Spec. Rpt. Front Matter - British Columbia · Steve and Sylvia Sharnoff, and Jan Zwicky all helped...

1 9 9 9

Ministry of Forests Research Program

THE LICHENS OF

BRITISH COLUMBIA

S P E C I A L R E P O R T

Illustrated Keys

Part 2 — Fruticose Species

by Trevor Goward

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THE LICHENS OF

BRITISH COLUMBIA

Ministry of Forests Research Program

Illustrated Keys

Part 2 — Fruticose Species

by Trevor Goward

(Illustrations by Trevor Goward)

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES ii

Copies of this report may be obtained, depending upon supply, from:Crown Publications521 Fort StreetVictoria, BC

(250) 386-4636www.crownpub.bc.ca

For more information on Forestry Division publications, visit our web site at:http://www.for.gov.bc.ca/hfd/pubs/index.htm

Canadian Cataloguing in Publication DataGoward, Trevor

The lichens of British Columbia, illustrated keys.Part 2, Fruticose Species

(Special report series, ISSN 0843–6452 ; 9)Includes bibliographical references, p.Includes index.ISBN 0–7726–3961–2

1. Lichens – British Columbia – Identification.2. Lichens – British Columbia – Geographicdistribution. I. British Columbia. Ministry of Forests.Research Branch. II. Title. III. Series: Special reportseries (British Columbia Ministry of Forests) ; 9

QK587.7.B7G68 1999 579.7’09711 C99 – 960259 – 4

Prepared byTrevor GowardHerbarium, Department of BotanyUniversity of British ColumbiaVancouver, BC

Mailing Address:Edgewood Blue, Box 131Clearwater, BC

forB.C. Ministry of ForestsResearch Branch712 Yates StreetVictoria, BC

© 1999 Province of British Columbia

ACKNOWLEDGEMENTS iii

ACKNOWLEDGEMENTS

This manual has been four years in themaking. During that time, it has benefit-ted from the kindness of many people. Inparticular, I wish to thank five colleagues—Teuvo Ahti, Irwin Brodo, StephenClayden, Pekka Halonen, and HiroyukiKashiwadani—for entering into co-authorship on treatments of Cladina,Cladonia, Stereocaulon, Usnea, andRamalina. I also wish to thank the follow-ing specialists for reviewing earlier draftsof other critical genus accounts: IrwinBrodo (Alectoria, Bryoria), Håkon Holien(Calicium, Chaenotheca, Microcalicium,Sclerophora), Scott Kroken (Letharia),Scott LaGreca (Ramalina), Leif Tibell(Chaenothecopsis), Tor Tønsberg (Gyali-deopsis, Microlychnus, Szczawinskia),and Mats Wedin (Bunodophoron,Sphaerophorus). In addition, severallichenologists have willingly shared theirtaxonomic expertise: Teuvo Ahti, OthmarBreuss, Irwin Brodo, Aino Henssen, PerMagnus Jørgensen, Scott Kroken, ScottLaGreca, Bruce McCune, Juoko Rikkinen,Roger Rosentreter, Steve Selva, Leif Tibell,Tor Tønsberg, and Mats Wedin.

In addition: Olivia Lee, of the Universityof British Columbia, packaged, labelled,sorted, and forwarded lichen specimensby the thousands; Dan Burgess, NathalieDjan-Chékar, David Miège, CarlaRydholm, and Patrick Williston helped to prepare the distribution maps; IrwinBrodo, Stephen Clayden, BernardGoffinet, and Tor Tønsberg subjected sev-eral specimens to thin-layer chromatogra-phy; Stephen Clayden, Bernard Goffinet,Stuart Harris, Olivia Lee, Vivian Miao,Roger Rosentreter, Wilf Schofield, andÉmannuël Sérusiaux responded to urgentpleas for literature; Tyler Innes and Carla

Rydholm tested the keys; Irwin Brodo,François Lutzoni, Eric B. Peterson, andBruce Ryan provided unpublished data onlichen distribution in the western UnitedStates; Mikhail Zhurbenko augmented myunderstanding of lichen distribution ineastern Eurasia; and André Arsenault,Robert Bringhurst, Irwin Brodo, DennisFafard, Bruce McCune, Don McKay,Roger Rosentreter, Evelyn Schimmel,Steve and Sylvia Sharnoff, and Jan Zwickyall helped to pry open the Pandora’s boxof common names. To all of the above, Iextend my deepest gratitude.

I also express thanks to: Lyle Ottenbreitand Dan Bashaw for bringing the mapsand figures to camera-ready; SusanBannerman, Bruce Maricle, Vivian Miao,Anna Roberts, and Steve Smith forEnglish edit and proofreading; DonnaLindenberg for typesetting; Rick Scharf forcarrying the manuscript through to publi-cation; and Paul Nystedt for cheerfullyjuggling many balls in the air at the sametime.

Evelyn Hamilton, of the BritishColumbia Ministry of Forests, deservesspecial thanks for supporting this projectin many ways throughout its gestation—not least by funding it through the BritishColumbia Forest Renewal Fund (FRBC).For supplementary funding through theBritish Columbia Conservation DataCentre (CDC), I also thank AndrewHarcombe.

Finally, this book is fondly dedicated totwo eminent lichenologists: Teuvo Ahti, ofthe University of Helsinki, and IrwinBrodo, of the Canadian Museum ofNature. Many thanks, my friends, for allyou have done.

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES iv

CONTENTS

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1Interpreting the Genus and Species Accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3Understanding Biogeoclimatic Zonation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5Identifying Lichens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10Making Use of Lichen Chemistry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20A Note on Common Names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Keys to Genera of Fruticose Macrolichens and Microlichens . . . . . . . . . . . . . . . . . . 22Key to Lichen Photobionts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23Key to Lichen Growth Forms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27Key A: Fruticose Macrolichens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29Key B: “Calicioid” Lichens (and others) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36Key C: Fruticose Microlichens (and others). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

Keys to Species of Fruticose Macrolichens and Microlichens . . . . . . . . . . . . . . . . . . 45Acroscyphus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45Agrestia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46Alectoria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47Baeomyces . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51Bryocaulon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53Bryoria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54Bunodophoron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69Calicium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70Chaenotheca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76Chaenothecopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89Cladina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101Cladonia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 107Coelocaulon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164Cornicularia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165Cystocoleus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166Dactylina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167Dibaeis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170Ephebe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171Evernia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173Gyalideopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176Leciophysma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178Lempholemma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179Leprocaulon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181Letharia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183Lichinella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185Lichinodium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186Loxosporopsis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189Microcalicium. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190Microlychnus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192Mycocalicium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193Niebla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194Nodobryoria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195Phaeocalicium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

