Flora of North America

Contributors to Volume 6

Kristin PierceCompositor and Editorial Assistant

Heidi H. SchmidtProduction Coordinator and

Managing Editor

María Mercedes ArboKerry A. Barringer

David M. BatesOrland J. Blanchard Jr.

Luc BrouilletGarrett E. CrowLaurence J. Dorr

Aaron FlodenPaul A. Fryxell†

Susannah B. FultonJohn F. Gaskin

Aaron Goldberg†

Douglas H. GoldmanMargaret M. Hanes

Steven R. HillWalter C. HolmesBrian R. KeenerJohn La Duke

David E. LemkeR. John Little

John M. MacDougalLandon E. McKinney†

T. Lawrence MellichampMeghan G. Mendenhall

Guy L. NesomLorin I. Nevling Jr.

C. Thomas PhilbrickNorman K. B. Robson

Janice G. SaundersBruce A. SorrieJohn L. StrotherJohn W. Thieret†Gordon C. TuckerLinda E. WatsonMolly A. Whalen

George Yatskievych

Editors for Volume 6

David E. BouffordTaxon Editor for Apodanthaceae

and Thymelaeaceae

Kanchi GandhiNomenclatural Editor

Ronald L. HartmanCo-Taxon Editor for Violaceae

Robert W. KigerBibliographic Editor, Lead

Editor, and Taxon Editor for Calophyllaceae, Cistaceae, Clusiaceae, Cucurbitaceae, Hypericaceae, Malvaceae,

Muntingiaceae, Passifloraceae, and Tamaricaceae

Nancy R. MorinTaxon Editor for Droseraceae

Jackie M. PooleTaxon Editor for Begoniaceae,

Cochlospermaceae, Datiscaceae, Frankeniaceae, and Turneraceae

Richard K. RabelerCo-Taxon Editor for Violaceae

Heidi H. SchmidtManaging Editor

Mary Ann SchmidtTechnical Editor

Leila M. ShultzTaxon Editor for Podostemaceae

John L. StrotherReviewing Editor

James L. ZarucchiEditorial Director

Volume Composition

NEW YORK OXFORD OXFORD UNIVERSITY PRESS 2015

Flora of North AmericaNorth of Mexico

FLORA OF NORTH AMERICA EDITORIAL COMMITTEEEdited by

VOLUME 6

Magnoliophyta: Cucurbitaceae to Droseraceae

· ·

Oxford University Press is a department of the University of Oxford.It furthers the University’s objective of excellence in research,

scholarship, and education by publishing worldwide.

Oxford New York

Auckland Cape Town Dar es Salaam Hong Kong Karachi

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With offices in

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Oxford is a registered trademark of Oxford University Press in the UK and certain other countries.

Published by Oxford University Press, Inc.198 Madison Avenue, New York, New York 10016

www.oup.com

Copyright © 2015 by Flora of North America AssociationFrontispiece art Dionaea muscipula by Linny Heagy © 3/07/2014 all rights reserved

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior written permission of the Flora of North America Association, or as expressly

permitted by law, by license, or under terms agreed with the appropriate reproduction rights organization. Inquiries concerning reproduction outside the scope of the above should be sent to the Flora of North America Association.

You must not circulate this work in any other form and you must impose this same condition on any acquirer.

Library of Congress Cataloging-in-Publication Data(Revised for Volume 6)

Flora of North America North of Mexicoedited by Flora of North America Editorial Committee.

Includes bibliographical references and indexes.Contents: v. 1. Introduction—v. 2. Pteridophytes and gymnosperms—

v. 3. Magnoliophyta: Magnoliidae and Hamamelidae—v. 22. Magnoliophyta: Alismatidae, Arecidae, Commelinidae (in part), and Zingiberidae—

v. 26. Magnoliophyta: Liliidae: Liliales and Orchidales—v. 23. Magnoliophyta: Commelinidae (in part): Cyperaceae—

v. 25. Magnoliophyta: Commelinidae (in part): Poaceae, part 2—v. 4. Magnoliophyta: Caryophyllidae (in part): part 1—v. 5. Magnoliophyta: Caryophyllidae (in part): part 2—

v. 19, 20, 21. Magnoliophyta: Asteridae (in part): Asteraceae, parts 1–3—v. 24. Magnoliophyta: Commelinidae (in part): Poaceae, part 1—

v. 27. Bryophyta, part 1—v. 8. Magnoliophyta: Paeoniaceae to Ericaceae—

v. 7. Magnoliophyta: Salicaceae to Brassicaceae—v. 28. Bryophyta, part 2—

v. 9. Magnoliophyta: Picramniaceae to Rosaceae—v. 6. Magnoliophyta: Cucurbitaceae to Droseraceae

