Flora of North America Contributors to Volume 28

Bruce H. Allen Masanobu Higuchi Terry J. O’Brien Tomotsugu Arikawa Michael S. Ignatov Paul L. Redfearn Jr. Jillian D. Bainard Robert R. Ireland Jr. William D. Reese† Mona Bourell David W. Jamieson Joseph R. Rohrer William R. Buck Robert E. Magill Inés Sastre-De Jesús John A. Christy Piers Majestyk Wilfred B. Schofeld† Steven P. Churchill Paul C. Marino A. Jonathan Shaw Howard A. Crum† Terry T. McIntosh James R. Shevock Patricia M. Eckel Norton G. Miller† John R. Spence Jean Faubert Gert Steen Mogensen Lloyd R. Stark Íñigo Granzow-de la Cerda Clayton Newberry Stephen L. Timme† Dana G. Griffn III Steven G. Newmaster Alain Vanderpoorten Judith A. Harpel Angela E. Newton Dale H. Vitt Terry A. Hedderson Daniel H. Norris William A. Weber Lars Hedenäs Richard H. Zander

Editors for Volume 28

Ariel S. Buback Pat Harris Lloyd R. Stark Technical Editor (2011–2014) Technical Editor (–2010) Taxon Editor

Marshall R. Crosby Martha J. Hill Barbara M. Thiers Taxon Editor Assisting Technical Editor Former Lead Editor

Claudio Delgadillo M. Robert W. Kiger Dale H. Vitt Taxon Editor Bibliographic Editor Taxon Editor

Patricia M. Eckel Terry T. McIntosh Richard H. Zander Taxon Editor Taxon Editor Lead and Taxon Editor

Kanchi Gandhi Robert E. Magill James L. Zarucchi Nomenclatural Advisor Nomenclatural Editor Editorial Director

Heidi H. Schmidt Managing Editor

Volume Composition

Kristin Pierce Heidi H. Schmidt Compositor and Editorial Assistant Production Coordinator and Managing Editor Flora of North America North of Mexico

Edited by FLORA OF NORTH AMERICA EDITORIAL COMMITTEE

VOLUME 28

Bryophyta, part 2

NEW YORK OXFORD ·· OXFORD UNIVERSITY PRESS 2014 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 Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto

With offces in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam

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 © 2014 by Flora of North America Association

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 28) Flora of North America North of Mexico edited 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

ISBN: 978-0-19-020275-0 (v. 28) 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 9 Printed in the United States of America on acid-free paper Contents

Founding Member Institutions vi Helodiaceae 320 Donors vii Hylocomiaceae 325 Art Sponsors viii Rhytidiaceae 338 Editorial Committee ix Leskeaceae 340 Project Staff x Pterigynandraceae 366 Contributors xi Thuidiaceae 373 Reviewers xiii Calliergonaceae 384 Acknowledgments xiv Brachytheciaceae 404 Preface xv Stereophyllaceae 469 Myriniaceae 474 Introduction xvi Fabroniaceae 477 Meteoriaceae 479 BRYOPHYTA, in part Plagiotheciaceae 483 The Impact of Molecular Data on our Understanding Fontinalaceae 489 of the Bryophyte Flora of North America 3 Entodontaceae 502 Splachnaceae 14 Climaciaceae 508 Meesiaceae 30 Pleuroziopsaceae 512 Catoscopiaceae 35 Hypnaceae 515 Orthotrichaceae 37 Sematophyllaceae 571 Hedwigiaceae 83 Cryphaeaceae 584 Aulacomniaceae 91 Leucodontaceae 590 Bartramiaceae 97 Pterobryaceae 597 Orthodontiaceae 113 Neckeraceae 602 Bryaceae 117 Lembophyllaceae 616 Roellobryaceae 186 Leptodontaceae 623 Pseudoditrichaceae 188 Anomodontaceae 629 Mielichhoferiaceae 190 Theliaceae 637 Mniaceae 215 Rhizogoniaceae 244 Key to the Moss Genera of North America North of Mexico 640 Racopilaceae 246 Errata to Volume 27 668 Hookeriaceae 248 Daltoniaceae 251 Pilotrichaceae 254 Rutenbergiaceae 259 Literature Cited 669 Hypopterygiaceae 261 Index 688 Amblystegiaceae 263 FOUNDING MEMBER INSTITUTIONS

Flora of North America Association

Agriculture and Agri-Food Canada Kansas State University University of Idaho Ottawa, Ontario Manhattan, Kansas Moscow, Idaho

