Lichens and lichenicolous fungi of the Queen

Lichens and lichenicolous fungi of the Queen Charlotte Islands, British Columbia, Canada. 2. The Cladoniaceae lrwin M. Brodo and Teuvo Ahti

Can. J. Bot. Downloaded from www.nrcresearchpress.com by 192.64.11.124 on 06/04/13 For personal use only.

Abstract: The Queen Charlotte Islands, off the west coast of British Columbia and with a hypermoist, oceanic climate, has 44 taxa in the Cladoniaceae: 5 species and I forma of Cladina, and 34 species with 3 additional subspecies and I variety of Cladonia. Two species and one subspecies are described as new to science: Cladonia albonigra Brodo & Ahti, Cladonia schofieldii Ahti & Brodo, and Cladonia ecmocyna Leight. ssp. occidentalis Ahti. In addition, one new combination is made: Cladonia novochlorophaea (Sipman) Brodo & Ahti. Cladonia tzotnosekekaica Nuno, although not part of the Queen Charlotte flora, is also described and discussed. Chemical variation in the Cladoniaceae is examined critically, and many taxa formerly recognized at the species or infraspecific levels are reduced to unnamed chemotypes. The following synonymies were made or confirmed: Cladina aberrans (Abbayes) Hale & W.L. Culb. =Cladina stellaris (Opiz) Brodo; Cladonia squarnosa var. subsquamosa (Nyl. ex Leight.) Vain. =Cladonia squamosa Hoffm.; Cladonia pseudostellata Asahina = Cladonia uncialis (L.) F.H. Wigg. ; Cladonia japonica Vain. = Cladonia crispata (Ach.) Flot.; Cladonia pseudorangiformis Asahina =Cladonia wainioi Savicz. A thamnolic acid chemotype of Cladonia bellidiflora (Ach.) Schaer. and a thamnolic and usnic acid containing chemotype of Cladonia umbricola Tpnsberg & Ahti are common on the Charlottes. Cladonia singularis S. Hammer is reported as new to Canada based on a specimen from Vancouver Island. Cladonia macroptera Rasanen, Cladonia polydactyla (Florke) Spreng., Cladonia pseudomacilenta Asahina, and Cladonia subsubulata Nyl. are excluded from the North American flora. Cladorzia kanewskii Oksner is reported as new to Norway and Europe.

Key words: Cladina, Cladonia, Cladoniaceae, British Columbia.

Resume : Les iles de la Reine Charlotte, au large de la cBte ouest de la Colombie Britannique, avec un c h a t ocCanique hyperhumide, comportent 4 4 taxons appartenant aux Cladoniaceae : 5 espbces et une forme de Cladina, et 34 espbces en plus de 3 sous-espkces additionnelles et une variCtC de Cladonia. Deux espbces et une sous-espbce sont dCcrites comme nouvelles pour la science : Cladonia albonigra Brodo & Ahti, Cladonia schofieldii Ahti & Brodo, et Cladonia ecmocyna Leight. ssp. occidentalis Ahti. De plus, on prCsente une nouvelle combinaison : Cladonia novochlorophaea (Sipman) Brodo & Ahti. Le Cladonia hornosekekaica Nuno, bien que n'appartenant pas aux iles de la Reine Charlotte, est Cgalement dCcrit et discutk. Les auteurs ont examink attentivement la variation chimique chez les Cladoniaceae, et plusieurs taxons prCcCdemment reconnus comme espkces ou comme entitCs infraspCcifiques ont MC rCduits au niveau de chCmotypes sans nom. 11s confirment les synonymes suivants : Cladina aberrans (Abbayes) Hale & W.L. Culb. =Cladina stellaris (Opiz) Brodo; Cladonia squamosa var. subsquarnosa (Nyl. ex Leight.) Vain. =Cladonia squamosa Hoffm.; Cladonia pseudostellata Asahina =Cladonia uncialis (L.) F.H. Wigg.; Cladonia japonica Vain. =Cladonia crispata (Ach.) Flot.; Cladonia pseudorangiforrnis Asahina =Cladonia wairzioi Savicz. Un chCmotype i acide thamnolique du Cladonia bellidiflora (Ach.) Schaer. et un chCmotype du Cladonia umbricola Tonsberg & Ahti, contenant de l'acide thamnolique et de l'acide usnique, sont frkquents sur les iles Charlottes. On rapporte comme nouveau pour le Canada, le Cladonia singularis S. Hammer, en se basant sur un spCcimen provenant de l'ile de Vancouver. Les auteurs excluent le Cladonia macroptera Rasanen, le Cladonia polydactyla (Florke) Spreng., le Cladonia pseridotnacilenta Asahina et le Cladonia subsubulata Nyl., de la flore nord-amkricaine. On rapporte le Cladonia kanewskii Oksner comme nouveau pour la Norvbge et 1'Europe.

Mots cl& : Cladina, Cladonia, Cladoniaceae, Colombie Britannique. [Traduit par la rCdaction]

I

Received September 15, 1995.

Introduction

I.M. Brodo.' Canadian Museum of Nature, Research

This is the second of a series of taxonomic treatments of the lichens of the Queen Charlotte Islands (QCI), the first being an introduction to the flora (Brodo 1995). This paper is one of the few that will cover a group of macrolichens; the remainder will focus on the crustose species. This is because major treatments have already been published or are in

Division, P.O. Box 3443, Station D, Ottawa, ON K I P 6P4, Canada. T. Ahti. Department of Ecology and Systematics, P.O. Box 47, FIN-00014, University of Helsinki, Finland.

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Author to whom all correspondence should be addressed.

Can. J. Bot. 74: 1147- 1180 (1996). Printed in Canada / Imprim6 au Canada


Can. J. Bot. Vol. 74, 1996

preparation by T. Goward and some co-workers that will cover all the macrolichens of British Columbia (e.g., Goward et al. 1994; T. Goward, B. McCune, and D. Meidinger, unpublished data). A manuscript on the Cladoniaceae was begun by the authors in 1980 but was delayed in no small part by the taxonomic difficulties presented by these taxa. S. Hammer addressed many of these problems in a series of papers on the Cladoniaceae of California, Oregon, and Washington (e.g., Hammer 1991, 1993a, 1993b, 1993c, 1995; Hammer and Ahti 1990). Although we have by no means solved all the remaining (and persisting) problems, we believe enough progress has been made to make a new treatment worthwhile. This treatment, moreover, will deal with the cooler parts of the Pacific coast, areas that differ significantly in their flora from the portions south of Vancouver Island. Because keys to the species of the Cladoniaceae of western United States (Hammer 1995) have recently appeared, and others covering all of British Columbia are in preparation (T. Goward, T. Ahti, and I. Brodo, unpublished data), we have not included keys to the Queen Charlotte Cladoniaceae here. Full descriptions and discussions of the Cladoniae, however, are provided because no modern descriptions of most have been published since Thomson (1968). Ahti (1961) can be consulted for species of Cladina.

The Queen Charlotte Islands Relatively few lichens have been gathered on the QC1 other than the collections made by the authors. The history of collecting is summarized by Brodo (1995). W. Spreadborough collected some Cladoniaceae in 1910 (specimens in CANL), and W.B. Schofield, D. Vitt, D. Horton, and H. Sjors have made some recent collections, many of them cited here. Almost the only published records of Cladoniae from the Charlottes, besides those listed as part of Lichenes Canadenses Exsiccati (Brodo 1971, 1977; Brodo and Wong 1993), or with reference to the British Columbia flora (Ahti et al. 1987; Brodo et al. 1987; Noble et al. 1987), were based on H. Persson's collections in 1957 now located in Stockholm (S). These 10 species of Cladonia were reported by Weber and Shushan (1959). Most of these records have been checked by us and annotated where necessary. The records of Cladonia chlorophaea (Persson No. 122) and Cladonia pyxidata (No. 99a) are, in fact, based on specimens of Cladonia albonigra. Persson's No. 104, listed as Cladonia digitata, is Cladonia transcendens; Cladonia digitata does not occur on the Charlottes. The specimen No. 99 named as "Cladonia gracilis, possibly var. dilatata" is a mixture of Cladonia cornuta ssp. groenlandica, Cladonia ochrochlora, and Cladonia albonigra. The reports on Cladina rnitis belong to Cladina portentosa ssp. pac$ca (Ahti 1961). Thomson's (1984) distribution maps of Cladoniaceae of northern North America include dots on the Charlottes for only five species: Cladonia bellidiflora, Cladonia chlorophaea, Cladonia cornuta, Cladonia gracilis var. gracilis, and Cladonia uncialis. Southeast Alaska Until recently, the lichen flora of southeastern Alaska was relatively unknown. In 1957, H. Krog visited 14 localities in the Eastern Pacific Coast District and recorded 34 species of Cladonia and seven taxa of Cladina (Krog 1968). Some

populations she regarded as "chemical strains" are recognized at the species level in our treatment. She also listed a number of earlier, minor collections. Krog's (1968) collections can be found at the University of Oslo herbarium (0). An inventory of the lichens of the Tongass National forest in southeast Alaska was undertaken by L. Geiser and co-workers between 1989 and 1993 as part of an air biomonitoring project funded by the USDA Forest Service (Geiser et al. 1994a, 1994b). The inventory was published too late to be cited in our discussions. In 1967, J.W. Thomson and T. Ahti made collections along the Haines Highway in Alaska and British Columbia. Most of their collections (H, WIS) are reported in Thomson and Ahti (1994). Other, mainly unpublished collections from the area include those of H.A. Imshaug (MSC), T. Tomberg (BG), R. O'Clair (private herbarium), and I.M. Brodo (CANL).

Geography, climate, and vegetation An account of the climate, general topography,,and flora of the QC1 was published in Brodo (1995), but some brief comments are appropriate.

Geography The QC1 form an archipelago of 138 islands ca. 275 km long and ca. 95 km across at its widest point (Fig. 1). A mountain range runs from the northwest corner of Graham Island to the tip of Moresby and Kunghit Islands, dividing the Charlottes into a hypermoist west coast and a relatively drier and flatter eastern region (the Queen Charlotte Lowlands). The western coastline is very rugged, with high, rocky headland bluffs, terraced bogs, and long fiords. The lowland areas in the northeastern sector of the archipelago contain many boggy lakes. Climate The QC1 receive abundant rainfall throughout much of the year (ca. 3000-4000 mmlyear along the western shore). These conditions reflect its true oceanic climate (mild wet winters and cool wet summers), although the eastern half, lying in a kind of rain shadow caused by the mountain range, have only one-third as much rain as the Pacific side (ca. 1000- 1500 mmlyear). Temperatures, by contrast, are quite similar on all parts of the islands at the same altitude (Williams in Calder and Taylor 1968). Vegetation The vegetation of the QC1 mainly consists of a tall, coniferous forest dominated by Tsuga heterophylla, Thuja plicata, and Picea sitchensis, with abundant Alnus rubra along lake shores and streams. Pinus contorta stands are in the bogs and exposed coastal terraces. At the higher elevations, one finds Tsuga rnertensiana and some Chaernaecyparis nootkatensis together with open alpine meadow. Materials and methods This treatment is based mainly on the collections of the authors on the QCI. Brodo's collecting efforts from 1967 through 1988 are described in Brodo (1995). Both of us made collections on the 1980 excursions to the QC1 as part of the International Congress of

Brodo and Ahti

Fig. 1. The Queen Charlotte Islands showing its physiographic regions and its position relative to western North America (inset). (From Sutherland Brown and Yorath 1989, reproduced with permission, @ 1989 Queen Charlotte Islands Museum Press, Skidgate, B.C.) m

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Langara I.

Systematic Biology meetings. The first set of lichens of Brodo are at CANL; those of Ahti are at H. The chemistry of almost all specimens of Cladonia and many specimens of Cladina was examined by means of thin-layer chromatography (TLC) using the standardized methods of Culberson (1972) and later modifications (e.g., Culberson and Johnson 1982). Thallus colour reactions were made using 10% KOH (K), a solution of a few crystals of para-phenylenediamine in a drop of 70% ethanol (PD), undiluted household bleaching solution (sodium hypochlorite) (C), and C following the soaking of thallus tissue with K (KC). All parts of the thallus were tested, but the most consistent results were usually obtained on the growing tips, the undersurface of podetial (or primary) squamules, or the medulla. The descriptions were compiled using DELTA software for MS-DOS developed by M. Dallwitz in Australia (Dallwitz and Payne 1986). Colour differences, although often subtle, are sometimes important. We attempted to use colour designations consistently by referring

to standardized colour charts (Kelly 1965), using both words and reference numbers. Although paying particular attention to the chemistry of our material, we did not adopt a mechanical "chemical strain = species" concept. We tried to consider each situation of morphological similarity and chemical difference independently, sometimes coming to what may seem to be an inconsistent use of chemistry in our taxonomy. Many of these situations are, indeed, still in flux and may undergo revision with the discovery of better material, new characters, or with opportunities for thorough field studies. The conclusions presented here are to be considered as the most workable conclusions given our present state of knowledge. Specimen citations are abbreviated to the essentials to save space, and for all but the rarest taxa, only collection numbers are given. Collection numbers of Brodo or Ahti are preceded by B- or A-, respectively. Other collectors are identified in full. Specimens collected by Brodo, Horton, and Vitt are all in CANL, those of Ahti


Can. J . Bot. Vol. 74, 1996

and Sjors are in H, and those by Schofield are at UBC, unless otherwise noted. Alaskan specimens gathered by L. Geiser (and C. Derr) as part of their U.S. Forest Service surveys, are housed at TFNS. (All herbarium abbreviations follow Holmgren et al. 1990). Complete lists of all specimens examined are given only for new or rare taxa; for most taxa, only selected specimens are cited. Distribution maps are presented for most species having more than a few localities on the Islands. These were generated using QUIKMap, a desktop computer mapping system in use at the Canadian Museum of Nature (Haber 1993). For those species not mapped, the location is described more fully in the text, often with reference to Fig. 1. The typification and bibliographic citation of each taxon is given in Ahti (1993), though some of the typifications need a repeated conservation proposal according to the Tokyo edition of thi Code of Botanical Nomenclature (T. Ahti, unpublished data).

The taxa Cladina Nyl. For description, synonymy, and nomenclatural discussion, see Ahti (1984). Cladina arbuscula (Wallr.) Hale & W.L. Culb. ssp. beringiana (Ahti) N.S. Golubk. DESCRIPTION: Podetia dense, forming mats or cushions, richly branched, 5O-80(- 100) mm high, 0.6- 1.6 mm wide; branching unequal (anisotomic), mainly in trichotomies but with dichotomies and tetrachotomies frequent, divergent, or infrequently falcate; axils open wide, not forming funnels. Podetia greenish yellow (104), fading in the herbarium, more or less uniform, or becoming brown at tips; colour of podetial base more or less the same as upper portions; podetial surface smooth, wall entire or perforated; surface layer 20 pm thick, stereome 40 -SO(- 100) pm thick. Apothecia rare. Pycnidia on tips of podetia; pycnidial jelly hyaline. CHEMISTRY: Thallus PD+ red, K-, UV-. Contains usnic and fumarprotocetraric acids,' Cph-2,3 sometimes also Cph- l . HABITAT: Growing in full sun, in dry or moist habitats usually in or near bogs or fens, less frequently over rocks. DISTRIBUTION: QCI: widespread in both Graham and Moresby Islands, at high and low elevations (Fig. 2). N. Am.: western, south to Colorado in Rockies. World: North America, eastern Asia. SELECTED SPECIMENS EXAMINED: A-38977, 391 70, 391 73; B-10204, 10979, 14222, 18003; Horton 1714 (CANL). DI~CUSSION: The coastal populations of Cladina arbuscula are variable in morphology. Many colonies clearly belong to ssp. beringiana, characterized by a smooth rather than areolate surface and rather robust podetia. Many others are thin and slender (e.g., A-39106, 39170) resembling Cladina ciliata (Stirt.) Trass, but the pycnidial jelly is hyaline. (Cladina ciliata has not yet been found on the Charlottes but can be expected to occur in the area.) Some colonies resemIn the genus Cladonia, fumarprotocetraric acid is almost always accompanied by at least traces of protocetraric acid, but this is not always mentioned in the chemical profiles. Cph-2 is confumarprotocetraric acid (Elix and Yu 1993), and Cph-l, according to J.A. Elix (personal communication), is convirensic acid, also characterized by Elix and Yu (1993). In most cases, we are using the short form in the text to save space.

Figs. 2-5. Distribution of some species of Cladina on the Queen Charlotte Islands. Fig. 2. Cladina arbuscitla ssp. beringiana. Fig. 3. Cladina portentosa ssp. pacifica. Fig. 4. Cladina rangiferina. Fig. 5 . Cladina srygia.

Zladina arbuscula subsp. beringiana +

2

8

C. portentosa subsp. pacifica 1

L . L-

ble Cladina mitis in the more open branching, but they contain fumarprotocetraric acid and no rangiformic acid and are therefore referred to Cladina arbuscula. The status of the subspecies beringiana requires further investigation. Cladina portentosa (Dufour) Follm. ssp. pacifica (Ahti) Ahti DESCRIPTION: Podetia dense, often forming cushions, richly branched, with somewhat curly appearance, 40 - 140 mm high, main axes 0.7 - 1.3(- 1.8) mm wide; branching anisotor&, mainly in trichotomies with dichotomies often frequent, sometimes with dominance of SO%, mostly divergent, frequently anastomosing; axils open to closed, not forming funnels. Podetia yellow-green or pale greenish yellow (121, 104) in f. pacifica (Ahti) Ahti and grey or yellowish to greenish white in f. decolorans (Ahti) Ahti; both forms can become brown at tips; colour of podetial base more or less the same as upper portions; podetial surface arachnoid, verruculose, or areolate, with very small, rounded, more or less flocculent patches of outer medulla distributed over a translucent inner medullary surface, even close to the podetial tips; podetial wall entire, or perforated; cortex 20-40 pm thick; stereome 80- 100 pm thick. Apothecia common, on podetial tips, pale brown, flat or somewhat convex, 0.3 -0.5 mm in diameter. Pycnidia on tips of podetia; pycnidial jelly hyaline. CHEMISTRY: Thallus PD-, K-, UV + bright blue-white in part; f. pacijica: KC+ yellow, contains usnic and perlatolic


Brodo and Ahti

acids (apparently lacking ursolic acid); f. decolorans: KC-, contains perlatolic and often ursolic acids. HABITAT: Growing in full sun in dry or moist habitats, especially luxuriant and often abundant in lowland bogs. DISTRIBUTION: QCI: low elevations; common throughout Alaska the islands (Fig. 3). N. Am.: west coast: f. paci'ca, to California; f. decolorans, Alaska to Washington. World: North America. SELECTED SPECIMENS EXAMINED: f. paci'ca: A-38544, 39103, 39181, 39184; B-10511, 12002, 12748, 14083, 14328, 17302, 18043, 18106; Chaatl I., Scho'eld & Boas 18704 (H); f. decolorans: A-39055, 39102, 39104, 39183; B-9745, 11184 (Lich. Can. Exs. 98), 10269, 10594, 11828, 12279, 12690, 12968, 17436; Horton 21402 (CANL); Sjors Q22, Q31, Q43, Q49 (H). DISCUSSION: The oceanic, coastal distribution of Cladina portentosa ssp. pacijca in the Pacific Northwest was emphasized by Ahti (1961) under the name Cladonia pacifica Ahti, reporting it for the first time from the QCI. It was subsequently reduced to a subspecific rank of Cladonia portentosa, described from Europe (Ahti 1978a), a rank maintained when the species was transferred to Cladina (Ahti 1984), but its taxonomic status is still not clear. In the field, ssp. pac$ca appears to be more yellow, slender, brown-tipped, and deflexed than ssp. portentosa, and it is smoother and more dichotomous under the microscope. However, most of the European populations are suffering from air pollution and grazing and hence are stunted, grey (even when usnic acid is present), and deformed, so that a comparison is difficult. The formapaciJica (Ahti) Ahti of Cladina portentosa ssp. pac$ca has usually, but incorrectly, been cited without an author citation, but because it does not include the type of the species (Cladina portentosa), an author citation is required (ICBN Art. 11.6). It seems to have been validated in Ahti (1978a, p. 9) through a page reference to the Latin description of Cladonia paci'ca and was referred to Cladina in Ahti (1984, p. 37). It is homotypic with ssp. pac$ca. The forma decolorans is an usnic acid-deficient chemotype of ssp. paci'ca and therefore conspicuously ash-grey rather than yellowish. It is usually easily recognized, especially in mixed colonies. Although usnic acid-free colonies are considered to be rare in Europe, where only ssp. portentosa occurs, in populations of ssp. paci'ca they are very common, often forming wide mosaic patches among f. pacifi c ~ If . ssp. paci'ca is not recognized, then f. decolorans must be called f. subimpexa (P.A. Duvign.) Ahti, described from Belgium. The type material o f f . decolorans (H, UPS) is from the QC1 (Graham I.: SW of Tow Hill, 1966, Sjors Q43) (Ahti 1 9 7 8 ~ ) . Cladina rangiferina (L.) Nyl. DESCRIPTION: Podetia dense, often forming cushions, richly branched, frequently with long internodes, 50- 120 mm high, 0.8-1.8 mm wide; branching anisotomic, mainly in trichotomies with frequent dichotomies and occasional tetrachotomies, falcate; axils open wide, not forming funnels. Podetia grey to yellowish white, mottled or becoming brown at tips; colour of podetial base usually more or less the same as upper portions; podetial surface arachnoid or verruculose; wall entire or perforated; cortex 20 pm thick, stereome