CONTENTS v

Pilophorus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199Polychidium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 202Pseudephebe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204Racodium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206Ramalina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207Sclerophora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215Siphula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217Sphaerophorus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218Spilonema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222Stenocybe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225Stereocaulon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 227Sticta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240Synalissa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241Szczawinskia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 242Teloschistes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 243Thamnolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244Thermutis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 245Tholurna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246Usnea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247Zahlbrucknerella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 265Unknown 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 266Unknown 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267Unknown 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 268

Appendix 1 Distribution maps of rare and infrequent fruticose lichens in British Columbia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269

Appendix 2 Excluded species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 292

Glossary and Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 295

References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312

TABLES

1 Distributional units and their definition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Summary information on the biogeoclimatic zones of British Columbia. . . . . . . . 7

FIGURES

1 First- and second-order lichen floristic studies in British Columbia to 1998. . . . . . 12 “Life zones” of British Columbia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 Biogeoclimatic zones of British Columbia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64 Thallus stratified/heteromerous (cross-section). . . . . . . . . . . . . . . . . . . . . . . . . . . . 115 Thallus nonstratified/homoiomerous (cross-section, in part). . . . . . . . . . . . . . . . . 116 Common photobionts (free-growing/unlichenized forms) . . . . . . . . . . . . . . . . . . . 127 Organs of attachment (cross-section). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138 Branching and degrees of attachment. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139 Growth forms (cross-section and surface view, in part). . . . . . . . . . . . . . . . . . . . . . 1410 Surface (and other) details: foliose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1511 Surface (and other) details: fruticose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1612 Spores and conidia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1713 Sexual and asexual reproductive structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1814 Vegetative reproductive structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

INTRODUCTION

Our knowledge of lichen floristics inBritish Columbia has advanced rapidly inrecent years. Only three decades ago, theknown British Columbia lichen florastood at 569 species (Otto and Ahti 1967).Today, by contrast, it stands at approxi-mately 1300 species (Noble et al. 1987;Goward, unpublished). Notwithstandingthis impressive figure, a comprehensiveinventory of the province’s lichen flora isstill a long way off. In support of thisclaim, consider that more than 20 lichenspecies, on average, are added to theprovincial lichen flora each year!

A significant number of lichen specieswarrant formal designation as “rare” or“infrequent” in British Columbia. Todate, four such species carry officialendangerment status in Canada (Gowardet al. 1998), while one species appears onthe International Union for Conservation

of Nature and Natural Resources (IUCN)“global redlist of lichens” (Thor 1996:www.dha.slu.se/guest/global.htm). Severalother species may already be at risk ofextirpation in the province (Goward1996). Reflecting these concerns, theBritish Columbia Conservation DataCentre has recently initiated a preliminarytracking list of the province’s “red-listed”and “blue-listed” lichens.

Few portions of the province havereceived serious attention from lichenolo-gists. To date, comprehensive lichen stud-ies have been carried out only onsoutheast Vancouver Island (Noble 1982)and the Queen Charlotte Islands (Brodo1995; Brodo and Ahti 1996; Brodo andSantesson 1997; Brodo and Wirth 1998).The lichen flora of Wells Gray Park is alsorelatively well documented (Goward andAhti 1992; Goward, unpublished).

INTRODUCTION

FIGURE 1 First- and second-order lichen floristic studies in British Columbia to 1998.

FIRST ORDER1. Brodo

(in prep.)2. Goward, Ahti

(1992)3. Noble (1982)

(Bird, Bird1973)(Ryan 1991)

SECOND ORDER4. Benton, Brodo, Richardson (1977)5. Brodo (unpublished)6. Goward (unpublished)7. Goward, Schofield (1983)8. Ohlsson (unpublished)9. Otto (unpublished)

10. Thomson, Ahti (1994)

6

6

6

6

6

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES

Important collections from other regionsdo exist (Figure 1), but have not beenpublished, and are widely scattered.

Lichen study in British Columbia hastraditionally been hampered by a lack ofcomprehensive keys to the species.Recently, however, Goward et al. (1994)published illustrated keys to 327 species offoliose and squamulose lichens. The pres-ent manual is a companion volume tothat work; it provides illustrated keys to309 species of fruticose lichens (and alliedfungi) that are known from or expected tooccur in British Columbia.

This manual adopts a broad interpreta-tion of the fruticose life form. Includedhere are all lichen genera—both macro-lichens and microlichens—in which amajority of species bear stalks or branchesthat are roughly circular in cross-section.Although fruticose microlichens are tradi-tionally regarded as crustose, they areincluded here owing to their dominantstalked sexual or asexual reproductivestructures. Examples of fruticose micro-lichens include Calicium, Chaenotheca,Gyalideopsis, and Microlychnus.

Two primary objectives have guidedthe preparation of this manual:1. to briefly summarize the ecology, dis-

tribution, and frequency status of allfruticose lichens known to occur inBritish Columbia.

2. to stimulate lichenological research bymaking these lichens accessible to abroad audience.In keeping with the presumed needs

and resources of ecologists, biologists,naturalists, teachers, and other beginningstudents of lichens, this manual emphasizesmorphological characters over chemicaland spore characters; it also avoids techni-cal terms as far as possible.1 It must beacknowledged, however, that some speciescannot be reliably identified withoutrecourse to thin-layer chromatography or

examination under a light microscope;fortunately, most such species are smalland inconspicuous, and are unlikely to beencountered by the beginner.

The genus and species concepts adoptedhere are often pragmatic. As a rule, theygive priority to “intuitive” morphologicalgroupings that do not always accord withthe latest findings of anatomical and molec-ular research. This approach proceeds fromthe assumption that laboratory researchwill continue to uncover evolutionary rela-tionships not readily perceived in the field.Looking ahead, two parallel approaches tolichen taxonomy can be expected to evolve:one that emphasizes taxonomic stability,and is suited to the requirements of lichenfloristics and field ecology; and one thatemphasizes phylogenetic relatedness, and issuited to continuing molecular, chemical,and ultrastructural studies. Although thismanual follows the first of these approach-es, alternative genus and species conceptsare given in the synonymy under theaccepted species.

Accompanying the keys are 320 linedrawings intended to convey species con-cepts based on typical material. The draw-ings emphasize specific charactersexpressed in the adjacent key, and are notintended to depict the entire lichen.Illustrations of whole lichens can be foundin various popular and semi-popular ref-erences, including Hale (1979), Kershaw et al. (1998), McCune and Geiser (1997),McCune and Goward (1995), MacKinnonet al. (1992), Parish et al. (1996), Pojar andMacKinnon (1994), Thomson (1984, 1997),and Vitt et al. (1988).

It is beyond the scope of this manual toprovide a comprehensive summary oflichen biology (see instead: Hale 1983;Hawksworth and Hill 1984; Lawrey 1984;Nash 1996). Effective identification does,however, require a basic understanding oflichen morphology and chemistry.

