Molecular Phylogenetics and Evolution 56 (2010) 768–783

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Molecular Phylogenetics and Evolution

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Phylogenetics and character evolution in the carnivorous A. St.-Hil. ()

Andreas Fleischmann a,*, Bastian Schäferhoff b,d, Günther Heubl a, Fernando Rivadavia c, Wilhelm Barthlott d, Kai F. Müller b,d a Department of Biology, Systematic Botany and Mycology, Ludwig-Maximilians-Universität München, Menzinger Straße 67, D-80638 Munich, Germany b Institute for Evolution and Biodiversity, Westfälische Wilhelms-Universität, Hüfferstraße 1, D-48149 Münster, Germany c Daniel Burnham Ct., San Francisco, CA 94109, USA d Nees-Institut für Biodiversität der Pflanzen, Friedrich-Wilhelms Universität Bonn, Meckenheimer Allee 170, D-53115 Bonn, Germany article info abstract

Article history: The genus Genlisea A. St.-Hil. (Lentibulariaceae) comprises at least 22 distrib- Received 19 December 2009 uted in South and Central America as well as in Africa (including Madagascar). It has only recently been Revised 1 March 2010 shown to be a true carnivore, specialized in and other small soil organisms. Here we present a Accepted 7 March 2010 statistically highly supported phylogeny of Genlisea based on three chloroplast loci. The most recent com- Available online 11 March 2010 mon ancestor of Genlisea most likely was of Neotropical origin and characterized by pedicels that are recurved in fruit, a strongly glandular inflorescence, and bivalvate capsule dehiscence. The further evolu- Keywords: tion of various morphological characters during the diversification of the genus is discussed. The two pre- Carnivorous viously suggested subgenera Tayloria and Genlisea correspond to the two major clades found in our Genlisea Lentibulariaceae analyses. In subgenus Genlisea, three clades can be clearly distinguished based on molecular and morpho- trnK/matK logical characters and on biogeographic patterns, which led us to propose a new sectional classification. rps16 Ó 2010 Elsevier Inc. All rights reserved. trnQ–rps16 Phylogeny

1. Introduction gions exhibit what is among the highest DNA substitutional rates in angiosperm chloroplasts (Müller et al., 2004). These attributes The genus Genlisea A. St.-Hil. belongs to the carnivorous plant make Genlisea a perfect model system for studying factors govern- family Lentibulariaceae (, I, APG III, 2009) and ing substitutional rate and size shifts. The evolution of comprises at least 22 species (a few more undiagnosed taxa await these ultrasmall is focus of ongoing research (Greilhuber their formal description (Fleischmann et al., in preparation), and et al., in preparation). unlike preceding treatments recognizing 21 species (Fischer et al., 2000; Taylor, 1991b), the authors do not consider G. subvir- idis Hutch. conspecific with G. africana Oliv.). Genlisea is distributed 1.1. Carnivory in Genlisea in South and Central America and in Africa (including Madagascar), with centers of highest species diversity in south-eastern , All species of Genlisea lack which have been replaced by rhi- the Guiana Highlands and the Flora Zambesia area (Fig. 2). zophylls (Fig. 1A–C) that are derived of and epiascidiate in Recently, the genus Genlisea has attended increased