Abstract
A new infrafamilial circumscription of the Verbenaceae with eight tribes: Casselieae, Citharexyleae, Duranteae, Lantaneae, Neospartoneae, Petreeae, Priveae, and Verbeneae, has been recently proposed, on the basis of molecular phylogenetic studies. Two genera, Dipyrena and Rhaphithamnus, remain unplaced. The aim of this work is to reconstruct the evolutionary history of morphological characters traditionally employed in the classification of the Verbenaceae, with special attention to tribes Verbeneae and Lantaneae. Twenty-one characters, related to habit and vegetative morphology, inflorescence and floral morphology, ovary and fruit morphology, as well as chromosome number, were optimized over a molecular phylogeny of Verbenaceae. All tribes are supported by at least one morphological trait except tribes Duranteae and Citharexyleae. Suffrutescent habit, sessile flowers, and four cluses are synapomorphies for tribe Verbeneae. Gynoecium with short style and entire stigma are synapomorphic traits for tribe Lantaneae. Sessile flowers and unicarpellate ovaries are morphological synapomorphies for the new tribe Neospartoneae. Suffrutescent habit is a synapomorphic trait for tribe Priveae. Homothetic pleiobotrya and absence of the adaxial staminode are synapomorphic traits for tribe Casselieae. Undivided fleshy fruits are probably a synapomorphic trait for tribe Petreeae. Putative plesiomorphies for the ancestor of the Verbenaceae are discussed as well as synapomorphic traits within other Verbenaceae clades. Many of the characters traditionally employed in classification have proven to be very homoplastic, or have been shown not to support relationships within the family. Moreover, traditional assumptions concerning character polarity have in some cases been shown to be incorrect.
Highlights
Twenty-one characters, related to habit and vegetative morphology, inflorescence and floral morphology, ovary and fruit morphology, as well as chromosome number, were optimized over a molecular phylogeny of Verbenaceae
Suffrutescent habit is a synapomorphic trait for tribe Priveae
The major objectives of this study are to identify morphological characters that support phylogenetic relationships within Verbenaceae as inferred on the basis of molecular data (Marx et al, 2010; Yuan et al, 2010; O’Leary et al, 2009; Lu-Irving and Olmstead, in press), including, where possible, those clades recognized as tribes, and to reconstruct the evolutionary history of those morphological characters traditionally used in the systematics of the family
Summary
Methods: Twenty-one characters, related to habit and vegetative morphology, inflorescence and floral morphology, ovary and fruit morphology, as well as chromosome number, were optimized over a molecular phylogeny of Verbenaceae. Verbenaceae, as recently circumscribed, comprises approximately 32 genera and 840 species (Cantino et al, 1992; Atkins, 2004; Marx et al, 2010). Recent molecular phylogenetic studies (Marx et al, 2010; Yuan et al, 2010) identified the major lineages in the family and resolved generic relationships throughout most of the group. This resulted in a revised classification for Verbenaceae, dividing the family into eight tribes (Marx et al, 2010).
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