phylogeny of cucurbitales inferred from nine chloroplast

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Phylogeny of Cucurbitales inferred from nine chloroplast, mitochondrial and nuclear loci LI-BING ZHANG and SUSANNE S. RENNER. Department of Biology, University of Missouri-St. Louis, St. Louis, MO 63121. Email: [email protected]; [email protected] Based on rbcL (Setoguchi et al. 1999; Schwarzbach & Ricklefs 2000), Cucurbitales comprise Anisophylleaceae, Begoniaceae, Coriariaceae, Corynocarpaceae, Cucurbitaceae, Datiscaceae, and Tetramelaceae. To further investigate the position of Anisophylleaceae and to approp- riately root a large molecular phylogeny of Cucurbitaceae, we are investigating family relationships in Cucurbitales. Previous molecular data have identified Fagales as the sister clade to Cucurbitales, but prior to our project, Anisophylleaceae had been weakly sampled and only sequenced for rbcL, which provided limited support for the monophyly of Cucurbitales. We have combined nuclear 18S, the chloroplast genes atpB, matK, ndhF, and rbcL, the trnL intron, the rpl20-rps12 and trnL-trnF spacers, and the mitochondrial nad1 b/c intron (together 9302 aligned bp) for 25 representatives of the ingroup families and two fagalean outgroups. Cucurbitales are strongly monophyletic, and Anisophylleaceae are sister to the remaining families, which form a trichotomy of (1) Cucurbitaceae, (2) Coriariaceae and Corynocarpaceae, and (3) Begoniaceae (Datiscaceae, Tetramelaceae). The relationship between the last three families has also been addressed with restriction site and ITS data (Rieseberg et al., 1992; Swensen et al., 1998),as well as flower morphology, which indicated a close relationship of Datiscaceae and Tetramelaceae. With the exception of a few Coriariaceae and Corynocarpaceae, Cucurbitales all have unisexual flowers, and plants are usually monoecious, with repeated shifts to dioecy in Begonia, Cucurbitaceae, Datisca cannabina, and Tetramelaceae. Because molecular data do not yet resolve family relationships in the Cucurbitales, it is unclear whether the few perfect- flowered Coriariaceae / Corynocarpaceae represent basal lines or reversals from unisexual to perfect flowers. The latter is exceedingly rare, and we are currently exploring additional DNA markers specifically to resolve their position in the order. Anisophylleaceae Photo: G. E. Schatz Anisophyllea fallax Datiscaceae Photo: P. Endress Datisca cannabina Tetramelaceae Octomeles sumatrana Photo: W. Takeuchi Coriariaceae Coriaria myrtifolia Photo: G. Gerlach & P. Endress Corynocarpaceae Photo: G. C. Linney Corynocarpus laevigatus Cucurbitaceae Photo: P. Endress Marah fabaceus Begoniaceae Begonia evansii Photo: B. Glick 96 Juglans nigra Fagus grandifolia Anisophyllea corneri Anisophyllea obtusifolia Anisophyllea fallax Combretocarpus rotundatus Begonia herbacea Begonia oxyloba Hillebrandia sandwicensis Datisca glomerata Datisca cannabina Octomeles sumatrana Tetrameles nudiflora Coriaria ruscifolia Coriaria sarmentosa Corynocarpus laevigatus Coccinia sessilifolia Lagenaria brevifolia Dendrosicyos socotranus Gurania megistantha Seyrigia humbertii Ecballium elaterium Marah macrocarpus Schizopepon bryoniifolius Gynostemma pentaphyllum Neoalsomitra sarcophylla Xerosicyos danguyi 50 changes 94 95 100 100 100 100 100 100 100 100 100 100 100 88 98 100 Anisophylleaceae Begoniaceae Datiscaceae Tetramelaceae Coriariaceae Corynocarpaceae Cucurbitaceae 100 (Single most parsimonious tree, CI = 0.72 RI =0.79 bootstraps at branches from 100 replicates)

