Trichome micromorphology in Alcea L. and allied genera (Malvaceae) and its systematic implication

Trichomes of 26 species of Alcea (Malvaceae) were investigated using light (LM) and scanning electron microscopy (SEM). The trichomes show a great micromorphological variation, which provides data useful for species delimitation in Alcea. Two basic types of trichomes could be distinguished in Alcea and allied genera: glandular and eglandular. The glandular trichomes could in turn be subdivided into two subtypes: capitate and clavate. The eglandular trichomes could be subdivided into five subtypes: simple, fascicled, stellate, fascicled-stellate and pluri-radiate. Characters of taxonomic interest are: trichome density (glabrous to dense), number of arms per trichome, orientation relative to the epidermal surface (appressed to erect) and presence/absence of a stalk. According to the results the species of Alcea can be divided into four informal groups based on trichome types. The results further support the exclusion of the annual Althaea species from the perennial ones and their close relationship to Malva. In addition, a close relationship between perennial Althaea and basal Alcea lineages is supported. Based on the evolutionary framework provided by recent molecular phylogenetic investigations, the following trends can be observed in the Malva alliance: long and narrow-armed trichomes are primitive in relation to short and thick-armed trichomes; dense indumentum coverage is primitive in relation to moderately dense or glabrous ones; presence of simple hairs on stem (particularly on leaves) is more advanced than their absence; spreading villous-stellate and fascicled trichomes are more advanced than appressed stellate ones. Clavate trichomes, which were found exclusively in a few species of Alcea, should be considered as a derived state in relation to capitate ones, and they may provide a synapomorphy for the crown group of Alcea.