 SPECIAL REPORT

 T HE THE LICHENS OF L CESOF ICHENS BRITISH COLUMBIA

1999 Illustrated Keys Part 2 — Fruticose Species

by Trevor Goward B RITISH C OLUMBIA

Ministry of Forests Research Program THE LICHENS OF BRITISH COLUMBIA

Illustrated Keys

Part 2 — Fruticose Species

by Trevor Goward

(Illustrations by Trevor Goward)

Ministry of Forests Research Program Canadian Cataloguing in Publication Data Goward, 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 – Geographic distribution. I. British Columbia. Ministry of Forests. Research Branch. II. Title. III. Series: Special report series (British Columbia Ministry of Forests) ; 9

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

Prepared by Trevor Goward Herbarium, Department of Botany University of British Columbia Vancouver, BC   Mailing Address: Edgewood Blue, Box 131 Clearwater, BC   for B.C. Ministry of Forests Research Branch 712 Yates Street Victoria, BC  

© 1999 Province of British Columbia

Copies of this report may be obtained, depending upon supply, from: Crown Publications 521 Fort Street Victoria, BC   (250) 386-4636 www.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

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES ii ACKNOWLEDGEMENTS

This manual has been four years in the Rydholm tested the keys; Irwin Brodo, making. During that time, it has benefit- François Lutzoni, Eric B. Peterson, and ted from the kindness of many people. In Bruce Ryan provided unpublished data on particular, I wish to thank five colleagues lichen distribution in the western United —Teuvo Ahti, Irwin Brodo, Stephen States; Mikhail Zhurbenko augmented my Clayden, Pekka Halonen, and Hiroyuki understanding of lichen distribution in Kashiwadani—for entering into co- eastern Eurasia; and André Arsenault, authorship on treatments of Cladina, Robert Bringhurst, Irwin Brodo, Dennis Cladonia, Stereocaulon, Usnea, and Fafard, Bruce McCune, Don McKay, Ramalina. I also wish to thank the follow- Roger Rosentreter, Evelyn Schimmel, ing specialists for reviewing earlier drafts Steve and Sylvia Sharnoff, and Jan Zwicky of other critical genus accounts: Irwin all helped to pry open the Pandora’s box Brodo (Alectoria, Bryoria), Håkon Holien of common names. To all of the above, I (Calicium, Chaenotheca, Microcalicium, extend my deepest gratitude. Sclerophora), Scott Kroken (Letharia), I also express thanks to: Lyle Ottenbreit Scott LaGreca (Ramalina), Leif Tibell and Dan Bashaw for bringing the maps (Chaenothecopsis), Tor Tønsberg (Gyali- and figures to camera-ready; Susan deopsis, Microlychnus, Szczawinskia), Bannerman, Bruce Maricle, Vivian Miao, and Mats Wedin (Bunodophoron, Anna Roberts, and Steve Smith for Sphaerophorus). In addition, several English edit and proofreading; Donna lichenologists have willingly shared their Lindenberg for typesetting; Rick Scharf for taxonomic expertise: Teuvo Ahti, Othmar carrying the manuscript through to publi- Breuss, Irwin Brodo, Aino Henssen, Per cation; and Paul Nystedt for cheerfully Magnus Jørgensen, Scott Kroken, Scott juggling many balls in the air at the same LaGreca, Bruce McCune, Juoko Rikkinen, time. Roger Rosentreter, Steve Selva, Leif Tibell, Evelyn Hamilton, of the British Tor Tønsberg, and Mats Wedin. Columbia Ministry of Forests, deserves In addition: Olivia Lee, of the University special thanks for supporting this project of British Columbia, packaged, labelled, in many ways throughout its gestation— sorted, and forwarded lichen specimens not least by funding it through the British by the thousands; Dan Burgess, Nathalie Columbia Forest Renewal Fund (FRBC). Djan-Chékar, David Miège, Carla For supplementary funding through the Rydholm, and Patrick Williston helped British Columbia Conservation Data to prepare the distribution maps; Irwin Centre (CDC), I also thank Andrew Brodo, Stephen Clayden, Bernard Harcombe. Goffinet, and Tor Tønsberg subjected sev- Finally, this book is fondly dedicated to eral specimens to thin-layer chromatogra- two eminent lichenologists: Teuvo Ahti, of phy; Stephen Clayden, Bernard Goffinet, the University of Helsinki, and Irwin Stuart Harris, Olivia Lee, Vivian Miao, Brodo, of the Canadian Museum of Roger Rosentreter, Wilf Schofield, and Nature. Many thanks, my friends, for all Émannuël Sérusiaux responded to urgent you have done. pleas for literature; Tyler Innes and Carla

ACKNOWLEDGEMENTS iii CONTENTS

Acknowledgements ...... iii Introduction ...... 1 Interpreting the Genus and Species Accounts ...... 3 Understanding Biogeoclimatic Zonation ...... 5 Identifying Lichens ...... 10 Making Use of Lichen Chemistry ...... 20 A Note on Common Names ...... 21 Keys to Genera of Fruticose Macrolichens and Microlichens ...... 22 Key to Lichen Photobionts ...... 23 Key to Lichen Growth Forms ...... 27 Key A: Fruticose Macrolichens ...... 29 Key B: “Calicioid” Lichens (and others)...... 36 Key C: Fruticose Microlichens (and others)...... 39 Keys to Species of Fruticose Macrolichens and Microlichens ...... 45 Acroscyphus ...... 45 Agrestia ...... 46 Alectoria ...... 47 Baeomyces ...... 51 Bryocaulon ...... 53 Bryoria ...... 54 Bunodophoron ...... 69 Calicium ...... 70 Chaenotheca ...... 76 Chaenothecopsis ...... 89 Cladina ...... 101 Cladonia ...... 107 Coelocaulon ...... 164 Cornicularia ...... 165 Cystocoleus ...... 166 Dactylina ...... 167 Dibaeis ...... 170 Ephebe ...... 171 Evernia ...... 173 Gyalideopsis ...... 176 Leciophysma ...... 178 Lempholemma ...... 179 Leprocaulon ...... 181 Letharia ...... 183 Lichinella ...... 185 Lichinodium ...... 186 Loxosporopsis ...... 189 Microcalicium...... 190 Microlychnus ...... 192 Mycocalicium ...... 193 Niebla ...... 194 Nodobryoria ...... 195 Phaeocalicium ...... 197