50-100 pm thick. Apothecia common, on podetial tips, brown, flat, somewhat or strongly convex to hemispherical. Pycnidia on tips of podetia; pycnidial jelly hyaline. CHEMISTRY: Thallus P D + red, K + pale yellow, UV-. Contains atranorin, fumarprotocetraric acid, and Cph-2. Atranorin is occasionally lacking (e.g., B-18090, B-9793); this chemotype was reported earlier by Krog (1968) from Alaska. HABITAT: Growing in full sun, in dry habitats, usually in patches of moss in forested habitats or on slopes. DISTRIBUTION: QCI: common at low and high elevations (Fig. 4). N. Am.: arctic to temperate, widespread. World: North and South America, Europe, Asia, Antarctica. SELECTED SPECIMENS EXAMINED: B-9793, 10118, 10256, 10508,10922, 12779, 12943,14219, 17593, 18090,18233; Vitt 12452 (CANL). DISCUSSION: Ahti (1961) pointed out that the western American populations of this species tend to be very pale, almost white, and have less branching and more dichotomous podetia. Similar variants in East Asia have been referred to ssp. grisea (Ahti) Ahti & Lai, and this name has occasionally been applied also in western North America (Krog 1968). However, it does not seem to be possible to recognize such a race as distinct, though further studies are required. Very white specimens with blackening stereome but apparently hyaline pycnidial jelly were collected (e.g., Brodo 10508, 14219, 17593); they were included here rather than in Cladina stygia based on the colour of the pycnidial jelly. Cladina stellaris (Opiz) Brodo syn. Cladina aberrans (Abbayes) Stuckenb.; Cladina stellaris var. aberrans ( ~ b b a ~ e Ahti s) DESCRIPTION: Podetia forming rounded heads, often in clumps, richly branched, 40-70 mm high, mature branches 0.7--0.8 mm wide; branching equal (Gotomic), mainly in tetrachotomies, dichotomies and trichotomies frequent, divergent; axils open wide, the branch tips flaring and starlike. Podetia greenish yellow (l@), more or less uniform; colour of podetial base more or less the same as upper portions; podetial surface arachnoid or verruculose; the tips having a thick, cottony appearance, with a verruculose surface occurring on older portions of the podetia; podetial wall entire; cortex 120- 150 pm thick; stereome 30-40 pm thick. Apothecia not produced, or rare. Pycnidia on tips of p ~ d e t i a ; ~ ~ c n i djelly i a l red. CHEMISTRY: Chemotype 1: usnic and perlatolic acids (thallus PD-, K-), not on QCI; chemotype 2: usnic, perlatolic, and psoromic acids (thallus PD+ yellow at the branch tips, K-); both chemotypes are UV+ blue-white. HABITAT: Growing in full sun in dry habitats. DISTRIBUTION: QCI: northern part of Queen Charlotte Range, high elevation, very rare. N. Am.: arctic-boreal, widespread; chemotype 2: frequent Alaska to western Keewatin, scattered in eastern regions. World: North America, Europe, and Asia; chemotype 2: North America, East Asia, very rare in Europe. SPECIMEN EXAMINED: Moresby Island: "Laing Point Mountain," 52"59'N, l32"@'W, subalpine bluffs near summit, exposed knoll at 760 m elev., Brodo 10840 (CANL). DISCUSSION: It is noteworthy that Cladina stellaris is

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represented only by its psoromic acid-containing chemotype (var. aberrans) on the QCI, although the widespread perlatolic acid strain (var. stellaris) is close by on the mainland. It is questionable whether var. aberrans has any diagnostic morphological characters, and it therefore can be treated as an unnamed chemotype. This chemotype is regarded as a coastal Beringian radiant but is also known in scattered localities in eastern North America (map in Thomson 1984). Cladina stygia (Fr.) Ahti DESCRIPTION: Podetia dense, often forming cushions, richly branched, 50-120 mm high, main axes 0.8-1.6(-2) mm wide; branching anisotomic, mainly in dichotomies, with frequent trichotomies, falcate; axils open wide, not forming funnels. Podetia grey to yellowish white, mottled, becoming brown at tips; podetial base strongly melanotic; podetial surface arachnoid or verruculose; wall entire or perforated; cortex 20 pm thick; stereome 50- 100 pm thick. Apothecia rare. Pycnidia on tips of podetia; pycnidial jelly red. CHEMISTRY: Thallus PD+ red, K + pale yellow, UV-. Contains atranorin and fumarprotocetraric acid, traces of Cph-2, and unidentified fatty acids. HABITAT: Growing in full sun, usually in or near bogs or fens, but also in grassy or mossy heath, elev. 10- 150 m. DISTRIBUTION: QCI: infrequent, scattered (Fig. 5). N. Am.: arctic-bored, widespread. World: North America, Asia, Europe. SPECIMENS EXAMINED: Graham Island: 24 km N of Queen Charlotte City, B-1 7997; Haida, B-2661 7. Moresby Island: "Blue Heron Bay" N of Sunday Inlet, B-14119, B-14111; Kootenay Inlet, B-12172; Tasu Sound, Fairfax Inlet, A-38514 (CANL, H, UBC). DISCUSSION: The distribution of this species is little known because it was only recently segregated from Cladina rangiferina by Ahti (1984) and Ahti and Hyvonen (1985). They also reported it from the QCI. It is recognized from its blackening stereome in the basal parts, presence of purple jelly in pycnidia, and usually strongly browned top branchlets. It has turned out to be locally very common in northern to central British Columbia, including the highly oceanic parts. Cladonia Hill ex P. Browne For description, see Thomson (1968).

Cladonia albonigra Brodo & Ahti, sp.nov. Fig. 6 A Cladonia merochlorophaea differt podetiis melanoticis et scyphis saepe prolificationes centrales producentibus. Semper acidum fumarprotocetraricum et vulgo acidum grayanicum vel acidum 4-0-methylcryptochlorophaeicum continens. TYPE:Canada. British Columbia. QCI: Graham Island: 2 mi SE of Port Clements, 53 "40fN, 132"09'W, open Pinus contorta bog, 1971, I.M. Brodo 18104, with P. Y. Wong (CANL, holotype; H, isotype). DESCRIPTION:Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dispersed, 0.5 2 mm long, 0.4- 1 mm broad, entire or crenulate, rounded or ligulate, grey-green to pale yellow-green (121); undersurface white, ecorticate, esorediate. Podetia loose or scattered, unbranched, 10-47 mm high, 1-2.5 mm wide, with broad scyphi, 2 - 8 mm across, closed, proliferating from the center or margins of the cups, producing up to four or five

Fig. 6. Cladonia albonigra, holotype, Brodo (& Wong) 18104 (CANL). Scale in millimetres.

tiers; margins entire or dentate. Podetia grey-green, yellowgreen, or brown (121) to reddish brown (47-43), mottled. Base strongly melanotic. Cortex present or absent. Podetial surface areolate; areoles thick, convex, greenish or brownish when fresh, turning almost white in old podetia and standing out against the blackened, decorticate stereome. Podetial squamules occasional: small, rounded, entire to crenulate, perpendicular to surface, strongly decumbent or peltate. Podetia coarsely granular to sorediate; soredia granular, diffuse, on upper half of podetia or covering podetia except for base, present within cups. Podetial wall entire. Apothecia rare, on the cup margins, brown, strongly convex to hemispherical, 1-2 mm i n diameter. Pycnidia sparingly produced, at cup margins, at first globular, sessile to subsessile, later short-stalked, shortly cylindrical to pyriform, redbrown, constricted at base. CHEMISTRY: Thallus PD red, K-, UV bright blue-white or UV-. Major compounds: chemotype 1: grayanic acid and often 4-0-methylgrayanic (with minor satellites) and always fumarprotocetraric acid with satellites (type); chemotype 2: 4-0-methylcryptochlorophaeic acid and fumarprotocetraric acid aggr.; chemotype 3: fumarprotocetraric acid aggr. (including protocetraric acid and Cph-2; see footnote 2) alone. HABITAT: Growing on mineral soil, wood or moss, wet or dry habitats, in full sun. DISTRIBUTION: QCI: throughout the islands at low and high elevations (Fig. 7). N. Am.: west coast, Alaska to California (Canadian and Alaskan distribution, Fig. 8). World: North America. (See comments on distribution of chemotypes below .)

+

+


Brodo and Ahti SPECIMENS EXAMINED: Chemotype 1: QC1 (selected specimens): A-39011, 39032; B-10946, 12705, 12926, 12782, 17612, 17795, 17937; Schojeld 25039 (UBC). U. S.A. : ALASKA: Juneau area, B-26222, 26173; Derr & Geiser L-3304; Prince of Wales Island, Geiser & Derr L-3305, L-3307. Chemotype 2: QCI: Graham Island: Haida, B-26622. Vancouver area, West Vancouver, B-23714. Tongass National Forest, Geiser 1021 (not U.S.A.: ALASKA: mapped). WASHINGTON: Clallam Co., Sequim, 7homson 14827 (H); Whatcom Co., Mt. Baker, 500-800 m, Hammer 4153 (FH, H), Sulfur Creek Lava Flow, Ahti & Rhoades 51008b (H). Chemotype 3: QCI: Graham Island: Mt. Raymond, B-26638. Moresby Island: Upper Victoria Lake, B-12281. U.S.A.: ALASKA: Juneau area, B-26200, Derr & Geiser L-3290; Chatham area, Berner 'S Bay Peninsula, Geiser L-3308; Yakutat, Geiser & Derr L-3306. DISCUSSION: Cladonia albonigra is a variable member of the Cladonia chlorophaea group. It is characterized by having a conspicuously melanotic base, the stereome gradually becoming entirely blackened except for the youngest proliferations. Over this melanotic, decorticate surface can be seen scattered, round, corticate granules to rounded peltate squamules or verrucae. The podetia are often tall and slender and frequently proliferate from the center as well as the margins of the cups. Although this morphologically defined population always contains fumarprotocetraric acid, it can apparently occur as three chemotypes (see Chemistry above). The chemotypes differ slightly in morphological details and distribution. Chemotype 1: grayanic acid; most strongly melanotic; often tall and slender but can be short and squat; central proliferations very common; granulose rather than squamulose (squamules on podetia infrequent); QC1 north into coastal Alaska, common. Chemotype 2: 4-0-methylcryptochlorophaeicacid; central proliferations rare; very squamulose; southern (California, common in Oregon and Washington, rare on QC1 and in SE Alaska (Geiser 1021); localities in conterminous U.S.A. mapped by Hammer (1995), sub Cladonia merochlorophaea var. merochlorophaea). Chemotype 3: no orcinol meta-depsides; less melanotic than grayanic strain; medium height; central proliferations present but not common; podetia with abundant, decumbent, rounded podetial squamules; QC1 north to coastal Alaska, especially common in Alaska. According to our morphological species concept, these strains are united because they share two very distinctive traits: strongly melanotic medulla and frequent production of central proliferations in scyphi. Both of these characters are unknown in the other members of the Cladonia chlorophaea group, although the species is undoubtedly very close to, for example, Cladonia merochlorophaea. In spite of the production of grayanic acid, Cladonia grayi of eastern North America is not very similar to the new species, although it may rarely produce central proliferations (f. centralis A. Evans). Cladonia albonigra is often very pale, even whitish, rather than brown, unlike all the other closely related members in the group, although it can also become browned in certain habitats, and especially with age. Occasionally the scyphi are crowded inside with abundant, small phyllidia. The chemotype with 4-0-methylcryptochlorophaeic acid as a major compound was first reported from Washington by Hennings (1983) (mainly under Clado-

nia merochlorophaea) and discussed by Hammer (1993~)(as Cladonia cfr. merochlorophaea). The latter also noted the essential diagnostic characters of this taxon, which appears to belong to Cladonia albonigra. Cladonia albonigra was listed without a validating description as Cladonia albonigra Brodo in Geiser et al. (1994a, 1994b). Cladonia asahinae J. W. Thomson DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dispersed, 2 -5 mm long, 0.8-5 mm broad, crenulate or deeply incised, pale yellow-green (121) to greyish greenish yellow (105), undersurface white or grey, ecorticate, esorediate. Podetia loose or scattered, unbranched, 10-25 mm high, 0.75-2 mm wide, with scyphi that are broad (goblet shaped), or narrow (trumpet shaped), 2-7 mm across, closed, not proliferating or proliferating from the margins of the cups, very rarely from the center. Margins entire or, more frequently, dentate. Podetia grey-green or yellow-green (121), sometimes almost white, more or less uniform. Base more or less the same as upper portions, i.e., where corticate; cortex present on lower half of podetia or up to cup, sometimes almost absent. Podetial surface verruculose or areolate where corticate or where cortex is breaking up. Podetial squamules occasional, mostly perpendicular to surface, or decumbent. Podetia sorediate, or with spherical, corticate granules. Soredia farinose or rarely granular around upper part of outside of cups, diffuse, mainly confined to tip of podetia, on upper half of podetia, or occasionally covering podetia except for base; present within cups. Podetial wall entire. Apothecia rare, on the cup margins, usually on short stipes, brown, strongly convex to hemispherical, 1-4 mm in diameter. Pycnidia at cup margins, black, subsessile, pyriform, constricted at base, containing hyaline slime. Thallus PD+ red, K-, UV- (fatty acid defiCHEMISTRY: cient), or UV+ dull (with fatty acids). Chemotype 1: fumarprotocetraric acid, rangiformic acid, and norrangiformic acid; chemotype 2: fumarprotocetraric acid and protolichesterinic acid (rarely with rangiformic); chemotype 3: fumarprotocetraric acid alone (rare). HABITAT: Growing on mineral soil, or on mossy rocks, in full sun. DISTRIBUTION: QCI: common at low elevations in northern Moresby Island (Fig. 9). N. Am.: west coast, Alaska to California. World: North and South America, Asia, Europe, and Antarctica. SPECIMENS EXAMINED: Chemotype 1: B-10956, 11776B, 11988, 13899, 139731, 14002, 14273A, 17370, 17873A, Tongass National Forest, 17932, 18013. U.S.A. : ALASKA: Yakutat, Geiser & Derr L-3299. Chemotype 3: QCI: Moresby Island: Sandspit, A-39111; B-17154. DISCUSSION: This large, goblet-producing Cladonia containing fatty acids and somewhat resembling Cladonia j m briata (L.) Fr. was named as a distinct species by Thomson (1977) based on material from Washington, Idaho, California, and British Columbia. Thomson (1977) regarded the common occurrence of granular soredia, the large size of the podetia, and the greyer colour as being diagnostic, in addition to chemistry. Hennings (1983) and Huovinen et al. (1990) commented on the chemistry of the type material. Hennings (1983) regarded both chemotypes 1 and 2 to be


Can. J. Bot. Vol. 74,

C. bellidiflora chemotype 1

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Q

C. cornuta subsp. groenlandica

Brodo and Ahti

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Figs. 7-15. Distribution of species of Cladonia on the Queen Charlotte Islands. Fig. 7. Queen Charlotte Islands distribution of Cladonia albonigra. 0 , chemotype 1 (grayanic and fumarprotocetraric acids); m, chemotype 2 (4-0-methylcryptochlorophaeic and fumarprotocetraric acids); A , chemotype 3 (fumarprotocetraric acid alone). Fig. 8. Canadian distribution of Cladonia albonigra. Symbols as in Fig. 7. Fig. 9. clado& asahinae. Figs. 10 and 11. Cladonia bellidflora. Fig. 10. Chemotype 1 (squamatic acid). Fig. 11. Chemotype 2 (thamnolic acid). Fig. 12. Cladonia carneola. Fig. 13. Cladonia coniocraea. Fig. 14. Cladonia cornuta ssp. groenlandica. Fig. 15. Cladonia crispata var. crispata. A , chemotype 1 (squamatic acid); 0 , chemotype 2 (thamnolic acid).

close to, if not conspecific with Cladoniajmbriata but said the strains differed slightly in morphology, with the protolichesterinic strain differing most significantly. We did not find the protolichesterinic acid-containing strain on the QCI. The rangiformic acid strain, on the other hand, is fairly common and was also reported from the Kenai Peninsula of Alaska by Krog (1968), and we saw specimens from southeast Alaska. It is also known from Oregon (Hammer 1991), Norway, Sweden, and Czechoslovakia (Holien and Tonsberg 1985), and Finland (Huovinen et al. 1990), and we saw material from Iceland and Siberia (H). It is also widespread in Tierra del Fuego, antarctic islands, and even on the Antarctic Peninsula (Ahti and Kashiwadani 1984; Stenroos et al. 1992; Stenroos 1993). It is therefore obviously widely distributed, especially in the north, although at least in Norway and in the southern hemisphere, it seems to have oceanic tendencies (Holien and Tonsberg 1985). Its distribution in western Washington (Hennings 1983) attests to this as well. From the few specimens we examined of the protolichesterinic strain, however, we can see no morphological divergence and we are therefore reluctant to name that strain. Several Queen Charlotte specimens apparently belong to a fatty acid deficient strain. They have tentatively been named as Cladonia asahinae by us because they appear to be indistinguishable morphologically. The species Cladonia asahinae is fairly well defined despite its great variability. Its tall, dentate cups, mixture of farinose and granular soredia with frequent granules and verrucae, its squamulose base, and its tendency to have a rather greyish or brownish grey colour, all make it relatively distinctive. The morphology, nevertheless, is sometimes difficult to interpret, and there can be a problem in distinguishing it from Cladonia chlorophaea and Cladoniajmbriata in particular. It is often entirely farinose sorediate and very pale, then resembling Cladonia jmbriata. Cladonia bellidiflora (Ach.) Schaer. DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dense or dispersed, 2-12 mm long, 1-7 mm broad, deeply incised, yellow-green (135-136) or maculate, undersurface white, or dark yellow to orange at dying base, ecorticate, esorediate. Podetia arising from center of squamules, scattered or clustered, unbranched, or once or twice branched, 10-50 mm high, 1-3 mm wide, lacking or with scyphi. Scyphi narrow or irregular, rarely broad, 2-7 mm across, closed (imperforate), proliferating from the margins of the cups. Margins dentate or squamulose. Podetia yellow-green (121), discolouring to yellowish or brownish, more or less uniform. Base turning yellow. Cortex present. Podetial surface areolate or broken into irregular "plates." Podetial squamules abundant, small and entire to very large and deeply incised, perpendicular to surface or strongly decumbent. Podetia

without soredia or granules, but tiny granulose podetial squamules (microsquamules) can appear to be granular. Podetial wall entire, or longitudinally split. Stereome cylindrical, entire, usually dark yellow. Apothecia common, on podetial tips or on the cup margins, red, somewhat convex, or strongly convex to hemispherical, 0.5-6 mm in diameter. Pycnidia on margins of scyphi. CHEMISTRY: Chemotype 1: thallus PD-, K-, UV+ bright blue-white: usnic and squamatic acids (type strain); chemotype 2: thallus PD+ yellow-orange, K + deep yellow, UV-: usnic and thamnolic acids. Usnic acid is sometimes difficult to demonstrate with TLC, but is always present in at least the specimens containing squamatic acid. HABITAT: Growing on mineral soil or moss, or on wood (especially stumps and logs), in full sun, partial shade, or in deeply shaded habitats. DISTRIBUTION: QCI: chemotype 1: common throughout the islands at all elevations (Fig. 10); chemotype 2: frequent, especially in the eastern sectors at low elevations (Fig. 11). N. Am.: west coast and arctic-boreal. World: North and South America, Europe, Asia, New Zealand. SELECTED SPECIMENS EXAMINED: Chemotype l : A-38959; B-l 01l l , 10260, 10837, 10974, 12399A, 14060, 14358, 18057; Schojeld 18703 (UBC); Slatechuck Mt., Christie, S.n. (UBC). Chemotype 2: A-38091, 38924; B-11658, 11875, 12399C, 17562, 18235, Schojeld 23705, 24221, 242 71 (UBC) . DISCUSSION: Cladonia bellidiflora is a widely distributed species found throughout the low-arctic region of the northern hemisphere with marginal extensions to the boreal zone (Goward and Ahti 1992), but disjunct populations also occur in the southern hemisphere (Stenroos and Ahti 1990). Unlike many other western European - western American disjunct~,the species is absent from temperate lowland regions of Europe and yet is common along the west coastal strip of North America from California to Alaska (Thomson 1984). Furthermore, although the European populations consistently produce usnic and squamatic acids, the chemistry of the species is more variable in the west coast and southern hemisphere material. In Tierra del Fuego, strains occur that contain fumarprotocetraric acid or thamnolic acid (Stenroos and Ahti 1990), and the strain containing thamnolic acid is common on the QCI, although rare elsewhere in the Pacific Northwest, e.g., southern Vancouver Island (specimens in CANL and H). It has also turned out that the scrappy, original type specimens of both Cladonia transcendens (Vain.) Vain. ("Oregon Boundary Commission," 1858, Lya11, syntype in TUR-V 14166 seen) and Cladonia pseudosipeana Gyeln. (Oregon: Lane Co.: Coburg Hills, on logs, Sipe 689; isotype in US seen) seem to represent the thamnolic acid strain of Cladonia bellidiflora (see discussion on typification under Cladonia transcendens). Although quite variable in morphology, Cladonia bellidi-

