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Growth form evolution in piperales and its relevance for understanding angiosperm diversification: An integrative approach combining plant architecture, anatomy, and biomechanics

Isnard S., Prospéri M.J., Wanke S., Wagner S.T., Samain M.C., Trueba S., Frenzke L., Neinhuis C., Rowe N.P.. 2012. International Journal of Plant Sciences, 173 (6) : p. 610-639.

DOI: 10.1086/665821

A striking feature of early angiosperm lineages is the variety of life forms and growth forms, which ranges from herbs, aquatic herbs, climbers, and epiphytes to woody shrubs and trees. This morphological and anatomical diversity is arguably one of the factors explaining how angiosperms dominate many ecosystems worldwide. However, just how such a wide spectrum of growth forms has evolved in angiosperms remains unclear. In this review, we investigate patterns of growth form diversification in Piperales, an early-diverging lineage (with stem age estimated at 201-128 Myr ago) and the most morphologically diverse clade among magnoliids. We outline patterns of growth form diversity and architecture as well as the biomechanical significance of developmental characters, such the organization, loss, and gain of woodiness. Asaroideae and Saururaceae are terrestrial as well as semiaquatic to aquatic herbaceous perennials bearing rhizomes. The Aristolochioideae and Piperaceae show higher levels of growth form diversity and biomechanical organization, with complex patterns of increasing or decreasing woodiness and architectural organization. The climbing habit has probably evolved independently in the Aristolochiaceae and Piperaceae, while mechanically unstable shrubs and, less frequently, treelets have evolved several times within these two most species-rich clades. A key developmental character underlying diversity in most Piperales-with the exception of the herbaceous Saruma (Asaroideae)-is the conserved development of the wood cylinder, in which fusiform initials are limited to fascicular cambial initials. The resulting large fraction of raylike tissue in the stem-a highly characteristic feature of woody species in the Piperales-potentially introduced mechanical constraints on the diversification of self-supporting architectures. This was possibly circumvented by the architectural development of repeated, large-diameter meristems in some shrublike habits via sympodial growt

Mots-clés : piperales; évolution; biodiversité; anatomie végétale; port de la plante; développement biologique; croissance; propriété mécanique

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