ISBN: 978-0-19-534027-3 (v. 6)1. Botany—North America.

2. Botany—United States.3. Botany—Canada.

I. Flora of North America Editorial Committee.QK110.F55 2002 581.97 92-30459

1 2 3 4 5 6 7 8 9Printed in the United States of America

on acid-free paper

Contents

Dedication vii

Founding Member Institutions viii

Donors ix

Art Sponsorship x

Editorial Committee xi

Project Staff xii

Contributors xiii

Taxonomic Reviewers xv

Regional Reviewers xv

Preface xviii

Introduction xix

Cucurbitaceae 3

Datiscaceae 59

Begoniaceae 61

Calophyllaceae 64

Clusiaceae 67

Podostemaceae 69

Hypericaceae 71

Violaceae 106

Turneraceae 165

Passifloraceae 170

Apodanthaceae 183

Muntingiaceae 185

Malvaceae 187

Cochlospermaceae 376

Thymelaeaceae 380

Cistaceae 386

Frankeniaceae 409

Tamaricaceae 413

Droseraceae 418

Literature Cited 426

Index 453

This volume of the

Flora of North America North of Mexico

is dedicated to the memory of

Paul Arnold Fryxell

1927 – 2011

Courtesy of Hunt Institute for Botanical Documentation, Carnegie Mellon University, Pittsburgh, PA Photograph by Walter Henricks Hodge.

FOUNDING MEMBER INSTITUTIONS

Flora of North America Association

Agriculture and Agri-Food CanadaOttawa, Ontario

Arnold ArboretumJamaica Plain, Massachusetts

Canadian Museum of Nature Ottawa, Ontario

Carnegie Museum of Natural History

Pittsburgh, Pennsylvania

Field Museum of Natural HistoryChicago, Illinois

Fish and Wildlife ServiceUnited States Department of

the InteriorWashington, D.C.

Harvard University Herbaria Cambridge, Massachusetts

Hunt Institute for Botanical Documentation

Carnegie Mellon UniversityPittsburgh, Pennsylvania

Jacksonville State UniversityJacksonville, Alabama

Jardin Botanique de MontréalMontréal, Québec

Kansas State UniversityManhattan, Kansas

Missouri Botanical GardenSt. Louis, Missouri

New Mexico State UniversityLas Cruces, New Mexico

The New York Botanical GardenBronx, New York

New York State MuseumAlbany, New York

Northern Kentucky UniversityHighland Heights, Kentucky

Université de MontréalMontréal, Québec

University of AlaskaFairbanks, Alaska

University of AlbertaEdmonton, Alberta

The University of British ColumbiaVancouver, British Columbia

University of CaliforniaBerkeley, California

University of CaliforniaDavis, California

University of IdahoMoscow, Idaho

University of Illinois Urbana-Champaign, Illinois

University of IowaIowa City, Iowa

The University of KansasLawrence, Kansas

University of MichiganAnn Arbor, Michigan

University of OklahomaNorman, Oklahoma

University of OttawaOttawa, Ontario

University of Southwestern Louisiana

Lafayette, Louisiana

The University of TexasAustin, Texas

University of Western OntarioLondon, Ontario

University of WyomingLaramie, Wyoming

Utah State UniversityLogan, Utah

viii

ix

For their support of the preparation of this volume,

we gratefully acknowledge and thank:

Franklinia Foundation

The Philecology Foundation

The Andrew W. Mellon Foundation

The David and Lucile Packard Foundation

an anonymous foundation

Chanticleer Foundation

The Stanley Smith Horticultural Trust

Chris Davidson and Sharon Christoph,

Botanical Research Foundation of Idaho

For sponsorship of illustrations included in this volume,

we express sincere appreciation to:

Erick Adams, Olympia, Washington Drosera intermedia, Droseraceae

David Bates, Ithaca, New York Malacothamnus palmeri, Malvaceae

Jennifer Bryant, Hobart, Oklahoma Callirhoë involucrata, Malvaceae

Dorothy King Young Chapter, California Native Plant Society, Gualala, California Viola adunca, Violaceae

Friends of University of North Carolina, Charlotte, Botanical Gardens, Charlotte, North Carolina Volume 6 Frontispiece, Dionaea muscipula, Droseraceae – in honor of T. Lawrence Mellichamp

Arthur V. Gilman, Marshfield, Vermont Viola rostrata, Violaceae

Douglas Goldman, Greensboro, North Carolina Passiflora tenuiloba, Passifloraceae – in memory of Hobbes Goldman

Nancy Morin, Point Arena, California Hypericum ascyron subsp. pyramidatum, Hypericaceae – in honor of Norman K. B. Robson Fremontodendron californicum, Malvaceae – in appreciation of the California Native Plant Society Melochia corchorifolia, Malvaceae – in memory of Aaron Goldberg

The Polly Hill Arboretum, West Tisbury, Massachusetts Dirca palustris, Thymelaeaceae

Jackie Poole, Austin, Texas Fryxellia pygmaea, Malvaceae – in memory of Paul Fryxell

Barbara and David Rice, Gualala, California Viola glabella, Violaceae

Jennifer H. Richards, Miami, Florida Tilia americana, Malvaceae Passiflora incarnata, Passifloraceae Piriqueta cistoides subsp. caroliniana, Turneraceae

Linda Watson, Stillwater, Oklahoma Amoreuxia palmatifida, Cochlospermaceae

x

FLORA OF NORTH AMERICA EDITORIAL COMMITTEE

VASCULAR PLANTS

Kerry A. BarringerTaxon EditorJersey City, New Jersey

David E. BouffordTaxon Editor and Regional

Coordinator,Northeastern United StatesHarvard University HerbariaCambridge, Massachusetts

Luc BrouilletTaxon Editor and Regional

Coordinator, Eastern Canada and GreenlandInstitut de recherche en biologie

végétaleUniversité de MontréalMontréal, Québec

Wayne J. ElisensUniversity of OklahomaNorman, Oklahoma

Bruce A. FordTaxon Editor and Regional

Coordinator,Western CanadaUniversity of ManitobaWinnipeg, Manitoba

Craig C. FreemanTaxon Editor and Regional

Coordinator,North Central United States R. L. McGregor HerbariumThe University of KansasLawrence, Kansas

Kanchi Gandhi Nomenclatural EditorHarvard University HerbariaCambridge, Massachusetts

David E. GiblinTaxon EditorUniversity of WashingtonSeattle, Washington

Lynn GillespieTaxon EditorCanadian Museum of NatureOttawa, Ontario

Ronald L. HartmanTaxon Editor and Regional

Coordinator, Rocky MountainsUniversity of WyomingLaramie, Wyoming

Steffi M. Ickert-BondRegional Coordinator, Alaska and YukonUniversity of Alaska,

Museum of the NorthFairbanks, Alaska

Robert W. Kiger Bibliographic and Taxon EditorHunt Institute for Botanical

DocumentationCarnegie Mellon UniversityPittsburgh, Pennsylvania

Thomas G. LammersTaxon EditorUniversity of Wisconsin

OshkoshOshkosh, Wisconsin

Geoffrey A. LevinTaxon EditorIllinois Natural History SurveyChampaign, Illinois

Barney L. LipscombBotanical Research Institute of

TexasFort Worth, Texas

Timothy K. LowreyTaxon EditorUniversity of New MexicoAlbuquerque, New Mexico

James A. MacklinAgriculture and Agri-Food

CanadaOttawa, Ontario

Gerry MooreLiason to Governmental AgenciesUnited States Department of

AgricultureGreensboro, North Carolina

Nancy R. MorinTaxon Editor and Regional

Coordinator, Southwestern United StatesPoint Arena, California

Robert F. C. NacziThe New York Botanical GardenBronx, New York

Jackie M. PooleTaxon Editor and Regional

Coordinator,South Central United StatesUniversity of Texas at AustinAustin, Texas