1 Technical terms are discussed in “Identifying Lichens” (page 10) and appear there in bold type. Additionalterms are defined in the keys, as well as in the Glossary (page 295).

INTRODUCTION

Interpreting theGenus and Species

Accounts

For convenience, lichens can be arrangedin several different growth forms, includ-ing crustose, squamulose, foliose, and fru-ticose (see “Identifying Lichens,” page 10).Because, however, these growth forms areunits of convenience, not biology, they donot always offer a perfect “fit” with thegenera they are supposed to circumscribe.For example, while all species of Bryoriaare fruticose, the genus Cladonia containsboth squamulose species and fruticosespecies. For convenience, this manualincorporates all lichen genera known tooccur in British Columbia in which amajority of species can be described asfruticose. In a few instances, fruticosespecies belonging to essentially nonfruti-cose genera are also included (in paren-theses) in the keys, but are not discussedin the species accounts. Species appearingin square brackets [...] are expected tooccur in British Columbia, but have yet tobe reliably recorded.

The body of the manual is arrangedalphabetically, first by genus and then byspecies within each genus.

The genus accounts include:1. the scientific name,2. a common name,3. a short description of the genus, with

diagnostic characters placed in bolditalic type,

4. pertinent references,5. the derivation of the common name,6. notes on global status, distribution,

taxonomy, chemistry, and points ofdistinction with similar genera.The species accounts are more com-

plex, and are organized under the follow-ing headings:

Species and Author Citation: Except incases of recent taxonomic or nomenclat-ural revision, species names and authorcitations follow Esslinger and Egan (1995).Synonyms: As a rule, only synonyms inrecent or widespread use are given.Distribution Maps: The map numberappearing to the right of some species iskeyed to Appendix 1, in which distribu-tion maps are provided for species judgedto be rare or infrequent in the province.Common Names: Common names areadopted, adapted, or introduced for alllichen species included in this manual.Alternative common names are given inparentheses (...). See also “A Note onCommon Names,” page 21.Habitat/Range: Information is providedon lichen frequency status, common sub-strates, site characteristics, provincial “lifezone” distribution, distribution in thenorthern hemisphere, and cordilleran dis-tribution. 1. Frequency status is given using the fol-

lowing terms: rare, infrequent, fre-quent, common.

2. Substrates include acid/base-rich/mossy/seasonally inundated rock,coniferous/deciduous trees or shrubs,and soil, moss, duff, bark, or wood.

3. Site characteristics are expressed as:exposed/open/ sheltered/shady/old-growth forests, steppe, depressions, oroutcrops, at lower/middle/upper/alpineelevations.

4. Provincial “life zone” distributions aregiven according to the terms listed incolumns I and II of Table 1; see alsoFigure 2. Biogeoclimatic units (columnIII of Table 1) are occasionally used,

2 Please direct comments to Trevor Goward, Edgewood Blue, Box 131, Clearwater, BC V0E 1N0.

Accordingly, the reader is invited to con-sult “Identifying Lichens” (page 10) priorto using the keys.

This manual represents a first attemptto provide comprehensive keys to the fru-ticose lichens of British Columbia. Though

every effort has been made to make thekeys as usable as possible, numerous errorsand oversights doubtless remain. Thereader is invited to bring these to theauthor’s attention for the benefit of futurestudents of British Columbia’s lichens.2


TABLE 1 Distributional units and their definition

General Life Biogeoclimatic Conrad’s Index ofRange Zone Equivalenta Continentality

(I) (II) (III) (IV)

COAST Hypermaritime CWH wh and vhb < 8Maritime 9 - 29

-dry CDF

-wet CWH (not wh and vh)-subalpine MH

INLAND Intermontane 30 - 39-semi-arid BG, PP

-dry (lowland) IDF

-dry (upland) SBPS

-moist (lowland) SBS

-moist (upland) MS

-humid (lowland) ICH

-subalpine ESSF

Boreal BWBS, SWB > 40SUBALPINE Throughout MH, ESSF variousALPINE Throughout AT variousWIDESPREAD Throughout “throughout” various

a See Table 2 or the Glossary for definitions of these biogeoclimatic zone codes.b Only the Wet Hypermaritime (wh) and Very Wet Hypermaritime (vh) subzones of the Coastal Western

Hemlock Zone (CWH) are included here.

FIGURE 2 “Life zones” of British Columbia.

I N T E R M O N T A N E

M A R I T I M E

B O R E A L

H Y P E R -M A R I T I M E

ki lometres

INTRODUCTION

and are mapped in Figure 3; see also“Understanding BiogeoclimaticZonation” (below). The continentalityunits in column IV are based onConrad’s Index of Continentality(Conrad 1946), and are included toenable ecoclimatic comparisons withother regions of the world (for furtherdetails, see Goward and Ahti 1992).

5. Distribution in the northern hemi-sphere is expressed relative to westernNorth America. The following distribu-tional units are used:

western N Amwestern N Am - eastern N Amwestern N Am - western Eurasiawestern N Am - eastern Eurasiaincompletely circumpolar (= any three

of the above distributional units)circumpolar

6. Cordilleran distribution is summarizedusing the following geographic units: Nto sAK (Alaska: Pacific coast only), AK(Alaska: boreal and arctic regionsonly), YU (Yukon), or wNT(Northwest Territories: west of theMackenzie River only); and S to AZ(Arizona), CA (California), CO(Colorado), ID (Idaho), MT(Montana), MX (Mexico), NM (NewMexico), NV (Nevada), OR (Oregon),

UT (Utah), WA (Washington), or WY(Wyoming); and AB (Alberta). Whilemost state records appearing in thismanual are based on published reports,a few unpublished records are alsogiven; these appear in parentheses.

Reactions: In most cases, only positivereactions are given, based on commonlyused chemical reagents and ultravioletlight; negative reactions are generallyomitted. For further details see “MakingUse of Lichen Chemistry,” page 20.Contents: In most cases, only dominantlichen substances are listed. Substancesgiven in parentheses are “accessory,” thatis, they do not occur in all specimens.Variability: Some lichen species varygreatly with habitat, whereas others aremore uniform. To assist in identification,each species has been assigned a variabilityrating of “low,” “medium,” or “high.”Species rated as “high” do not necessarilyconform with all characters given in thekeys.Notes: Included here are comments ontaxonomic problems, points of distinctionwith similar species, chemistry, and keysto subspecies and varieties. In general,detailed notes are reserved for taxonomi-cally difficult genera.

UnderstandingBiogeoclimatic

Zonation

British Columbia is a highly diverse prov-ince in which hundreds of ecosystems canbe recognized. Maintaining these in theface of increasing pressure for resourcedevelopment represents an enormouschallenge—and involves, as a first step,classifying the province’s ecosystems indetail.