Arnold Arboretum Missouri Botanical Garden University of Illinois Jamaica Plain, Massachusetts St. Louis, Missouri Urbana-Champaign, Illinois

University of Iowa Canadian Museum of Nature New Mexico State University Iowa City, Iowa Ottawa, Ontario Las Cruces, New Mexico

The New York Botanical Garden The University of Kansas Carnegie Museum of Bronx, New York Lawrence, Kansas Natural History Pittsburgh, Pennsylvania New York State Museum University of Michigan Albany, New York Ann Arbor, Michigan Field Museum of Natural History Chicago, Illinois Northern Kentucky University University of Oklahoma Highland Heights, Kentucky Norman, Oklahoma Fish and Wildlife Service University of Ottawa United States Department of Université de Montréal the Interior Ottawa, Ontario Montréal, Québec Washington, D.C. University of Southwestern University of Alaska Louisiana Harvard University Herbaria Fairbanks, Alaska Cambridge, Massachusetts Lafayette, Louisiana

University of Alberta The University of Texas Hunt Institute for Botanical Edmonton, Alberta Austin, Texas Documentation Carnegie Mellon University The University of British Columbia Pittsburgh, Pennsylvania University of Western Ontario Vancouver, British Columbia London, Ontario

Jacksonville State University University of California University of Wyoming Jacksonville, Alabama Berkeley, California Laramie, Wyoming

Jardin Botanique de Montréal University of California Utah State University Montréal, Québec Davis, California Logan, Utah vi For their support of the preparation of this volume, we gratefully acknowledge and thank:

The Andrew W. Mellon Foundation The David and Lucile Packard Foundation an anonymous foundation Chanticleer Foundation National Science Foundation

(Award DEB-0072754) The Fairweather Foundation ChevronTexaco ESRI Chris Davidson and Sharon Christoph Botanical Research Foundation of Idaho

vii For sponsorship of illustrations included in this volume, we express sincere appreciation to:

American Bryological and Lichenological Society, Madison, Wisconsin – Hylocomium splendens, Hylocomiaceae Ellen M. Anderson, Juneau, Alaska – Paludella squarrosa, Meesiaceae – in memory of Wilfred B. Schofeld Yelena Kosovich-Anderson, Cheyenne, Wyoming – Tomentypnum falcifolium, Amblystegiaceae & Plagiobryoides vinosula, Bryaceae – in memory of Siberian bryologist Leonid V. Bardunov James P. Bennett, Madison, Wisconsin – Bartramia pomiformis, Bartramiaceae Marshall R. Crosby & Shelley L. Pryor, Sugar Grove, North Carolina – Bryoandersonia illecebra, Brachytheciaceae Sandra J. Davis, Calgary, Alberta – Cratoneuron flicinum, Amblystegiaceae & Orthotrichum obtusifolium, Orthotrichaceae Janice M. Glime, Houghton, Michigan – Fontinalis neomexicana, Fontinalaceae Phillip Henderson, Fort Langley, British Columbia – Ctenidium schofeldii, Hypnaceae – in honor of Wilfred B. Schofeld Dale A. Kruse, College Station, Texas – Donrichardsia macroneuron, Brachytheciaceae Stephen Lodder, Watsonville, California – Alsia californica, Leptodontaceae, Claopodium whippleanum, Leskeaceae & Antitrichia californica, Leucodontaceae Bruce McCune, Corvallis, Oregon – Plagiomnium ciliare, Mniaceae Wynne Miles, Victoria, British Columbia – Brotherella canadensis, Sematophyllaceae – in honor of Wilfred B. Schofeld Harvey A. Miller, Palm Bay, Florida [general donation] Montana Native Plant Society, Kelsey Chapter, Helena, Montana – Scorpidium scorpioides, Calliergonaceae Montana Native Plant Society, Missoula, Montana – Elodium blandowii, Helodiaceae Nancy R. Morin, Point Arena, California – Thamnobryum allegheniense, Neckeraceae – in honor of Marshall R. Crosby Northwest Lichenologists, Corvallis, Oregon – Tetraplodon mnioides, Splachnaceae Stephen P. Rae, Napa, California – Anacolia menziesii, Bartramiaceae & Mnium ambiguum, Mniaceae James R. Shevock, San Francisco, California – Bryocrumia vivicolor & Dacryophyllum falcifolium, Hypnaceae, Bestia longipes, Lembophyllaceae & Porotrichum bigelovii, Neckeraceae John R. Spence, Page, Arizona – Rosulabryum canariense, Bryaceae David Toren, San Francisco, California – Bryum chryseum, Bryaceae Paul Wilson, Northridge, California – Pseudoleskeella serpentinensis, Leskeaceae & Warnstorfa fuitans, Calliergonaceae Gerhard Winter, Herbarium Senckenbergianum, Frankfurt am Main, Germany – Fontinalis antipyretica, Fontinalaceae – in memory of Jan-Peter Frahm Carl Wishner, Chicago Park, California – Meesia longiseta, Meesiaceae