Can. J. Bot. Vol. 74. 1996


j7ora can generally be recognized by its abundantly squamulose podetia usually ending in a narrow cup, its thick cortex becoming disrupted, leaving thick areoles or plates on the otherwise decorticate upper portions of the podetia and its large, deeply incised primary squamules. The podetial squamules can be large and lobed, or they can be extremely small and scale-like. In extreme cases, the squamules can become almost granulose (i.e., constituting microsquamules), and the specimens can resemble Cladonia transcendens (containing thamnolic acid) or Cladonia vulcani Savicz (containing thamnolic or squamatic acid; Huovinen et al. 1 9 8 9 ~ )The . latter two species, however, generally always show at least some unambiguous patches of granular soredia. Cladonia borealis S. Stenroos DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending, dense or dispersed, 3 - 10 mm long, 3 - 10 mm broad, crenulate, greyish to pale greenish yellow (105-104) to yellow green (121), maculate; undersurface white, or brownish black at base, ecorticate, esorediate. Podetia arising from center of squamules, scattered or clustered, unbranched, 7-40 mm high, with broad scyphi, 4- 10 mm across, closed, not proliferating or proliferating from the margins of the cups; margins entire. Podetia pale to greyish greenish yellow (104-105) or yellow green (121), more or less uniform. Base more or less the same as upper portions or turning yellow or brownish. Cortex present. Podetial surface smooth or areolate, forming rounded or angular cortical plates. Podetial squamules occ&ional at very base, mostly perpendicular to surface. Podetia without soredia or granules. Podetial wall entire, 260-400 pm thick; cortex 20-80 um thick; medulla 160-240 um thick. Apothecia common, on thk cup margins on s h o i stipes or proliferations, red, somewhat convex or strongly convex to hemispherical, 1.5 -5 mm in diameter. Pycnidia on margins of scyphi. Pycnidial jelly red. CHEMISTRY: Thallus PD-, K-, UV-. Contains usnic, barbatic, and 4-0-demethylbarbatic acid. HABITAT: Growing on sandy soil or directly on rock, alpine, in full sun. DISTRIBUTION: QCI: northern part of San Cristoval Range, at 1000 m elev.; very rare. N. Am.: arctic-boreal, widespread. World: North and South America, Europe, Asia, Antarctica. SPECIMENS EXAMINED: Moresby Island: Tasu Mt., 52"45'N, 132"03'W, Brodo 14273B. U.S.A.: ALASKA:Tongass National Forest, Juneau area, mouth of Herbert Glacier, Geiser L-3292. DISCUSSION: Cladonia borealis is actually a new name given by Stenroos (1989~)to what was called Cladonia coccifera (L.) Willd. s.str. by Thomson (1968) and others, i.e., the barbatic acid containing, esorediate taxon in the Cladonia coccifera complex. The true Cladonia coccifera contains zeorin but is also esorediate, although granulose. Older herbarium specimens can be recognized under the microscope by the presence of needle crystals on the podetial surface (see under Cladonia pleurota) . Cladonia carneola (Fr.) Fr. DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dispersed, 2 -5 mm

long, 0.4 - 8 mm broad, deeply incised, often thickened and lobulate, greenish yellow (105) to pale yellow-green (121), undersurface white, or brownish at point of attachment, ecorticate, esorediate. Podetia arising from center of squamules, loose or clustered, unbranched, 7-30 mm high, with broad or narrow scyphi, 1.5-5 mm across, closed, proliferating once or twice from the margins of the cups, rarely from the center; margins entire or often dentate, rarely squamulose. Podetia greenish yellow (104) to very pale yellowish green (121), more or less uniform, or mottled where ecorticate. Base more or less the same as upper portions. Cortex present, areolate, at least at base, absent from most of podetia. Podetial squamules occasional, mostly perpendicular to surface. Podetia sorediate; soredia farinose or granular, diffuse, on upper half of podetia or covering podetia except for base, present within cups. Podetial wall entire. Apothecia common, on the cup margins, pale to greyish to light brown, or light orange (61-57, 53), flat, or somewhat convex, 0.5-5 mm in diameter. Pycnidia on margins of scyphi. CHEMISTRY: Thallus PD-, K-, UV-. Contains usnic acid and zeorin; barbatic and isousnic acids are present in most specimens. HABITAT: Growing on rotting wood and logs, humus, or on bark, in full sun, in partial shade, or in deeply shaded habitats. DISTRIBUTION: QCI: common throughout the islands at low and high elevations (Fig. 12). N. Am.: arctic-boreal, widespread. World: North and South America, Europe, Asia, Antarctica. SELECTED SPECIMENS EXAMINED: A-38967, 39031; B-9946, 10236, 10696B, 10943, 14263, 14315. DISCUSSION: This is widespread in the QCI, although Goward and Ahti (1992) indicated that its general distribution shows continental tendencies. Cladonia cervicornis (Ach.) Flot. ssp. cervicornis DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending, dense, 3 - l 0 mm long, 2 -6 mm broad, entire, crenulate or deeply incised to lobed or ligulate, greyish green to pale yellow-green (122-121) to greyish olive (109) or light brown (76-77), undersurface white, or brownish black at base, ecorticate, esorediate. Podetia arising from center of squamules, scattered or dense, unbranched, 10-22 mm high, 0.8- 1.3 mm wide, with broad or narrow, more or less flat scyphi, 2 -4 mm across, imperforate, proliferating from the center or the margins of the cups; margins entire or dentate. Podetia yellowish green (122-121), becoming brown (76-77), more or less uniform (in Scandinavian specimens) or mottled. Base more or less the same as upper portions outside of QCI, but weakly to strongly melanotic in QC1 material. Cortex present. Podetial surface verruculose or areolate, areoles becoming convex and conspicuous. Podetial squamules occasional, mostly perpendicular to surface. Podetia without soredia or granules. Podetial wall entire. Apothecia not produced, or rare, brown. Pycnidia on margins of scyphi. CHEMISTRY: Thallus PD+ red, K-, UV-. Contains fumarprotocetraric and protocetraric acids, and Cph-2. HABITAT: Growing on mineral soil or directly on rock, at 760 m elev. in full sun. DISTRIBUTION: QCI: E side of Masset Inlet, and the Queen

Brodo and Ahti


Charlotte Range; rare. N.Am.: west coast; range incompletely known. World: North America, Europe, poorly known. SPECIMENS EXAMINED: Moresby Island: alpine slope above Takakia Lake, 52"56'N, 132"02'W, B-26795. DISCUSSION: The taxonomy of Cladonia cervicornis in the Pacific Northwest needs closer study, being very incompletely known (Hammer and Ahti 1990, Hammer 1 9 9 3 ~ ) . Much of the material does not fit the European concepts of the taxa. Cladonia chlorophaea (Florke ex Sommerf.) Spreng. DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dense or dispersed, 2 -5 mm long, 2 -6 mm broad, crenulate or deeply incised, yellow-green (135), olivaceous or brown, undersurface white, or brownish to black at base, ecorticate, esorediate. Podetia arising from center of squamules, or arising from edge of squamules, scattered or clustered, unbranched, 7-35 mm high, with broad or rarely narrow scyphi, 3 -8 mm across, closed, not proliferating or proliferating from the margins of the cups, especially when fertile; margins mostly entire. Podetia pale yellowish white to yellow-green (92-121), becoming brown, more or less uniform. Base more or less the same as upper portions. Cortex usually present, at least on lower third to half. Podetial surface verruculose or areolate on lower half. Podetial squamules absent, occasional, or rarely abundant, granuliform, mostly perpendicular to surface. Podetia sorediate, or with spherical, corticate granules. Soredia granular, diffuse, on upper half of podetia, present within cups. Podetial wall entire. Apothecia common, on the cup margins on short proliferations, brown, somewhat convex, or strongly convex to hemispherical, finally compound, 1-8 mm in diameter. Pycnidia common, on margins of scyphi, pyriform, sessile to stalked, finally constricted at base. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-. Contains fumarprotocetraric and protocetraric acids, Cph-2; atranorin was found once as a trace. HABITAT: Growing on sandy soil or directly on rock, but mainly on wood, especially on stumps, logs, and rotting wood, in full sun or in partial shade. DISTRIBUTION: QCI: infrequent; scattered, mostly in lowland localities in the NE sector; also in northern San Cristoval Range. N. Am.: arctic to temperate, widespread. World: North and South America, Europe, Asia, Africa, Australasia, Antarctica. SPECIMENS EXAMINED: Graham Island: Sandspit, B-10082, 17371; N of Skidegate Mission, B-1 7969; Skonun Pt., E of Masset, B-18196. Skidegate Inlet: Robbers Island, B-11MO; Torrens Island, B-17299. Moresby Island: Tasu, A-38974; Mountain E of Sunday Inlet, B-26840. DISCUSSION: Some Queen Charlotte material we named as Cladonia chlorophaea may not be conspecific with common eastern North American morphs. Cladonia coniocraea (Florke) Spreng. DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dispersed, 1 -6 mm long, 1-6 mm broad, entire, or crenulate, rounded, or ligulate, grey-green or olivaceous, undersurface white, ecorticate, esorediate. Podetia arising from center of squamules,

scattered or clustered, unbranched, or once or twice branched, 12 -28 mm high, 0.5 - 1.5 mm wide, with or without scyphi. Scyphi narrow, 0.4- 1 mm across, closed, not proliferating; margins entire or dentate. Podetia grey-green, olivaceous, or brown in decorticate areas. Base more or less the same as upper portions. Podetial squamules absent, or occasional at base, mostly perpendicular to surface. Podetia sorediate. Soredia farinose, rarely granular, diffuse, covering podetia except for base, or completely covering podetia, absent from cup interior, or present within cups. Podetial wall entire. Apothecia rare, on podetial tips or on the cup margins, brown, strongly convex to hemispherical, 0.3-0.5 mm in diameter. Pycnidia on tips of podetia. CHEMISTRY: Thallus PD+ red, K-, UV-. Contains fumarprotocetraric and protocetraric acids, Cph-2, and often Cph-l. HABITAT: Mainly on rotting or mossy logs on the Charlottes, but elsewhere, also on mineral soil or bark. DISTRIBUTION: QCI: infrequent; scattered, at low elevations (Fig. 13). N. Am. : boreal -temperate, widespread. World: North America, Europe and Asia. SPECIMENS EXAMINED: QCI. Graham Island: Tow Hill, B-9895; Moresby Island: along the Deena River, B-10857B, Sunday Inlet, B-12201; Louise Island: Cumshewa Inlet, B-l 7931. Cladonia coniocraea was retypified by Ahti DISCUSSION: (1993) and proposed for conservation (T. Ahti, unpublished data). Thus, the citation of the authors "(Florke) Spreng.," which was replaced by "auct." by Ahti (1980b) for nomenclatural reasons and adopted by many authors (e.g., Egan 1987; Santesson 1984), can again be used. Although this species is very common throughout the Canadian forests, in the study area it is less common and certainly less abundant than the very similar Cladonia ochrochlora. Juvenile specimens of the two species can be very difficult to distinguish. Cladonia cornuta (L.) Hoffm. ssp. cornuta DESCRIPTION: Primary thallus squamulose, evanescent, or rarely persistent. Primary squamules horizontal, dispersed or scattered, 2-3 mm long, 2-5 mm broad, crenulate, medium olive-brown (95) to yellow-brown (74-75), undersurface white, ecorticate, esorediate. Podetia arising from center of squamules, scattered, clustered or dense, unbranched, or rarely once or twice branched, 25 -40 mm high, 12.2 mm wide, without or rarely with scyphi. Scyphi narrow, 1-3 mm across, closed, not proliferating; margins entire or dentate. Podetia shiny brown to olive-brown (95), or greyish yellowish green (122) when shaded, the decorticatk parts becoming brown (77, 74-75); becoming brown at tips. Base somewhat blackened. Cortex present over the greater part of the ~odetia.Podetial surface smooth where conicate.- odetial squamules absent or very rare. Podetia sorediate. Soredia farinose, in discrete patches on upper half of podetia, or rarely covering podetia except for base. Podetial wall entire. Apothecia not produced. Pycnidia not observed. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-. Contains fumarprotocetraric and protocetraric acids, Cph-2. HABITAT:row in^ on mineral soil or on wood, in full sun or in partial shade. DISTRIBUTION: QCI: Queen Charlotte Range at 700- 1000 m elev.; rare. N. Am.: boreal, widespread. World: North and South America, Europe, Asia, Australasia. SPECIMENS EXAMINED: Graham Island: Carew Bay Moun-

Can. J . Bot. Vol. 74. 1996


tain, 53" 18'N, 132"37'W, B-10246; Moresby Island: Tasu, 52"401N, 132"02'W, A-39029. Cladonia cornuta ssp. cornuta is a continental DISCUSSION: lichen, found only twice on the QCI. It differs from ssp. groenlandica in having tall, long subulate podetia, a blackened base, and in usually lacking primary squamules; ssp. groenlandica is paler and almost always has distinct cups and some granular soredia, and has large, persistent primary squamules. Cladonia cornuta ( L . ) Hoffm. ssp. groenlandica (A. E. Dahl) Ahti syn. Cladonia groenlandica (A. E. Dahl) Trass DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dispersed, 1-6 mm long, 2-8 mm broad, crenulate or deeply incised, dark greenish yellow to greyish greenish yellow (103-105), or very pale brownish, undersurface white, ecorticate, esorediate. Podetia arising from center of squamules, loose, scattered or clustered, unbranched, or once or twice branched, 12-40 mm high, 1-2.5 mm wide, with or rarely without scyphi. Scyphi narrow or irregular, somewhat "inflated," 1.5 -5 mm across, closed, not proliferating or proliferating from the margins of the cups; margins entire or dentate. Podetia yellow-green, olivaceous, or brown: same as squamules where corticate, more or less uniform or becoming brown at tips. Base more or less the same as upper portions. Cortex present. Podetial surface smooth, or showing some longitudinal ridges in a few specimens. Podetial squamules absent, or occasionally present at the base, mostly perpendicular to surface. Podetia sorediate. Soredia farinose or granular, in discrete patches very irregular in size, finally coalescing on upper half of podetia, or rarely covering podetia except for base, at least partly absent from cup interior. Podetial wall entire, 260 - 370 pm thick. Cortex 25-40 pm thick. Medulla 70- 160 pm thick. Stereome cylindrical, entire (about equal in thickness to outer medulla and cortex), 90- 190 pm thick. Apothecia rare, on the cup margins, brownish orange to dark brown (54-56), somewhat convex, or strongly convex to hemispherical, 1-2 mm in diameter. Pycnidia not observed. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-. Contains fumarprotocetraric and protocetraric acids, and Cph-2. HABITAT:Growing mostly on rotting, mossy logs, but sometimes on humus; in full sun, partial shade, or in deeply shaded habitats. DISTRIBUTION: QCI: frequent throughout the archipelago, mostly at low elevations (Fig. 14). N. Am.: west and east coasts. World: North America, Europe, Asia. SELECTED SPECIMENS EXAMINED: A-39005; B-10088B, 11487, 11791, 17514A, 18196, 23665, 26762. DISCUSSION:Cladonia cornuta ssp. groenlandica and Cladonia ochrochlora are extremely similar on the QC1 (see discussions in Ahti 1980a; Hammer and Ahti 1990). Both have tall, tapering, almost unbranched podetia, continuously corticate on at least the basal half in most specimens, and producing orbicular or irregular patches of soredia on the upper half. Cladonia cornuta ssp. groenlandica tends to be more frequently cupped, and the cups are broader, slightly

more regular or inflated, with the inner surface usually corticate, at least partially. The soredia can be farinose or granulose, but it is the coarseness of the soredia that is supposed to distinguish ssp. groenlandica from ssp. cornuta. The basal squamules often disappear but tend to be large (up to 10 mm broad) and crenulate to entire. Ahti ( 1 9 8 0 ~ )says that the inner cartilaginous medulla ("stereome") is thinner than the outer medullary and cortical tissues making the wall softer and easier to cut than in Cladonia ochrochlora, where the stereome is thick and tough, thicker than the outer tissues. This does not seem to be true in every case. Colour is also a character that has been used and perhaps is still the best of all the poor ways of separating the taxi. Cladonia ochrochlora tends to be pale yellow-green, to pale greyish yellowish-green, although it can "brown up" enough to be olive-brown (94) in some thalli. Cladonia cornuta s s ~ . . groenlandica, on the other hand, always tends to be brown at least in part, even when pale. According to Hammer (1991), Cladonia ochrochlora has basal squamules "entirerounded, with incisions less than '/z the length of squamule," but he has annotated specimens in H as Cladonia ochrochlora that have deeply incised squamules, exactly as he describes for his Cladonia cornuta ssp. groenlandica. Therefore, either the identifications are incorrect or the character is unreliable. Specimens of both Cladonia cornuta ssp. cornuta and ssp. groenlandica in H have thickish, almost entire squamules in many cases. Habitat and substrate are slightly different according to Ahti ( 1 9 8 0 ~ ) Cladonia . ochrochlora tends to be restricted to rotting wood, and Cladonia cornuta ssp. groenlandica is often found on mineral soil or over rocks as well as on wood. If the combination of colour, stereome thickness, and cup development fails, the species can be indistinguishable. Over most'of their total ranges, the taxa do not meet. In the study area, however, both of them are common. ,

Cladonia crispata (Ach.) Flot. var. cetrariifortnis (Delise) Vain. DESCRIPTION: Primary thallus squamulose, evanescent, or persistent. Primary squamules ascending or horizontal, dispersed, 1-2 mm long, 1-2 mm broad, crenulate, yellowgreen or olivaceous, undersurface white, brownish at base, ecorticate, esorediate. Podetia clustered or .dense, once or twice branched. Branching in dichotomies. Axils open, wide, not forming funnels, 30-45 mm high, 0.5- 1.4 mm wide, lacking scyphi. Podetia mottled, medium yellow-green (120) to olive-brown ( 9 3 , or becoming brown at tips (74-55) or on upper half; base paler. Cortex present. Podetial surface smooth, matt to weakly shiny. Podetial squamules occasional or abundant, very small, entire, mostly perpendicular to surface. Podetia without soredia or granules. Podetial wall perforated with oval holes. Apothecia not produced. Pycnidia not observed. CHEMISTRY: Thallus PD-, K-, UV+ bright blue-white. Contains squamatic acid and sometimes zeorin. HABITAT: Growing in heath at 800- 1000 m elevation, in full sun. DISTRIBUTION: QCI: San Cristoval Range; rare. N. Am.: west and east coasts; poorly known. world: North America, Europe, Asia, Australasia.