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES iv Pilophorus ...... 199 Polychidium ...... 202 Pseudephebe ...... 204 Racodium ...... 206 Ramalina ...... 207 Sclerophora ...... 215 Siphula ...... 217 Sphaerophorus ...... 218 Spilonema ...... 222 Stenocybe ...... 225 Stereocaulon ...... 227 Sticta ...... 240 Synalissa ...... 241 Szczawinskia ...... 242 Teloschistes ...... 243 Thamnolia ...... 244 Thermutis ...... 245 Tholurna ...... 246 Usnea ...... 247 Zahlbrucknerella ...... 265 Unknown 1 ...... 266 Unknown 2 ...... 267 Unknown 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 ...... 4 2 Summary information on the biogeoclimatic zones of British Columbia...... 7

FIGURES 1 First- and second-order lichen floristic studies in British Columbia to 1998...... 1 2 “Life zones” of British Columbia ...... 4 3 Biogeoclimatic zones of British Columbia ...... 6 4 Thallus stratified/heteromerous (cross-section)...... 11 5 Thallus nonstratified/homoiomerous (cross-section, in part)...... 11 6 Common photobionts (free-growing/unlichenized forms)...... 12 7 Organs of attachment (cross-section)...... 13 8 Branching and degrees of attachment...... 13 9 Growth forms (cross-section and surface view, in part)...... 14 10 Surface (and other) details: foliose ...... 15 11 Surface (and other) details: fruticose ...... 16 12 Spores and conidia ...... 17 13 Sexual and asexual reproductive structures ...... 18 14 Vegetative reproductive structures ...... 19

CONTENTS v

INTRODUCTION

Our knowledge of lichen floristics in of Nature and Natural Resources (IUCN) British Columbia has advanced rapidly in “global redlist of lichens” (Thor 1996: recent years. Only three decades ago, the www.dha.slu.se/guest/global.htm). Several known British Columbia lichen flora other species may already be at risk of stood at 569 species (Otto and Ahti 1967). extirpation in the province (Goward Today, by contrast, it stands at approxi- 1996). Reflecting these concerns, the mately 1300 species (Noble et al. 1987; British Columbia Conservation Data Goward, unpublished). Notwithstanding Centre has recently initiated a preliminary this impressive figure, a comprehensive tracking list of the province’s “red-listed” inventory of the province’s lichen flora is and “blue-listed” lichens. still a long way off. In support of this Few portions of the province have claim, consider that more than 20 lichen received serious attention from lichenolo- species, on average, are added to the gists. To date, comprehensive lichen stud- provincial lichen flora each year! ies have been carried out only on A significant number of lichen species southeast Vancouver Island (Noble 1982) warrant formal designation as “rare” or and the Queen Charlotte Islands (Brodo “infrequent” in British Columbia. To 1995; Brodo and Ahti 1996; Brodo and date, four such species carry official Santesson 1997; Brodo and Wirth 1998). endangerment status in Canada (Goward The lichen flora of Wells Gray Park is also et al. 1998), while one species appears on relatively well documented (Goward and the International Union for Conservation Ahti 1992; Goward, unpublished).

FIRST ORDER 1. Brodo (in prep.) 2. Goward, Ahti 6 (1992) 3. Noble (1982) (Bird, Bird 6 1973) 6 (Ryan 1991) 6 SECOND ORDER 4. Benton, Brodo, Richardson (1977) 5. Brodo (unpublished) 6 6. Goward (unpublished) 7. Goward, Schofield (1983) 8. Ohlsson (unpublished) 9. Otto (unpublished) 10. Thomson, Ahti (1994)

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

INTRODUCTION  Important collections from other regions examination under a light microscope; do exist (Figure 1), but have not been fortunately, most such species are small published, and are widely scattered. and inconspicuous, and are unlikely to be Lichen study in British Columbia has encountered by the beginner. traditionally been hampered by a lack of The genus and species concepts adopted comprehensive keys to the species. here are often pragmatic. As a rule, they Recently, however, Goward et al. (1994) give priority to “intuitive” morphological published illustrated keys to 327 species of groupings that do not always accord with foliose and squamulose lichens. The pres- the latest findings of anatomical and molec- ent manual is a companion volume to ular research. This approach proceeds from that work; it provides illustrated keys to the assumption that laboratory research 309 species of fruticose lichens (and allied will continue to uncover evolutionary rela- fungi) that are known from or expected to tionships not readily perceived in the field. occur in British Columbia. Looking ahead, two parallel approaches to This manual adopts a broad interpreta- lichen taxonomy can be expected to evolve: tion of the fruticose life form. Included one that emphasizes taxonomic stability, here are all lichen genera—both macro- and is suited to the requirements of lichen lichens and microlichens—in which a floristics and field ecology; and one that majority of species bear stalks or branches emphasizes phylogenetic relatedness, and is that are roughly circular in cross-section. suited to continuing molecular, chemical, Although fruticose microlichens are tradi- and ultrastructural studies. Although this tionally regarded as crustose, they are manual follows the first of these approach- included here owing to their dominant es, alternative genus and species concepts stalked sexual or asexual reproductive are given in the synonymy under the structures. Examples of fruticose micro- accepted species. lichens include Calicium, Chaenotheca, Accompanying the keys are 320 line Gyalideopsis, and Microlychnus. drawings intended to convey species con- Two primary objectives have guided cepts based on typical material. The draw- the preparation of this manual: ings emphasize specific characters 1. to briefly summarize the ecology, dis- expressed in the adjacent key, and are not tribution, and frequency status of all intended to depict the entire lichen. fruticose lichens known to occur in Illustrations of whole lichens can be found British Columbia. in various popular and semi-popular ref- 2. to stimulate lichenological research by erences, including Hale (1979), Kershaw making these lichens accessible to a et al. (1998), McCune and Geiser (1997), broad audience. McCune and Goward (1995), MacKinnon In keeping with the presumed needs et al. (1992), Parish et al. (1996), Pojar and and resources of ecologists, biologists, MacKinnon (1994), Thomson (1984, 1997), naturalists, teachers, and other beginning and Vitt et al. (1988). students of lichens, this manual emphasizes It is beyond the scope of this manual to morphological characters over chemical provide a comprehensive summary of and spore characters; it also avoids techni- lichen biology (see instead: Hale 1983; cal terms as far as possible.1 It must be Hawksworth and Hill 1984; Lawrey 1984; acknowledged, however, that some species Nash 1996). Effective identification does, cannot be reliably identified without however, require a basic understanding of recourse to thin-layer chromatography or lichen morphology and chemistry.

