title: Phylogenetics, ancestral state reconstruction, and a new infrafamilial classification of the pantropical Ochnaceae (Medusagynaceae, Ochnaceae s.str., Quiinaceae) based on five DNA regions.
abstract: Ochnaceae s.str. (Malpighiales) are a pantropical family of about 500 species and 27 genera of almost exclusively woody plants. Infrafamilial classification and relationships have been controversial partially due to the lack of a robust phylogenetic framework. Including all genera except Indosinia and Perissocarpa and DNA sequence data for five DNA regions (ITS, matK, ndhF, rbcL, trnL-F), we provide for the first time a nearly complete molecular phylogenetic analysis of Ochnaceae s.l. resolving most of the phylogenetic backbone of the family. Based on this, we present a new classification of Ochnaceae s.l., with Medusagynoideae and Quiinoideae included as subfamilies and the former subfamilies Ochnoideae and Sauvagesioideae recognized at the rank of tribe. Our data support a monophyletic Ochneae, but Sauvagesieae in the traditional circumscription is paraphyletic because Testulea emerges as sister to the rest of Ochnoideae, and the next clade shows Luxemburgia+Philacra as sister group to the remaining Ochnoideae. To avoid paraphyly, we classify Luxemburgieae and Testuleeae as new tribes. The African genus Lophira, which has switched between subfamilies (here tribes) in past classifications, emerges as sister to all other Ochneae. Thus, endosperm-free seeds and ovules with partly to completely united integuments (resulting in an apparently single integument) are characters that unite all members of that tribe. The relationships within its largest clade, Ochnineae (former Ochneae), are poorly resolved, but former Ochninae (Brackenridgea, Ochna) are polyphyletic. Within Sauvagesieae, the genus Sauvagesia in its broad circumscription is polyphyletic as Sauvagesia serrata is sister to a clade of Adenarake, Sauvagesia spp., and three other genera. Within Quiinoideae, in contrast to former phylogenetic hypotheses, Lacunaria and Touroulia form a clade that is sister to Quiina. Bayesian ancestral state reconstructions showed that zygomorphic flowers with adaptations to buzz-pollination (poricidal anthers), a syncarpous gynoecium (a near-apocarpous gynoecium evolved independently in Quiinoideae and Ochninae), numerous ovules, septicidal capsules, and winged seeds with endosperm are the ancestral condition in Ochnoideae. Although in some lineages poricidal anthers were lost secondarily, the evolution of poricidal superstructures secured the maintenance of buzz-pollination in some of these genera, indicating a strong selective pressure on keeping that specialized pollination system.
authors.affiliation: Senckenberg Research Institute and Natural History Museum Frankfurt, Department of Botany and Molecular Evolution, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany; Goethe-University, Department of Ecology, Evolution and Diversity, Max-von-Laue-Str. 13, D-60439 Frankfurt am Main, Germany; Biodiversity and Climate Research Centre (BiK-F), Senckenberganlage 25, D-60325 Frankfurt am Main, Germany. Electronic address: Julio.Schneider@senckenberg.de.
authors.forename: Julio V
authors.initials: JV
authors.lastname: Schneider
authors.affiliation: Naturalis Biodiversity Center (Section NHN), Darwinweg 2, 2333CR Leiden, The Netherlands; Wageningen University, Biosystematics Group, Droevendaalsesteeg 1, NL-6708 PB Wageningen, The Netherlands; Herbier National du Gabon (IPHAMETRA-CENAREST), B.P. 1156 Libreville, Gabon.
authors.forename: Pulcherie
authors.initials: P
authors.lastname: Bissiengou
authors.affiliation: Department of Plant Biology, Institute of Biology, CP 6109, University of Campinas - UNICAMP, 13083-970 Campinas, SP, Brazil.
authors.forename: Maria do Carmo E
authors.initials: Mdo C
authors.lastname: Amaral
authors.affiliation: Senckenberg Research Institute and Natural History Museum Frankfurt, Department of Botany and Molecular Evolution, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany; Goethe-University, Department of Ecology, Evolution and Diversity, Max-von-Laue-Str. 13, D-60439 Frankfurt am Main, Germany; Biodiversity and Climate Research Centre (BiK-F), Senckenberganlage 25, D-60325 Frankfurt am Main, Germany.
authors.forename: Ali
authors.initials: A
authors.lastname: Tahir
authors.affiliation: Jodrell Laboratory, Royal Botanic Gardens Kew, Richmond, Surrey TW9 3DS, United Kingdom.
authors.forename: Michael F
authors.initials: MF
authors.lastname: Fay
authors.affiliation: Senckenberg Research Institute and Natural History Museum Frankfurt, Department of Botany and Molecular Evolution, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany; Goethe-University, Department of Ecology, Evolution and Diversity, Max-von-Laue-Str. 13, D-60439 Frankfurt am Main, Germany; Biodiversity and Climate Research Centre (BiK-F), Senckenberganlage 25, D-60325 Frankfurt am Main, Germany.
authors.forename: Marco
authors.initials: M
authors.lastname: Thines
authors.affiliation: Naturalis Biodiversity Center (Section NHN), Darwinweg 2, 2333CR Leiden, The Netherlands; Wageningen University, Biosystematics Group, Droevendaalsesteeg 1, NL-6708 PB Wageningen, The Netherlands; Botanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium.
authors.forename: Marc S M
authors.initials: MS
authors.lastname: Sosef
authors.affiliation: Senckenberg Research Institute and Natural History Museum Frankfurt, Department of Botany and Molecular Evolution, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany; Goethe-University, Department of Ecology, Evolution and Diversity, Max-von-Laue-Str. 13, D-60439 Frankfurt am Main, Germany; Biodiversity and Climate Research Centre (BiK-F), Senckenberganlage 25, D-60325 Frankfurt am Main, Germany.
authors.forename: Georg
authors.initials: G
authors.lastname: Zizka
authors.affiliation: Wageningen University, Biosystematics Group, Droevendaalsesteeg 1, NL-6708 PB Wageningen, The Netherlands.
authors.forename: Lars W
authors.initials: LW
authors.lastname: Chatrou
mesh_terms: D001499:Bayes Theorem; D018744:DNA, Plant / Q000737:chemistry; D043842:Ochnaceae / Q000145:classification* / Q000235:genetics; D010802:Phylogeny; D017422:Sequence Analysis, DNA
chemical_list: D018744:DNA, Plant
keywords: Ancestral state reconstruction; Bayesian hypothesis testing; Classification; ITS; Malpighiales; Plastid markers