Brodo and Ahti


SPECIMENS EXAMINED: CANADA: BRITISH COLUMBIA: Moresby Island: near Mt. Laysen, E of Sunday Inlet, B-26843; Tasu, "Mine Mt.," 52"401N, 132'00-04'W, Juneau area, A-38992, 39013, 39036; U.S.A. : ALASKA: Gastineau Peak, B-26341A. DISCUSSION: Similar to Cladonia subjkrcata, which has a shiny brown colour and lacks podetial squamules, and is conspicuously melanotic at the base. Var. cetrariiformis is a very distinct ascyphose race in many oceanic or arctic regions, but its taxonomic status is insufficiently known. It is easily confused with Cladonia gracilis ssp. gracilis, which always reacts P D + orange-red rather than P-.

Cladonia crispata (Ach.) Flot. var. crispata syn. Cladonia japonica Vain. in Hue, Nouv. Archiv. Mus. Ser. 3, 10: 265. 1898, syn.nov. DESCRIFTION: Primary thallus squamulose, evanescent, or persistent. Primary squamules ascending or horizontal, dispersed, 1-4 mm long, 0.5- 1.5 mm broad, crenulate, or deeply incised, yellow-green, olivaceous, or brown (121, 95, 79, 57-55), undersurface white, or brownish at base, ecorticate, esorediate. Podetia clustered or dense, unbranched, or once or twice branched from cup or axil margins. Axils wide open, forming funnels, 15 -70 mm high, 0.6-3.0 mm wide, with, or rarely without scyphi. Scyphi narrow or irregular, 1-3 mm across, opening by a gaping hole, proliferating from the margins of the cups; margins dentate. Podetia yellow-green, olivaceous, or brown (121, 95, 79, 57-55), browner in crispata s.str., greener in japonica morph, mottled, or becoming brown at tips. Base more or less the same as upper portions, or blackened weakly melanotic. Cortex present. Podetial surface smooth; or areolate, with paler patches on a dark brown to black background. Podetial squamules absent, occasional, or rarely abundant, especially on base, mostly perpendicular to surface, ca. 2 X 0.5 mm. Podetia without soredia or granules. Podetial wall entire, or rarely longitudinally split, 160-250 pm thick. Cortex 20-40 pm thick; medulla 60- 120 pm thick; stereome cylindrical, entire, 40- 120 pm thick. Apothecia common, on the cup margins, greyish red-brown (46-47), flat, emarginate, or with a thin margin even with the disk, 0.15-0.8 mm in diameter. Pycnidia abundant, on margins of scyphi; pycnidial jelly red. CHEMISTRY: Chemotype 1: squamatic and often consquamatic acids (thallus PD-, K-, UV+ bright blue-white); chemotype 2: thamnolic acid, often with traces of decarboxythamnolic and barbatic acids (thallus PD orange, K persistent deep yellow, and UV-). HABITAT:Growing on mineral soil, over rocks, or on moss, generally in or around bogs, in full sun, or in partial shade. DISTRIBUTION: QCI: both chemotypes frequent, although the thamnolic acid chemotype is most common in the northeastern lowlands (Fig. 15). N. Am. : boreal, widespread. World: North and South America, Europe, Asia, Africa. SELECTED SPECIMENS EXAMINED: Chemotype l : A-38513; B-10286, 12781, 14117, 18126, 23680. Chemotype 2: B-9752, 9795 (Lich. Can. Exs. 27), B-12698,12293,17804, 18092, 18232B. Cladonia crispata is generally one of the more DISCUSSION:

+

+

easily identified and well-understood species of Cladonia, with its cuplike, gaping axils, frequently squamulose, but well corticate podetia, occasional branches, brownish colour, and production of squamatic acid. On the west coast of North America and eastern Asia, however, and in scattered localities elsewhere, a thamnolic acid producing strain exists that has caused a considerable amount of confusion. Most centers around the South American species, Cladonia carassensis Vain. and Cladonia subsubulata Nyl., described from Campbell Island in New Zealand; see discussions in Krog (1968), Stenroos (1988, 1993), Stenroos and Ahti (1990), Hammer (19936, 1 9 9 3 ~ ) The . latter species, which can look similar to some forms of Cladonia cris~ata. ' . differs in being generally more slender and taller, paler (i.e., less browned), lacking any primary thallus (i.e., having indeterminate growth), lacking podetial squamules, and being commonly subulate. The axils do not broaden enough to be called cups. Cladonia aueri Rasanen, described from Tierra del Fuego at the tip of South America (type seen in H), is identical with this material. Indeed, Archer and Bartlett (1986) and Stenroos (1993) sy nony mized Cladonia aueri under Cladonia subsubulata, although Stenroos and Ahti (1990) maintained them as distinct^ based on broader scyphi and the frequent presence of barbatic acid in the latter. Cladonia carassensis s.str. is short, broadly cupped, often with areoles or squamules developing in the inside surface of the cups, and has a persistent squamulose primary thallus. Cladonia japonica Vain. (syn. Cladonia carassensis ssp. japonica (Vain.) Asahina) seems to belong to the Asian western American, thamnolic-producing population of Cladonia crispata (see Huovinen and Ahti 1988). Cladonia pseudohondoensis Asahina, described from the mountains of Japan, is another thamnolic-producing member of this complex and seems to be closest to Cladonia subsubulata in its branched, cupless, smooth podetia, and apparently indeterminate growth. In his comparison of "Cladonia carassensis ssp. japonica" and Cladonia pseudohondoensis, however, Asahina (1959) regarded the former as "whitish" and the latter as "browner or variegate . . . ," which would not be in accord with conspecificity of pseudohondoensis and subsubulata. Judging from the type material (TNS) it is possible that Cladonia pseudohondoensis represents the thamnolic acid strain of Cladonia crispata var. cetrariiformis. Several specimens on the QC1 resemble Cladonia subsubulata very closely, having indistinct, oblique cups and producing barbatic as well as thamnolic acids. These specimens might be referable to Cladonia artuata S. Hammer, a species described from California (Hammer 1993b), but we prefer to recognize the British Columbia material as morphotypes of Cladonia crispata. Although it is possible that the barbatic acid is located exclusively in the apothecia, S. Hammer (personal communication) confirms that it occurs in all parts of the podetia. Barbatic acid was already reported for Cladonia crispata by Huovinen and Ahti (1988). Another related, more southern taxon is Cladonia poroscypha S. Hammer. It has frequent, slender, not-at-all divergent, dichotomous branches, lacks cups (although Hammer 19936 described it as "branching around scyphus-like openings"), and frequently bears podetial squamules. It is known as far north as southern Vancouver Island (Hammer 1 9 9 3 ~ ) .

1160

Can. J . Bot. Vol. 74. 1996

Table 1. Some thamnolic acid containing members of Cladonia sect. Perviae in the Pacific Northwest of North America, compared with the Japanese C. pseudohondoensis. Cladonia crispata var. crispata (thamnolic strain) Distribution

California -Alaska

Colour

Scyphi (cups)

Olivaceous; often strongly browned Mostly unbranched except from cup margins Regular to oblique

Axil openings

Wide open

Lateral perforations

Uncommon; rarely split

Podetial squamules

Absent or occasional, rarely abundant Smooth, continuous to areolate


Branching

Podetial cortex

Podetial height (mm) Podetial width (mm) Podetial uniformity

15-70 0.6-3.0 Strongly dilated at axils and cups

Cladonia pseudohondoensis

Cladonia nrtuata

Cladonia poroscypha

California - Vancouver Island Greenish or olive to chestnut brown Unbranched except from cup margins

California - Vancouver Island Yellowish grey, browned only at tips 1-3 times, di- or trichotomies

Japan

Irregular, from dilated axils Wide open, sometimes sieve-like Common and conspicuous Infrequent, especially close to apices Thin and breaking up, areolate

Rare or absent

Absent

Often closed, or narrow and irregular Absent

Wide open, oblique

Small, frequent, or sometimes absent Mostly continuous, or breaking up if squamulose (6-)14-45 0.5-2.0 Quite uniform and slender

Infrequent (?)

(5-)12-75 Up to 6 Very irregular

The distinctions between some of these thamnolic acid producing taxa is given by Hammer (1993~)in a key, and we summarize the data, adding other taxa, in Table 1.

Cladonia ecmocyna Leight. ssp. occidentalis Ahti, ssp. nov. Fig. 16 Sicut subspecies ecmocyna sed podetiis semper tenuioribus (0.5- 1.6 mm crassis), pallide glaucescentibus, apice facile violascente fuscescentibus, escyphosis vel scyphis angustis, esquamosis vel ad basin versus parce squamosis. TYPE: U.S. A. : Washington: Skamania Co. : along eastern edge of Big Lava Bed in Gifford Pinchot National Forest, 10 km NW of Willard, 45"511N, 121°43'W, 750 m, on detritus, 1991, Nash 29914, Nash: Lich. Exs. ASU 106 (holotype, H; isotype, CANL). DESCRIPTION: Primary thallus squamulose, evanescent, or rarely persistent. Primary squamules ascending or horizontal, dispersed, 1 - 3 mm long, 0.5 - 1.5 mm broad, crenulate, medium to greyish yellow-green (120-122), undersurface white, ecorticate, esorediate. Podetia arising from center of squamules, clustered, often dense, unbranched or very irregularly once or twice branched. Axils closed. Podetia 30-80 mm high, 0.5- 1.6 mm wide, mostly lacking scyphi, or with infrequent, oblique (1.0 -3.6 mm) scyphi. Podetia light to greyish yellow-green (121-122, 120), dark yellowbrown at tips, or whitish to greenish grey, with bluish hue caused by pruinosity. Necrotic base turning yellowish grey or pale grey, rarely blackened (see Discussion). Cortex present. Podetial surface smooth to white-checkered. Podetial squamules absent or occasional, especially close to base. Podetia without soredia or granules. Podetial wall entire or

Ashy brown, variegated Not, o r infrequently branched

Common (?)

Continuous, smooth to areolate 30-80(- 100) 1.0-2.5 Slightly dilated at axils

rarely perforate (see Discussion). Apothecia often produced along the cup margins, dark brown. Pycnidia common, often solitary at tips of podetia, pyriform, constricted at base. CHEMISTRY: Thallus PD+ red, K + pale yellow, UV-, or UV + dull. Contains fumarprotocetraric and protocetraric acids, atranorin, and Cph-2. HABITAT: Usually growing on mossy rocks near sea level in lowland areas over most of its range, but all Queen Charlotte collections were on mountains above 700 m and may occur at elevations up to 1500 m in Washington. DISTRIBUTION: QCI: infrequent, at high elevations (Fig. 17). N. Am.: western, mainly coastal, mid British Columbia to Oregon (Fig. 18). World: North American endemic. SELECTED SPECIMENS EXAMINED: CANADA: BRITISH COLUMBIA: QCI: Moresby Island: 2.5 km SW of Tasu, 800- 1000 m, 1980, Ahti 39037 (CANL, H); also from QCI: Moresby Islands: B-10918, 11040, 12813, 23622, 26716, 26770. Non-QCI: Kitimat, Mt. Claque, 850 -900 m, 1970, Ohlsson 2776 (H); S of Powell R., 15 mi NE of Stillwater, 1350 m, 1969 Ohlsson 929 (H); Vancouver Island: Sutton Pass between Port Alberni and Ucluelet, 280 m, 1983, Goward 83-263 (H, UBC), 400 m, 1984, A-42976 & Noble (H); Little Qualicum Falls Prov. Park, 1961, A. & A-15328 (CANL, H, UBC); Goldstream, Mt. Wells, 1958, A-14965 (H). Gulf Islands: Saltspring Island, Mt. Maxwell Prov. Park, 600 m, 1968, B-13843 & Schoj?eld (CANL). Garibaldi Prov. Park, 360 m, 1966, B-8187 (CANL, H). U.S.A.: MONTANA: Flathead Co.: 12 mi NE of West Glacier, Mt. Cannon, 1100 m, 1985, DeBolt 542 (H); OREGON: Lane Co.: hwy. 242, 2-5 mi S of hwy. 126, McKenzie Bridge, 1989, Hammer 3598 (FH, H); WASHINGTON: What-

Brodo and Ahti


Fig. 16. Cladorzia ecrrzocyna ssp. occidentalis, isotype, Nash 29914 (CANL). Scale in millimetres.

com Co.: Mt. Baker National Forest, Sulfur Creek Lava Flow, ca. 500 m, 1992, Ahti & Rhoades 5101 7, 51020 (H); hwy. 20, East Creek Trail, 1500 m, 1989, Hammer 4070 (FH, H). Numerous specimens from Idaho, Montana, Oregon, and especially Washington are listed by Hammer ( 1 9 9 3 ~ )as " Cladonia ecmocyna, unnamed subspecies." DISCUSSION: This taxon was included in ssp. intermedia by Ahti ( 1 9 8 0 ~ )based on preliminary observations. His field studies on the QCI, Vancouver Island, and Washington State, and the examination of collections by Hammer, indicated that there is a third subspecies to be recognized in Cladonia ecmocyna. It essentially embraces the temperate lowland populations in Pseudotsuga forests and elsewhere at lower elevations of the Pacific coast referred to by Ahti (1980a), but it ascends up to 1500 m on coastal mountains. On the U.S. side (Hammer 1993a), from central Oregon to Puget Sound and east to the Rocky Mts., it is even more common than ssp. intermedia. The new subspecies resembles ssp. ecmocyna in being largely ascyphous, but it differs in having virtually no podetial squamules and tends to be thinner (see Table 2). Subspecies intermedia, which is often without podetial squamules and is about the same size, is typically richly cupped and is much more conspicuously pruinose. These and other subulate Cladoniae of the gracilis group are compared in Table 2. Unlike the other subspecies, which are extremely variable in morphology, ssp. occidentalis is often very uniform, espe-

cially on old lava beds in various localities in Washington, where it can be a dominant component of the vegetation. Hammer ( 1 9 9 3 ~ )reported that in a few localities where he saw ssp. intermedia and ssp. occidentalis growing together, they were morphologically distinct. On the other hand, he considered that intergradation between them may occur, and therefore they are treated here as subspecies. Cladonia fimbriata (L.) Fr. DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dense or dispersed, 1-5 mm long, 1 -6 mm broad, crenulate or deeply incised, medium to pale yellow-green (120- 12 l), becoming brownish, undersurface white, ecorticate, esorediate. Podetia arising from center of squamules, scattered or clustered, unbranched; 6-23 mm high, 0.5-2 mm wide, with narrow (trumpet-shaped) scyphi, rarely broad and goblet shaped, 3-8 mm across, closed, not or rarely proliferating from the margins of the cups; margins entire or dentate. Podetia pale yellow-green (121), or pale olivaceous to greyish olive (109) where corticate; yellowish white to yellowish green where sorediate, more or less uniform. Base more or less the same as upper portions. Cortex absent except at very base. Podetial squamules absent. Podetia sorediate. Soredia farinose or rarely granular, diffuse, covering podetia except for base or completely covering podetia, present within cups. Podetial wall entire. Apothecia rare, on the cup margins, stalked, brown, somewhat convex, or strongly convex to hemispherical, finally compound. Pycnidia relatively scarce, black, on cup margins, pyriform, not or slightly constricted at base. CHEMISTRY: Thallus P D + red, K- (or brownish), UV-. Contains fumarprotocetraric and protocetraric acids, and Cph-2. Cph-l can be present or absent. HABITAT: Growing on mineral (sandy) soil, but usually on wood, especially logs, in full sun or in partial shade. DISTRIBUTION: QCI: infrequent, mostly in lowland Graham Island (Fig. 19). N. Am. : primarily boreal- temperate, widespread. World: North and South America, Europe, Asia, Australasia, Antarctica. SPECIMENS EXAMINED: B-10714, 11143, 12076, 18185, 18188, 26660. (affin. fimbriata): A-391 10; B-9975, 9969, 12509. DISCUSSION: Cladonia fimbriata is much less common on the Charlottes than on the mainland. Unlike the "typical" Cladonia fimbriata, the dominant morphotype is not truly farinose sorediate but rather somewhat granulose, especially towards the base, and the bare medulla easily turns brown. The Queen Charlotte population includes a variant, possibly a distinct species, with elongate podetia but wide cups (="affin. fimbrata" above). It tends to produce large, rounded basal squamules that turn bluish grey in exposed localities. It is also known from Vancouver Island and Del Norte Co., California. This morphotype deserves closer study in the field. Cladonia fircata (Huds.) Schrad. DESCRIPTION: Primary thallus squamulose, evanescent, or persistent. Primary squamules ascending, dispersed, 25 mm long, 1-2.5 mm broad, deeply incised, grey-green or yellow-green, undersurface white, ecorticate, esorediate.

Table 2. Esorediate, mostly subulate members of the Cladot~iagracilis group in western North America.

A 2

m


Cladonia rnaxirna

Cladonia gracilis ssp. elongata

Cladonia gracilis ssp. vulnerata

Presence in west

Rare, coastal; possibly absent

Rare on coast, more continental

Common in coastal region

Atranorin

Rare

Present in small quantity or absent

Very rare

Colour of necrotic base of podetia

Yellow

Strongly melanotic

Height (mm) Width (mm) Perforations in podetia

(30-)70- 120(- 180) 1-3 Infrequent

Cups

Usually present on some podetia, oblique, 1 -6 mm

(30-)70- loo(- 150) 0.8-2 Scattered lateral holes, slits, or cracks; sometimes abundant Lacking to common, 1-2 mm, narrow

Weakly or not at all melanotic (i.e., brownish, grey) 25-65(- 140) 1 -2(-3) Common, conspicuous, especially slits at base Lacking or sparse, irregular

Podetial squamules

Exceedingly rare

Branching (no. of times) Pruinosity

0(-3) None

General colour

Greenish to greyish green, hardly browned

None, or a few close to base O(- 1) None

Dark, brownish

None 0-1 None

Yellowish green to greenish grey, browned

Cladonia ecmoc)~na ssp. ecn~ocyna

Cladonia ecmocyna ssp. inrertnedia Common, esp. at high elevations

Cladonia ecmocynn ssp. occidenralis Common in coastal region, esp. at low elevations Present in large quantity

Probably absent, or rare at high elevations Present in large quantity

Present in large quantity

Yellow

Yellow

Yellow or grey (to melanotic)

60- 100 2 -4 Rare

30-60(- 100) 0.5 -4 Rare or absent

25 -SO(- 80) 0.5-1.6 Usually absent

Frequent, 0-3 per podetium, 2-5 mm, with secondary scyphi Sometimes scattered, esp. at base (0-)l-3 Minutely white pruinose, esp. at tips, or epruinose Greyish to greenish grey

Very frequent, 1.5- 10 mm, with secondary scyphi, but sometimes absent Usually densely squamulose up to top 0(-2) Heavily white pruinose over most of podetium Greyish to greenish grey

Infrequent, very narrow (1-3.5 mm)

Esquamulose except at base W- 1) Very slight, sometimes apparently absent Brownish to greenish grey, dark, or bluish grey

