SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 66 

A Monograph of the Lichen Genus 

Parmelia Acharius 

sensu strict0 (Ascomycotina: 

Pameliaceae) 

Mason E. Hale, Jr. 

SMITHSONIAN INSTITUTION PRESS 

Washington, D.C. 

1987 
ABSTRACT 

Hale, Mason E., Jr. A Monograph of the Lichen Genus Parmelia Acharius sensu stricto 

(Ascomycotina: Parmeliaceae). Smithsonian Contributions to Botany, number 66, 55 pages, 

25 figures, 1987.--The 38 species of Parmelia Acharius sensu stricto are revised at the world 

level. The genus is characterized by punctate or effigurate pseudocyphellae, a black lower 

surface with simple, furcate or squarrose rhizines, simple hyaline spores, and cylindrical to 

weakly bifusiform conidia. The most frequent secondary metabolites are salazinic acid and 

protocetraric acid. Parmelia is an extremely conservative genus, most common in 

temperate-boreal and austral regions, with a low level of vegetative morph formation and 

speciation. The greatest concentrations of species are found in Japan and New Zealand. Seven 

new species are described: P. neodiscordans Hale, P. norcrambidiocarpa Hale, P. 

novae-zelandiae Hale, P. queemlanderuis Hale, l? salcrambidiocarpa Hale, P. skultii Hale, 

and P. subtestacea Hale. 

O~CIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf clearing from 

the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Hale, Mason E., Jr. 

A monograph of the lichen genus Pannelia Acharius sensu stricto (Ascomycotha: Parmeliaceae) 

(Smithsonian contributions to botany ; no. 66) 

Bibliography: p. 

Includes index. 

Supt of Docs. no. SI1.29:66. 

1. Parmelia-Classification. 2. Lichens-Classification. I. Title. 11. Series. QKl.SZ747 no. 66 581 s 86- 

607945 

[QK585.P2] [589.1] 
Contents 

Page 

Introduction ........................................ 1 

Acknowledgments ................................... 2 

Cortical and Internal Structure .............................. 2 

Vegetative Structures ................................... 2 

Reproductive Structures ................................. 12 

Chemistry ........................................ 14 

Chemotypes ....................................... 14 

Phytogeography ..................................... 16 

Key to the Species of Parmelia ............................. 16 

Taxonomic Treatment .................................. 18 

Literature Cited ...................................... 52 

Index ........................................... 55 

... 

111 

A Monograph of the Lichen Genus 

Parmel ia Ac harius 

sensu strict0 (Ascomycotina: 

Parmeliac e ae) 

Mason E. Hale, Jr. 

Introduction 

The genus Parmelia was proposed by Acharius in 1803. His 

circumscription was very broad and encompassed a large 

number of foliose species with lecanorine apothecia, species 

now recognized in such diverse genera as Cetraria, Heterodermia, Hypogymnia, Lobaria, Pannaria, Parmelia sensu lato, 

Parmeliella, Parmeliopsis, Physcia, Physconia, and Xanthoria. 

Early 19th century authors continued to use the name in this 

broad sense for almost any lichen with a thalline apothecial 

rim, from Lecanora to Usnea. 

By the end of the 19th century Parmelia had begun to 

assume its more modernsand familiar circumscription as a 

foliose rhizinate genus with laminal apothecia and simple 

spores (Fries, 1871-1874). At the same time, recognizing clear 

morphological discontinuities, lichenologists began to describe 

a number of segregate genera. Early ones, with their dates of 

publication, included Menegazzia Massalongo (1 854), Parmotrema Massalongo (1860), Anzia Stizenberger (1861), Parmeliopsis (Nylander) Nylander (1 869), Hypogymnia (Nylander) 

Nylander (1896), Pseudevernia Zopf (1903), Pannoparmelia 

(Miiller Argoviensis) Darbishire (1912), and Pseudoparmelia 

Lynge (1914). Of these genera only Anzia and Parmeliopsis 

were ultimately adopted by Zahlbruckner in his catalogue 

(1929), the others being synonymized under Parmelia. 