In recent years, researchers with theB.C. Ministry of Forests have describedmedium-scale ecosystems according to theprinciples of biogeoclimatic ecosystemclassification (Pojar et al. 1987). They havealso arranged these ecosystems into a hier-archical system of biogeoclimatic zones,subzones, and variants.

Collectively, the zones, subzones, and

variants of the biogeoclimatic system arereferred to as biogeoclimatic units. Eachunit is characterized by a unique set of cli-matic variables, and supports—and is forpractical purposes defined by—a uniquevegetation. In biogeoclimatic ecosystemclassification, the defining vegetation foreach unit occurs on moderately well-drained sites. Such sites are said to be“zonal.”

The most encompassing of the biogeo-climatic units is the biogeoclimatic zone.Fourteen biogeoclimatic zones are recog-nized for British Columbia and many ofthese are used here to describe lichen dis-tribution. They are briefly characterized inTable 2 and mapped in Figure 3. For a

Research Branch, Ministry of Forests, Victoria, BC

Biogeoclim

atic zones of British Colum

bia.

INTRODUCTION

more detailed summary, see Ecosystems ofBritish Columbia (Meidinger and Pojar1991).

Lichen distribution may also beexpressed using more generalized classifi-cation systems such as the “life zone sys-tem” (see Figure 2) and “general rangesystem” adopted here. These systems arecompared with their biogeoclimatic coun-terparts in Table 1. The comparison is

made mostly at the zonal level, thoughtwo biogeoclimatic subzones have alsobeen used: the Wet Hypermaritime (wh)and Very Wet Hypermaritime (vh) sub-zones of the Coastal Western HemlockZone (CWH). These subzones occur in thehypermaritime or outer coastal areas ofBritish Columbia (see Figure 2). See Table2 for the full names of other biogeoclim-atic zones.

TABLE 2 Summary information on the biogeoclimatic zones of British Columbia (Source: Lavender et al. 1990)

Selected climatic characteristicsa

Monthly °days °days May–Sept. Oct.–AprilZone Code Zonal vegetation Zonal soils temp. range >5°C <0°C ppt (mm) ppt (mm)

AlpineTundra

BorealWhite and BlackSpruce

Bunch-grass

CoastalDouglas-fir

AT

BWBS

BG

CDF

Cassiope spp., Phyllodoce spp.,Luetkea pectinata, Loiseleuriaprocumbens, Dryas spp., Salixspp., Silene acaulis, Poa spp.,Festuca spp., Carex spp.,Cetraria spp., Stereocaulonspp., Polytrichum piliferum

White spruce, lodgepole pine,black spruce, Rosa acicularis,Viburnum edule, Mertensiapaniculata, Pyrola asarifolia,Cornus canadensis, Vacciniumvitis-idaea, Ptilium crista-cas-trensis, Pleurozium schreberi

Agropyron spicatum, Artemisiatridentata, Artemisia frigida,Poa sandbergii, Koeleriamacrantha, Festuca scabrella,Festuca idahoensis,Chrysothamnus nauseous

Douglas-fir, grand fir, bigleafmaple, western flowering dog-wood, Holodiscus discolor,Gaultheria shallon, Mahonianervosa, Rosa gymnocarpa,Symphoricarpos albus,Trientalis latifolia, Rubus ursi-nus, Pteridium aquilinum,Kindbergia oregana, Rhytidia-delphus triquetrus

Regosols,HumicRegosols,Brunisols,Humo-FerricPodzols

Gray Luvisols,Dystric andEutricBrunisols

Brown andDark BrownChernozems

DystricBrunisols

-11.1–9.5 427 1763 287 469

-24.5–16.6 709–1268 1692–2742 145–305 182–198

-10.8–22.4 1771–2516 230–878 98–175 108–208

1.8–18.0 1794–2121 9–43 107–238 540–1107


CoastalWesternHemlock

Engel-mannSpruce–SubalpineFir

InteriorCedar–Hemlock

InteriorDouglas-fir

CWH

ESSF

ICH

IDF

Western hemlock, amabilis fir,Sitka spruce, yellow-cedar,Vaccinium alaskaense,Vaccinium parvifolium,Menziesia ferruginea,Gaultheria shallon, Polystichummunitum, Pteridium aquil-inum, Blechnum spicant,Clintonia uniflora,Rhytidiadelphus loreus,Hylocomium splendens

Subalpine fir, Engelmannspruce, Rhododendron albiflo-rum, Menziesia ferruginea,Vaccinium (membranaceum,ovalifolium, scoparium), Rubuspedatus, Gymnocarpium dry-opteris, Tiarella unifoliata,Valeriana sitchensis, Orthiliasecunda, Streptopus roseus,Veratrum viride, Barbilophozialycopodioides, Pleuroziumschreberi, Rhytidiopsis robusta

Western hemlock, western red-cedar, hybrid white spruce,Douglas-fir, subalpine fir,Vaccinium ovalifolium,Oplopanax horridus, Vacciniummembranaceum, Rubus parvi-florous, Paxistima myrsinites,Smilacina racemosa, Streptopus(amplexifolius, roseus),Chimaphila umbellata,Goodyera oblongifolia,Gymnocarpium dryopteris,Ptilium crista-castrensis,Pleurozium schreberi,Hylocomium splendens,Rhytidiadelphus triquetrus

Douglas-fir, lodgepole pine,ponderosa pine, Spiraea betuli-folia, Amelanchier alnifolia,Juniperus communis,Symphoricarpos albus,Mahonia aquifolium, Paxistimamyrsinites, Calamagrostisrubescens, Arctostaphylos uva-ursi, Agropyron spicatum,Pleurozium schreberi

Ferro-Humicand Humo-Ferric Podzols

Humo-FerricPodzols

Humo-FerricPodzols, GrayLuvisols, andDystricBrunisols

Gray Luvisols,Eutric andDystricBrunisols

-6.6–18.7 1059–2205 5–493 159–1162 695–3225

-10.9–13.3 629–801 879–1189 205–425 271–1597

-10.7–20.8 1267–2140 238–820 200–439 294–1098

-13.1–21.3 903–2366 235–1260 107–291 149 –1022



TABLE 2 Continued

INTRODUCTION

MontaneSpruce

MountainHemlock

PonderosaPine

Spruce–Willow–Birch

Sub-BorealPine–Spruce

MS

MH

PP

SWB

SBPS

Hybrid white spruce, sub-alpine fir, lodgepole pine,Douglas-fir, Vaccinium scopari-um, Lonicera utahensis,Shepherdia canadensis,Paxistima myrsinites,Vaccinium membranaceum,Alnus viridis, Linnaea borealis,Empetrum nigrum,Calamagrostis rubescens,Pleurozium schreberi

