoma

Studies on embryology and seed morphology are complementary to molecular phylogenetics and of special value at the genus level. This paper discusses the delimitation and evolutionary relationships of genera within the tribe Hydrophylleae of the Boraginaceae. The seven Nemophila species characterized by a conspicuous seed appendage are similar in embryology and seed structure. The ovule is tenuinucellate and unitegmic with a meristematic tapetum. The embryo sac penetrating the nucellar apex is of the Polygonum type, has short-lived antipodal cells, and an embryo sac haustorium. The endosperm is cellular, producing two terminal endosperm haustoria, of which the chalazal has a lateral branch. Embryogeny is of the Chenopodiad type (as in Pholistoma). The seed coat is formed from the small-celled inner epidermis of the integument. The large-celled outer epidermis of the integument disintegrates into scattered cells. Seed pits evolve from irregularly placed inner epidermal cells of the integument. The chalazal part of the ovule produces a cucullus, that functions as an ant-attracting elaiosome. Those species of Nemophila with a conspicuous cucullus form a natural genus. Nemophila is most closely related to Pholistoma. The integumentary seed pits of Nemophila might have evolved from ovular seed pits similar to those in Pholistoma.