However, since 1950, many workers have adopted Hypogymnia, Menegazzia, Pannoparmelia, and Pseudevernia. 

I recently subdivided Parmelia sensu lato into two groups 

on the basis of cortical structure newly discovered with the 

scanning electron microscope (Hale, 1973), resurrecting some 

of the old, neglected segregates as well as describing several 

Mason E. Hale, Jr., Department of Botany, National Murewn of Natural 

History, Smithonian Institution. Washington, D.C. 20560 

new genera. One group is represented by the epicorticate 

species found in Arctoparmelia Hale, Bulbothrix Hale, 

Canoparmelia Elix and Hale, Cetrariastrum Sipman, Concamerella Culberson, Everniastrum Hale ex Sipman, Flavoparmelia Hale, Hypotrachyna (Vainio) Hale, Neofuscelia Esslinger, Paraparmelia Elix and Johnston, Parmelina Hale, 

Parmotrema Massalongo, Pseudoparmelia Lynge, Relicina 

(Hale and Kurokawa) Hale, Relicinopsis Elix and Verdon, 

Xanthomaculina Hale, and Xanthoparmelia (Vainio) Hale. 

A second group consists of the pseudocyphellate species of 

Parmelia (Hale, 1975, 1976a, 1981). This group has been 

further subdivided into Flavopunctelia (Krog) Hale (the P. 

flaventior group) (Hale, 1980, 1984), Melanelia Esslinger (the 

brown Parmeliae) (Esslinger, 1978), and Punctelia Krog 

(Krog, 1982) (the P. borreri group), now leaving in Parmelia 

a small, apparently irreducible assemblage of 38 species 

typified by P. saxatilis. 

The following characters may be used to delimit Parmelia: 

adnate, sublinear to subirregular lobes without cilia; upper 

surface effigurate-pseudo-cyphellate, less commonly punctatepseudocyphellate (in three Asian species); lower surface black, 

rhizinate, the rhizines simple, furcate, or squarrosely branched; 

microconidia cylindrical or weakly bifusiform, less than 8.0 

pm long; spores simple, 8 per ascus. Chemically it is 

distinguished by the presence of atranorin and chloroatranorin 

and lack of usnic acid in the cortex. The closest relative in the 

family is Punctelia, which Krog (1982) characterizes by 

punctate pseudocyphellae and cylindrical, filiform, or unciform 

microconidia. I am excluding from Punctelia and Parmelia the 

unique usnic acid-containing American endemic P. 

sphaerosporella Muller Argoviensis, now recognized as the 

monotypic genus Ahtiana Goward (Goward, 1985), closer to 

Cetraria Acharius than to Parmelia. 

1 
2 SMITHSOhTAN CONTRIBUlTONS TO BOTANY 

ACKSOWLEDG.MEXS.-~ amindebted to Dr. Ove Almborn 

for reviewing the nomenclature and providing valuable notes 

on citations and literature. I have also benefited from 

discussions with Trevor Goward, Dr. J.A. Elix, and Dr. H. 

Skult. Lastly I must give thanks to the curators of the museums 

and institutions who sent loans of types and other specimens. 

Cortical and Internal Structure 

Parmelia has a typical paraplectenchymatous cortex (Hale, 

1973, 1981). It consists of densely packed cells forming a layer 

4-6 cells thick (20-30 p). Anglesea et al. (1982) recently 

discovered a technique of digesting the intracellular polysaccharides, giving new insights into cortical organization. The 

cortical layer consists of densely packed hyphae that branch 

in a coralloid pattern, forming 3-5 short branches lying in the 

same plane. The lower cortex is thinner, less than 20 pm thick. 

The algal layer and the medulla occupy 100-200 pm of the 

total thallus thickness. 

The upper cortex of Parmelia species is perforated by 

pseudocyphellae. These pores originate as the cortex disintegrates to form a narrow tube about 10 pm in diameter, which 

enlarges and becomes filled with medullary hyphae (Figure 

la) (Hale, 1981). This structure is quite different from the 

epicortex, which is a pored polysaccharide sheet overlaying a 

more or less loosely packed, continuous palisade parenchymatous layer (Hale, 1973, 1981). These two structures differ 

considerably in size: pseudocyphellae are essentially macroscopic and visible with low power magnification (0.1-2 mm 

long), but the pores in the epicortex can be seen only with 

SEM, being of the order of 15-40 pm in diameter. 