Richard K. RabelerTaxon EditorUniversity of MichiganAnn Arbor, Michigan

Yury RoskovIllinois Natural History SurveyChampaign, Illinois

Heidi H. SchmidtManaging EditorMissouri Botanical GardenSt. Louis, Missouri

Leila M. Shultz Taxon EditorUtah State UniversityLogan, Utah

John L. StrotherReviewing and Taxon EditorUniversity of CaliforniaBerkeley, California

Janet R. SullivanTaxon EditorUniversity of New HampshireDurham, New Hampshire

Debra K. TrockTaxon EditorCalifornia Academy of

SciencesSan Francisco, California

Emeritus/a Members of the Editorial Committee

xi

Gordon C. TuckerTaxon EditorEastern Illinois UniversityCharleston, Illinois

Michael A. VincentTaxon EditorMiami UniversityOxford, Ohio

Alan S. WeakleyTaxon Editor and Regional

Coordinator,Southeastern United StatesUniversity of North CarolinaChapel Hill, North Carolina

James L. ZarucchiEditorial Director Missouri Botanical GardenSt. Louis, Missouri

BRYOPHYTES

Terry T. McIntoshTaxon EditorUniversity of British ColumbiaVancouver, British Columbia

Dale H. VittTaxon EditorSouthern Illinois UniversityCarbondale, Illinois

David H. WagnerTaxon EditorNorthwest Botanical InstituteEugene, Oregon

Alan T. WhittemoreTaxon EditorUnited States National

ArboretumWashington, D.C.

Richard H. Zander Bryophyte Coordinator and

Taxon EditorMissouri Botanical GardenSt. Louis, Missouri

(as of 13 October 2014)

George W. Argus Merrickville, Ontario

Mary E. BarkworthLogan, Utah

Marshall R. CrosbySt. Louis, Missouri

Claudio Delgadillo M.Mexico City, Mexico

Marie L. HicksMoab, Utah

Marshall C. JohnstonAustin, Texas

Aaron I. ListonCorvallis, Oregon

John McNeillEdinburgh, Scotland

Barbara M. MurrayFairbanks, Alaska

David F. MurrayFairbanks, Alaska

J. Scott PetersonBaton Rouge, Louisiana

Alan R. SmithBerkeley, California

Richard W. Spellenberg Las Cruces, New Mexico

Lloyd R. StarkLas Vegas, Nevada

Barbara M. ThiersBronx, New York

Rahmona A. Thompson Ada, Oklahoma

Frederick H. Utech Pittsburgh, Pennsylvania

Project Staff — past and present

involved with the preparation of Volume 6

Barbara Alongi, IllustratorBurgund Bassüner, Information CoordinatorAriel S. Buback, Editorial AssistantTrisha K. Distler, GIS AnalystPat Harris, Editorial AssistantLinny Heagy, Illustrator Ruth T. King, Editorial AssistantJohn Myers, Illustrator and Illustration CompositorKristin Pierce, Editorial Assistant and CompositorMary Ann Schmidt, Senior Technical EditorHong Song, Programmer Yevonn Wilson-Ramsey, Illustrator

xii

Contributors to Volume 6

xiii

María Mercedes ArboInstituto de Botánica del NordesteCorrientes, Argentina

Kerry A. BarringerJersey City, New Jersey

David M. BatesCornell UniversityIthaca, New York

Orland J. Blanchard Jr.Florida Museum of Natural

HistoryUniversity of FloridaGainesville, Florida

Luc BrouilletUniversité de MontréalMontréal, Québec

Garrett E. CrowUniversity of New HampshireDurham, New Hampshire andMichigan State UniversityEast Lansing, Michigan