Mountain hemlock, amabilisfir, yellow-cedar, Vaccinium(ovalifolium, membranaceum,alaskaense), Menziesia ferrug-inea, Rhododendron albiflorum,Rubus pedatus, Phyllodoceempetriformis, Rhytidiopsisrobusta, Rhytidiadelphus loreus,Hylocomium splendens

Ponderosa pine, Agropyron spi-catum, Balsamorhiza sagittata,Festuca (saximontana, ida-hoensis), Koeleria macrantha,Lithosperum ruderale, Achilleamillefolium

White spruce, subalpine fir,Salix glauca, Betula glandulosa,Potentilla fruticosa, Shepherdiacanadensis, Festuca altaica,Lupinus arcticus, Pedicularislabradorica, Epilobium angusti-folium, Empetrum nigrum,Vaccinium (vitis-idaea, caespi-tosum), Hylocomium splendens,Cladina spp., Nephromaarcticum

Lodgepole pine, white spruce,Shepherdia canadensis, Spiraeabetulifolia, Rosa acicularis,Calamagrostis rubescens,Arctostaphylos uva-ursi,Vaccinium caespitosum,Linnaea borealis, Pleuroziumschreberi, Peltigera spp.,Cladina spp.

DystricBrunisols andHumo-FerricPodzols

Ferro-HumicPodzols andFolisols

Eutric andDystricBrunisols

Eutric orDystricBrunisols,Humo-FerricPodzols

Gray Luvisolsand DystricBrunisols

-12.5–17.4 891–1310 847–890 158–252 223–469

-2.3–13.2 919–933 307–352 694–707 1857–2260

-8.6–21.6 1505–2442 258–861 86–270 170–334

-19.2–14.0 534–933 2036–2298 275–280 179–424

-13.8–14.3 697–1044 1140–1405 243–300 218–222



TABLE 2 Continued


Sub-BorealSpruce

SBS Hybrid white spruce, sub-alpine fir, lodgepole pine,Vaccinium membranaceum,Rubus parviflorus, Viburnumedule, Lonicera involucrata,Spiraea betulifolia, Rosa acicu-laris, Aralia nudicaulis, Cornuscanadensis, Linnaea borealis,Arnica cordifolia, Clintoniauniflora, Aster conspicuus,Osmorhiza chilensis, Oryzopsisasperifolia, Smilacina racemosa,Gymnocarpium dryopteris,Pleurozium schreberi, Ptiliumcrista-castrensis, Hylocomiumsplendens, Dicranum polyse-tum, Rhytidiadelphus tri-quetrus, Peltigera spp.

Gray Luvisolsand DystricBrunisols,Humo-FerricPodzols

-14.6–16.9 884–1510 792–1369 189–353 250–1383



TABLE 2 Concluded

The vast majority of lichens are classifiedas Ascomycetes (cup fungi), and hence arerelated to morels and elf saddles. Whilemost fungi, however, draw their nourish-ment from sources external to themselves(e.g., decaying leaves or logs), lichen fungi“cultivate” their foodstuff among the fun-gal threads of which they themselves arecomposed. This foodstuff consists of tiny,photosynthesizing algal or cyanobacterialcells, or both. Lichens can therefore beviewed as living greenhouses supportedlargely by carbohydrates derived from thephotosynthetic “crops” growing withinthem. This in part accounts for theexposed lifestyle adopted by most lichens:whereas a majority of fungi pass their lives(except when fruiting) within the thingsthey feed on, lichens colonize the surfacesof rocks, trees, logs, duff, and soil.

When a fungus enters into a stable,enduring relationship with a microscopicalga or cyanobacterium, both partners aresaid to be lichenized. In general appear-ance, most lichenized fungi, algae, andcyanobacteria do not closely resemble

their unlichenized relatives. Rather, theyform a composite “plant,” or thallus(Figures 4–5). Under the microscope, atypical lichen thallus resembles a kind ofsandwich in which the fungal partner(mycobiont) and the “algal” partner (photobiont) are stratified in distinct lay-ers (Figure 4). In many conspicuouslichen species, four such layers can be dis-cerned: a protective rind or upper cortex(Figure 4a); an “algal” or photobiont layer(Figure 4b); a pale, usually whitish regionof loosely interwoven fungal threads,called the medulla (Figure 4c); and anoth-er protective covering or lower cortex(Figure 4d).

As already mentioned, the photobiontsin nearly all lichens consist of green algaeor cyanobacteria. When exposed by arazor blade and viewed under a dissectingmicroscope, most algae are easily recog-nized by their characteristic single-celledhabit, as well as by their bright grass-greencolour (but yellowish green in Trente-pohlia). Cyanobacteria are much morevariable in form—single-celled, cluster-

Identifying Lichens

a Selected climatic characteristics summarized from Atmospheric Environment Service (AES) long-term stations. Prepared by D. Meidinger.

INTRODUCTION

FIGURE 4 Thallus stratified/heteromerous (cross-section): (a) upper cortex; (b) algal orcyanobacterial layer / photobiont layer; (c) medulla; (d) lower cortex; and (e) acephalodium.

(a)

(d)

(b)

(c) (e)

celled, or strandlike—but they are nevergrass-green, although at least one genuscan be yellowish green. In some speciesdominated by a green algal photobiont,cyanobacteria are also present as scattered,localized colonies called cephalodia(Figure 4e). These can occur internally or over the upper or lower surface(Figures 10d, 11l).

In some groups having a cyanobacterialphotobiont, the photobiont cells are inter-mingled throughout with fungal threads,and the thallus is uniformly dark bothinside and out. Such lichens are said to benonstratified (Figure 5); viewed from theoutside, they are typically brownish,blackish, or bluish grey. They can assumea gelatinous consistency when wet, andare then popularly referred to as “gellichens.” Most nonstratified lichens lack acortex (Figure 5a), though a primitive cel-lular cortex is present in Leptogium and

Polychidium (Figure 5b). Strandlike rhi-zoids (Figure 5c) can also occur in somecyanobacterial species.

Many different groups of algae and cy-anobacteria have entered into associationwith lichen fungi. Common cyanobacteri-al groups include the Chroococcales(Figure 6a), Nostoc (Figure 6b), theRivulariaceae (Figure 6c), Scytonema(Figure 6d), and Stigonema (Figure 6e).The most common algal groups includeStichococcus (Figure 6f), Trebouxia andrelated genera (Figure 6g), and Trente-pohlia (Figure 6h). For keys to these andother photobiont genera, see page 23.

