The Relationship Between Anisosperma and Fevillea (Cucurbitaceae), and a New Species of Fevillea from Bolivia
Total Page：16
File Type：pdf, Size：1020Kb
Systematic Botany (2009), 34(4): pp. 704–708 © Copyright 2009 by the American Society of Plant Taxonomists The Relationship Between Anisosperma and Fevillea (Cucurbitaceae), and a New Species of Fevillea from Bolivia Michael Nee, 1 , 4 Hanno Schaefer, 2 and Susanne S. Renner 3 1 Institute of Systematic Botany, New York Botanical Garden, Bronx, New York 10458 U.S.A. 2 Ecology and Evolutionary Biology, Imperial College London, Silwood Park Campus, Ascot, SL5 7PY, U.K. 3 Systematic Botany, University of Munich, Menzinger Strasse 67, D-80638 Munich, Germany 4 Author for correspondence ( [email protected] ) Communicating Editor: Andrea Schwarzbach Abstract— We describe and illustrate the new species Fevillea anomalosperma from the semideciduous forests of northwestern Bolivia. Analysis of chloroplast DNA sequences representing most genera of Cucurbitaceae, including Fevillea with four of its seven species, supports placing the new species in Fevillea despite its small (10–11.5 × 7.5–10 mm, 2–3 mm thick), flattened, and highly sculptured seeds, which are unusual in the genus. The molecular phylogeny also revealed that the recent sinking of the monotypic genus Anisosperma into Fevillea is not justified. Resumen— Se ilustra y describe Fevillea anomalsperma , una nueva especie de los boques semi-deciduos del noroeste de Bolivia. Aunque las hojas y las flores son típicas de Fevillea , las semillas pequeñas (10–11.5 x 7.5–10 mm, 2–3 mm de grueso), aplanadas y muy esculpidas de esta nueva especie no lo son. La posición de esta nueva especie dentro de Fevillea ha sido apoyado por análisis filogenéticos de secuencias de ADN del cloroplasto, en los cuales se ha incluido representantes de la mayoría de los géneros de Cucurbitaceae, junto con quatro especies de Fevillea . Filogenia molecular demuestra que la inclusión del género monotípico Anisosperma en Fevillea no es justificada. Keywords— Anisosperma , dry forests of Bolivia , Fevilleeae , molecular phylogenetics , oil-rich fruits. The cucurbit genus Fevillea, named after the French clergy- DNA extraction kit (NucleoSpin, Macherey-Nagel, Düren, Germany), man and explorer L. É. Feuillée (1660–1732), comprises tall following the manufacturer’s instructions. Thermal reaction products were purified with a PCR clean-up kit (PCR Wizard, Promega GmbH, lianas of moist or wet forests from southern Mexico and the Mannheim, Germany), and cycle sequencing was performed with the Caribbean to northern Argentina. A recent revision ( Robinson BigDye Terminator cycle sequencing kit on an ABI Prism 3100 Avant auto- and Wunderlin 2005b ) recognized seven species in two sub- mated sequencer (Applied Biosystems, Foster City, California). Twelve genera, one with six species, the other containing Fevillea passi- sequences were newly generated for this study, and their Genbank acces- flora Vell., the former Anisosperma passiflora (Vell.) Silva Manso. sion numbers and vouchers are listed in Appendix 1. Accession numbers for all other sequences used for the analyses are given in Kocyan et al. The oil-rich seeds of some Fevillea species can weigh three to (2007 ) and Schaefer et al. (2009 ). nine g when dry ( Robinson and Wunderlin 2005b ) and are Phylogenetic analyses were conducted with a 147-taxon subset of the among the largest seeds in the Cucurbitaceae, exceeded only family dataset of Schaefer et al. (2009 ), which includes all but one of the by those of the Old World genera Telfairia Hook. and Hodgsonia genera of Cucurbitaceae and 25% of the 960 species. Besides two accessions of the new species, we included four species of Fevillea , including the type Hook. f. & Thomson. Indigenous South Americans apparently species, and six species representing the other three genera of Fevilleeae. have used the oil of F. cordifolia L. and F. trilobata L., species As outgroups, we used 15 species of Anisophylleaceae, Begoniaceae, locally known as nhandiroba, for burning and as medicine Coriariaceae, Corynocarpaceae, Datiscaceae, and Tetramelaceae based on for centuries ( Gentry and Wettach 1986 ; www.cnpf.embrapa. Zhang et al. (2006 ). The only genus of Cucurbitaceae that has not yet been br/publica/folders/Nhandiroba_2004.pdf ). The same is true sequenced is Khmeriosicyos W. J. de Wilde & Duyfjes, a monotypic genus from Cambodia, known only from the type. Judging from floral morphol- of Anisosperma passiflora ( Fevillea passiflora ), known in Brazil ogy, it is not closely related to Fevillea . as castanha-mineira, andiroba, castanha de bugre, or castanha Sequence Alignment and Phylogenetic Analyses— Sequences were de jatoba ( Silva Manso 1836 ; Martius 1843 , 1854 ). edited with Sequencher (4.6; Gene Codes, Ann Arbor, Michigan) and A collection made in 1987 in northwestern Bolivia in the aligned by eye. The aligned plastid matrix comprises 4,973 nucleotides after exclusion of a TATATA microsatellite region in the trnL-F intergenic Departamento La Paz, Provincia Nor Yungas, and others spacer. The data matrix and trees