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

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES  Accordingly, the reader is invited to con- every effort has been made to make the sult “Identifying Lichens” (page 10) prior keys as usable as possible, numerous errors to using the keys. and oversights doubtless remain. The This manual represents a first attempt reader is invited to bring these to the to provide comprehensive keys to the fru- author’s attention for the benefit of future ticose lichens of British Columbia. Though students of British Columbia’s lichens.2

Interpreting the For convenience, lichens can be arranged Species and Author Citation: Except in Genus and Species in several different growth forms, includ- cases of recent taxonomic or nomenclat- Accounts ing crustose, squamulose, foliose, and fru- ural revision, species names and author ticose (see “Identifying Lichens,” page 10). citations follow Esslinger and Egan (1995). Because, however, these growth forms are Synonyms: As a rule, only synonyms in units of convenience, not biology, they do recent or widespread use are given. not always offer a perfect “fit” with the Distribution Maps: The map number genera they are supposed to circumscribe. appearing to the right of some species is For example, while all species of Bryoria keyed to Appendix 1, in which distribu- are fruticose, the genus Cladonia contains tion maps are provided for species judged both squamulose species and fruticose to be rare or infrequent in the province. species. For convenience, this manual Common Names: Common names are incorporates all lichen genera known to adopted, adapted, or introduced for all occur in British Columbia in which a lichen species included in this manual. majority of species can be described as Alternative common names are given in fruticose. In a few instances, fruticose parentheses (...). See also “A Note on species belonging to essentially nonfruti- Common Names,” page 21. cose genera are also included (in paren- Habitat/Range: Information is provided theses) in the keys, but are not discussed on lichen frequency status, common sub- in the species accounts. Species appearing strates, site characteristics, provincial “life in square brackets [...] are expected to zone” distribution, distribution in the occur in British Columbia, but have yet to northern hemisphere, and cordilleran dis- be reliably recorded. tribution. The body of the manual is arranged 1.Frequency status is given using the fol- alphabetically, first by genus and then by lowing terms: rare, infrequent, fre- species within each genus. quent, common. The genus accounts include: 2.Substrates include acid/base-rich/ 1. the scientific name, mossy/seasonally inundated rock, 2. a common name, coniferous/deciduous trees or shrubs, 3. a short description of the genus, with and soil, moss, duff, bark, or wood. diagnostic characters placed in bold 3.Site characteristics are expressed as: italic type, exposed/open/ sheltered/shady/old- 4. pertinent references, growth forests, steppe, depressions, or 5. the derivation of the common name, outcrops, at lower/middle/upper/alpine 6. notes on global status, distribution, elevations. taxonomy, chemistry, and points of 4.Provincial “life zone” distributionsare distinction with similar genera. given according to the terms listed in The species accounts are more com- columns I and II of Table 1; see also plex, and are organized under the follow- Figure 2. Biogeoclimatic units (column ing headings: III of Table 1) are occasionally used,

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

INTRODUCTION  BOREAL

INTERMONTANE

HYPER- MARITIME MARITIME

kilometres

FIGURE 2 “Life zones” of British Columbia.

TABLE 1 Distributional units and their definition

General Life Biogeoclimatic Conrad’s Index of Range Zone Equivalenta Continentality (I) (II) (III) (IV)

COAST Hypermaritime CWH wh and vhb < 8 Maritime 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 > 40 SUBALPINE Throughout MH, ESSF various ALPINE Throughout AT various WIDESPREAD 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.

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES  and are mapped in Figure 3; see also UT (Utah), WA (Washington), or WY “Understanding Biogeoclimatic (Wyoming); and AB (Alberta). While Zonation” (below). The continentality most state records appearing in this units in column IV are based on manual are based on published reports, Conrad’s Index of Continentality a few unpublished records are also (Conrad 1946), and are included to given; these appear in parentheses. enable ecoclimatic comparisons with Reactions: In most cases, only positive other regions of the world (for further reactions are given, based on commonly details, see Goward and Ahti 1992). used chemical reagents and ultraviolet 5.Distribution in the northern hemi- light; negative reactions are generally sphere is expressed relative to western omitted. For further details see “Making North America. The following distribu- Use of Lichen Chemistry,” page 20. tional units are used: Contents: In most cases, only dominant western N Am lichen substances are listed. Substances western N Am - eastern N Am given in parentheses are “accessory,” that western N Am - western Eurasia is, they do not occur in all specimens. western N Am - eastern Eurasia Variability: Some lichen species vary incompletely circumpolar (= any three greatly with habitat, whereas others are of the above distributional units) more uniform. To assist in identification, circumpolar each species has been assigned a variability 6.Cordilleran distribution is summarized rating of “low,” “medium,” or “high.” using the following geographic units: N Species rated as “high” do not necessarily to sAK (Alaska: Pacific coast only), AK conform with all characters given in the (Alaska: boreal and arctic regions keys. only), YU (Yukon), or wNT Notes: Included here are comments on (Northwest Territories: west of the taxonomic problems, points of distinction Mackenzie River only); and S to AZ with similar species, chemistry, and keys (Arizona), CA (California), CO to subspecies and varieties. In general, (Colorado), ID (Idaho), MT detailed notes are reserved for taxonomi- (Montana), MX (Mexico), NM (New cally difficult genera. Mexico), NV (Nevada), OR (Oregon),