Specialized holdfasts, or rhizines(Figure 7a–e), occur in many species hav-ing a lower cortex. Rhizines anchor thelichen to the colonized surface or sub-strate, and can be simple (Figure 7a),forking (Figure 7b), squarrose (Figure 7c),tufted (Figure 7d), or flocculent

FIGURE 5 Thallus nonstratified/homoiomerous (cross-section, in part): (a) noncorticate, (b)corticate(←), (c) bearing rhizoids(←).

(a) (b) (c)


FIGURE 6 Common photobionts (free-growing/unlichenized forms; for lichenized forms, see the key on page 23), a–e = cyanobacteria; f–h = algae: (a) Chroococcales; (b) Nostoc; (c) Rivulariaceae; (d) Scytonema; (e) Stigonema; (f) Stichococcus; (g) trebouxioid; and (h) Trentepohlia.

(a) (b) (c) (d)

(e)

(f)

(g)

(h)

(Figure 7e). In some foliose species (seebelow), rhizines are replaced by a singlethickened point of attachment, or umbili-cus (Figure 7f). In others, the rhizines arereplaced by a dark, woolly hypothallus(Figure 7g) that can sometimes extendbeyond the margins of the lichen.Rhizine-like structures that occur alongthe margins of leaflike lobes are calledcilia (Figure 10i).

Two broad categories of branching arerecognized in this manual. Branching issaid to be even when the arms of eachbranch pair are predominantly equal(Figure 8a), and uneven when they are notequal (Figure 8b). Similarly, five degreesof attachment can be distinguished:appressed (Figure 8c), decumbent

(Figure 8d), semi-erect (Figure 8e), erect(Figure 8f), and pendent (Figure 8g).

Lichens have traditionally been dividedinto three growth forms (crustose, foliose,and fruticose: see below), though otherclassifications are possible. The one adopt-ed here recognizes seven growth forms:1. Dust/leprose lichens (Figure 9a) lack

both an upper and lower cortex, thelower surface being attached directly tothe substrate, and the upper surfacebearing a continuous covering of pow-dery or granular soredia.

2. Crust/crustose lichens (Figure 9b) arealso attached directly to the substrate,but have a hard, protective upper cor-tex; viewed from above, they oftenresemble paint stains. Some crust

INTRODUCTION

FIGURE 7 Organs of attachment (cross-section): (a) rhizines (simple); (b) rhizines (forking); (c)rhizines (squarrose); (d) rhizines (tufted); (e) rhizines (flocculent, confluent); (f) holdfast(umbilicus); and (g) hypothallus.

FIGURE 8 Branching and degrees of attachment: (a) even/isotomic branching; (b) uneven/anisotomicbranching; (c) appressed(←); (d) decumbent; (e) semi-erect; (f) erect; and (g) pendent.

(a)

(b) (c)

(d)

(e) (f)

(g)

(a) (b) (c) (d) (e) (f) (g)


lichens give rise to elongate isidia (seebelow), while others bear stalked fruit-ing structures termed podetia andpseudopodetia; such species can beclassified as club lichens or shrublichens (see below). Other crust lichensintergrade with scale and leaf lichens(see below).

3. Scale/squamulose lichens (Figure 9c)are similar to dust and crust lichens inlacking a lower cortex (and rhizines).The thallus, however, consists of small,often partly raised, and usually overlap-ping/imbricate scales/squamules(Figure 9c), the lower surface of whichis often white and cottony (check underhand lens). In Cladonia, the squamulesoften give rise to hollow, stalked fruit-ing structures called podetia; see alsoclub and shrub lichens, below.

4. Leaf/foliose lichens (Figure 9d) more orless resemble leaves—at least in thesense that their thalli consist of flat-tened lobes that typically have an upperand lower cortex. The lobes can be nar-row or broad, and short or elongate.The degree of attachment to the sub-strate varies from appressed throughsemi-erect or even unattached. This isthe only growth form in which well-developed rhizines occur.

5. Club/fruticose lichens (Figure 9e) are atleast partly round in cross-section; theyhave no “true” lower surface and there-fore no lower cortex and rhizines. Mostclub lichens consist of thickened, up-

right, unbranched, or sometimes spar-sely branched stems; when hollow, theyare usually referred to as podetia; whensolid, they are called pseudopodetia.

6. Shrub/fruticose lichens (Figure 9f)resemble club lichens in having some-what thickened stems that are more orless circular in cross-section. Here,however, the stems are stronglybranched. Occasionally the stems arisefrom a basal crust or basal scales. Whenhollow, they are generally termed pode-tia; whereas, when solid, they are againcalled pseudopodetia. Shrub lichensvary from decumbent to semi-erect orerect.

7. Hair/fruticose lichens (Figure 9g) differfrom shrub lichens in having muchfiner, and proportionately much longer,branches. In habit, hair lichens aresemi-erect or pendent.The upper cortex of most lichens is

smooth and naked. However, in somespecies it can be minutely roughened (i.e., scabrid), or covered in pustules(Figure 10a) or a fine network of ridges (areticulum: Figure 10b). Other species beara thin whitish frosting (or pruina), whileothers still have a fine nap of tiny, erect,or appressed glasslike hairs (or tomentum:Figure 10c). A woolly tomentum is alsosometimes present over the lower surface.

The lower surface of most lichens isalso smooth, though raised veins (Figure10h) occur in some foliose species. In oth-ers, the lower surface can be sparsely

FIGURE 9 Growth forms (cross-section and surface view, in part): (a) dust/leprose; (b)crust/crustose; (c) scale/squamulose (with squamules); (d) leaf/foliose (with lobes); (e)club/fruticose; (f) shrub/fruticose; and (g) hair/fruticose.

(a) (b) (c) (d) (e) (f) (g)

INTRODUCTION

speckled with tiny pits through which the medulla is exposed. When rimmedand craterlike, these pits are termedcyphellae (Figure 10f); otherwise they arecalled pseudocyphellae (Figure 10g). Insome lichens, pseudocyphellae occur overthe upper cortex, and must then be care-fully distinguished from maculae (Figure10e): pale areas of the upper surface inwhich the cortex is unbroken. Fruticoselichens can also produce pseudocyphellae.

Surface details can provide important

clues to identification. In some species, forexample, the cortex is pocked by broaddepressions (or foveoles: Figure 11a), whilein others it bears conspicuous longitudi-nal striations (Figure 11b). Yet otherspecies carry numerous tile-like areoles(Figure 11c) which may or may not bepeltate (i.e., attached by a short centralstalk: Figure 11d). The presence of tiny,scale-like microsquamules (Figure 11f) isdiagnostic for some Cladonia species,whereas some species of Usnea bear

FIGURE 10 Surface (and other) details: foliose: (a) pustules(←); (b) reticulum(←); (c) tomentum(←); (d) cephalodia(←); (e) maculae(←); (f) cyphellae(←) (lower); (g) pseudocyphellae(←); (h) veins(←) (lower); and (i) cilia(←).