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Phylogeny of Cucurbitales inferred from nine chloroplast, mitochondrial and nuclear loci

LI-BING ZHANG and SUSANNE S. RENNER. Department of Biology, University of Missouri-St. Louis, St. Louis, MO 63121. Email: [email protected]; [email protected]

Based on rbcL (Setoguchi et al. 1999; Schwarzbach & Ricklefs 2000), Cucurbitales comprise Anisophylleaceae, Begoniaceae, Coriariaceae, Corynocarpaceae, Cucurbitaceae, Datiscaceae, and Tetramelaceae. To further investigate the position of Anisophylleaceae and to approp- riately root a large molecular phylogeny of Cucurbitaceae, we are investigating family relationships in Cucurbitales. Previous molecular data have identified Fagales as the sister clade to Cucurbitales, but prior to our project, Anisophylleaceae had been weakly sampled and only sequenced for rbcL, which provided limited support for the monophyly of Cucurbitales. We have combined nuclear 18S, the chloroplast genes atpB, matK, ndhF, and rbcL, the trnL intron, the rpl20-rps12 and trnL-trnF spacers, and the mitochondrial nad1 b/c intron (together 9302 aligned bp) for 25 representatives of the ingroup families and two fagalean outgroups. Cucurbitales are strongly monophyletic, and Anisophylleaceae are sister to the remaining families, which form a trichotomy of (1) Cucurbitaceae, (2) Coriariaceae and Corynocarpaceae, and (3) Begoniaceae (Datiscaceae, Tetramelaceae). The relationship between the last three families has also been addressed with restriction site and ITS data (Rieseberg et al., 1992; Swensen et al., 1998),as well as flower morphology, which indicated a close relationship of Datiscaceae and Tetramelaceae. With the exception of a few Coriariaceae and Corynocarpaceae, Cucurbitales all have unisexual flowers, and plants are usually monoecious, with repeated shifts to dioecy in Begonia, Cucurbitaceae, Datisca cannabina, and Tetramelaceae. Because molecular data do not yet resolve family relationships in the Cucurbitales, it is unclear whether the few perfect- flowered Coriariaceae / Corynocarpaceae represent basal lines or reversals from unisexual to perfect flowers. The latter is exceedingly rare, and we are currently exploring additional DNA markers specifically to resolve their position in the order.

Anisophylleaceae

Photo: G. E. Schatz

Anisophyllea fallax�

Datiscaceae

Photo: P. Endress

Datisca cannabina

Tetramelaceae

Octomeles sumatrana

Photo: W. TakeuchiCoriariaceae

Coriaria myrtifolia

Photo: G. Gerlach & P. Endress

Corynocarpaceae

Photo: G. C. Linney

Corynocarpus laevigatus

Cucurbitaceae

Photo: P. Endress�

Marah fabaceus

Begoniaceae

Begonia evansii

Photo: B. Glick

96

Juglans nigra

Fagus grandifolia

Anisophyllea corneri

Anisophyllea obtusifolia

Anisophyllea fallax

Combretocarpus rotundatus

Begonia herbacea

Begonia oxyloba

Hillebrandia sandwicensis

Datisca glomerata

Datisca cannabina

Octomeles sumatrana

Tetrameles nudiflora

Coriaria ruscifolia

Coriaria sarmentosa

Corynocarpus laevigatus

Coccinia sessilifolia

Lagenaria brevifolia

Dendrosicyos socotranus

Gurania megistantha

Seyrigia humbertii

Ecballium elaterium

Marah macrocarpus

Schizopepon bryoniifolius

Gynostemma pentaphyllum

Neoalsomitra sarcophylla

Xerosicyos danguyi

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Anisophylleaceae

Begoniaceae

Datiscaceae

Tetramelaceae

Coriariaceae

Corynocarpaceae

Cucurbitaceae

100

(Single most parsimonious tree, CI = 0.72 RI =0.79 bootstraps at branches from 100 replicates)