Understanding British Columbia is a highly diverse prov- variants of the biogeoclimatic system are Biogeoclimatic ince in which hundreds of ecosystems can referred to as biogeoclimatic units. Each Zonation be recognized. Maintaining these in the unit is characterized by a unique set of cli- face of increasing pressure for resource matic variables, and supports—and is for development represents an enormous practical purposes defined by—a unique challenge—and involves, as a first step, vegetation. In biogeoclimatic ecosystem classifying the province’s ecosystems in classification, the defining vegetation for detail. each unit occurs on moderately well- In recent years, researchers with the drained sites. Such sites are said to be B.C. Ministry of Forests have described “zonal.” medium-scale ecosystems according to the The most encompassing of the biogeo- principles of biogeoclimatic ecosystem climatic units is the biogeoclimatic zone. classification (Pojar et al. 1987). They have Fourteen biogeoclimatic zones are recog- also arranged these ecosystems into a hier- nized for British Columbia and many of archical system of biogeoclimatic zones, these are used here to describe lichen dis- subzones, and variants. tribution. They are briefly characterized in Collectively, the zones, subzones, and Table 2 and mapped in Figure 3. For a

INTRODUCTION    Biogeoclimatic zones ofBritishColumbia.

Research Branch, Ministry of Forests, Victoria, BC more detailed summary, see Ecosystems of made mostly at the zonal level, though British Columbia (Meidinger and Pojar two biogeoclimatic subzones have also 1991). been used: the Wet Hypermaritime (wh) Lichen distribution may also be and Very Wet Hypermaritime (vh) sub- expressed using more generalized classifi- zones of the Coastal Western Hemlock cation systems such as the “life zone sys- Zone (CWH). These subzones occur in the tem” (see Figure 2) and “general range hypermaritime or outer coastal areas of system” adopted here. These systems are British Columbia (see Figure 2). See Table compared with their biogeoclimatic coun- 2 for the full names of other biogeoclim- terparts in Table 1. The comparison is 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.–April Zone Code Zonal vegetation Zonal soils temp. range >5°C<0°C ppt (mm) ppt (mm)

Alpine AT Cassiope spp., Phyllodoce spp., Regosols, -11.1–9.5 427 1763 287 469 Tundra Luetkea pectinata, Loiseleuria Humic procumbens, Dryas spp., Salix Regosols, spp., Silene acaulis, Poa spp., Brunisols, Festuca spp., Carex spp., Humo-Ferric Cetraria spp., Stereocaulon Podzols spp., Polytrichum piliferum

Boreal BWBS White spruce, lodgepole pine, Gray Luvisols, -24.5–16.6 709–1268 1692–2742 145–305 182–198 White black spruce, Rosa acicularis, Dystric and and Viburnum edule, Mertensia Eutric Black paniculata, Pyrola asarifolia, Brunisols Spruce Cornus canadensis, Vaccinium vitis-idaea, Ptilium crista-cas- trensis, Pleurozium schreberi

Bunch- BG Agropyron spicatum, Artemisia Brown and -10.8–22.4 1771–2516 230–878 98–175 108–208 grass tridentata, Artemisia frigida, Dark Brown Poa sandbergii, Koeleria Chernozems macrantha, Festuca scabrella, Festuca idahoensis, Chrysothamnus nauseous

Coastal CDF Douglas-fir, grand fir, bigleaf Dystric 1.8–18.0 1794–2121 9–43 107–238 540–1107 Douglas- maple, western flowering dog- Brunisols fir wood, Holodiscus discolor, Gaultheria shallon, Mahonia nervosa, Rosa gymnocarpa, Symphoricarpos albus, Trientalis latifolia, Rubus ursi- nus, Pteridium aquilinum, Kindbergia oregana, Rhytidia- delphus triquetrus

INTRODUCTION  TABLE 2 Continued

Selected climatic characteristicsa Monthly °days °days May–Sept. Oct.–April Zone Code Zonal vegetation Zonal soils temp. range >5°C<0°C ppt (mm) ppt (mm)

Coastal CWH Western hemlock, amabilis fir, Ferro-Humic -6.6–18.7 1059–2205 5–493 159–1162 695–3225 Western Sitka spruce, yellow-cedar, and Humo- Hemlock Vaccinium alaskaense, Ferric Podzols Vaccinium parvifolium, Menziesia ferruginea, Gaultheria shallon, Polystichum munitum, Pteridium aquil- inum, Blechnum spicant, Clintonia uniflora, Rhytidiadelphus loreus, Hylocomium splendens

Engel- ESSF Subalpine fir, Engelmann Humo-Ferric -10.9–13.3 629–801 879–1189 205–425 271–1597 mann spruce, Rhododendron albiflo- Podzols Spruce– rum, Menziesia ferruginea, Subalpine Vaccinium (membranaceum, Fir ovalifolium, scoparium), Rubus pedatus, Gymnocarpium dry- opteris, Tiarella unifoliata, Valeriana sitchensis, Orthilia secunda, Streptopus roseus, Veratrum viride, Barbilophozia lycopodioides, Pleurozium schreberi, Rhytidiopsis robusta

Interior ICH Western hemlock, western red- Humo-Ferric -10.7–20.8 1267–2140 238–820 200–439 294–1098 Cedar– cedar, hybrid white spruce, Podzols, Gray Hemlock Douglas-fir, subalpine fir, Luvisols, and Vaccinium ovalifolium, Dystric Oplopanax horridus, Vaccinium Brunisols membranaceum, 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

Interior IDF Douglas-fir, lodgepole pine, Gray Luvisols, -13.1–21.3 903–2366 235–1260 107–291 149 –1022 Douglas- ponderosa pine, Spiraea betuli- Eutric and fir folia, Amelanchier alnifolia, Dystric Juniperus communis, Brunisols Symphoricarpos albus, Mahonia aquifolium, Paxistima myrsinites, Calamagrostis rubescens, Arctostaphylos uva- ursi, Agropyron spicatum, Pleurozium schreberi