(a)(b)

(c)

(d)

(e) (f)

(g)(h)

(i)


numerous goosefleshlike papillae (Figure11g), or copious short side branches (orfibrils: Figure 11h). Similarly, mostStereocaulon species support a dense“foliage” of phyllocladia (Figure 11j),while small spine-like spinules (Figure 11i)occur in some species of Bryoria.

As a group, lichens reproduce both

vegetatively (in which case the fungus andalga/cyanobacterium are dispersed togeth-er, as a functional unit), and by sexualmeans (in which case only the spores ofthe fungus are dispersed: see below).Vegetative reproduction sometimes occursas a result of mechanical fragmentation(wear and tear), but more often it involves

FIGURE 11 Surface (and other) details: fruticose: (a) foveoles(←); (b) striations(←); (c) plates/areoles(←); (d) peltate areoles(←); (e) pseudocyphellae(←); (f) microsquamules(←); (g) papillae(←); (h) fibrils(←); (i) spinules(←); (j) phyllocladia(←); (k) central cord(←); and (l) cephalodia(←).

(a) (b) (c)

(d)

(e) (f)

(g)

(h)

( i )( j ) (k)

( l )

INTRODUCTION

transport (as by birds) of specialized out-growths called soredia and isidia. Soredia(Figure 14a–d) are masses of soft, powderygranules that have erupted through cracksand other areas of weakness in the cortex.When very fine (i.e., resembling talcumpowder), they are said to be powdery/fari-nose (Figure 14b); otherwise they can bedescribed as more or less granular (Figure14c). Though commonly borne in well-delimited soralia (Figure 14a), soredia canalso be broadcast over the surface; theyare then said to be diffuse (Figure 14d). Bycontrast, isidia (Figure 14e–i) are tiny out-growths of the upper cortex; their hard-ened outer surface is usually readilydistinguished from the powdery appear-ance of soredia. Similar to soredia, isidia

contain both a fungus and an alga/cyano-bacterium. They can be granular/globose(Figure 14e), barrel-shaped (Figure 14f),fingerlike/cylindrical (Figure 14g), coral-loid (Figure 14h), or scale-like (Figure 14i).

The primary function of sexual fruitingbodies is to produce sexual spores (Figure12a–l). Sexual spores come in manyshapes, and can be threadlike/filiform(Figure 12a), spindle-shaped/fusiform(Figure 12b), ellipsoid (Figure 12c), club-like/clavate (Figure 12d), globose (Figure12e), or peanut-shell-like (Figure 12f).They can also be 1-celled (Figure 12g), ordivided by a narrow cross-wall or septum(Figure 12h, i) into 2-celled (Figure 12h)or multi-celled spores (Figure 12i). Whenthe septum is very broad, the spores are

FIGURE 12 Spores and conidia: (a) threadlike/filiform; (b) spindle-shaped/fusiform; (c) ellipsoid; (d)clublike/clavate; (e) globose; (f) peanut-shell-shaped; (g) 1-celled/simple; (h) 2-celled(with narrow septum(←)); (i) many-celled/multi-septate; (j) polarilocular (with broadseptum(←)); (k) submuriform; and (l) muriform.

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termed polarilocular (Figure 12j). Sporeshaving both crosswise and lengthwisesepta are called submuriform (Figure 12k)or, when well developed, muriform(Figure 12l).

For most species covered in this manu-al, sexual fruiting bodies take the form ofapothecia (Figure 13a–g). These are smallsaucerlike, buttonlike, or hemisphericalstructures usually readily observed overthe thallus surface. Apothecia are gen-

erally unstalked (Figure 13a–e), but in afew genera they are borne at the ends oflong, brittle stalks as expanded “heads,”or capitula (Figure 13f, g).

Apothecia contain both fertile and ster-ile tissues. The former comprise the cen-tral disc: a compact cluster of tiny vial-likeasci, each bearing its complement of one toseveral spores (Figure 13a–e). Surroundingthe disc is a sterile rim, or excipulum(Figure 13a, d, e). When photobiont cells

FIGURE 13 Sexual and asexual reproductive structures (surface view unless indicated): (a) unstalked apothecia (showing discsand well-developed apothecial rims/excipula(←)); (b) unstalked apothecium (with a buttoned disc/omphalo-disc(←)); (c) unstalked apothecium (with fissured disc/gyrodisc); (d) unstalked apothecium (cross-section: withnonthalline rim/excipulum and spore-containing asci(←)); (e) unstalked apothecium (cross-section: with thallinerim/excipulum: note presence of algae(←)); (f) stalked apothecium (with excipulum and disc [i.e., capitulum(←)]);(g) stalked apothecium (showing mazaedium(←)); (h) perithecia; (i) perithecia (cross-section: note spore-containing asci); (j) pycnidia; (k) pycnidia (cross-section): note absence of asci; (l and m) hyphophores (showingstalks and conidial heads(←)).

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INTRODUCTION

FIGURE 14 Vegetative reproductive structures: (a) soredia in a soralium(←); (b) soredia (powdery/farinose(←)); (c) soredia (granular(←)); (d) soredia (diffuse); (e) isidia (granular/globose); (f) isidia (barrel-shaped); (g) isidia (fingerlike/cylindrical); (h) isidia (coralloid);and (i) isidia (scale-like(←)).

are present in the excipulum, it is said tobe thalline (Figure 13e); when they arelacking, it is nonthalline (Figure 13d). InCalicium, Sphaerophorus, and related gen-era, the disc is replaced by a distinctlypowdery mazaedium (Figure 13g) thatreadily smudges when rubbed. In otherlichens, sexual fruiting bodies take theform of perithecia (Figure 13h, i): minuteflask-like, ascus-bearing structures that areimmersed in the thallus, and are visiblefrom above as blackish or brownish dots.

Perithecia must be carefully distin-guished from some forms of pycnidia

(Figure 13j, k) which, though also dotlike,bear reproductive cells called conidia.Conidia come in many shapes but, unlikesexual spores, they never develop in vial-like asci. They are also usually muchsmaller than sexual spores, often measur-ing less than 4–5 µm long. In some genera,conidia are produced at the tips of stalkedfruiting bodies called hyphophores(Figure 13l, m), while in others they areborne directly over the cortex in well-demarcated, black, sooty patches; theseare termed thalloconidia.