Laurence J. DorrNational Museum of Natural

HistorySmithsonian InstitutionWashington, DC

Aaron FlodenUniversity of TennesseeKnoxville, Tennessee

Paul A. Fryxell†Claremont, California

John F. GaskinNorthern Plains Agricultural

Research LaboratoryUnited States Department of

AgricultureSidney, Montana

Aaron Goldberg†National Museum of Natural

HistorySmithsonian InstitutionWashington, DC

Douglas H. GoldmanU.S.D.A. Natural Resources

Conservation ServiceGreensboro, North Carolina

Margaret M. HanesUniversity of WisconsinMadison, Wisconsin

Steven R. HillIllinois Natural History SurveyChampaign, Illinois

Walter C. HolmesBaylor UniversityWaco, Texas

Susannah B. Johnson-FultonShasta CollegeRedding, California

Brian R. KeenerThe University of West AlabamaLivingston, Alabama

John La DukeUniversity of Nebraska at KearneyKearney, Nebraska

David E. LemkeSouthwest Texas State UniversitySan Marcos, Texas

R. John LittleSacramento, California

John M. MacDougalHarris-Stowe State UniversitySt. Louis, Missouri

Landon E. McKinney†Environmental Quality

Management, Inc.Cincinnati, Ohio

T. Lawrence MellichampUniversity of North CarolinaCharlotte, North Carolina

Meghan G. MendenhallTexas Department of Public Safety Austin, Texas

Guy L. NesomFort Worth, Texas

xiv CONTRIBUTORS TO VOLUME 6

Lorin I. Nevling Jr.Illinois Natural History SurveyChampaign, Illinois

C. Thomas PhilbrickWestern Connecticut State

UniversityDanbury, Connecticut

Norman K. B. RobsonThe Natural History MuseumLondon, England

Janice G. SaundersInstituto de Botánica DarwinionBuenos Aires, Argentina

Bruce A. SorrieWhispering Pines, North Carolina

John L. StrotherUniversity of CaliforniaBerkeley, California

John W. Thieret†Northern Kentucky UniversityHighland Heights, Kentucky

Gordon C. TuckerEastern Illinois UniversityCharleston, Illinois

Linda E. WatsonOklahoma State UniversityStillwater, Oklahoma

Molly A. WhalenFlinders UniversityAdelaide, Australia

George YatskievychMissouri Botanical GardenSt. Louis, Missouri

Taxonomic Reviewers

xv

ALASKA / YUKON

Bruce BennettYukon Department of

EnvironmentWhitehorse, Yukon

Robert LipkinAlaska Natural Heritage ProgramUniversity of AlaskaAnchorage, Alaska

David F. MurrayUniversity of Alaska Museum of the NorthUniversity of Alaska FairbanksFairbanks, Alaska

Carolyn ParkerUniversity of Alaska Museum of the NorthUniversity of Alaska FairbanksFairbanks, Alaska