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES  TABLE 2 Continued

Selected climatic characteristicsa Monthly °days °days May–Sept. Oct.–April Zone Code Zonal vegetation Zonal soils temp. range >5°C<0°C ppt (mm) ppt (mm)

Montane MS Hybrid white spruce, sub- Dystric -12.5–17.4 891–1310 847–890 158–252 223–469 Spruce alpine fir, lodgepole pine, Brunisols and Douglas-fir, Vaccinium scopari- Humo-Ferric um, Lonicera utahensis, Podzols Shepherdia canadensis, Paxistima myrsinites, Vaccinium membranaceum, Alnus viridis, Linnaea borealis, Empetrum nigrum, Calamagrostis rubescens, Pleurozium schreberi

Mountain MH Mountain hemlock, amabilis Ferro-Humic -2.3–13.2 919–933 307–352 694–707 1857–2260 Hemlock fir, yellow-cedar, Vaccinium Podzols and (ovalifolium, membranaceum, Folisols alaskaense), Menziesia ferrug- inea, Rhododendron albiflorum, Rubus pedatus, Phyllodoce empetriformis, Rhytidiopsis robusta, Rhytidiadelphus loreus, Hylocomium splendens

Ponderosa PP Ponderosa pine, Agropyron spi- Eutric and -8.6–21.6 1505–2442 258–861 86–270 170–334 Pine catum, Balsamorhiza sagittata, Dystric Festuca (saximontana, ida- Brunisols hoensis), Koeleria macrantha, Lithosperum ruderale, Achillea millefolium

Spruce– SWB White spruce, subalpine fir, Eutric or -19.2–14.0 534–933 2036–2298 275–280 179–424 Willow– Salix glauca, Betula glandulosa, Dystric Birch Potentilla fruticosa, Shepherdia Brunisols, canadensis, Festuca altaica, Humo-Ferric Lupinus arcticus, Pedicularis Podzols labradorica, Epilobium angusti- folium, Empetrum nigrum, Vaccinium (vitis-idaea, caespi- tosum), Hylocomium splendens, Cladina spp., Nephroma arcticum

Sub- SBPS Lodgepole pine, white spruce, Gray Luvisols -13.8–14.3 697–1044 1140–1405 243–300 218–222 Boreal Shepherdia canadensis, Spiraea and Dystric Pine– betulifolia, Rosa acicularis, Brunisols Spruce Calamagrostis rubescens, Arctostaphylos uva-ursi, Vaccinium caespitosum, Linnaea borealis, Pleurozium schreberi, Peltigera spp., Cladina spp.

INTRODUCTION  TABLE 2 Concluded

Selected climatic characteristicsa Monthly °days °days May–Sept. Oct.–April Zone Code Zonal vegetation Zonal soils temp. range >5°C<0°C ppt (mm) ppt (mm)

Sub- SBS Hybrid white spruce, sub- Gray Luvisols -14.6–16.9 884–1510 792–1369 189–353 250–1383 Boreal alpine fir, lodgepole pine, and Dystric Spruce Vaccinium membranaceum, Brunisols, Rubus parviflorus, Viburnum Humo-Ferric edule, Lonicera involucrata, Podzols Spiraea betulifolia, Rosa acicu- laris, Aralia nudicaulis, Cornus canadensis, Linnaea borealis, Arnica cordifolia, Clintonia uniflora, Aster conspicuus, Osmorhiza chilensis, Oryzopsis asperifolia, Smilacina racemosa, Gymnocarpium dryopteris, Pleurozium schreberi, Ptilium crista-castrensis, Hylocomium splendens, Dicranum polyse- tum, Rhytidiadelphus tri- quetrus, Peltigera spp. a Selected climatic characteristics summarized from Atmospheric Environment Service (AES) long-term stations. Prepared by D. Meidinger.

Identifying Lichens The vast majority of lichens are classified their unlichenized relatives. Rather, they as Ascomycetes (cup fungi), and hence are form a composite “plant,” or thallus related to morels and elf saddles. While (Figures 4–5). Under the microscope, a most fungi, however, draw their nourish- typical lichen thallus resembles a kind of ment from sources external to themselves sandwich in which the fungal partner (e.g., decaying leaves or logs), lichen fungi (mycobiont) and the “algal” partner “cultivate” their foodstuff among the fun- (photobiont) are stratified in distinct lay- gal threads of which they themselves are ers (Figure 4). In many conspicuous composed. This foodstuff consists of tiny, lichen species, four such layers can be dis- photosynthesizing algal or cyanobacterial cerned: a protective rind or upper cortex cells, or both. Lichens can therefore be (Figure 4a); an “algal” or photobiont layer viewed as living greenhouses supported (Figure 4b); a pale, usually whitish region largely by carbohydrates derived from the of loosely interwoven fungal threads, photosynthetic “crops” growing within called the medulla (Figure 4c); and anoth- them. This in part accounts for the er protective covering or lower cortex exposed lifestyle adopted by most lichens: (Figure 4d). whereas a majority of fungi pass their lives As already mentioned, the photobionts (except when fruiting) within the things in nearly all lichens consist of green algae they feed on, lichens colonize the surfaces or cyanobacteria. When exposed by a of rocks, trees, logs, duff, and soil. razor blade and viewed under a dissecting When a fungus enters into a stable, microscope, most algae are easily recog- enduring relationship with a microscopic nized by their characteristic single-celled alga or cyanobacterium, both partners are habit, as well as by their bright grass-green said to be lichenized. In general appear- colour (but yellowish green in Trente- ance, most lichenized fungi, algae, and pohlia). Cyanobacteria are much more cyanobacteria do not closely resemble variable in form—single-celled, cluster-

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES  (a) (b)

(c) (e)

(d)

