The seeds of Loasaceae subfam. Loasoideae (Cornales) II: Seed morphology of “South Andean Loasas” (Loasa, Caiophora, Scyphanthus and Blumenbachia)

South Andean Loasas (Blumenbachia, Caiophora, Loasa, Scyphanthus) are a monophyletic group of taxa within Loasaceae subfam. Loasoideae, comprising some 100 species, 49 of which are investigated here. They retain a many-layered testa in the mature seeds and usually have passive transfer testas with complex, spongiose wall outgrowths. Additional modifications concern the undulations of the testa epidermis, presence or absence of the outer periclinal wall, secondary sculpturing, the presence or absence of spines, warts and finally spongiose structures on the anticlinal walls of the testa epidermis and the inner periclinal wall. Seeds of the widespread “deeply pitted” type are plesiomorphic, while various subclades of South Andean Loasas have derivations underscoring their relationships and confirming the relationships found with molecular markers and other morphological characters. The genus Blumenbachia has either seeds with a many-layered testa forming longitudinal lamellae (sect. Angulatae), or balloon seeds with a loose outer testa layer and spongiose wall outgrowths on the inner periclinal walls (sect. Blumenbachia and sect. Gripidea) and is clearly monophyletic. Loasa s.str. (ser. Loasa, ser. Macrospermae, ser. Floribundae, ser. Deserticolae) is characterized by the presence of a subterminal hilum or hilar scar and one subgroup (ser. Loasa, ser. Macrospermae) by very large and heavy seeds with a collapsed testa. L. ser. Pinnatae, ser. Acaules, ser. Volubiles, Scyphanthus and Caiophora share more or less one seed types with minor modifications. Within Caiophora various derivations are observed, of which the gradual loss of the secondary sculpture of the inner periclinal wall is the most striking one. Anemochoria is the most widespread dispersal mechanism in South Andean Loasas and is achieved in at least five structurally different ways.