For sponsorship of illustrations included in the previous moss volume, we also thank:

Botanical Club of Wisconsin, Markesan, Wisconsin – Schistostega pennata, Schistostegaceae Olivia L. Bottum, Ann Arbor, Michigan – Sphagnum steerei, Sphagnaceae – in memory of Gladys Bottum Patricia M. Eckel, St. Louis, Missouri – Grimmia brittoniae, Grimmiaceae A. Virginia Freire, Stevens Point, Wisconsin – Timmia megapolitana subsp. megapolitana, Timmiaceae Ohio Moss and Lichen Association, Columbus, Ohio – Polytrichastrum ohioense, Polytrichaceae Stephen P. Rae, Napa, California – Meiotrichum lyallii, Polytrichaceae & Scouleria aquatica, Scouleriaceae Richard H. Zander & Patricia M. Eckel, St. Louis, Missouri – Erpodium acrifolium, Erpodiaceae – in memory of Ronald A. Pursell viii FLORA OF NORTH AMERICA EDITORIAL COMMITTEE (as of 14 October 2013)

VASCULAR PLANTS

Kerry A. Barringer Ronald L. Hartman Robert F. C. Naczi Michael A. Vincent Taxon Editor Taxon Editor and Regional The New York Botanical Garden Taxon Editor Jersey City, New Jersey Coordinator, Rocky Bronx, New York Miami University Mountains Oxford, Ohio David E. Boufford Jackie M. Poole University of Wyoming Taxon Editor and Regional Alan S. Weakley Laramie, Wyoming Taxon Editor and Regional Coordinator, Coordinator, Taxon Editor and Regional Northeastern United States Steff M. Ickert-Bond South Central United States Coordinator, Harvard University Herbaria Regional Coordinator, Alaska Texas Parks and Wildlife Southeastern United States Cambridge, Massachusetts and Yukon Department University of North Carolina University of Alaska, Austin, Texas Chapel Hill, North Carolina Luc Brouillet Museum of the North Taxon Editor and Regional Richard K. Rabeler Elizabeth H. Zacharias Fairbanks, Alaska Coordinator, Taxon Editor Berkeley, California Eastern Canada and Greenland Robert W. Kiger University of Michigan James L. Zarucchi Institut de recherche en biologie Bibliographic and Taxon Editor Ann Arbor, Michigan Editorial Director végétale Hunt Institute for Botanical Jay A. Raveill Missouri Botanical Garden Université de Montréal Documentation St. Louis, Missouri Montréal, Québec Carnegie Mellon University Taxon Editor Pittsburgh, Pennsylvania University of Central Wayne J. Elisens Missouri BRYOPHYTES Taxon Editor Geoffrey A. Levin Warrensburg, Missouri University of Oklahoma Taxon Editor Terry T. McIntosh Heidi H. Schmidt Norman, Oklahoma Illinois Natural History Survey Taxon Editor Managing Editor Champaign, Illinois University of British Columbia Bruce A. Ford Missouri Botanical Garden Vancouver, British Columbia Taxon Editor and Regional Barney L. Lipscomb St. Louis, Missouri Coordinator, Botanical Research Institute of Dale H. Vitt Leila M. Shultz Western Canada Texas Taxon Editor Taxon Editor University of Manitoba Fort Worth, Texas Southern Illinois University Utah State University Winnipeg, Manitoba Carbondale, Illinois Timothy K. Lowrey Logan, Utah Craig C. Freeman Taxon Editor David H. Wagner John L. Strother Taxon Editor and Regional University of New Mexico Taxon Editor Reviewing and Taxon Editor Coordinator, Albuquerque, New Mexico Northwest Botanical Institute University of California North Central United States Eugene, Oregon R. L. McGregor Herbarium James A. Macklin Berkeley, California Harvard University Herbaria Alan T. Whittemore The University of Kansas Janet R. Sullivan Cambridge, Massachusetts Taxon Editor Lawrence, Kansas Taxon Editor United States National Gerry Moore University of New Hampshire Kanchi Gandhi Arboretum Liason to Governmental Agencies Durham, New Hampshire Nomenclatural Editor Washington, D.C. United States Department of Harvard University Herbaria Debra K. Trock Agriculture Richard H. Zander Cambridge, Massachusetts Taxon Editor Greensboro, North Carolina Bryophyte Coordinator and California Academy of David E. Giblin Taxon Editor Nancy R. Morin Sciences Taxon Editor Missouri Botanical Garden Taxon Editor and Regional San Francisco, California University of Washington St. Louis, Missouri Coordinator, Seattle, Washington Gordon C. Tucker Southwestern United States Lynn J. Gillespie Point Arena, California Taxon Editor Taxon Editor Eastern Illinois University Canadian Museum of Nature Charleston, Illinois Ottawa, Ontario