N


Brodo and Ahti Podetia loose, clustered or dense, once or twice branched, or much branched especially on upper half of podetia; branching isotomic, in dichotomies. Axils partially or wide open, not forming funnels. Podetia 25 - 120 mm high, 1-3 mm wide, lacking scyphi. Podetia pale yellow-green to quite dark brown and shiny, more or less uniform or mottled, becoming brown at tips. Base more or less the same as upper portions or somewhat paler. Cortex mainly continuous, but can become discontinuous and areolate above. Podetial surface smooth or areolate. Podetial squamules occasional or abundant, mostly perpendicular to surface. Podetia without soredia or granules, but decorticate, areolate areas at tip can sometimes appear to be granular. Podetial wall longitudinally split and perforated. Apothecia not produced, or rare, on podetial tips, brown. Thallus P D + red, K- (or brownish), UV-. CHEMISTRY: Contains fumarprotocetraric and protocetraric acids, Cph-2, and sometimes Cph- l . HABITAT: Growing on mineral soil or on moss, in full sun, in partial shade, or in deeply shaded habitats. DISTRIBUTION: QCI: common, mainly at low elevations throughout the islands, rarely in subalpine heath (Fig. 20). N. Am.: southern boreal to temperate, western and eastern regions, but not transcontinental. World: North and South America, Europe, Asia, Australasia. SELECTED SPECIMENS EXAMINED: A-39052; B-9734, 9824, 10001B, 11103A, 12795, 14207, 14327 (Lich. Can. Exs. 105), 17216, 17434, 23683, 26776. DISCUSSION: Cladonia furcata is a central species of a problematic complex in the Pacific Northwest. Other members are ~ l a d o n i a scabriuscula, Cladonia multiformis G. Merr., Cladonia herrei Fink ex J. Hedrick, and Cladonia macroptera Rasanen. They were discussed by Noble (1982), Brodo et al. (1987), and most recently by Hammer ( 1 9 9 3 ~ ) . Cladonia furcata is very common, but its morphological variability does not entirely match either eastern North American or European populations, which tend to be highly branched, uniformly corticate, and often abundantly squamulose. Cladonia scabriuscula is also present along the Pacific coast, but as elsewhere where Cladonia fircata and Cladonia scabriuscula are sympatric, intermediate morphs can be found having decorticate, microareolate tips difficult to classify as sorediate or nonsorediate. In the QCI, there are robust, little-branched morphs producing large, laciniate squamules and having slightly eroded tips. Such a morph was reported as Cladonia macroptera by Brodo et al. (1987) as new to North America. However. we are now of the opinion that before the taxonomy of c l a d h a furcata is treated on a worldwide basis, it is prudent to treat Cladonia macroptera (in Canada, as well as the scrappy type material from Japan) as a luxuriant morph of Cladonia furcata of moist, shady habitats. Cladonia scabriuscula can also produce very similar, luxuriant morphs (e.g., Ahti 38991), and the possibility that Cladonia macroptera actually represents Cladonia scabriuscula is not totally excluded. It seems clear that Cladonia multiformis G. Merr., which can sometimes closely resemble Cladoniafurcata, is essentially lacking on the coast, as pointed out by Hammer ( 1 9 9 3 ~ ) . If the Pacific morphs of Cladonia furcata are to be recognized taxonomically, it should be noted that the type material of Cladonia herrei (a name used by Noble 1982) has

an uncertain disposition (Hammer 1993b), and the name may not be applicable at all in this group. Cladonia gracilis (L.) Willd. ssp. elongata (Jacq.) Vain. syn. Cladonia gracilis ssp. nigripes (Nyl.) Ahti (See Ahti 1980a for citations.) DESCRIPTION: Primary thallus squamulose, evanescent. Podetia clustered or dense, unbranched, or rarely once or twice branched; branching in dichotomies. Axils closed. Podetia 20-45 mm high, 0.6-1 mm wide, without scyphi or infrequently with some narrow scyphi, 1-3 mm across, closed, not proliferating or proliferating from the margins; margins dentate. Podetia yellow-green, or olivaceous, or yellowish brown, more or less uniform or mottled, or becoming brown at tips. Base strongly melanotic. Cortex present. Podetial surface smooth, verruculose or areolate. Podetial squamules absent or occasional, mostly perpendicular to surface. Podetia without soredia or granules. Podetial wall longitudinally split, or with small and infrequent, oval perforations. Apothecia rare, on the cup margins, brown, somewhat convex or strongly convex to hemispherical, 0.5 - 1 mm in diameter. Pycnidia on margins of scyphi. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-. Contains fumarprotocetraric and protocetraric acids, Cph-2, and usually atranorin in small amounts. HABITAT: Growing on mineral soil, directly on rock, or on moss, in full sun. DISTRIBUTION: QCI: rare, high elevations (Fig. 21). N. Am.: arctic to northern boreal, widespread. World: North and South America, Europe, Asia, Antarctica. SPECIMENS EXAMINED: CANADA: BRITISH COLUMBIA: QCI: Graham Island: Carew Bay Mountain, on exposed mossy rock, elev. 700 m, Brodo 10246; Moresby Island: mountain at W end of Mosquito Lake, tundra ridge top, Schoj?eld 25461 (UBC, CANL). U.S.A. : ALASKA: Juneau area, Gastineau Peak, alpine ridge on ground, Brodo 26346. DISCUSSION: This widespread, arctic -bored and antarcticantiboreal race was called ssp. nigripes by Ahti ( 1 9 8 0 ~ ) As . the result of subsequent studies (Stenroos and Ahti 1990), it became clear that the Fuegian type material of ssp. elongata is indistinguishable from ssp. nigripes, and the former name was reinstated. It is easily identified when the podetia are rather robust (although they are never as robust as those of ssp. vulnerata) and strongly melanotic at base, with many subulate tips, and when they contain atranorin. One specimen from the QCI, and one from southeast Alaska, both collected at the timber line, are included here, although they do not contain atranorin. Similar specimens from lower elevations are usually clearly perforated and are hardly melanotic; they are included in ssp. vulnerata. Cladonia gracilis ssp. turbinata (Ach.) Ahti DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending, dense or dispersed, 1 -5 mm long, 1-5 mm broad, deeply incised, yellow-green, olivaceous, or brown (like podetia), undersurface white, or brownish at base, ecorticate, esorediate. Podetia scattered or clustered, unbranched or once or twice branched; branching in dichotomies. Axils closed. Podetia 20-40 mm high, 0.6-1.5 mm wide, with broad or narrow scyphi, 3-8 mm across, closed or partially perforate, especially on the outside


Can. J. Bot. Vol. 74, 1996 -

subsp. occidentalis

C. furcata

20 I l

21 ,

% '

0

C. gracilis

I I

Brodo and Ahti


Figs. 17-25. Distribution of species of Cladonia on the Queen Charlotte Islands. Figs. 17 and 18. Cladonia ecmocyna ssp. occidentalis. Fig. 17. Queen Charlotte Islands distribution. Fig. 18. World distribution. Fig. 19. Cladoniafimbriata. Fig. 20. Cladonia&rcata. Fig. 21. Cladonia gracilis. a, ssp. elongata; m, ssp. turbinata; 0 , ssp. vulnerata. Fig. 22. Cladonia kanewskii. Fig. 23. Cladonia macilenta. Fig. 24. Cladonia metacorallifera. Fig. 25. Cladonia norvegica, western North American distribution.

rim of the cups, proliferating from the margins of the cups, rarely from the centers; margins entire or dentate. Podetia greyish olive (109) or greyish greenish yellow (105) becoming pale brownish, rarely dark brown, more or less uniform or mottled. Base more or less the same as upper portions. Cortex present. Podetial surface smooth. Podetial squamules occasional or abundant, mostly perpendicular to surface. Podetia without soredia or granules. Podetial wall entire. Apothecia common, on the cup margins, brown, somewhat convex or strongly convex to hemispherical, 0.5-3 mm in diameter. Pycnidia on margins of scyphi. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-. Contains fumarprotocetraric and protocetraric acids. HABITAT:Growing on mineral soil, wood, or moss, in full sun. DISTRIBUTION: QCI: very rare, sea level (Fig. 21). N. Am.: boreal to temperate, widespread. World: North America, Europe, Asia. SPECIMEN EXAMINED: QCI: Graham Island: near Port Clements, on a log in a salt marsh, Brodo 18156. DISCUSSION: Although this subspecies is widespread and circumpolar, it is rarely encountered in highly oceanic areas, and therefore its rarity on the Charlottes is understandable. Cladonia gracilis (L.) Willd. ssp. vulnerata Ahti DESCRIPTION: Primary thallus squamulose, evanescent. Podetia loose, scattered or clustered, unbranched or rarely once or twice branched; axils closed; 25 - loo(- 180) mm high, 0.6-2(-5) mm wide, with or without scyphi. Scyphi rare, narrow, 2-5 mm across, closed, not proliferating or proliferating from the margins of the cups; margins dentate. Podetia pale to medium yellow-green (119-122) to olive (106) or dark brown to brownish green, more or less uniform or mottled, or becoming brown at tips. Base more or less the same as upper portions, or sometimes weakly melanotic or brown (not yellowish). Cortex present. Podetial surface smooth, without squamules, soredia, or granules. Podetial wall longitudinally split or perforated with wide open oval perforations; typical material: total wall 120- 190 pm, cortex 25-40 pm, medulla 50- 100 pm thick; stereome 40-90 pm thick; robust morphotype: total wall 140250 pm, cortex 20 -50 pm, medulla 60 - 160 pm, stereome 30-60 pm thick; medulla not clearly differentiated from stereome. Apothecia rare, on the cup margins, brown, somewhat convex or strongly convex to hemispherical, 0.51 mm in diameter. Pycnidia on margins of scyphi. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-, or UV+ dull. Contains fumarprotocetraric and protocetraric acids, and Cph-2, and sometimes Cph-l. HABITAT: Growing on moss or in heath, often boggy, in full sun. DISTRIBUTION: QCI: frequent and widespread, at low and high elevations (Fig. 21). N. Am.: west coast, Alaska to Washington. World: North America, eastern Asia.

SELECTED SPECIMENS EXAMINED: A-39069; B-10509, 10770, 10819, 10983, 11998A, 12271, 12669, 17673, 17801, 26740, 26781, 26955; Schojield 24297 (UBC). DISCUSSION: Ahti ( 1 9 8 0 ~ )described this taxon, with an Alaskan type, to accommodate the essentially coastal population~of Cladonia gracilis in the Pacific Northwest that produce perforate, rather stout podetia with pale or slightly blackened base and no atranorin. It can be particularly difficult to distinguish this subspecies from Cladonia maxima (Asahina) Ahti, which is typically a uniform greenish grey or grey-green to grey becoming yellow at the necrotic bases, not strongly browning, usually producing some well-formed, narrow cups, and is not particularly perforate. Cladonia maxima rarely has the bent and otherwise contorted podetia characteristic of Cladonia gracilis ssp. vulnerata, and the latter rarely produces cups. The QC1 specimens with very stout and tall podetia were first referred to Cladonia maxima, but they appear to represent extremely well-developed Cladonia gracilis ssp. vulnerata. More field work is required, however, for an analysis of their differences in various habitat conditions. Cladonia maxima is a suboceanic species that is definitely known from southeast Alaska (e.g., Thompson Pass, Chugach Mts., Ahti 23636: H, WIS) and Vancouver Island (e.g., Parson's Mountain, Macoun 274C: CANL), but its distribution along the coast is unclear because of confusion with Cladonia gracilis ssp. vulnerata. The British Columbia records of Cladonia maxima in Ahti ( 1 9 8 0 ~ need ) confirmation, and at present there is no definite record from the QCI, though the species is expected to occur there.

Cladonia cfr. homosekikaica Nuno DESCRIPTION: Primary thallus squamulose, evanescent, or persistent. Primary squamules ascending, dense or dispersed, 1 -3 mm long, 1- 1.5 mm broad, entire or crenulate, thick, rounded, olivaceous; undersurface white, ecorticate, esorediate. Podetia scattered or clustered, unbranched, 10-45 mm high, 1-4 mm wide, with broad or narrow scyphi, 3-5 mm across, closed, not proliferating or proliferating from the margins of the cups; margins dentate, irregular. Podetia grey-green, yellow-green, brownish, or grey (79) (greyer in Alaska). Colour of upper part of podetia more or less uniform. Colour of podetial base more or less the same as upper portions. Cortex present or absent. Podetial squamules occasional or abundant, especially at base, mostly perpendicular to surface. Podetia sorediate; soredia farinose or granular, diffuse or in discrete patches mainly confined to tipof podetia, or covering podetia except for base, rarely completely covering podetia; present within cups. Podetial wall entire. Apothecia rare, on the cup margins on short proliferations, strong brown (55), strongly convex to hemispherical, 2.54 mm in diameter. Pycnidia not observed. CHEMISTRY: Thallus PD+ red, K-. Contains fumarprotocetraric and homosekikaic acids.


Can. J. Bot. Vol. 74, 1996

HABITAT: Growing on mineral soil in full sun. DISTRIBUTION: QCI: absent! (See below.) N. Am.: rare, Alaska, Illinois (McKnight et al. 1987). World: North America, Europe, Asia, Australasia (Archer 1983). SPECIMEN EXAMINED:U.S.A.: ALASKA: Tongass National Forest, Chatham area, north of Juneau at mouth of Herbert Glacier, on moss covered rocks and gravel, elev. 30 m, Derr & Geiser L-3293 (TNFS, CANL). DISCUSSION: Although not part of the Queen Charlotte flora, Cladonia homosekikaica is being included in this treatment because of its discovery in nearby Alaska and because of its rarity. This specimen is only tentatively called Cladonia homosekikaica because of its chemistry. It may actually represent an undescribed species, but our material is too meager to demonstrate it conclusively. In her discussion of Cladonia homosekikaica, Nuno (1975) regarded it as morphologically resembling species in the Cladonia chlorophaea group, but those species generally have coarsely granular soredia or corticate granules covering the cups. The true Cladonia homosekikaica is actually very close to Cladonia fimbriata, if not its chemotype. Our single coastal specimen (from southeast Alaska) deviates by producing rather tall, slender podetia, with fairly narrow scyphi and patches of cortex inside the scyphi; otherwise, it is heavily farinose sorediate. It is, in fact, more like Cladonia ochrochlora but is more robust and sorediate. Cladonia kanewskii Oksner DESCRIPTION: Primary thallus evanescent. Podetia dense, much branched, but side branches short; branching anisotomic, in dichotomies; axils closed or partially to wide open, not forming funnels. Podetia 10-70 mm high, 1.5-7 mm wide, lacking scyphi; pale greenish yellow (104), becoming brown or greyish red-brown at the tips, with tiny colourless crystals forming on older specimens, base more or less the same colour as upper portions. Cortex present. Podetial surface smooth to irregular and bumpy. Podetial squamules absent. Podetia without soredia or granules. Stereome broadly fibrose, almost entire. Apothecia not produced. Pycnidia not observed with certainty. CHEMISTRY: Thallus PD-, K-, UV-. Contains usnic acid, f isousnic acid, and some unidentified triterpenes and fatty acids. HABITAT: Growing on mineral soil, in heath, or directly on rock mainly on exposed bluffs and outcrops, in full sun. DISTRIBUTION: QCI: Moresby Island at high elevations (Fig. 22). N. Am.: northwestern, mainly coastal. World: North America, Europe, Asia. SPECIMENS EXAMINED: A-39012, 39097; B-10839, 10923, 10984,12028,12790,26712,26817; Horton 1563 (CANL); Schofield 24940, 249 70 (UBC). DISCUSSION:Cladonia kanewskii was regarded as an Amphiberingian radiant, which extends down to the Vancouver region along the Pacific coast (Ahti and Brodo 1981; their record from Vancouver not shown on their published map!). It is abundant in some localities on the QCI, but very few other records are known of this species south of Alaska. It was recently detected on the coast of central Norway as new to Europe (specimens in BG, H).

Cladonia macilenta Hoffm. syn. Cladonia bacillaris Genth (see discussion below) DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dense or dispersed, 1-1.5 mm long, 0.8- 1 mm broad, entire or crenulate, rounded, thick, convex, greenish grey to grey, undersurface white, ecorticate, esorediate or occasionally sorediate; soredia granular, marginal. Podetia arising from center of squamules, scattered, clustered or dense, unbranched, clavate to pin-shaped, 10-20 mm high, 1 - 1.5 mm wide, lacking scyphi, grey-green or grey, more or less uniform. Base more or less the same as upper portions. Cortex present at base of some podetia, but on most, being replaced by soredia. Podetial surface smooth, or verruculose on thickly corticate parts of base. Podetial squamules occasional at base, strongly decumbent or peltate, rarely perpendicular; rounded and often convex. ~ o d e t i asorediate; ~ o r e d i afarinose to granular, diffuse, covering podetia except for base, or completely covering podetia. Podetial wall entire. Apothecia rare, on podetial tips singly, or in a ring at the tip of the blunt podetium, red, strongly convex to hemispherical, 0.3-0.5 mm in diameter. Pycnidia on tips of podetia. CHEMISTRY: Chemotype 1: contains thamnolic and decarboxythamnolic acids, sometimes with barbatic acid (type) (thallus PD+ yellow to orange, K + persistent deep yellow, UV-); or chemotype 2: barbatic, f 4-0-demethylbarbatic, squamatic (trace) acids (= "bacillaris" chemotype) (PD-, K-, UV+) (not found on QCI). HABITAT: Growing on mineral soil, directly on rock, or on wood, especially logs and fence rails, in full sun. DISTRIBUTION: QCI: infrequent, eastern lowlands (Fig. 23). N. Am.: widespread. World: North and South America, Europe, Asia, Australasia, Africa. SPECIMENS EXAMINED (all chemotype l): QCI: Graham Island: S of Tlell, spruce forest and adjoining field close to shore, on exposed logs, B-9741; 17 km N of Port Clements, nuja-Tsuga forest, very open, on exposed roots at roadside, B-9815; Moresby I.: Sandspit, along roadsides, on exposed fence post, B-10086, B-12441 (= Lich. Can. Exs. 104). Christensen (1987) concluded that the thamDISCUSSION: nolic acid strain (Cladonia macilenta s.str.) and the barbatic acid strain (Cladonia bacillaris) are morphologically indistinguishable. We agree, although other authors (Stenroos 1986; Hammer 1993a; Tonsberg 1994) claimed that they are distinguishable in some regions. In our area, Cladonia macilenta is not particularly common and is exclusively represented by the thamnolic acid strain. The barbatic acid chemotype is rare in British Columbia. Cladonia macilenta may be difficult to distinguish from Cladonia umbricola, but it never forms cups even at maturity and has thicker, less finely divided basal squamules. Regarding the author citation of Cladonia bacillaris, see Ahti (1993).

+

Cladonia metacorallifera Asahina DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules mostly horizontal, dense or dispersed, 0.7-2 X 0.7-2.5 mm, entire or crenulate, quite thick, somewhat convex, rounded, pale greenish yellow (104-102);


Brodo and Ahti maculate, undersurface white, or brownish at point of attachment to rhizoid bundle entering substrate, ecorticate, esorediate. Podetia arising from center of squamules, loose or clustered, unbranched, 10-26 mm high, with broad or sometimes narrow scyphi, 3-7 mm across, closed, not proliferating; margins entire. Podetia yellow-green, same as primary squamules where corticate; yellowish white elsewhere, more or less uniform. Base more or less the same as upper portions. Cortex present over most of podetium or podetial squamules. Podetial surface verruculose or areolate to granulose-squamulate. Podetial squamules occasional or abundant, strongly decumbent, convex, at first closely adnate to podetial surface. Podetia with spherical, corticate granules mainly confined to tip of podetia or on upper half of podetia, or rarely covering podetia except for base, present within cups. Podetial wall entire. Apothecia rare, on the cup margins, red, somewhat convex or strongly convex to hemispherical to linear along cup margin, 0.5-3.5 mm in diameter. CHEMISTRY: Thallus PD-, K-, UV+ bright blue-white. Contains usnic, didymic, and squamatic acids; condidymic, barbatic, and 4-0-demethylbarbatic acids. HABITAT: Growing on mineral soil, or on mossy rocks, in full sun. DISTRIBUTION: QCI: rare; at high elevations in the San Cristoval Range (Fig. 24), but collected in a lowland bog in SE Alaska. N. Am.: western (especially Alaska, Yukon, NWT: map in Thomson 1984), and Newfoundland and New Brunswick in the east; arctic to boreal, suboceanic. World (map in Stenroos 1989b): North America, Europe, Asia. SPECIMENS EXAMINED: QCI: Moresby Island: San Cristoval Range, "Monday Mountain," on soil at summit, elev. 945 m, B-26845; Tasu Mountain, subalpine meadows, sandy ridge, elev. 930- 1000 m, A-38981a, 38545, B-14267; 3 km SE of Tasu, 0-50 m, A-39058. DISCUSSION: This species is expected to be widespread along the Pacific coast although there are still few reports or specimens of it.