FIGURE 4 Thallus stratified/heteromerous (cross-section): (a) upper cortex; (b) algal or cyanobacterial layer / photobiont layer; (c) medulla; (d) lower cortex; and (e) a cephalodium. celled, or strandlike—but they are never Polychidium (Figure 5b). Strandlike rhi- grass-green, although at least one genus zoids (Figure 5c) can also occur in some can be yellowish green. In some species cyanobacterial species. dominated by a green algal photobiont, Many different groups of algae and cy- cyanobacteria are also present as scattered, anobacteria have entered into association localized colonies called cephalodia with lichen fungi. Common cyanobacteri- (Figure 4e). These can occur internally al groups include the Chroococcales or over the upper or lower surface (Figure 6a), Nostoc (Figure 6b), the (Figures 10d, 11l). Rivulariaceae (Figure 6c), Scytonema In some groups having a cyanobacterial (Figure 6d), and Stigonema (Figure 6e). photobiont, the photobiont cells are inter- The most common algal groups include mingled throughout with fungal threads, Stichococcus (Figure 6f), Trebouxia and and the thallus is uniformly dark both related genera (Figure 6g), and Trente- inside and out. Such lichens are said to be pohlia (Figure 6h). For keys to these and nonstratified (Figure 5); viewed from the other photobiont genera, see page 23. outside, they are typically brownish, Specialized holdfasts, or rhizines blackish, or bluish grey. They can assume (Figure 7a–e), occur in many species hav- a gelatinous consistency when wet, and ing a lower cortex. Rhizines anchor the are then popularly referred to as “gel lichen to the colonized surface or sub- lichens.” Most nonstratified lichens lack a strate, and can be simple (Figure 7a), cortex (Figure 5a), though a primitive cel- forking (Figure 7b), squarrose (Figure 7c), lular cortex is present in Leptogium and tufted (Figure 7d), or flocculent

(a) (b) (c)

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

INTRODUCTION  (a) (b) (c) (d)

(e)

(f) (h)

(g)

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.

(Figure 7e). In some foliose species (see (Figure 8d), semi-erect (Figure 8e), erect below), rhizines are replaced by a single (Figure 8f), and pendent (Figure 8g). thickened point of attachment, or umbili- Lichens have traditionally been divided cus (Figure 7f). In others, the rhizines are into three growth forms (crustose, foliose, replaced by a dark, woolly hypothallus and fruticose: see below), though other (Figure 7g) that can sometimes extend classifications are possible. The one adopt- beyond the margins of the lichen. ed here recognizes seven growth forms: Rhizine-like structures that occur along 1. Dust/leprose lichens (Figure 9a) lack the margins of leaflike lobes are called both an upper and lower cortex, the cilia (Figure 10i). lower surface being attached directly to Two broad categories of branching are the substrate, and the upper surface recognized in this manual. Branching is bearing a continuous covering of pow- said to be even when the arms of each dery or granular soredia. branch pair are predominantly equal 2. Crust/crustose lichens (Figure 9b) are (Figure 8a), and uneven when they are not also attached directly to the substrate, equal (Figure 8b). Similarly, five degrees but have a hard, protective upper cor- of attachment can be distinguished: tex; viewed from above, they often appressed (Figure 8c), decumbent resemble paint stains. Some crust

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES  (a) (b) (c) (d) (e) (f) (g)

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.

(b) (c)

(a)

(d)

(e) (f) (g)

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

INTRODUCTION  lichens give rise to elongate isidia (see right, unbranched, or sometimes spar- below), while others bear stalked fruit- sely branched stems; when hollow, they ing structures termed podetia and are usually referred to as podetia; when pseudopodetia; such species can be solid, they are called pseudopodetia. classified as club lichens or shrub 6. Shrub/fruticose lichens (Figure 9f) lichens (see below). Other crust lichens resemble club lichens in having some- intergrade with scale and leaf lichens what thickened stems that are more or (see below). less circular in cross-section. Here, 3. Scale/squamulose lichens (Figure 9c) however, the stems are strongly are similar to dust and crust lichens in branched. Occasionally the stems arise lacking a lower cortex (and rhizines). from a basal crust or basal scales. When The thallus, however, consists of small, hollow, they are generally termed pode- often partly raised, and usually overlap- tia; whereas, when solid, they are again ping/imbricate scales/squamules called pseudopodetia. Shrub lichens (Figure 9c), the lower surface of which vary from decumbent to semi-erect or is often white and cottony (check under erect. hand lens). In Cladonia, the squamules 7. Hair/fruticose lichens (Figure 9g) differ often give rise to hollow, stalked fruit- from shrub lichens in having much ing structures called podetia; see also finer, and proportionately much longer, club and shrub lichens, below. branches. In habit, hair lichens are 4. Leaf/foliose lichens (Figure 9d) more or semi-erect or pendent. less resemble leaves—at least in the The upper cortex of most lichens is sense that their thalli consist of flat- smooth and naked. However, in some tened lobes that typically have an upper species it can be minutely roughened and lower cortex. The lobes can be nar- (i.e., scabrid), or covered in pustules row or broad, and short or elongate. (Figure 10a) or a fine network of ridges (a The degree of attachment to the sub- reticulum: Figure 10b). Other species bear strate varies from appressed through a thin whitish frosting (or pruina), while semi-erect or even unattached. This is others still have a fine nap of tiny, erect, the only growth form in which well- or appressed glasslike hairs (or tomentum: developed rhizines occur. Figure 10c). A woolly tomentum is also 5. Club/fruticose lichens (Figure 9e) are at sometimes present over the lower surface. least partly round in cross-section; they The lower surface of most lichens is have no “true” lower surface and there- also smooth, though raised veins (Figure fore no lower cortex and rhizines. Most 10h) occur in some foliose species. In oth- club lichens consist of thickened, up- ers, the lower surface can be sparsely

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

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.