Emeritus/a Members of the Editorial Committee

George W. Argus Marie L. Hicks Barbara M. Murray Alan R. Smith Barbara M. Thiers Merrickville, Ontario Moab, Utah Fairbanks, Alaska Berkeley, California Bronx, New York Mary E. Barkworth Marshall C. Johnston David F. Murray Richard W. Spellenberg Rahmona A. Thompson Logan, Utah Austin, Texas Fairbanks, Alaska Las Cruces, New Mexico Ada, Oklahoma Marshall R. Crosby Aaron I. Liston J. Scott Peterson Lloyd R. Stark Frederick H. Utech Sugar Grove, North Carolinia Corvallis, Oregon Baton Rouge, Louisiana Las Vegas, Nevada Pittsburgh, Pennsylvania Claudio Delgadillo M. John McNeill Mexico City, Mexico Edinburgh, Scotland ix Project Staff — past and present involved with the preparation of Volume 28

Burgund Bassüner, Information Coordinator Ariel S. Buback, Technical Editor Trisha K. Distler, GIS Analyst Patricia M. Eckel, Illustrator Pat Harris, Technical Editor Martha J. Hill, Assisting Technical Editor Cassandra L. Howard, Editorial Assistant John Myers, Illustration Compositor Kristin Pierce, Compositor and Editorial Assistant Hong Song, Programmer

x Contributors to Volume 28

Bruce H. Allen Íñigo Granzow-de la Cerda Paul C. Marino Missouri Botanical Garden University of Michigan Memorial University St. Louis, Missouri Ann Arbor, Michigan St. John’s, Newfoundland

Tomotsugu Arikawa Dana G. Griffn III Terry T. McIntosh Keio University University of Florida University of British Columbia Yokohama, Japan Gainesville, Florida Vancouver, British Columbia

Jillian D. Bainard Judith A. Harpel Norton G. Miller† University of Saskatchewan Brush Prairie, Washington New York State Museum Saskatoon, Saskatchewan Albany, New York Terry A. Hedderson Mona Bourell University of Cape Town Gert Steen Mogensen California Academy of Sciences Rondebosch, South Africa University of Copenhagen San Francisco, California Copenhagen, Denmark Lars Hedenäs William R. Buck Swedish Museum of Natural History Clayton Newberry The New York Botanical Garden Stockholm, Sweden Bureau of Land Management Bronx, New York Vernal, Utah Masanobu Higuchi John A. Christy National Science Museum Steven G. Newmaster Oregon Natural Heritage Tsukuba, Japan University of Guelph Information Center Guelph, Ontario Oregon State University Michael S. Ignatov Portland, Oregon Main Botanical Garden Angela E. Newton Moscow, Russia The Natural History Museum Steven P. Churchill London, England Missouri Botanical Garden Robert R. Ireland Jr. St. Louis, Missouri Stittsville, Ontario Daniel H. Norris University of California Howard A. Crum† David W. Jamieson Berkeley, California University of Michigan Fort Lewis College Ann Arbor, Michigan Durango, Colorado Terry J. O’Brien Rowan University Patricia M. Eckel Robert E. Magill Glassboro, New Jersey Missouri Botanical Garden Missouri Botanical Garden St. Louis, Missouri St. Louis, Missouri Paul L. Redfearn Jr. Powell Gardens Jean Faubert Piers Majestyk Kansas City, Missouri Société québécoise de bryologie Duke University Saint-Valerian-de-Rimouski, Québec Durham, North Carolina

xi xii CONTRIBUTORS TO VOLUME 28

William D. Reese† James R. Shevock Alain Vanderpoorten University of Louisiana California Academy of Sciences University of Liége Lafayette, Louisiana San Francisco, California Liége, Belgium