+

Cladonia norvegica Tonsberg & Holien DESCRIPTION (based on QC1 material; see discussion below): Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dense or dispersed, 1-3 mm long, 2-4 mm broad, deeply incised or rarely crenate, pale yellow-green (134) or brownish (105, 88, 74), undersurface white, or dark yellow to orange at the base, KOH + reddish purple, ecorticate, esorediate, or sorediate. Soredia granular, marginal or on undersurface. Podetia arising from center of squamules, loose or scattered, unbranched, or rarely once or twice branched. Axils closed. Podetia 10-30 mm high, 0.5 - 1 mm wide, lacking scyphi. Podetia pale yellowish to yellow-green (134-1 19) or brown where corticate; variegated brown in part where sorediate or more or less uniform. Base more or less the same as upper portions. Cortex present on lower 10-50%; absent where sorediate. Podetial surface smooth where corticate. Podetial squamules absent, or occasional at base, mostly perpendicular to surface. Podetia sorediate. Soredia farinose, diffuse or rarely in discrete patches, on upper half of podetia, covering podetia except for base, or rarely completely covering

podetia. Podetial wall entire. Apothecia rare, on podetial tips, waxy pale brown, flat, 0.3-2 mm in diameter; in one specimen, dark to pale brown, strongly convex to hemispherical. Pycnidia on tips of podetia. CHEMISTRY: Thallus PD-, K-, UV+ bright blue-white. Contains barbatic and 4-0-demethylbarbatic acids, and rarely traces of squamatic acid. HABITAT: Growing on bark, usually on conifers such as Picea, or on mossy wood, in partial shade at low or high elevations. DISTRIBUTION: QCI: rare (see below). N. Am. : west coast, Alaska to Washington (Fig. 25). World: North America, southern South America, Europe, Asia. SPECIMENS EXAMINED: CANADA: BRITISH COLUMBIA: QCI: Graham Island: Port Lewis, 53'41 'N, 132'55'W, along Coates river and adjoining shore, B-10490B; Moresby Island: Mt. Moresby, 53'011N, 132'05'W, elev. 640 m, B-26746. U.S.A.: ALASKA: Juneau area, Auke Bay, along shady wet trail, B-26231; Tongass National Forest, Ketchikan area, Prince of Wales Island, old growth stand, slope with large spaces between trees and tall dense understory, on moss on Tsuga heterophylla, elev. 60 m, Geiser & Derr L-3297 (TNFS, CANL) . DISCUSSION: Although originally thought to be restricted to Norway (Tonsberg and Holien 1984), Cladonia norvegica is a widespread oceanic lichen now known from Europe, western and eastern North America, and southern South America (Tonsberg and Goward 1992, and references therein) and Japan (Stenroos and Ahti 1994). It was collected in southeast Alaska (see above) and was reported from the Prince Rupert area just east of the Charlottes (Tonsberg and Goward 1992). There are two additional specimens from the Hazelton, B.C. area in CANL (Goward L1 6864, L1 6895) (see Fig. 25). The two specimens from the Charlottes differ in some respects from the typical populations and may represent a distinct taxon. They form soralia that are brownish in part, they produce traces of squamatic acid in addition to the abundant barbatic acid, and the basal squamules are thicker and less finely divided. In addition, the apothecia on the fertile specimen (B-26746) are large and much darker brown than normal. Nevertheless, we are provisionally placing them under Cladonia norvegica pending the collection of better material. Cladonia novochlorophaea (Sipman) Brodo & Ahti, comb.nov. syn. Cladonia merochlorophaea Asahina var. novochlorophaea Sipman, Acta Bot. Neerl. 22: 496. 1973. DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dense, 1-2.5 mm long, 1 - 1.5 mm broad, crenulate, thick, pale green to yellowish green to olive-brown or yellowish brown, undersurface white, ecorticate, esorediate. Podetia clustered or dense, unbranched or 1-2 times branched, 5-7 mm high, 1.5-3 mm wide, with broad scyphi, 1.5-7 mm across, closed, proliferating from the margins of the cups; margins entire. Podetia yellow-green, olivaceous, or brown, more or less uniform, or becoming brown at tips. Base more or less the same as upper portions. Cortex present. Podetial surface


Can. J. Bot. Vol. 74, 1996

areolate, or with rounded, peltate squamules. Podetial squamules occasional or abundant, strongly decumbent or peltate. Podetia without soredia or granules. Podetial wall entire. Apothecia common, dark brown, 0.5-2.5 mm in diameter, on the cup margins, often on short thick stipes, strongly convex. Pycnidia often scarce, inconspicuous, on cup margins, black, sessile. CHEMISTRY: Thallus PD-, K-, UV+ weak blue-white. Contains sekikaic and homosekikaic acids. HABITAT: Growing on mossy rocks in full sun. DISTRIBUTION: QCI: Queen Charlotte Range, high elevation; very rare. N. Am.: western (see below). World: North and South America, Europe, Australasia, Antarctica. SPECIMEN EXAMINED: QCI: Moresby Island: Mt. Moresby, 53"001N, 132"05'W, on rock on ridge top, elev. 760 m, B-26694. DIsCuss~oN:Although commonly regarded as a chemotype of Cladonia rnerochlorophaea, there is no need to regard it as closer to that than to Cladonia cryptochlorophaea, as noted by Paus (1992). It is therefore recognized here as a distinct species. Leuckert et al. (l97 1) and Paus (1992) actually informally used the epithet novochlorophaea at the species level. It is chemically distinct by producing homosekikaic and sekikaic acids and only rarely fumarprotocetraric acid. In spite of some claims (e.g., Ahti 1966; Holien and Tonsberg 1985) that it is morphologically indistinguishable from Cladonia rnerochlorophaea s.str., it seems to be at least as distinct in morphology as the other members of the group. It is always very decidedly nonsorediate and nongranulose, although verrucose and strongly areolate - corticate throughout, whereas Cladonia rnerochlorophaea can be granulose or even sorediate. On the other hand, we admit that the status of the taxon may change, since the "Cladonia chlorophaea group" is undergoing intense study, even at the genetic level (e.g., Culberson et al. 1988; DePriest 1994) and is still very imperfectly understood on a worldwide basis. The North American distribution of Cladonia novochlorophaea is very incompletely known. In addition to the west coast (e.g., Hennings 1983), it is perhaps only known from the Great Slave Lake region (Culberson and Kristinsson 1969). In Europe, southern South America, and Antarctica (Stenroos 1993), Cladonia novochlorophaea is very common in some regions, often in cool-oceanic coastal regions. Along the west coast, Cladonia rnerochlorophaea s.str. is known from the Juneau area in SE Alaska (Krog 1968) and Vancouver Island, but not from the QCI. Cladonia ochrochlora Flijrke DESCRIPTION:Primary thallus squamulose, persistent. Primary squamules ascending, dispersed, 3 - 8 mm long, 3 - 11 mm broad, crenulate or deeply incised, greyish greenish yellow (105) or pale yellow-green (121), undersurface white, dark yellow to orange (and KOH+ red-violet), or brownish at base, ecorticate, esorediate. Podetia arising from center of squarnules, loose, scattered or clustered, unbranched, 10-35 mm high, 0.5-2 mm wide, with or without scyphi. Scyphi irregular, 1-3 mm across, closed, not proliferating; margins entire, or dentate. Podetia, including the base, greyish greenish yellow (105), pale yellow-green (121), or olive brown (94), more or less uniform, but whitish where decorticate. Cortex present usually for about half the length

of the podetium. Podetial surface smooth. Podetial squamules absent, or occasional (at base), mostly perpendicular to surface. Podetia sorediate. Soredia farinose or granular, diffuse or in discrete patches on upper half of podetia, or covering podetia except for base, absent from cup interior. Podetial wall entire, 230-550 pm thick; cortex 25-40 pm thick; medulla 50-70 pm thick; stereome cylindrical, entire, very thick and hard, more than half of entire wall, 120450 pm thick. Apothecia rare, on podetial tips or on the cup margins, brown, somewhat convex, or strongly convex to hemispherical, 1-2 mm in diameter. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-. Contains fumarprotocetraric and protocetraric acids, Cph-2, and usually Cph-l as well. HABITAT: Growing on wood, mostly on rotting logs, in full sun, partial shade, or in deeply shaded habitats. DISTRIBUTION: QCI: frequent on Moresby and adjacent islands, less common on Graham Island (Fig. 27). N. Am. : boreal to temperate, incompletely known. World: North and South America, Europe, Asia, Australasia, Africa, Oceania. SELECTED SPECIMENS EXAMINED: A-38975; B-9925A, 10823, 10885, 10992A, 12050A, 12280, 17514B, 18313, 2671 7. DISCUSSION: See Discussion under C. cornuta. Cladonia phyllophora Hoffm. s.lato DESCRIPTION: Primary thallus squamulose, evanescent, or persistent. Primary squamules ascending or horizontal, scattered, 2-4 mm long, 1-4 mm broad, crenulate or deeply incised, light yellowish green (134-135) to light green (144), somewhat scabrose. undersurface white or brownish black at point of attachment, ecorticate, esorediate. Podetia arising from center of squamules, scattered or clustered, unbranched or once or twice branched. 15 -50 mm high. 0.5 -2.5 mm wide, with or without scyphi. Scyphi irregular, 1-2.5 mm across, closed, proliferating from the margins of the cups. Podetia yellow-green, or brown to black between areoles (like primary squamules where corticate), mottled. Base strongly melanotic (blackened) and "spotted" with pale corticate areoles. Cortex present as areoles or absent below. Podetial surface dull, appearing tomentose when welldeveloped, areolate, with occasional podetial squamules, mostly perpendicular to surface, entire to crenulate; without soredia or granules. Podetial wall longitudinally split or perforated, 200-300 pm thick; cortex 30-35 pm thick; medulla 50-80 pm thick; stereome cylindrical, entire, 80-200 pm thick. Apothecia not produced, or rare, brown. Pycnidia not observed. CHEMISTRY: Thallus PD+ red, K- or brownish, UV-. Contains fumarprotocetraric and protocetraric acids, Cph-2, and occasionally atranorin. HABITAT: Growing on sandy soil. DISTRIBUTION: QCI: northern San Cristoval Range; very rare. N. Am.: widespread, low arctic to boreal for the species as a whole; Washington to Yukon and Alaska, and subarctic Asia for this morphotype (see below). World: circumboreal, southern South America, Subantarctic Islands (Stenroos 1993). SPECIMENS EXAMINED: CANADA: BRITISH COLUMBIA: Moresby Island: near Mt. Laysen, E of Sunday Inlet, 52"411N, 131°50'W, at summit, elev. 945 m, B-26839. U

.

Brodo and Ahti


U.S.A.: ALASKA:Juneau, near Mendenhall Glacier, East Glacier Trail, on sandy ground under Alnus sinuata, B-26062. DISCUSSION: This is a slender, often ascyphous morphotype of Cladonia phyllophora. It is common in Alaska (Thomson and Ahti 1994) and Siberia and may be a distinct species. It is morphologically very similar to Cladonia alaskana A. Evans but is lacking usnic acid. Cladonia pleurota (Florke) Schaer. DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dispersed, 2 - 6 mm long, 2 - 8 mm broad, crenulate or deeply-incised, yellowgreen (122) to greyish greenish yellow (105), undersurface white, or dark yellow to orange at base, ecorticate, esorediate. Podetia arising from center of squamules, scattered or clustered, unbranched, or rarely once or twice branched from sides of podetia, 6-25 mm high, with broad scyphi relative to podetial length, 2 - 8 mmacross, closed, not or rarely proliferating from the margins of the cups; margins entire, or dentate, or rarely squamulose. Podetia greenish yellow (104-105) to yellow-green (1 19-121), more or less uniform. Base more or less the same as upper portions. Cortex present at least at very base, sometimes for much of podetium. Podetial surface verruculose, or areolate where corticate. Podetial squamules occasional, mostly perpendicular to surface. Podetia sorediate, or with spherical, corticate granules. Soredia granular, diffuse, on upper half of podetia, or covering podetia except for base, sometimes confined to edge of cups, present within cups. Podetial wall entire. Apothecia common, on the cup margins, red, strongly convex (hemispherical), 1-4 mm in diameter. ~ ~ c n i dbina margins of scyphi. CHEMISTRY: Thallus PD-, K-, UV-. Contains usnic and isousnic acids and zeorin. After several years, fine needle crystals are formed on the thallus surface; they are thought to be zeorin. HABITAT: Growing on mineral soil, directly on rock, or on moss, in full sun, or in partial shade. DISTRIBUTION: QCI: northern Graham Island; very rare, as it is elsewhere along the British Columbia coast. N. Am.: boreal-temperate, continental, widespread. World: North and South America, Europe, Asia, Australasia, Antarctica (see Stenroos 1989a for map). SPECIMEN EXAMINED: ~ r a h a mIsland: Naden Harbour on the west shore, 54"001N, 132"38'W, on wood, B-10691D (pp) (with C. umbricola). DISCUSSION: Cladonia pleurota is a widespread, sorediate species very close to Cladonia coccifera (Stenroos 1 9 8 9 ~ ) . - -

Cladonia prolijca Ahti & S. Hammer DESCRIPTION: Primary thallus squamulose, persistent, or evanescent. Primary squamules ascending and curled over, showing undersurface colour, dispersed, or scattered, 28 mm long, 0.5 -4 mm broad, entire, or crenulate, or rarely deeply incised, ligulate, yellowish brown to olive brown (74-94), undersurface white to bluish grey, or brownish at base, ecorticate, esorediate. Podetia scattered or clustered, usually unbranched except from cup margins, or rarely branched once or twice, 15-35 mm high, 1-4 mm wide,

with irregular or regular scyphi, 1.5-5 mm across, closed or rarely partially perforate, usually flat, repeatedly proliferating from the margins of the cups; margins dentate or squamulose. Podetia dark yellow (88) to light yellowish brown (76) or yellowish white (92), mottled. Base more or less the same as upper portions. Cortex present. Podetial surface smooth at base, or verruculose to areolate above. Podetial squamules occasional or abundant, strongly decumbent. Podetia without soredia or granules. Podetial wall entire or perforated, 180-235 pm; cortex 8- 18 pm; medulla 63 -88 pm; stereome 88 - 130 pm thick. Apothecia rare (not on QC1 material), on the cup margins, brown, 1-3 mm in diameter. Pycnidia on margins of scyphi, brownblack, strongly constricted at the base. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-. Contains fumarprotocetraric acid and traces of protocetraric acid and Cph-2. HABITAT: Growing on mineral soil over rocks or sand dunes in lowland localities, in full sun. DISTRIBUTION: QCI: scattered; rare. N. Am.: west coast, California to Queen Charlottes. World: North America, Europe (see below). SPECIMENS EXAMINED: Moresby Island: 3 km SE of Tasu, 52"401N, 132"001W,A-39090 (H); Portland Bay, 52"48'N, 132" 11'W, B-14078; Graham Island: White Creek Bog area (E of Tow Hill, 54"04'N, l 3 1"43'W), Schojield 23792, with Vaarama (UBC, CANL) . DISCUSSION: Hammer and Ahti (1990) described this "nondescript" species from material collected on the coast from California to Vancouver Island. The present records represent a considerable extension of the range. It resembles Cladonia phyllophora Hoffm., but the senescent parts do not turn black, the cortex is better developed, only slightly arachnoid, and the cups develop characteristic proliferations. The species has recently been reported from Spain and Greece in Europe (Burgaz and Ahti 1994), so that it should no longer be considered as a North American endemic. Cladonia pyxidata (L.) Hoffm. DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending, dense or dispersed, 1-7 mm long, 0.5-4 mm broad, entire or crenulate, thin to somewhat thick, greyish olive (109) to olive-brown (94) or yellow brown (74), undersurface white or grey, ecorticate, esorediate. Podetia arising from center of squamules, scattered, unbranched, 7- 16 mm high, with broad or rarely narrow scyphi, 3-7 mm across, closed, not proliferating, or rarely proliferating from the margins of the cups; margins entire. Podetia olivaceous, or brown, same as primary squamules or somewhat greyer where decorticate, mottled. Base more or less the same as upper portions. Podetial surface smooth, or areolate at base. Podetial squamules occasional, or abundant, mostly perpendicular to surface, or peltate. Podetia without soredia or granules. Podetial wall entire. Apothecia not produced, or rare, on the cup margins, brown. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-. Contains fumarprotocetraricand protocetraric acids and Cph-2. HABITAT: Growing on mineral soil, rock, wood, or moss, in full sun. DISTRIBUTION: QCI: infrequent, mainly in lowland localities in the NE sector (Fig. 28). N. Am.: arctic to temperate,

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Can. J . Bot. Vol. 74, 1996

Figs. 27-35. Distribution of species of Cladonia on the Queen Charlotte Islands. Fig. 27. Cladonia ochroch10ra. Fig. 28.

Cladonia pyxidara. Fig. 29. Cladonia scabriuscula. Fig. 30. Cladonia schofieldii. Figs. 31 and 32. Cladonia squamosa. Fig. 31. Chemotype 1 (squamatic acid). Fig. 32. Chemotype 2 (thamnolic acid). Fig. 33. Cladonia rranscendens. Figs. 34 and 35. Cladonia umbricola. Fig. 34. a , chemotype 1 (squamatic acid); A , chemotype 2 (squamatic and usnic acids). Fig. 35. a, chemotype 3 (thamnolic acid); A , chemotype 4 (thamnolic and usnic acids).


Fig. 26. Cladonia schofieldii. (a) Large primary thallus, Ahri 38996 (H). (b) Podetium on primary thallus, holotype, Ahri 38989 (H). (c and d) Smaller primary thalli, holotype. ~. Scale bar = 10 mm.

widesuread. World: North and South .America, Europe, Asia, .~ustralasia,Africa, subantarctic islands. SPECIMENS EXAMINED: Graham Island: Kumdis Bay, NE of Port Clements, on moss on tree base in open salt marsh, B-18154; Moresby Island: Gray Bay, on rocky ledge on beach, B-12605; on shaded rock at edge of woods, B-12607; Sandspit, sandy pasture and road bank, A-391 13, A-391 15; Tasu, rock outcrop by sea, A-38543. ,

Cladonia scabriuscula (Delise) Nyl. DESCRIPTION: Primary thallus squamulose, evanescent, or persistent. Primary squamules ascending, dispersed, 2 -5 mm long, 1-2.5 mm broad, deeply incised, grey-green or yellow-green, undersurface white, ecorticate, esorediate.

Podetia loose or clustered, up to four-times branched; branching isotomic in dichotomies; at least some axils open, often incomdetelv. Podetia 20-65 mm high, 1 - 1.5 mm wide, lacking x i p h i . Podetia grey-green, or very light yellowish green (134-121), especially at base or in shady habitats, white to yellow-brown on decorticate areas, especially at tips, mottled, becoming brown at tips. Base paler green or yellowish green. Cortex present in patches, absent on at least upper third to half of podetia, and sometimes almost entirely decorticate. Podetial surface smooth, or areolate where corticate. Podetial squamules occasional or abundant, mostly perpendicular to surface. At least some podetia sorediate; soredia granular, diffuse, or in discrete patches, mainly confined to tip of podetia. Podetial wall longitudinally split, or perforated. Apothecia rare, on podetial tips, brown. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-. Contains fumarprotocetraric and protocetraric acids and Cph-2. HABITAT: Growing on mineral soil or moss, or mossy logs, in full sun, partial shade, or in deeply shaded habitats. DISTRIBUTION: QCI: frequent, at coastal localities throughout the islands, rarely at high elevations (Fig. 29). N. Am.: boreal to temperate, with oceanic tendencies. World: North and South America, Europe, Asia, Australasia. SELECTED SPECIMENS EXAMINED: A-38512, 38991, 39112; B-9762, I O44OB, 12181, 14038, 26587, 26777; Schofield 23797, 24190 (UBC). DISCUSSION: It is interesting that both Cladonia scabriuscula and Cladoniafurcata are found almost exclusively at or close to the shoreline on the Charlottes. In the east, these species are found in a variety of shaded or sunny habitats, none particularly associated with lakes or the sea, although at least Cladonia scabriuscula has oceanic tendencies (Krog 1968). (Also see Discussion under Cladonia furcata.)

