Evolution of shrub-like growth forms in the lianoid subfamily Secamonoideae (Apocynaceae s.l.) of Madagascar: phylogeny, biomechanics, and development.

Lianas are common in the Apocynaceae s.l. and are predominant in the subfamily Secamonoideae. Shrub-like taxa are rare within this subfamily but occur in Malagasy genera such as Secamone, Secamonopsis, and Pervillaea. We explored the evolutionary appearance of shrub-like growth forms in Malagasy Secamonoideae through a molecular phylogeny using chloroplastic sequences. The phylogeny revealed several independent appearances of shrub-like growth forms within the Secamonoideae. Biomechanics and development of the shrub-like growth form were detailed in one species, Secamone sparsiflora, which has upright and self-supporting young stems that become procumbent in older stages of development. Biomechanical investigations revealed characteristics atypical of both lianas and self-supporting shrubs. Anatomical development in S. sparsiflora is initially similar to lianas in the same clade but shows potentially neotenic retention of juvenile wood development for most of the growth trajectory. The results suggest that evolution of lianescence can carry a high degree of specialization and developmental burden that might limit evolution back to self-supporting growth forms. Under certain geographic and ecological conditions, such as geographic isolation, xeric conditions and/or reduced biotic competition, escapes from lianescence to other growth forms can occur in some angiosperm groups via relatively simple heterochronic shifts of mechanically significant growth processes.