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Revised Classification of the Onagraceae

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Revised Classification of the Onagraceae Chylismia brevipes subsp. brevipes [Illustration by Alice Tangerini, based on Beatley 7918 (US) and colored in Adobe PhotoShop CS using images from CalPhoto (J. Andre, image 01040887; C. Cloud-Hughes, images 07040686 and 07040690)]. SYSTEMATIC BOTANY MONOGRAPHS VOLUME 83 Revised Classification of the Onagraceae Warren L. Wagner Peter C. Hoch Peter H. Raven THE AMERICAN SOCIETY OF PLANT TAXONOMISTS 17 September 2007 SYSTEMATIC BOTANY MONOGRAPHS ISSN 0737-8211 Copyright © 2007 The American Society of Plant Taxonomists All rights reserved ISBN 0-912861-83-5 ISBN 978-0-912861-83-8 Printed in the United States of America Editor CHRISTIANE ANDERSON University of Michigan Herbarium 3600 Varsity Drive Ann Arbor, Michigan 48108-2287 Editorial Committee BRUCE G. BALDWIN JOHN V. F REUDENSTEIN University of California, Berkeley The Ohio State University LYNN BOHS DAVID M. JOHNSON University of Utah Ohio Wesleyan University DAVID E. BOUFFORD DONALD H. LES Harvard University University of Connecticut GREGORY K. BROWN THOMAS A. RANKER University of Wyoming University of Colorado REVISED CLASSIFICATION OF THE ONAGRACEAE Warren L. Wagner Department of Botany, MRC 166 Smithsonian Institution, P. O. Box 37012 Washington, DC 20013-7012 Peter C. Hoch Peter H. Raven Missouri Botanical Garden P. O. Box 299 St. Louis, Missouri 63166-0299 ABSTRACT. Recent molecular phylogenetic analyses in the plant family Onagraceae support the need for revisions in the family classification. In this paper we briefly survey the history of generic and suprageneric clas- sification in Onagraceae, summarize our knowledge of the morphological and molecular variation in the family in a phylogenetic context, and propose a revised classification that reflects that phylogeny. We include a species- level synopsis of the family, incorporating all nomenclatural changes and combinations but not full species-level synonymy. We provide descriptions of all taxa recognized for the first time, as well as tribes, genera, sections, subsections, and series. In this treatment, we recognize 22 genera in the family, subdivided into two subfamilies, subfam. Ludwigioideae (only Ludwigia) and subfam. Onagroideae (the other genera), and the latter into six tribes, two with only one genus each, three with two genera each, and one (tribe Onagreae) with 13 genera. Chro- mosome numbers and breeding system information are summarized, as is the geographical and ecological dis- tribution of each taxon. For each group we list the included taxa. Many changes involve the tribe Onagreae, from which we have segregated Gongylocarpus as its own tribe, sister to the tribes Epilobieae and Onagreae, and within which we propose changes in the delimitation of Camissonia and Oenothera. Camissonia as currently de- fined is broadly paraphyletic; our new classification recognizes nine generic lineages (Camissonia, Camisso- niopsis, Chylismia, Chylismiella, Eremothera, Eulobus, Holmgrenia, Taraxia, and Tetrapteron), which in part form a grade at the base of Oenothera. Each of these lineages is well-supported by morphological and molecu- lar data. In contrast, molecular and morphological data both suggest the need to broaden the delimitation of Oenothera to include Calylophus, Gaura, and Stenosiphon. This redefined Oenothera, strongly supported by molecular data, is marked by at least two morphological synapomorphies: the presence of an indusium on the style, and a lobed or peltate stigma. We summarize these new generic alignments and review the morphological characters used to diagnose all tribes, genera, and sections. New taxa recognized include subfamilies Ludwigioideae and Onagroideae, two genera (Camissoniopsis and Holmgrenia), three sections (Epilobium sect. Macrocarpa, Oenothera sect. Leucocoryne, and O. sect. Xan- thocoryne), and one subspecies (O. macrocarpa subsp. mexicana). All other nomenclatural novelties involve new combinations or new names; these include three genera (Chylismiella, Eremothera, and Tetrapteron), 23 sections or subsections (Chylismia sect. Lignothera; Clarkia sect. Phaeostoma subsect. Connubium, C. sect. Phaeostoma subsect. Heterogaura; Epilobium sect. Epilobiopsis; Ludwigia sect. Isnardia; Oenothera sect. Anogra, O. sect. Calylophus, O. sect. Calylophus subsect. Calylophus, O. sect. Calylophus subsect. Salpingia, O. sect. Gaura, O. sect. Gaura subsect. Campogaura, O. sect. Gaura subsect. Gaura, O. sect. Gaura subsect. Gauridium, O. sect. Gaura subsect. Schizocarya, O. sect. Gaura subsect. Stenosiphon, O. sect. Gaura subsect. Stipogaura, O. sect. Gaura subsect. Xenogaura, O. sect. Gaura subsect. Xerogaura, O. sect. Hartmannia, O. sect. Lavauxia, O. sect. Lavauxia subsect. Lavauxia, O. sect. Megapterium, and O. sect. Oenothera