have been deposited in TreeBASE study made in 2005 and 2006 in the adjacent Provincia Tamayo, have number S2210 in connection with Schaefer et al. (2009 ). brought to light a new species that is described here. The new Maximum likelihood (ML) analyses and bootstrap searches were per- entity appeared to be a Fevillea , except for its unusual fruit formed using RAxML-VI-HPC v7.0.4 ( Stamatakis et al. 2008 , available at and seeds. We therefore tested the phylogenetic affinities of http://phylobench.vital-it.ch/raxml-bb/ ). RAxML searches relied on the GTR + G + I model. Model parameters were estimated over the duration the new species with a chloroplast data set that included two of specified runs. accessions of the new species, four other species of Fevillea (including the type species, F. trilobata L.), representatives of all genera of Fevilleeae, and other species of Cucurbitaceae Results from all but one of the genera of the family. Morphologically, the new species belongs to the tribe Fevilleeae Benth. & Hook. f., which apart from Fevillea Materials and Methods comprises Pteropepon (Cogn.) Cogn. (probably to include Morphology— The first author studied all relevant specimens in the Pseudosicydium Harms) and Sicydium Schltdl. (including herbaria MO, NY, and LPB and the other authors specimens in BM, K, and Chalema Dieterle) from the New World, and Cyclantheropsis M. Measurements are from herbarium specimens. Harms, which has two species in East and South tropi- Sequencing and Taxon Sampling— DNA extraction and sequencing cal Africa and one endemic in Madagascar (Schaefer and followed standard procedures, using the rbcL , matK , rpl20-rps12 , and trnL and trnL-F primers and polymerase chain reaction (PCR) protocols Renner 2009). These other genera all have single-seeded listed in Kocyan et al. (2007 ). Total genomic DNA was isolated from sil- fruits, and based on chloroplast DNA, are not close to the ica-dried leaves or from herbarium specimens with a commercial plant new species ( Fig. 1 ). 704 2009] NEE ET AL.: A NEW FEVILLEA FROM BOLIVIA 705 Fig. 1. Condensed version of a maximum likelihood phylogram for 147 taxa (only 24 shown), including representatives of all but one genus of Cucurbitaceae ( Schaefer et al. 2009 ; this study). Likelihood bootstrap values >60% are shown at the nodes. The two accessions of Fevillea anomalosperma M. Nee are nested within the Fevillea clade, while the accession of Anisosperma ( Fevillea ) passiflora is not. The tendrils of Fevillea anomalosperma are simple, and 2005b ). Another species in subgenus Fevillea , F. cordifolia , has hence the characteristic of tendrils spiraling both above flowers of exactly the same form and dull reddish-brown and below the point of branching, typical of many species color as those of F. anomalosperma , but differs as indicated in of Fevilleeae, is not applicable. The presence of five free, the key. bilocular anthers in Fevillea (and the new species) is unique Three species of Fevillea overlap the range of F. anoma- for the family ( Robinson and Wunderlin 2005b ), but with- losperma in Bolivia (Nee unpubl. data): Fevillea cordifolia , F. per- out pistillate flowers, we cannot assess whether the new spe- gamentacea (Kuntze) Cogn., and F. pedatifolia (Cogn.) C. Jeffrey. cies has the three free styles characterizing early-diverging All three have subglobose fruits 6–13 cm in diameter and Cucurbitaceae. The apparently capsular fruits of F. anoma- 6–16 cm long with lenticular seeds 3.5–6 cm wide and losperma are similar to those of Siolmatra Baill. ( Robinson and 1–2.5 cm thick ( Robinson and Wunderlin 2005b ), different Wunderlin 2005a ), a genus of two species that have fruits from the fruits of F. anomalosperma. The distinct testa structure, with three to six flattened, winged seeds and that occur in however, seems to be similar in all four species. The fruits of southern Amazonian forests as well as the dry forest to the F. pedatifolia (as F. peruviana (Huber) C. Jeffrey) were described south. Molecular data support Siolmatra as being sister to as dry capsules ( Jeffrey 1962 ) based on herbarium material of the monotypic Asian Zanonia L., with both then sister to a Krukoff 8466 , and in Robinson and Wunderlin (2005b ) there is clade of African and Madagascan genera ( Kocyan et al. 2007 ; a personal communication from A. Gentry indicating that the Schaefer et al. 2009 ). fruits may have circumscissile dehiscence around the hypan- Robinson and Wunderlin (2005b ) divided Fevillea into the thium lip scar. In the carpological collection at Kew, only typical subgenus Fevillea , with six species, and a monotypic fruits of F. cordifolia show these openings, but not the collec- subgenus Anisosperma (Silva Manso) G. Rob. & Wunderlin. tions of F. pedatifolia . The fruits of F. passiflora might open by Our new species most closely resembles species in subgenus longitudinal splitting of the fruit walls, but this is more likely Fevillea . The leaves and male inflorescences of F. anomalosperma an artifact of drying. The fruit of F. anomalosperma appears to are almost identical to those of F. trilobata from eastern Brazil.