Cladonia schofieldii Ahti & Brodo, sp.nov. Fig. 26 Thallus primarius persistens, squamosus; squamae 4 -50 mm longae, 1 - 5 mm latae, incisae; superficie pallide virides, scabridae, subtus albae vel cinerascentes sed apice violascentes et basin versus melanoticae, passim corticatae; podetia pallide cinerea, sparsa, simplicia, scyphifera, scyphis irregularibus, corticata, superficie laevia. Pycnidia terminalia. Apothecia non visa. Acidum fumarprotocetraricum et interdum etiam atranorinam continens. TYPE: British Columbia. QCI: Moresby Island, Tasu Sound, ca. 2.5 km SW of Tasu, N slope of the summit of "Mine Mountain", alt. 800- 1000 m, timber-line grassland with Tsuga mertensiana scrub, 1980, Ahti 38989 (H, holotype; CANL, UBC, isotypes). DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending, dense, 4 -50 mm long, 1-5 mm broad, deeply incised (lobate to branched in the best developed squamules), light green (143) to light yellowish green


Brodo and Ahti

C. chemotype squamosa k 2

32

C

I

transcendens

I

I

1 I

B


Can. J. Bot. Vol. 74. 1996

(135-136), surface scabrose; undersurface white, greenish white (153), grey (265-266) at melanotic base, or entirely brown (77) to purplish violet (214); ecorticate or commonly corticate, sometimes developing an algal layer; esorediate. Podetia rarely present, scattered, unbranched, 5-10 mm high, 0.5- 1.5 mm wide, with irregular scyphi, 1.5-3 mm across, imperforate, flat, brown, not proliferating; margins dentate or squamulose. Podetia yellow-green (same as upper surface of squamules). Colour of upper part of podetia more or less uniform. Colour of podetial base more or less the same as upper portions or somewhat melanotic. Cortex present. Podetial surface very smooth or areolate. Podetial squamules occasional, mostly perpendicular to surface. Podetia without soredia or granules. Podetial wall entire. Apothecia rare, on the cup margins, brown. Pycnidia on margins of scyphi, scarce. CHEMISTRY: Thallus PD+ red, K- or K + pale yellow. Chemotype 1 (most common): fumarprotocetraric and protocetraric acids (type); chemotype 2 (infrequent): fumarprotocetraric acid and abundant atranorin (no protocetraric acid). Convirensic acid (Cph-l) and confumarprotocetraric acid (Cph-2) or other unidentified trace compounds are sometimes present as accessories in all chemotypes. Cph-l, however, was only found twice. Huovinen et al. (1990, p. 235, as "Cladonia sp. 2") reported 2.2% of secondary products consisting of fumarprotocetraric acid and trace amounts of protocetraric acid, Cph-2, and an unknown compound detected with high performance liquid chromatography (HPLC). HABITAT: Growing on mineral soil in rocky heath, or directly on rock, in full sun. DISTRIBUTION: QCI: rare, mostly at high elevations in the Moresby Range, once at sea level on the east side of the islands (Fig. 30). N. Am.: QC1 and coastal southern Alaska. World: western North American endemic. ADDITIONAL SPECIMENS EXAMINED: Chemotype l: QCI: Tanu Island: on mossy CANADA: BRITISH COLUMBIA: rocks, sea level, B-17490; Moresby Island: Takakia Lake, talus slopes and subalpine meadows, in heath, 600 m, B-10944; Tasu, timber-line grassland, 800-1000 m, A-38501 & Schofield (H), A-38996 (H). U.S.A. : ALASKA: Prince William Sound, Knight Island, small tundra, 1940, Eyerdam 308 p.p. (with Cladina rangiferina) (FH). ChemoQCI, Mt. Moresby, type 2: CANADA: BRITISH COLUMBIA: open subalpine meadow and talus slope, on boulder, 670 m, B-26764. U.S.A.: ALASKA: Tongass National Forest, Petersburg Ranger District, mainland, Thunder Mt. E of summit, elev. 1036 m, Geiser 656 (TFNS). DISCUSSION: With its very large, elongate, often lobate and even branched primary squamules, Cladonia schofieldii is easily recognized in the field. Based on podetial morphology and chemistry, the new species should be placed in sect. Cladonia. Indeed, the large basal squamules and the grey colour would suggest Cladonia rnacrophyllodes Nyl., but the podetia do not seem to have the ability to produce central proliferations, and atranorin is infrequently produced. Podetia are rare, but when present, show clear affinities to Cladonia cervicornis (Ach.) Flot. The large squamules and chemistry also bring to mind Cladonia turgida Hoffm., especially in the chemical strain containing both fumarprotocetraric acid and atranorin. An important feature of Cladonia schofieldii, present in almost all specimens, is the partially corticate lower surface of the squamules, which can even

develop a secondary vegetative layer containing algae. Cladonia schofieldii was published as "Cladonia sp. 2" by Huovinen et al. (1990) in a chemotaxonomic screening of Cladonia. The report is based on the specimen Ahti & Schofield 38501 (H), collected on the same mountain as the type material. The former material has almost no podetia and was therefore less suitable as a type, though it came from a site where the species was forming a large patch in a rocky mountain meadow and was recognized as an undescribed species in the field because of its very large squamules. The presence of the new species in the eastern and central Pacific Coast districts of Alaska (e.g., near Petersburg and near the Kenai Peninsula) as well as on the Charlottes makes it likely that it will be found elsewhere along the cool Pacific coast, perhaps including the Aleutians. The species is named in honour of W.B. Schofield, Vancouver, who has made many important contributions to our knowledge of the phytogeography and bryology of the QCI, and whose keen eye in the field makes him one of our most outstanding collectors of bryophytes and lichens. Cladonia squamosa Hoffm. sy n. Cladonia squamosa var. subsquamosa (Nyl. ex Leight.) Vain. DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dispersed or scattered, 1-3 mm long, 0.5 -3 mm broad, deeply incised, thin, pale to greyish yellow-green (121-122), undersurface white, or brownish at base, ecorticate, esorediate. Podetia arising from center or edges of squamules, clustered or dense, unbranched or once or twice branched. Axils open, wide, not forming funnels. Podetia 10-70 mm high, 1-3 mm wide, with or without scyphi. Scyphi narrow, 2.5 -3.5 mm across, opening by a gaping hole, proliferating from the margins of the cups; margins dentate or squamulose. Podetia yellowgreen (121) where squamulose, brown or white where decorticate, more or less uniform, mottled, or becoming brown at tips. Base more or less the same as upper portions. Cortex absent from most of podetium. Podetial surface smooth, verruculose, or areolaie and decorticate. Podetial squamules abundant, mostly perpendicular to surface, or strongly decumbent, tending to be narrow and small, even granulose (microsquamulose). Podetia without soredia or granules, but granulose microsquamules often present, mainly confined to tip of podetia. Podetial wall perforated. Apothecia common, on podetial tips, or on the cup margins, pale to dark reddish brown, flat at first, later somewhat convex or hemispherical and often clustered, 0.4- 1.5 mm in diameter. Pycnidia on tips of podetia, or on margins of scyphi. consquamatic CHEMISTRY: Chemotype 1: squamatic acids (type chemotype) (thallus PD-, K-, UV+ bright bluewhite); chemotype 2: thamnolic and decarboxythamnolic acids (including "var. subsquamosa") (thallus PD + orange, K+ bright yellow, UV-). HABITAT: Growing on mineral soil, rock, bark, wood, or moss; the thamnolic acid chemotype is mainly on rotting wood, and is especially associated with swamps; in full sun or in partial to deep shade. DISTRIBUTION: QCI: although the squamatic acid chemotype is common and widespread through the archipelago at various elevations (Fig. 31), the thamnolic acid chemotype seems to be largely restricted to eastern lowland localities

+


Brodo and Ahti (Fig. 32). N. Am.: arctic to southern temperate, widespread. World: North and South America, Europe, Asia, Australasia, Africa, Antarctica. SELECTED SPECIMENS EXAMINED: Chemotype 1: B-9921, 10516, 12021, 12224, 14301, 18488, 13969, 26778 (Lich. Can. Exs. 233); Skidegate, Spreadborough 83158 (CANL); Schojield 23 724 (UBC). Chemotype 2: A-39064; B-9739A, 11983, 12964A, 17372, 17746, 17983; Schojield 24233 (UBC) . DISCUSSION: There are many species of Cladonia having alternative thamnolic versus squamatic acid chemotypes (especially in the Cocciferae, for example; see Discussion under Cladonia bellidiflora and Cladonia umbricola). There are no morphological distinctions between the chemotypes of Cladonia squamosa, and although they show slightly different habitat preferences and distributions on the Charlottes (see above), both are widely distributed on the Pacific coast. We therefore do not recognize the thamnolic acid chemotype as a named variety. The population containing thamnolic acid, however, is largely coastbound here as it is in Europe and elsewhere. Hammer ( 1 9 9 3 ~ )maps the distribution of both chemotypes for northwestern United States. The QC1 material of both strains, like material from other parts of the Pacific Northwest, often becomes entirely decorticate, translucent, and finally melanotic, covered with microsquamules. Hammer ( 1 9 9 3 ~ segregated ) the squamatic acid containing Cladonia singularis S. Hammer (in Washington State) from this group on the basis of the way the podetia develop from the squamules ("phyllopodially from the distal tip of the squamules"), the more frequently closed axils, and the tendency to have subulate podetia. The only Canadian material of it that we have seen is from Vancouver Island (Sutton Pass, 400 m, 1984, Ahti 42969 & Noble (CANL, H, UBC); new to Canada). Cladonia subfurcata (Nyl.) Arnold DESCRIPTION: Primary thallus squamulose, evanescent. Podetia loose or clustered, unbranched or rarely once or twice branched; branching isotomic in dichotomies; axils partially open. Podetia 20-35 mm high, 0.7- 1.5 mm wide, lacking scyphi but occasionally with open, cuplike axils. Podetia olivaceous, or brown (107) to dark brown, mottled with black areas. Base strongly melanotic. Cortex present over most of podetia. Podetial surface smooth and shiny. Podetial squamules absent, or rarely with a few basal squamules. Podetia without soredia or granules. Podetial wall longitudinally split or perforated. Apothecia not produced. Pycnidia not observed. Thallus PD-, K-, UV bright blue-white. CHEMISTRY: Contains squamatic acid + zeorin. HABITAT: Growing on the ground in heath and in sloping, Pinus contorta - Chamaecyparis nootkatensis fen, in full sun. DISTRIBUTION: QCI: NW Graham Island; very rare. N. Am.: arctic to northern boreal. World: North America, Asia, Europe. SPECIMEN EXAMINED: QCI: Graham Island: Eden Lake, 53"5l 'N, 132"44'W, B-12952.

+

Cladonia symphycarpia (Florke) Fr. DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending, dense, 2 - 8 mm long, 1.5 6 mm broad, deeply incised, curled up showing under-

surface, pale to greyish yellow-green (1 19- 121, 122), undersurface white to brownish at base, ecorticate, esorediate. Podetia absent or rare, arising from edge of squamules, unbranched or little branched, 15 mm high, 0.5 - 1 mm wide, lacking scyphi; yellow-green where corticate, or white elsewehere, more or less uniform. Base more or less the same as upper portions. Cortex absent except for scattered corticate verrucae. Podetial surface verruculose or areolate, with scattered convex areolae, almost granule size. Podetial squamules absent. Podetia without soredia or granules. Podetial wall longitudinally split or deeply grooved. Apothecia uncommon, on podetial tips (light brown primordia on primary thallus!), brown, somewhat convex, or strongly convex to hemispherical, 2.5 mm in diameter. Pycnidia on primary thallus, globular to subpyriform, constricted at base. Thallus PD-, K + pale yellow, UV-, or UV+ CHEMISTRY: dull. Contains atranorin, a fatty acid, and two unidentified compounds. HABITAT: Growing on mineral soil or directly on rock, in full sun; is generally regarded as an indicator of rich, usually calcareous soil. DISTRIBUTION: QCI: northern San Cristoval Range; very rare. N. Am.: boreal, widespread. World: North and South America (Stenroos and Ahti 1990), Europe, Asia, Australasia, Africa. SPECIMEN EXAMINED: Moresby Island: Tasu, "Mine Mountain," 52"40fN, 132"03'W, subalpine, A-39026 (CANL., H). , DISCUSSION: Cladonia cervicornis ssp. cervicornis, when epodetiate, is similar to Cladonia symphycarpia but has smaller squamules and contains fumarprotocetraric acid. Through most of its range Cladonia symphycarpia also contains norstictic acid, but in western North America, there is a strain having only atranorin (and large squamules, helping to distinguish it from Cladonia cariosa (Ach.) Spreng.) (Huovinen et al. 1989b). The author citations follow Ahti (1993), but the spelling must be corrected to symphycarpia to replace "symphycarpa," which is based on the illegitimate (superfluous) Lichen symphycarpus Ach. Cladonia transcendens (Vain.) Vain. Fig. 36 DESCRIPTION: Primary thallus squamulose, persistent. Primary squamules ascending or horizontal, dense or dispersed, 2-7 mm long, 1-5 mm broad, deeply incised, yellow-green, often maculate, or discoloured to shades of yellow or olive; undersurface white, or pale yellowish to dark yellow to orange (skyrin) at the dying base, ecorticate, esorediate. Podetia arising from the center, or more commonly from the edge of squamules, scattered or clustered, unbranched or once or twice branched. Podetia 15 -40 mm high, 0.8-2.5 mm wide, with or without scyphi, rarely lacking cups entirely. Scyphi narrow or irregular, rarely broad, 1.5 -4 mm across, closed, proliferating from the margins of the cups; margins dentate. Podetia yellow-green, discoloured to yellow or brownish in places, more or less uniform. Base more or less the same as upper portions, or more frequently coloured a deeper yellow or orange. Cortex present, but breaking down into squamules and soredia. Podetial squamules occasional or abundant, mostly perpendicular to surface. Podetia sorediate, or with spherical, corticate granules. Soredia granular or rarely farinose, diffuse, on upper half of podetia, occasionally only at tips, or more rarely covering

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Fig. 36. Clador~iutranscendens, proposed conserved type, Brodo 13003 (CANL). ( a ) General habit. Scale in millimetres. (6) Detail of sorediate portions. Scale bar = 5 mm.

podetia except for base, present within cups. Podetial wall entire, or longitudinally split, 190-250 pm thick; cortex 30-70 pm thick. Apothecia common, on the cup margins, red, flat, somewhat convex, or strongly convex to hemispherical, 0.5-3 mm in diameter. Pycnidia on margins of scyphi. Thallus PD orange, K persistent deep CHEMISTRY: yellow, UV-. Contains thamnolic and decarboxythamnolic acids, usually with usnic acid; skyrin produced at the dying base. HABITAT: Growing on conifer bark, or on wood, especially logs, in full sun, partial shade, or in deeply shaded habitats. SELECTED SPECIMENS EXAMINED: A-39067; B13003 (proposed conserved type), 13968, 14311B, 17120, 17950, 17990, 23618; Tasu Inlet, Christie, s.n. (UBC). DISTRIBUTION: QCI: frequent at low elevations throughout the islands, widespread (Fig. 33). N. Am.: west coast, Alaska to California. World: North American endemic. DISCUSSION: Cladonia rranscendens is central in a complex of red-fruited taxa including the morphologically and chemically diverse species Cladonia umbricola and Cladonia bellidiflora. In its more typical or unambiguous state, it forms robust, usually fertile podetia with abundant granular soredia and podetial micro- and macro-squamules. It often produces scyphi, usually narrow relative to the length of the podetia, which have a tendency to emerge from the edges of the large, deeply incised, primary squamules. Podetial cortex is usually well-developed and often forms plates with slightly raised margins, especially at the base.

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Unfortunately, Cladonia transcendens is a very variable species. When it is sparsely sorediate and abundantly squamulose, it strongly resembles the thamnolic acid containing strain of Cladonia bellidiflora. Being unaware of the latter, Evans (1951) and many collectors confused the two species. The thamnolic acid chemotype of Cladonia bellidiflora is common on the QC1 but is perhaps rare elsewhere in the Pacific Northwest (see Discussion under Cladonia bellidiflora). In fact, the original type material of Cladonia transcendens seems to represent the same chemotype (cited above under Cladonia bellidiflora), and Ahti (1993) attempted to neotypify Cladonia transcendens with a new specimen from the QC1 (Brodo 13003). The proposal, however, must formally be repeated with a conserved type in accordance with the Tokyo Code (T. Ahti, unpublished data). If the conservation proposal is not accepted, the species must be called Cladonia sipeana Gyeln., which certainly belongs to the present species (type: Oregon, Lane Co., Coburg Hills near Eugene, 193, Sipe 690 (US, isotype)). Cladonia transcendens can, however, vary in the other direction as well. Some specimens are abundantly sorediate with at least some of the soredia becoming fine, and on some specimens, podetial squamules are hardly produced. Here, we must rely on the extent and thickness of the cortex, place of origin of the podetia, and general stature to distinguish the species from Cladonia umbricola (see Discussion below). Cladonia umbricola TGnsberg & Ahti DESCRIPTION: Primary thallus squamulose, persistent.


Brodo and Ahti Primary squamules ascending or horizontal, dense or dispersed, 1.5-7 mm long, 2 -7 mm broad, deeply incised and lobulate, usually very thin, greenish yellow to greyish greenish yellow or olive (103-105, 109-120-121), undersurface white, pale yellowish to dark yellow or orange, or brownish at the base, ecorticate, esorediate or sorediate. Soredia farinose or granular, on undersurface close to margins. Podetia arising from center of squamules, loose, scattered or clustered, unbranched or rarely branched once, 6-27 mm high, 0.5-2 mm wide, with or without scyphi. Scyphi narrow, 1 -3.5 mm across, closed, not proliferating or proliferating from the margins of the cups; margins entire or dentate. Podetia yellow-green or greenish yellow (119-121, 104), or yellowish white where decorticate. Base more or less the same as upper portions, or darker, even weakly melanotic, when corticate. Cortex absent on podetia. Podetial squamules absent, or occasional at very base, mostly perpendicular to surface. Podetia sorediate. Soredia farinose or less frequently granular, diffuse, covering podetia except for base, or rarely completely covering podetia, present within cups. Podetial wall entire. Apothecia common, on podetial tips or on the cup margins, red, flat, margins persistent, very thin, sometimes flexuose, 0.5- 1 mm in diameter. Pycnidia on tips of podetia, or on margins of scyphi. CHEMISTRY: Chemotype 1: squamatic acid alone (as in type); chemotype 2: squamatic and usnic acids (both PD-, K-, UV+ bright blue-white); chemotype 3: thamnolic acid alone; chemotype 4: thamnolic and usnic acids (both PD+ orange, K + persistent deep yellow, UV-); decarboxythamnolic acid accompanies the thamnolic. HABITAT: Growing on rotten wood or on bark, especially of conifers; in full sun, or in partial or deep shade. DISTRIBUTION: QCI: common throughout the islands; chemotypes 1 and 2 at low and sometimes high elevations (Fig. 34); chemotypes 3 and 4 at low elevations (Fig. 35). N. Am.: western, Alaska to California, mostly coastal. World: North America and western Europe. SELECTED SPECIMENS EXAMINED: Chemotype 1: B-10461, 10691C(pp), 11416, 12450. Chemotype 2: B-9850, 9890 (Lich. Can. Exs. 29), 10545, 17376(pp) (for HPLC analysis, see Huovinen et al. 1989a, as "Cladonia sp. 2 ' 7 , 26652, 26743. Chemotype 3: A-38925; B-1 0290, 10691C(pp), 12441 (Lich. Can. Exs. 104), B-1 7376(pp), 17820. Chemotype 4: A-39108; B-10116, 10691D(pp), 11067, 12129, 17849, 17936. DISCUSSION: Much of the farinose sorediate material from the west coast, especially the QCI, is morphologically identical to the Norwegian Cladonia umbricola also cited from western North America (Tonsberg and Ahti 1980). In North America, however, it is much more variable chemically than was mentioned in the original and subsequent publications in which squamatic acid is the only major substance given (Huovinen et al. 1989a). In western North America, the typical chemotype is found from the California-Oregon area (Hammer 1991) to the Queen Charlottes, and eastward to Montana (McCune 1982) and the interior wet belt of British Columbia (Goward and Ahti 1992). Along the coast from Washington to southeast Alaska, morphologically indistinguishable populations containing thamnolic instead of squamatic acid occur, and both the thamnolic acid and squamatic acid containing strains commonly contain usnic acid as well. The usnic acid is usually abundant enough to give the thalli

a very yellow appearance, but sometimes it can only be found in traces. Because ranges of these chemotypes overlap completely on the west coast, and because there seem to be no correlating morphological features, we are reluctant to recognize any of these populations taxonomically. With a thamnolic acid containing strain now known, comparisons with Cladonia polydactyla (Florke) Spreng. are even more appropriate than they were when Cladonia umbricola was described (see Tonsberg and Ahti 1980). In brief, Cladonia polydactyla has broader, more dentate and proliferating cups, is taller, and is more corticate, as well as being greyer in colour. Cladonia pseudomacilenta Asahina, a name that was used for the western American material of Cladonia umbricola, represents a squamatic acid containing, ascyphose, apically branching species in East Asia, still unknown from North America (Stenroos and Ahti 1994) (see comments below). The morphology of Cladonia umbricola is no less variable than its chemistry. Some specimens are very robust, producing more granular than farinose soredia, and greatly resemble Cladonia transcendens (especially when containing thamnolic and usnic acids). Hammer (1993a), in fact, regarded all sorediate, thamnolic acid containing specimens as Cladonia transcendens, regardless of the type of soredia. Some European material shows the same variation in the direction of Cladonia transcendens, becoming somewhat granular sorediate and corticate at the base of the podetia. There is a tendency for the podetia of true Cladonia umbricola to emerge from the centers of the primary squamules, whereas those of Cladonia transcendens tend to develop from the squamule edges, but this is not an infallible difference. (See isc cuss ion under Cladonia transcendens.) There is also variation in the extent of cup production. Specimens consisting entirely of subulate podetia, as well as some having only cupped podetia, have been seen both from Europe and the west coast. The amount of cortex at the podetial base also varies: no cortex to corticate over half the length of the podetium. The cortex is generally thin, however, rarely forming the thick cortical plates seen in Cladonia transcendens. The California to Oregon material in H, however, seems to be less farinose and more corticate, almost always emerges from the edge of the primary squamules, and lacks soredia on the primary squamules. Farther north, the population is more heavily farinose sorediate. Cupped specimens with granular soredia and producing squamatic and usnic acids require comparisons with other taxa. Cladonia yunnana (Vain.) Abbayes ex J.C. Wei & Y.M. Jiang is an Asian species with well-corticate, cupped podetia having patches of granular soredia on the upper half (much as in Cladonia cornuta ssp. groenlandica). It is discussed and illustrated by Stenroos (1986), who points out that the North American records from ldaho (Schroeder et al. 1973) are incorrect; in fact, they are based on the usnic squamatic acid containing chemotype of Cladonia umbricola (specimens seen in H). Cladonia granulans Vain., which was reported from Alaska (Krog 1968), the Mackenzie Delta region, N.W.T. (Ahti et al. 1973), Wyoming, as well as Kamchatka and Japan (Yoshimura 1968), has broad cups more like Cladonia coccifera (L.) Willd. It was discussed and mapped by Stenroos (1989a). The report from Wyoming, however, is based on Cladonia blakei Robbins, once considered a synonym of Cladonia granulans, although