subsect. Can- dela), and 97 species or subspecies (Camissoniopsis bistorta, C. cheiranthifolia, C. cheiranthifolia subsp. suf- fruticosa, C. confusa, C. guadalupensis, C. guadalupensis subsp. clementiana, C. hardhamiae, C. hirtella, C. ig- nota, C. intermedia, C. lewisii, C. luciae, C. micrantha, C. pallida, C. pallida subsp. hallii, C. proavita, C. robusta; Chylismia atwoodii, C. brevipes subsp. arizonica, C. brevipes subsp. pallidula, C. cardiophylla subsp. cedrosensis, C. cardiophylla subsp. robusta, C. claviformis subsp. aurantiaca, C. claviformis subsp. cruciformis, C. claviformis subsp. funerea, C. claviformis subsp. integrior, C. claviformis subsp. lancifolia, C. claviformis subsp. peeblesii, C. claviformis subsp. peirsonii, C. claviformis subsp. rubescens, C. claviformis subsp. wiggin- sii, C. claviformis subsp. yumae, C. confertiflora, C. eastwoodiae, C. exilis, C. megalantha, C. munzii, C. scapoidea subsp. brachycarpa, C. scapoidea subsp. macrocarpa, C. scapoidea subsp. utahensis, C. specicola, C. specicola subsp. hesperia, C. walkeri subsp. tortilis; Chylismiella pterosperma; Epilobium campestre, 1 2 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 83 E. subdentatum; Eremothera boothii, E. boothii subsp. alyssoides, E. boothii subsp. condensata, E. boothii subsp. decorticans, E. boothii subsp. desertorum, E. boothii subsp. intermedia, E. chamaenerioides, E. gouldii, E. minor, E. nevadensis, E. pygmaea, E. refracta; Eulobus angelorum, E. crassifolius, E. sceptrostigma; Holm- grenia andina, H. hilgardii; Ludwigia speciosa; Oenothera arida, O. berlandieri subsp. pinifolia, O. boquillen- sis, O. calcicola, O. cinerea, O. cinerea subsp. parksii, O. coloradensis, O. coloradensis subsp. neomexicana, O. curtiflora, O. demareei, O. filiformis, O. filipes, O. gaura, O. glaucifolia, O. hartwegii subsp. fendleri, O. hartwegii subsp. filifolia, O. hartwegii subsp. maccartii, O. hartwegii subsp. pubescens, O. hexandra, O. hex- andra subsp. gracilis, O. lindheimeri, O. mckelveyae, O. patriciae, O. simulans, O. sinuosa, O. suffrutescens, O. suffulta, O. suffulta subsp. nealleyi, O. triangulata, O. tubicula subsp. strigulosus, O. xenogaura; Tetrapteron graciliflorum, and T. palmeri). INTRODUCTION Recent publication of phylogenetic analyses of the Onagraceae, particularly several in-depth studies using nucleotide sequence analysis (Levin et al. 2003, 2004), demonstrate the need for changes in the classification of the family. Raven (1979, 1988) provided the most recent summaries of the classification of Onagraceae, but those papers appeared be- fore most of the molecular analyses were available. In this study we briefly review the his- tory of the classification of Onagraceae and summarize the available data that underlie our new classification. We do not include a formal re-analysis of the extensive morphological data available for the family, which is beyond the scope of this paper, but we refer to these data throughout in reference to the taxa we have delimited. We recognize two subfamilies, Ludwigioideae (Ludwigia) and Onagroideae. The lat- ter comprises six tribes: Hauyeae (Hauya), Circaeeae (Circaea, Fuchsia), Lopezieae (Lopezia, Megacorax), Gongylocarpeae (Gongylocarpus), Epilobieae (Chamerion, Epilobium), and Onagreae (Camissonia, Camissoniopsis, Chylismia, Chylismiella, Clarkia, Eremothera, Eulobus, Gayophytum, Holmgrenia, Oenothera, Taraxia, Tetra- pteron, Xylonagra). We include four appendices. The first provides the accepted new names and new combinations at the specific and infra-specific level proposed in this paper along with the type information. The second provides new synonymy for names no longer accepted in Oenothera. The third lists taxa of Onagraceae that meet the criteria of IUCN for species of conservation concern, including rare species and those at risk of extinction. The fourth appendix is a list of taxa that are known to occur outside of their native ranges, with in- dication of those with greatest invasive potential. TAXONOMIC HISTORY The following account traces major developments and innovations in the history of Onagraceae classification, and is not intended to be comprehensive for all treatments. There is some emphasis on treatments that encompass western North America, for the simple reason that many genera in the family are restricted to that region or have their major distribution there; these also are the taxa whose taxonomic assignments have changed most dramatically over time. Table 1 summarizes some of the most influential treatments of the family, in comparison to the one presented in this work. Linnaeus (1753) described eight genera that eventually would be included in Ona- graceae; in our current classification, we still recognize five of these—Circaea L., Epilo- bium L., Fuchsia L., Ludwigia L. (now including Isnardia L. and Jussiaea L.), and 2007 ONAGRACEAE 3
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