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES  speckled with tiny pits through which clues to identification. In some species, for the medulla is exposed. When rimmed example, the cortex is pocked by broad and craterlike, these pits are termed depressions (or foveoles: Figure 11a), while cyphellae (Figure 10f); otherwise they are in others it bears conspicuous longitudi- called pseudocyphellae (Figure 10g). In nal striations (Figure 11b). Yet other some lichens, pseudocyphellae occur over species carry numerous tile-like areoles the upper cortex, and must then be care- (Figure 11c) which may or may not be fully distinguished from maculae (Figure peltate (i.e., attached by a short central 10e): pale areas of the upper surface in stalk: Figure 11d). The presence of tiny, which the cortex is unbroken. Fruticose scale-like microsquamules (Figure 11f) is lichens can also produce pseudocyphellae. diagnostic for some Cladonia species, Surface details can provide important whereas some species of Usnea bear

(a) (b) (c)

(d)

(e) (f)

(g) (h) (i)

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(←).

INTRODUCTION  numerous goosefleshlike papillae (Figure vegetatively (in which case the fungus and 11g), or copious short side branches (or alga/cyanobacterium are dispersed togeth- fibrils: Figure 11h). Similarly, most er, as a functional unit), and by sexual Stereocaulon species support a dense means (in which case only the spores of “foliage” of phyllocladia (Figure 11j), the fungus are dispersed: see below). while small spine-like spinules (Figure 11i) Vegetative reproduction sometimes occurs occur in some species of Bryoria. as a result of mechanical fragmentation As a group, lichens reproduce both (wear and tear), but more often it involves

(a) (b) (c)

(d)

(g)

(e) (f)

(h)

(j) (k) (i) (l)

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(←).

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES  transport (as by birds) of specialized out- contain both a fungus and an alga/cyano- growths called soredia and isidia. Soredia bacterium. They can be granular/globose (Figure 14a–d) are masses of soft, powdery (Figure 14e), barrel-shaped (Figure 14f), granules that have erupted through cracks fingerlike/cylindrical (Figure 14g), coral- and other areas of weakness in the cortex. loid (Figure 14h), or scale-like (Figure 14i). When very fine (i.e., resembling talcum The primary function of sexual fruiting powder), they are said to be powdery/fari- bodies is to produce sexual spores (Figure nose (Figure 14b); otherwise they can be 12a–l). Sexual spores come in many described as more or less granular (Figure shapes, and can be threadlike/filiform 14c). Though commonly borne in well- (Figure 12a), spindle-shaped/fusiform delimited soralia (Figure 14a), soredia can (Figure 12b), ellipsoid (Figure 12c), club- also be broadcast over the surface; they like/clavate (Figure 12d), globose (Figure are then said to be diffuse (Figure 14d). By 12e), or peanut-shell-like (Figure 12f). contrast, isidia (Figure 14e–i) are tiny out- They can also be 1-celled (Figure 12g), or growths of the upper cortex; their hard- divided by a narrow cross-wall or septum ened outer surface is usually readily (Figure 12h, i) into 2-celled (Figure 12h) distinguished from the powdery appear- or multi-celled spores (Figure 12i). When ance of soredia. Similar to soredia, isidia the septum is very broad, the spores are

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

(k) (i) (h) (g) (j)

(l)

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 broad septum(←)); (k) submuriform; and (l) muriform.

INTRODUCTION  termed polarilocular (Figure 12j). Spores erally unstalked (Figure 13a–e), but in a having both crosswise and lengthwise few genera they are borne at the ends of septa are called submuriform (Figure 12k) long, brittle stalks as expanded “heads,” or, when well developed, muriform or capitula (Figure 13f, g). (Figure 12l). Apothecia contain both fertile and ster- For most species covered in this manu- ile tissues. The former comprise the cen- al, sexual fruiting bodies take the form of tral disc: a compact cluster of tiny vial-like apothecia (Figure 13a–g). These are small asci, each bearing its complement of one to saucerlike, buttonlike, or hemispherical several spores (Figure 13a–e). Surrounding structures usually readily observed over the disc is a sterile rim, or excipulum the thallus surface. Apothecia are gen- (Figure 13a, d, e). When photobiont cells

(a) (c) (e)

(d) (b)

(f)

(g) (h) (i)

(l) (j) (k) (m)

FIGURE 13 Sexual and asexual reproductive structures (surface view unless indicated): (a) unstalked apothecia (showing discs and 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: with nonthalline rim/excipulum and spore-containing asci(←)); (e) unstalked apothecium (cross-section: with thalline rim/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 (showing stalks and conidial heads(←)).

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES  are present in the excipulum, it is said to (Figure 13j, k) which, though also dotlike, be thalline (Figure 13e); when they are bear reproductive cells called conidia. lacking, it is nonthalline (Figure 13d). In Conidia come in many shapes but, unlike Calicium, Sphaerophorus, and related gen- sexual spores, they never develop in vial- era, the disc is replaced by a distinctly like asci. They are also usually much powdery mazaedium (Figure 13g) that smaller than sexual spores, often measur- readily smudges when rubbed. In other ing less than 4–5 µm long. In some genera, lichens, sexual fruiting bodies take the conidia are produced at the tips of stalked form of perithecia (Figure 13h, i): minute fruiting bodies called hyphophores flask-like, ascus-bearing structures that are (Figure 13l, m), while in others they are immersed in the thallus, and are visible borne directly over the cortex in well- from above as blackish or brownish dots. demarcated, black, sooty patches; these Perithecia must be carefully distin- are termed thalloconidia. guished from some forms of pycnidia

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

(f) (e) (i)

(h) (g)

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(←)).