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Lichens produce a diverse array of chemi-cal substances. Reflecting this, lichenchemistry provides a useful tool for theidentification of many species. The pres-ence of specific chemical substances canbe established through the use of spottests, ultraviolet lamps, and thin-layerchromatography.1. Spot Tests A spot test is performedwhen a small quantity of liquid reagent isapplied to one or more lichen tissues. Thiscan be done using a fine brush or capillarypipette that has been drawn to a pointover a flame. The resulting colour change(or lack of it) is often diagnostic for oneor more of several chemical substancespresent in the lichen. Five reagents arecalled for in this manual: calciumhypochlorite (C),3 nitric acid (H), potassi-um iodide (I), potassium hydroxide (K),and paraphenyldiamine (PD). Of these,only C, K, and PD are used routinely. Allof the reagents are toxic and should becarefully stored in small, tightly sealedglass bottles. Chemical reactions can beobserved using a hand lens (10x orstronger), but a dissecting microscopeallows more detailed observation. Workthe lichen using a stiff, single-edged razorblade and a pair of fine forceps or tweez-ers. Never apply a reagent directly to anherbarium specimen; instead, break off atiny fragment that can later be discarded.Carefully record all colour changes, takingcare to record both positive and negativereactions (e.g., Cortex K+ yellow, C-, PD-;medulla K-, C-, PD+ yellow becomingorange).

Calcium hypochlorite (C): Thisreagent, chlorine bleach (e.g., Javex), canbe purchased from most grocery outlets.The reaction (a reddish or pinkish col-oration) is often fleeting, and must beobserved carefully. In some cases, a morevivid reaction can be obtained by using a“KC” test, in which K is followed by C.Calcium hypochlorite is unstable and

should be discarded (usually after two orthree months) when it no longer smellsstrongly of chlorine.

Nitric acid (H): This reagent is rarelyused, but is helpful in distinguishingbetween the foliose genera Neofuscelia andMelanelia. The expected reaction (inNeofuscelia) is a rapid darkening of theupper cortex, with a blue-green tinge. It isalso used to discriminate among certainspecies of Chaenothecopsis. Use at 50%concentration.

Potassium iodide (I): Iodine solutionsreact with a variety of starches. For exam-ple, when applied to the spore-producingapothecial layer (or hymenium) in thefoliose genera Fuscopannaria andPannaria, the tissues become blue, violet,or even bluish black (check under a lightmicroscope). This reagent also gives abluish or purplish reaction when appliedto the medulla of some Sphaerophorusspecies. The preferred formula is Lugol’siodine solution: 0.5 g iodine, 1.5 g potassi-um iodide, and 100 ml distilled water.

Potassium hydroxide (K): This is a10–35% solution of potassium hydroxidein water. The reagent can be purchased(in pellet form) from most drugstores.The usual colour reactions are: yellow,yellow changing to orange or red, and red.If tightly stoppered, a solution of K willoften remain active for up to six months.

Potassium hydroxide / calciumhypochlorite (KC): In this test, K isapplied first, and then C. A positive reac-tion yields instantaneous pinks or redsthat often fade quickly. Useful in the iden-tification of a wide range of lichens.

Calcium hypochlorite / potassiumhydroxide (CK): This is a seldom-usedtest, in which C is applied first, followedby K. A positive reaction gives a deep yel-low or orangish colour, and is useful pri-marily in the recognition of barbatic acid(diagnostic, for example, in Usnea cerati-na), although salazinic acid also yields a

Making Use ofLichen Chemistry

3 The abbreviations used here for calcium hypochlorite (C), nitric acid (H), potassium iodide (I), and potassi-um hydroxide (K) should not be confused with the standard symbols for the chemical elements carbon,hydrogen, iodine, and potassium. Alternative abbreviations include HNO3 for H, and KOH for K.

INTRODUCTION

CK+ yellowish to orangish reaction.Paraphenylenediamine (PD): This

reagent is most safely used as Steiner’sStable PD Solution: 1 g PD crystals, 10 gsodium sulphite, 5 ml detergent (e.g.,Photo-flo), 100 ml distilled water. Analternative solution (preferred forStereocaulon) can also be prepared by dis-solving a few crystals of PD in two orthree drops of 70% ethyl alcohol (note:rubbing alcohol works well, but avoid theuse of isopropyl alcohol). The resultingsolution is highly unstable, and deterio-rates after only a few minutes. By contrast,Steiner’s Solution lasts a month or more,especially if stored in a dark bottle; itshould be discarded when it turns a darkpink. PD is suspected of being carcino-genic; it must be handled carefully, as it isabsorbed through the skin, and stainscloth, books, and specimens. Reactionsoften develop slowly (e.g., 30–60 seconds),and result in a yellow, orange, or red col-oration. The crystals can be difficult toobtain.2. Ultraviolet (UV) Lamps Ultravioletfluorescence provides an effective means

of detecting many lichen substances.Long-wave UV is preferred (i.e., 365 µm).The technique involves exposing themedulla of the specimen with a razorblade, and then examining it with a UV

lamp in a darkened room. A positive UV

reaction yields a distinct bluish, yellowish,or whitish incandescence. Because UV

light is damaging to the eyes, protectivegoggles should be worn when conductingthese tests. Avoid using UV lamps forextended periods, and never look directlyinto the lamp. Ultraviolet lamps can beobtained from scientific and geologicalsupply outlets.3. Thin-layer Chromatography (TLC)Thin-layer chromatography is moreexpensive and time consuming than spottests or UV tests, but is also more discrimi-nating. In fact, many chemical substancescan be detected in no other way (i.e.,without the use of even more sophisticat-ed techniques). The technique is not diffi-cult to learn, but instruction in themethod is beyond the scope of this manu-al. White and James (1985) provide a goodintroduction.

In this manual, common names are pro-posed for all fruticose macrolichens andmicrolichens known to occur in BritishColumbia.

Some of these names have been adopt-ed from Lichens of North America (Brodoet al. [2001]), though most are originalwith this publication. Names in parenthe-ses have been used by earlier authors—for example, Alvin (1977), Benton andUnderhill (1977), Bland (1971), Bolton(1960), Brodo (1988), Casselman (1993),Hawksworth et al. (1995), Johnson et al.(1995), Kershaw et al. (1998), McCune andGeiser (1997), McGrath (1977), MacKin-non et al. (1992), Nearing (1947), Parish etal. (1996), Perez-Llano (1944), Pojar andMacKinnon (1994), Richardson (1975),Smith (1921), and Vitt et al. (1988)—butfor various reasons are not accepted here.

Most of the accepted common names

record observable attributes of the speciesand genera to which they apply, thoughother names are intentionally fanciful. Inmost cases, the same “root” name appliesto all members of a given genus (e.g., allspecies of Bryoria bear the name “horse-hair”), but in some cases this name isapplied also to the species of other similargenera.

Common names are perhaps most satis-factorily viewed as vehicles of communica-tion for those who are unwilling to usescientific names. Scientific names are in-tended to be universal and stable; commonnames are by nature regional and highlyplastic. The common names introducedhere are intended primarily for use by thenaturalist community of British Columbia.While some may gain currency elsewhere inNorth America, alternative names willprobably be coined for many species.

A Note onCommon Names

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