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Can. J . Bot. Vol. 74, 1996 it actually belongs under Cladonia coccifera (Stenroos 1 9 8 9 ~ ) . Cladonia vulcani Savicz (Cladonia theiophila Asahina) was shown to have two chemotypes, one containing usnic and squamatic acids and the other usnic and thamnolic acids (Stenroos 1986; Huovinen et al. 1989a; Stenroos and Ahti 1994), much like the Queen Charlotte Cladonia umbricola. Although it is also a narrowly cupped species with soredia, its soredia are coarsely granular to microsquamulose, and unlike Cladonia umbricola, the interior surface of the cups are corticate as are much of the podetia. The species was reported from the Vancouver area (Bird and Bird 1973), but the records should be checked (Stenroos 1986). Allowing for a certain variation in soredial size, we are including granular sorediate specimens under Cladonia umbricola. In the southern hemisphere (South America - Antarctica Australasia), there is a vicarious species, Cladonia ustulata (Hook. f. & Taylor) Leight., that displays a similar chemical variation (thamnolic or squamatic acids, with usnic acid present or absent), and the soredia range from farinose to granular (Stenroos and Ahti 1990; Stenroos 1993). It, however, has broader cups (up to l cm), but it is possible that some of the Chilean material is actually referable to Cladonia umbricola. Cladonia uncialis (L.) F.H. Wigg. syn. Cladoniapseudostellata Asahina, J. Jpn. Bot. 18: 620. 1942, syn.nov. DESCRIPTION: Primary thallus evanescent. Podetia dense or forming cushions, much to richly branched. Branching mostly anisotomic in dichotomies, or frequently in trichotomies. Axils wide open, not forming funnels. Podetia 2060 mm high, 0.7-4 mm wide, lacking scyphi. Podetia pale greenish yellow (104) or greenish white (153) with patches of pale yellow-green (12 1-119) or light yellow-green (134), often with irregular clear areas ("windows"), more or less uniform, or mottled, or rarely becoming brown at tips. Base more or less the same as upper portions. Cortex present. Podetial surface smooth, with no podetial squamules, soredia, or granules. Stereome cylindrical, uniform and smooth, or slightly fibrose in places. Apothecia not produced. Pycnidia on tips of podetia. CHEMISTRY: Thallus PD-, K-, UV+ bright blue-white (especially where podetia are broken) or UV-. Chemotype 1: usnic acid alone (type); chemotype 2: usnic and hypothamnolic acids, rarely with traces of squamatic acid (corresponding to the type of C. pseudostellata); chemotype 3: usnic and squamatic acids. (For quantitative data on the QC1 specimen B-14261 of this chemotype, see Huovinen and Ahti 1986, pp. 184-185.) HABITAT: Growing on mineral soil, in patches of moss, or directly on rock mainly in low elevation terraced bogs, bluffs, and outcrops; in full sun. DISTRIBUTION: QCI: chemotype 1: not on QCI, but elsewhere in British Columbia; chemotype 2: infrequent, in the Moresby Range at high and low elevations (Fig. 37); chemotype 3: common, widespread on the archipelago (Fig. 38). N. Am.: arctic to temperate, widespread; Brodo (1968) comments on the eastern American distributions of the chemotypes. World: chemotype 1: circumboreal, widespread in interior regions of North America and Eurasia; chemotype 2 is amphiberingian: Japan, Russian Far East,

Alaska, QCI; chemotype 3: circumboreal in northern hemisphere, perhaps incompletely, with mainly coastal localities: North America, Europe, Asia. See also comments by Evans (1944). SELECTED SPECIMENS EXAMINED: Chemotype 2: Moresby Island: "Blue Heron Bay" N of Sunday Inlet, elev. 150 m, in grass, B-14113, 14116; Mount Moresby, open subalpine meadow and talus slope, elev. 686 m, on soil over exposed boulder, B-26775; Moresby Lake, forest to subalpine, elev. 95 -610 m, Vitt 12253 (CANL); Tasu Sound, Fairfax Inlet, Pinus contorta - Chamaecyparis nootkatensis woodland on siliceous outcrops by sea, elev. 100-200 m, A-38511, 391 76 (CANL, H). Chemotype 3: A-39101; B-9945,10436, 12299, 12935, 14157, 17676B, 17789, 18231; SchoJield 23701, 24168 (UBC). DISCUSSION: There are several morphotypes of what has been named here as "Cladonia uncialis": morph 1: very robust, irregularly branched and rough-textured; morph 2: very smooth, regularly dichotomously (rarely trichotomously) branching, with fairly long internodes and branches of more or less even diameters; and morph 3: much like morph 2 but with conspicuously flaring axillary junctures. Morph 3 frequently contains hypothamnolic acid instead of, or rarely in addition to, squamatic acid, and corresponds to what has been called Cladonia pseudostellata Asahina. In Japan, the type locality, Cladonia pseudostellata does not look like morph 3, being rather darker (discoloured?) and more Cladina-like in branching. In the Russian Far East, however, a very similar morph of Cladonia pseudostellata was frequently collected by M. Andreev (specimens in H). There, it is even more fenestrate than the B.C. material. but otherwise is very similar. Cladonia pseudostellata is therefore included here as a new synonym of Cladonia uncialis. There do not seem to be fully comparable populations of Cladonia uncialis in Europe or eastern North America. Much of the coastal material from those regions has a very smooth inner medulla lined with a white, granulose or pruinose coating, and then also squamatic acid. In Europe such a variant was called ssp. biuncialis (Hoffm.) M. Choisy (Ahti 1978a), and it (or a very similar variant) seems to be widespread in the eastern U.S. and adjacent Canada and is occasionally very robust, e.g., in the bogs of Newfoundland. Nothing comparable is found in the Charlottes. The inner surface is egranulose as in the widespread boreal to arctic ssp. uncialis, which normally has no squamatic acid and is not particularly robust. Because of the lack of correspondence between morphology and chemistry in Cladonia uncialis from the Pacific Northwest, we are treating all coastal material under one name pending a revision of the entire section Unciales. Cladonia verruculosa (Vain.) Ahti DESCRIPTION: Primary thallus squamulose, evanescent, or persistent. Primary squamules ascending, or horizontal, dispersed, 1-2 mm long, 1-2 mm broad, crenulate, olivaceous (olive brown), undersurface white, ecorticate, esorediate. Podetia scattered or clustered, unbranched, or once or twice branched with branching in dichotomies; axils closed. Podetia 20-70 mm high, 0.8-3 mm wide, at first subulate, finally producing narrow scyphi, 1-4 mm across, imperforate, sometimes proliferating from the center of the cups,

Brodo and Ahti


Figs. 37-40. Distribution of species of Cladonia on the Queen Charlotte Islands. Figs. 37 and 38. Cladonia uncialis. Fig. 37. Chemotype 2 (usnic and hypothamnolic acids). Fig. 38. Chemotype 3 (usnic and squamatic acids). Fig. 39. Cladonia verruculosa. Fig. 40. Cladonia wainioi.

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or proliferating from the margins; margins dentate or squamulose. Podetia olivaceous, brown, or mottled with corticate granules and squamules over a black stereome. Base strongly melanotic. Cortex absent. Podetial surface verruculose or areolate. Podetial squamules abundant, peltate, sometimes granule-like. Podetia sorediate, or with spherical, corticate granules. Soredia granular, diffuse. Typically, if granular soredia are present, they are on the upper half of the podetia in patches, or cover the outside of cups if cups are present, and grade into cortical granules. Soredia are also present within cups. Podetial wall entire. Apothecia common, on the cup margins, brown, strongly convex to hemispherical, 1-3 mm in diameter. Pycnidia at cup margins, broadly pyriform. CHEMISTRY: Thallus PD+ red, K- (or brownish), UV-. Contains fumarprotocetraric and protocetraric acids, Cph-2, and sometimes Cph- l . HABITAT: Growing on mineral soil, in full sun. DISTRIBUTION: QCI: frequent, mainly lowlands in the NE sector (Fig. 39). N. Am.: California to Alaska, east to Idaho and interior British Columbia. World: North America, Central and South America (Andes). SELECTED SPECIMENS EXAMINED: A-39020a, 39059; B-10031, 10692B, 11749, 12628B, 14285, 18150, 18173B, 18416, 23667, 26541 (Lich. Can. Exs. 236); Moresby

Island: Skidegate, Spreadborough 83155 (CANL); Graham Island: White Creek Bog, Schofield 23791 (UBC). U.S.A.: ALASKA: Juneau area, Derr & Geiser L-3296. DISCUSSION: Dispersed pale granules and verrucae on a black decorticate surface and tall, slender, subulate to narrowly cupped podetia distinguish this common western American species from all others (Ahti 1978b). It was most recently discussed by Hammer and Ahti (1990) and Hammer (1993a), and its range has been extended from the coast to the interior wet belt in British Columbia (Goward and Ahti 1992). Cladonia wainioi Savicz Bull. Jard. Imp. Bot. St.-PCtersbourg, 14: 125. 1914, 'wainii'. Type: Russia: Kamchatka Region: between Malka and Ganal, 1909 Savicz 5244 (LE, holotype; FH, H, LE, UPS, isotypes). Synonymy (note new synonyms): Cladonia rangiferina f. leucosticta G. Merr., Bryologist 11: 109. 1908, syn.nov. Type: Canada: Yukon: in bogs around Hunker Creek, Macoun, Can. Lich. 112 (FH, lectotype, here selected; CANL, 0 , isolectotypes). Cladonia rangiformis var. versicolor Elenkin, Acta Horti Petropol. 31: 254. 1912. Type: Russia: Primor'e Terr. (FH, LE, syntypes). Cladonia peltastica var. verruculifera Vain., Bot. Mag. 35: 65. 1921, syn.nov. Type: Japan: Kotsuke: Mt. Kasadake, 1912 Yasuda 204 (TUR-V 15023, holotype). Cladonia pseudorangiformis Asahina, J. Jpn. Bot. 18: 549. 1942. Type: Japan: Mt. Shirouma, Lake Oike, 1936 Asahina 36288 (US, lectotype by Ahti 1962: 38; TNS, isolectotype). DESCRIPTION: Primary thallus evanescent. Podetia forming cushions, richly branched. Branching isotomic or anisotomic, mainly in dichotomies, with some trichotomies. Axils open, wide, not forming funnels. Podetia 40-60 mm high, 0.4-2.5(-4) mm wide, lacking scyphi. Podetia very pale yellowish green (121) to pale brown ("pale orange-yellow," 73), mottled, or becoming brown at tips. Base more or less the same as upper portions. Cortex present in patches, discontinuous, absent between areoles. Podetial surface areolate with thin, flat, smooth, irregular areoles lying over a transluscent stereome, without podetial squamules, soredia, or granules. Podetial wall entire or perforated. Apothecia not produced. Pycnidia on tips of podetia. CHEMISTRY: Thallus PD-, K yellow, UV-. Contains atranorin, merochlorophaeic acid, and 4-0-methylcryptochlorophaeic acid. For HPLC analysis of the QC1 specimen B-10989, see Huovinen et al. (1990). HABITAT: Growing on mineral soil, or over rocks and in heath at low and high elevations, or in patches of moss, in full sun. DISTRIBUTION: QCI: infrequent; scattered (Fig. 40). N. Am.: mainly northern boreal (Ahti 1964). World: North America, eastern Asia. SPECIMENS EXAMINED: A-39100, 39177; B-10423, 10985, 10989, 12157, 12934; Schofield 24867, 34513 (UBC). DISCUSSION: Cladonia wainioi, first reported from the area by Brodo et al. (1987), was described from volcanic soil in Kamchatka. It was regarded as an enigmatic species for a long time because the type material consists of strongly

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swollen, highly deformed podetia. Its chemistry, however, turned out to be identical with that of Cladonia pseudorangifortnis, and therefore Huovinen et al. (1990, p. 231) united these species. Indeed, in the perhumid conditions of the QCI, robust specimens very similar to the type material of Cladonia wainioi were collected growing close to patches that would correspond to "ordinary" Cladonia pseudorangiformis, both apparently being part of the same clone. It is therefore confirmed here that the two taxa should be regarded as different morphotypes of the same species. Cladonia wainioi is a widespread lichen in North America and East Asia (Ahti 1962, 1964, sub Cladonia pseudorangiformis). Here, two new synonyms, viz. Cladonia rangiferina f. leucosticta G. Merr. and Cladonia peltastica var. verruculifera Vain., are added to the list of synonyms of Cladonia wainioi.

Conclusions and summary (i) Although the Queen Charlotte archipelago is characterized by its very moist, highly oceanic climate and relatively low mountains (none above 1100 m), it was still somewhat surprising that numerous widespread, extremely common boreal to temperate species are rare there, or are completely lacking. Examples are Cladina stellaris (i.e., its normal psoromic acid-deficient chemotype), Cladina tnitis, Cladonia amaurocraea, Cladonia cariosa, Cladonia cenotea, Cladonia coniocraea, Cladonia cervicornis ssp. verticillata, Cladonia chlorophaea, Cladonia deformis, Cladonia pleurota, Cladonia phyllophora, Cladonia gracilis subspecies (other than the western ssp. vulnerata), and Cladonia pyxidata. This would seem to indicate that the mild, oceanic climate is not better for lichens in general and that many boreal taxa may have a real requirement for a continental climate with its cold, dry winters and relatively hot summers. (ii) Many populations along the hypermoist coast, especially on the Charlottes, even if they cannot presently justify nomenclatural recognition, have a different appearance than conspecific populations farther inland. Among these are Cladonia chlorophaea, Cladonia fimbriata, Cladonia furcata, Cladonia scabriuscula, Cladonia uncialis, and Cladina rangiferina. (iii) The more typical arctic-boreal flora can be found close by on the adjacent mainland: in the mountains of southeast Alaska and coastal British Columbia, or even close to sea level when adjacent to glaciers. (iv) Besides the morphological novelties encountered on the Charlottes, we see a surprising increase in chemical variation, especially with respect to the presence of thamnolic and usnic acids. Many taxa that are chemically fairly uniform elsewhere, have alternative chemotypes with or without usnic acid (e.g., Cladina portentosa, Cladonia umbricola, and even, to some extent, Cladonia bellidiflora) or thamnolic acid (e.g., Cladonia umbricola, Cladonia squamosa, Cladonia bellidijlora, and Cladorzia crispata). Note that in the thamnolic acid chemotypes, both sections Cocciferae and Perviae are represented. It would be intriguing to speculate on how a cool, oceanic climate might influence the biosynthetic pathways leading to the production of thamnolic acid, presumably from its precursor, squamatic acid, in these unrelated taxa.

(v) The Cladoniae on the QC1 and southeast Alaska are significantly different from those of the temperate Pacific coast from southern Vancouver Island to California. That is, although a number of western Cladoniae extend from California to Alaska (e.g., Cladonia verruculosa and the coastal morphotype of Cladonia furcata), many of them do not, or barely have their northern limits including the Charlottes, e.g., Cladonia dimorpha, Cladonia poroscypha, Cladonia artuata, Cladonia prolifica, Cladonia singularis, and Cladonia asahinae (protolichesterinic acid chemotype). Thus, the phytogeographic boundary just north of Vancouver and encompassing many of the Gulf Islands, characterized by the presence of Pseudotsuga menziesii (the Coastal South Pacific Cordilleran ecoclimatic region (Ecoregions Working Group 1989)), appears to be supported by the distributions of the Cladoniaceae. (vi) A biogeographic analysis of the Queen Charlotte Cladoniaceae indicates that about 40% of the species are essentially oceanic in distribution, usually being disjunctively distributed in several widely separated areas of high precipitation in the northern or southern hemispheres, e.g., western North America, Newfoundland, mountains of Japan, northwestern Europe, Tierra del Fuego, and New Zealand. Five taxa (out of 44) are restricted to western North America, and 2 have a northern amphipacific distribution (i.e., they are also in Asia). Some lichens have geographic chemotypes that have similar distributions. The rest of the QC1 species represent widespread, usually continuously circumpolar elements, although many of them are remarkably rare on the QCI, as mentioned under (i) above. (vii) Within the QCI, certain distribution patterns can be recognized from the maps: (i) lowland species, especially of muskeg, largely found in the Queen Charlotte Lowlands: e.g., Cladonia bellidiflora, thamnolic acid chemotype (Fig. 1l), Cladonia fimbriata (Fig. 19), Cladonia macilenta (Fig. 23), Cladonia pyxidata (Fig. 28), and Cladonia verruculosa (Fig. 39); (ii) high elevation species largely confined to Queen Charlotte and San Cristoval Ranges: e.g., Cladonia ecmocyna ssp. occidentalis (Fig. 17), Cladonia kanewskii (Fig. 22), Cladonia metacorallijera (Fig. 24), and Cladonia schofieldii (Fig. 30); (iii) widespread lowland species: e.g., Cladonia crispata var. crispata (Fig. 15) and Cladonia umbricola (Figs. 34 and 35). Other lichens are widespread at all elevations on the QCI, as they are elsewhere, at least in western North America: e.g., Cladina portentosa ssp. pacifica (Fig. 3), Cladina rangiferina (Fig. 4), Cladonia bellidiflora (Fig. 10), Cladonia cornuta ssp. groenlandica (Fig. 14), Cladonia squamosa (Fig. 31), and Cladonia furcata (Fig. 20).

Acknowledgements We gratefully acknowledge the help of Kati Lohtander and Leena Myllys in Helsinki and Pak Yau Wong in Ottawa for help in running TLC analyses; Wilfred Schofield, Vancouver, for collaboration in the field and for providing so many interesting specimens; Laurie Consaul for compiling most of the Cladonia specimen data and creating the computerized maps; Sampsa Lommi for drawings; and the many individuals on the Charlottes, especially Neil and Betty Carey and the Canadian Fisheries and Oceans Ministry, who

Brodo and Ahti

provided logistic support to Brodo and his assistants over several seasons. We are grateful to Samuel Hammer for his many useful comments on an early draft of this paper. The study is also part of a long-term project of Ahti on Cladoniaceae, financially supported by the Academy of Finland. Brodo's work on the QC1 lichen flora has been supported for many years by research grants from the Canadian Museum of Nature.


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