INTRODUCTION  Making Use of Lichens produce a diverse array of chemi- should be discarded (usually after two or Lichen Chemistry cal substances. Reflecting this, lichen three months) when it no longer smells chemistry provides a useful tool for the strongly of chlorine. identification of many species. The pres- Nitric acid (H): This reagent is rarely ence of specific chemical substances can used, but is helpful in distinguishing be established through the use of spot between the foliose genera Neofuscelia and tests, ultraviolet lamps, and thin-layer Melanelia. The expected reaction (in chromatography. Neofuscelia) is a rapid darkening of the 1. Spot Tests A spot test is performed upper cortex, with a blue-green tinge. It is when a small quantity of liquid reagent is also used to discriminate among certain applied to one or more lichen tissues. This species of Chaenothecopsis. Use at 50% can be done using a fine brush or capillary concentration. pipette that has been drawn to a point Potassium iodide (I): Iodine solutions over a flame. The resulting colour change react with a variety of starches. For exam- (or lack of it) is often diagnostic for one ple, when applied to the spore-producing or more of several chemical substances apothecial layer (or hymenium) in the present in the lichen. Five reagents are foliose genera Fuscopannaria and called for in this manual: calcium Pannaria, the tissues become blue, violet, hypochlorite (C),3 nitric acid (H), potassi- or even bluish black (check under a light um iodide (I), potassium hydroxide (K), microscope). This reagent also gives a and paraphenyldiamine (PD). Of these, bluish or purplish reaction when applied only C, K, and PD are used routinely. All to the medulla of some Sphaerophorus of the reagents are toxic and should be species. The preferred formula is Lugol’s carefully stored in small, tightly sealed iodine solution: 0.5 g iodine, 1.5 g potassi- glass bottles. Chemical reactions can be um iodide, and 100 ml distilled water. observed using a hand lens (10x or Potassium hydroxide (K): This is a stronger), but a dissecting microscope 10–35% solution of potassium hydroxide allows more detailed observation. Work in water. The reagent can be purchased the lichen using a stiff, single-edged razor (in pellet form) from most drugstores. blade and a pair of fine forceps or tweez- The usual colour reactions are: yellow, ers. Never apply a reagent directly to an yellow changing to orange or red, and red. herbarium specimen; instead, break off a If tightly stoppered, a solution of K will tiny fragment that can later be discarded. often remain active for up to six months. Carefully record all colour changes, taking Potassium hydroxide / calcium care to record both positive and negative hypochlorite (KC): In this test, K is reactions (e.g., Cortex K+ yellow, C-, PD-; applied first, and then C. A positive reac- medulla K-, C-, PD+ yellow becoming tion yields instantaneous pinks or reds orange). that often fade quickly. Useful in the iden- Calcium hypochlorite (C): This tification of a wide range of lichens. reagent, chlorine bleach (e.g., Javex), can Calcium hypochlorite / potassium be purchased from most grocery outlets. hydroxide (CK): This is a seldom-used The reaction (a reddish or pinkish col- test, in which C is applied first, followed oration) is often fleeting, and must be by K. A positive reaction gives a deep yel- observed carefully. In some cases, a more low or orangish colour, and is useful pri- vivid reaction can be obtained by using a marily in the recognition of barbatic acid “KC” test, in which K is followed by C. (diagnostic, for example, in Usnea cerati- Calcium hypochlorite is unstable and na), although salazinic acid also yields a

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.

LICHENS OF BRITISH COLUMBIA, PART 2 – FRUTICOSE SPECIES  CK+ yellowish to orangish reaction. of detecting many lichen substances. Paraphenylenediamine (PD): This Long-wave UV is preferred (i.e., 365 µm). reagent is most safely used as Steiner’s The technique involves exposing the Stable PD Solution: 1 g PD crystals, 10 g medulla of the specimen with a razor sodium sulphite, 5 ml detergent (e.g., blade, and then examining it with a UV Photo-flo), 100 ml distilled water. An lamp in a darkened room. A positive UV alternative solution (preferred for reaction yields a distinct bluish, yellowish, Stereocaulon) can also be prepared by dis- or whitish incandescence. Because UV solving a few crystals of PD in two or light is damaging to the eyes, protective three drops of 70% ethyl alcohol (note: goggles should be worn when conducting rubbing alcohol works well, but avoid the these tests. Avoid using UV lamps for use of isopropyl alcohol). The resulting extended periods, and never look directly solution is highly unstable, and deterio- into the lamp. Ultraviolet lamps can be rates after only a few minutes. By contrast, obtained from scientific and geological Steiner’s Solution lasts a month or more, supply outlets. especially if stored in a dark bottle; it 3. Thin-layer Chromatography (TLC) should be discarded when it turns a dark Thin-layer chromatography is more pink. PD is suspected of being carcino- expensive and time consuming than spot genic; it must be handled carefully, as it is tests or UV tests, but is also more discrimi- absorbed through the skin, and stains nating. In fact, many chemical substances cloth, books, and specimens. Reactions can be detected in no other way (i.e., often develop slowly (e.g., 30–60 seconds), without the use of even more sophisticat- and result in a yellow, orange, or red col- ed techniques). The technique is not diffi- oration. The crystals can be difficult to cult to learn, but instruction in the obtain. method is beyond the scope of this manu- 2. Ultraviolet (UV) Lamps Ultraviolet al. White and James (1985) provide a good fluorescence provides an effective means introduction.

A Note on In this manual, common names are pro- record observable attributes of the species Common Names posed for all fruticose macrolichens and and genera to which they apply, though microlichens known to occur in British other names are intentionally fanciful. In Columbia. most cases, the same “root” name applies Some of these names have been adopt- to all members of a given genus (e.g., all ed from Lichens of North America (Brodo species of Bryoria bear the name “horse- et al. [2001]), though most are original hair”), but in some cases this name is with this publication. Names in parenthe- applied also to the species of other similar ses have been used by earlier authors— genera. for example, Alvin (1977), Benton and Common names are perhaps most satis- Underhill (1977), Bland (1971), Bolton factorily viewed as vehicles of communica- (1960), Brodo (1988), Casselman (1993), tion for those who are unwilling to use Hawksworth et al. (1995), Johnson et al. scientific names. Scientific names are in- (1995), Kershaw et al. (1998), McCune and tended to be universal and stable; common Geiser (1997), McGrath (1977), MacKin- names are by nature regional and highly non et al. (1992), Nearing (1947), Parish et plastic. The common names introduced al. (1996), Perez-Llano (1944), Pojar and here are intended primarily for use by the MacKinnon (1994), Richardson (1975), naturalist community of British Columbia. Smith (1921), and Vitt et al. (1988)—but While some may gain currency elsewhere in for various reasons are not accepted here. North America, alternative names will Most of the accepted common names probably be coined for many species.

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