Joseph R. Rohrer John R. Spence Dale H. Vitt University of Wisconsin Glen Canyon National Recreation Southern Illinois University Eau Claire, Wisconsin Area Carbondale, Illinois National Park Service Inés Sastre-De Jesús Page, Arizona William A. Weber University of Puerto Rico University of Colorado Museum Mayaguez, Puerto Rico Lloyd R. Stark Boulder, Colorado University of Nevada Wilfred B. Schofeld† Las Vegas, Nevada Richard H. Zander University of British Columbia Missouri Botanical Garden Vancouver, British Columbia Stephen L. Timme† St. Louis, Missouri Pittsburg State University A. Jonathan Shaw Pittsburg, Kansas Duke University Durham, North Carolina Reviewers

Bruce H. Allen David W. Jamieson Wilfred B. Schofeld† Missouri Botanical Garden Fort Lewis College University of British Columbia St. Louis, Missouri Durango, Colorado Vancouver, British Columbia

Lewis E. Anderson† Robert E. Magill A. Jonathan Shaw Duke University Missouri Botanical Garden Duke University Durham, North Carolina St. Louis, Missouri Durham, North Carolina

William R. Buck Piers Majestyk James R. Shevock The New York Botanical Garden Duke University California Academy of Sciences Bronx, New York Durham, North Carolina San Francisco, California

Benjamin E. Carter Kenneth D. McFarland John R. Spence University of California University of Tennessee Glen Canyon National Recreation Berkeley, California Knoxville, Tennessee Area National Park Service Howard A. Crum† Terry T. McIntosh Page, Arizona University of Michigan University of British Columbia Ann Arbor, Michigan Vancouver, British Columbia Benito C. Tan University of California Claudio Delgadillo M. Norton G. Miller† Berkeley, California Universidad Nacional Autónoma New York State Museum de México Albany, New York Barbara M. Thiers Mexico City, Mexico The New York Botanical Garden Angela E. Newton Bronx, New York Jennifer Doubt The Natural History Museum Canadian Museum of Nature London, England Steven L. Timme† Ottawa, Ontario Pittsburg State University Terry J. O’Brien Pittsburg, Kansas Isabel Draper Rowan University Universidad Autónoma de Madrid Glassboro, New Jersey Andries Touw Madrid, Spain Leiden University Ryszard Ochyra Leiden, Netherlands Janice M. Glime Botanical Institute Michigan Technological University Polish Academy of Sciences Dale H. Vitt Houghton, Michigan Kraców, Poland Southern Illinois University Carbondale, Illinois Bernard Goffnet Ronald A. Pursell† University of Connecticut Missouri Botanical Garden William A. Weber Storrs, Connecticut St. Louis, Missouri University of Colorado Museum Boulder, Colorado Lars Hedenäs Paul L. Redfearn Jr. Swedish Museum of Natural History Powell Gardens Alan T. Whittemore Stockholm, Sweden Kansas City, Missouri U.S. National Arboretum Washington, DC Robert R. Ireland Jr. William D. Reese† Stittsville, Ontario University of Louisiana Richard H. Zander Lafayette, Louisiana Missouri Botanical Garden St. Louis, Missouri xiii Acknowledgments

Members of the Flora of North America Association (FNAA), especially those involved in the preparation and production of this volume, extend special thanks to:

Barney L. Lipscomb, editor of Journal of the Botanical Research Institute of Texas [formerly Sida], and Guy L. Nesom, editor of Phytoneuron, who ensured that new species and combinations were published promptly, such that these taxa could be included in the published bryophyte volumes.

The U.S. Fish and Wildlife Service (USFWS) is acknowledged for having provided Federal assistance in the form of a joint agreement to share Flora of North America- and USFWS- funded illustrations used in this volume for the Illustrated Moss Flora of the Aleutian Islands, an identifcation manual edited by Wilfred B. Schofeld† and Stephen S. Talbot.

We appreciate the efforts of many individuals who provided advice, corrections, encouragement, and support for the bryophyte team of authors and editors. Special thanks go to Pat Harris, technical editor for the moss volumes through 2010, and to Ariel S. Buback in her capacity as technical editor for Volume 28 since the beginning of 2011; we extend special thanks to both of them for their dedication to the Flora project. We single out James L. Zarucchi, whose unrelenting insistence on clarity, consistency, and completeness improved the books immeasurably.

To the authors who contributed treatments to this volume, we extend our deepest appreciation for their hard work, scholarly contributions, and patience throughout the editing and composition process.

Finally, we thankfully acknowledge the dedication and organizational skills of Heidi H. Schmidt, especially in overseeing the fnal and exacting process of volume composition.

xiv