Mary StensvoldU.S.D.A. Forest ServiceSitka, Alaska

PACIFIC NORTHWEST

Edward R. AlversonThe Nature ConservancyEugene, Oregon

Adolf CeskaBritish Columbia Conservation

Data CentreVictoria, British Columbia

Richard R. HalseOregon State UniversityCorvallis, Oregon

Eugene N. KozloffShannon Point Marine CenterAnacortes, Washington

Jim PojarBritish Columbia Forest ServiceSmithers, British Columbia

Cindy RochéTalent, Oregon

Peter F. ZikaUniversity of WashingtonSeattle, Washington

Regional Reviewers

Laurence J. DorrNational Museum of Natural

HistorySmithsonian InstitutionWashington, DC

Margaret M. HanesUniversity of WisconsinMadison, Wisconsin

Steven R. HillIllinois Natural History SurveyChampaign, Illinois

Peter M. JørgensenMissouri Botanical GardenSt. Louis, Missouri

Peter H. RavenMissouri Botanical GardenSt. Louis, Missouri

Zachary S. RogersMissouri Botanical GardenSt. Louis, Missouri

REGIONAL REVIEWERSxvi

SOUTHWESTERN UNITED STATES

H. David Hammond†Northern Arizona UniversityFlagstaff, Arizona

G. Frederic Hrusa Courtland, California

Charles T. Mason Jr.† University of ArizonaTucson, Arizona

James D. MorefieldNevada Natural Heritage ProgramCarson City, Nevada

Donald J. Pinkava Arizona State UniversityTempe, Arizona

Jon P. RebmanSan Diego Natural History

MuseumSan Diego, California

Margriet WetherwaxUniversity of CaliforniaBerkeley, California

WESTERN CANADA

William J. Cody†Agriculture and Agri-Food CanadaOttawa, Ontario

Lynn GillespieCanadian Museum of NatureOttawa, Ontario

A. Joyce GouldAlberta Tourism, Parks and

RecreationEdmonton, Alberta

Vernon L. HarmsUniversity of SaskatchewanSaskatoon, Saskatchewan

Elizabeth PunterUniversity of ManitobaWinnipeg, Manitoba

ROCKY MOUNTAINS

Curtis R. BjörkUniversity of IdahoMoscow, Idaho

Bonnie HeidelUniversity of WyomingLaramie, Wyoming

B. E. NelsonUniversity of WyomingLaramie, Wyoming

NORTH CENTRAL UNITED STATES

William T. BarkerNorth Dakota State UniversityFargo, North Dakota

Anita F. CholewaUniversity of MinnesotaSt. Paul, Minnesota

Neil A. HarrimanUniversity of Wisconsin OshkoshOshkosh, Wisconsin

Bruce W. HoaglandUniversity of OklahomaNorman, Oklahoma

Robert B. KaulUniversity of NebraskaLincoln, Nebraska

Deborah Q. LewisIowa State UniversityAmes, Iowa

Ronald L. McGregor†The University of KansasLawrence, Kansas

Lawrence R. StritchU.S.D.A. Forest ServiceShepherdstown, West Virginia

George YatskievychMissouri Botanical GardenSt. Louis, Missouri

SOUTH CENTRAL UNITED STATES

David E. LemkeSouthwest Texas State UniversitySan Marcos, Texas

EASTERN CANADA

Sean BlaneyAtlantic Canada Conservation

Data CentreSackville, New Brunswick

Jacques CayouetteAgriculture and Agri-Food CanadaOttawa, Ontario

Frédéric CoursolMirabel, Québec

William J. CrinsOntario Ministry of Natural

ResourcesPeterborough, Ontario

John K. Morton†University of WaterlooWaterloo, Ontario

Marian MunroNova Scotia Museum of Natural

HistoryHalifax, Nova Scotia

Michael J. OldhamNatural Heritage Information

CentrePeterborough, Ontario

GREENLAND

Geoffrey HallidayUniversity of LancasterLancaster, England

NORTHEASTERN UNITED STATES

Ray AngeloNew England Botanical ClubCambridge, Massachusetts

Tom S. CooperriderKent State UniversityKent, Ohio

Arthur HainesBowdoin, Maine

Edward G. Voss†University of MichiganAnn Arbor, Michigan

Kay YatskievychMissouri Botanical GardenSt. Louis, Missouri

SOUTHEASTERN UNITED STATES

Mac H. AlfordUniversity of Southern MississippiHattiesburg, Mississippi

J. Richard Carter Jr.Valdosta State UniversityValdosta, Georgia

L. Dwayne EstesAustin Peay State UniversityClarksville, Tennessee

Paul J. HarmonWest Virginia Natural Heritage

ProgramCharleston, West Virginia

W. John HaydenUniversity of RichmondRichmond, Virginia

John B. NelsonUniversity of South CarolinaColumbia, South Carolina

Bruce A. SorrieUniversity of North CarolinaChapel Hill, North Carolina

R. Dale ThomasSeymour, Tennessee

Lowell E. UrbatschLouisiana State UniversityBaton Rouge, Louisiana

Thomas F. WieboldtVirginia Polytechnic Institute and

State UniversityBlacksburg, Virginia

B. Eugene WoffordUniversity of TennesseeKnoxville, Tennessee

FLORIDA

Loran C. AndersonFlorida State UniversityTallahassee, Florida

Bruce F. HansenUniversity of South FloridaTampa, Florida

Richard P. WunderlinUniversity of South FloridaTampa, Florida

REGIONAL REVIEWERS xvii