Each species of Parmelia has a characteristic orientation of 

pseudocyphellae as illustrated in Figures 2-8. The most 

frequent type appears as an irregular effigurate white marking 

under low power. These so-called maculate markings are 

marginal and lamina1 and often fuse into a loose reticulate 

network as they grow beyond 1 mm in length Figure lc). At 

maturity the cortex tends to crack open along the pseudocyphellae, which may then serve as points of origin for isidia 

and soredia. 

Under SEM the maculate appearance can be explained as a 

thin persistent, densely pored polysaccharide layer acting as a 

roof over the disintegrating pore area below. This layer is part 

of the polysaccharide layer covering the rest of the surface. 

This roof may persist (Figure Id) or soon disintegrate, leaving 

an open pore (Figure lb) (see Hale (1981) for additional 

figures). 

Pseudocyphellae may also originate exclusively along the 

lobe margins, just back from the edge. In these cases they form 

a continuous white line, a rim around the lobes, as in P. 

neodiscordans, P. niitakana (Figure le), P. pseudoshinanoana, 

P. sectilis, P. shinanoana, P. ricasolioides, and in part P. 

cochleata, P. crambidiocarpa, P. norcrambidiocarpa, P. 

salcrambidiocarpa, P. subtestacea, and P. testacea, and some 

narrow-lobed forms of P. discordans and P. omphalodes, and 

in P. skultii. 

Two Asian species, P. meiophora and P. submutata, are 

distinctive in having very small effigurate pseudocyphellae not 

much more than 0.1-0.2 mm in size (Figure lb), so small that 

neither Hue, Nylander, nor Zahlbruckner noticed them when 

describing the species involved. As far as I can tell, they are 

identical in origin with the larger pseudocyphellae described 

above. 

A third type of pseudocyphella is characteristic of a small 

group of three Asian species, P. isidioclada, P. laevior, and P. 

pseudolaevior. The pores form along the lobe edge as round, 

widely spaced, and rather inconspicuous white spots (Figure 

If). These pores do not have the persistent roof of other 

pseudocyphellae in the genus and in fact appear to be identical 

with the punctiform pores of Punctelia. The lobe surface itself 

is continuous and has neither pseudocyphellae nor reticulated 

markings. These species are placed more appropriately in 

Parmelia rather than Punctelia, because of the presence of 

salazinic acid and the furcate to squarrosely branched rhizines. 

Vegetative Structures 

Most Parmelia species have rather narrow, sublinear, adnate 

lobes, They grow on rocks and trees in forested areas and on 

rocks and humus or mosses in arctic-alpine areas, forming 

adnate colonies 4-20 cm broad (up to 60 cm in P. tenuirima! ). 

All of the species are whitish or greenish mineral gray but 

some, such as P. discordans, P. omphalodes, P. signifera, and 

P. skultii, have a tendency to turn brown to nearly black in 

exposed habitats. 

SL~FACE FATmES.-The surface features of several Parmelia species have been studied by Hale (1973) using the 

scanning electron microscope (SEW. The genus is characterized by a smooth and featureless or nodular surface, typical of 

lichens with a paraplectenchymatous cortex (Figure 9a). 

Many species have a hoary white pruina on the surface. 

Under SEM this pruina is revealed as masses of crystals. I 

have not investigated the chemical composition of these 

crystals, but they are reported to consist of oxalates (Wilson 

et al., 1980), often weddelite (calcium oxalate tetragonal 

dihydrate) (CaC204(2+x).H,0), and less commonly whewellite 

(calcium oxalate monoclinic monohydrate, CaC,O,.H,O) 

(Jackson, 1981; Wadsten and Moberg, 1985). 

The crystals fall into three general shapes: bi-pyramidal 

weddelite (P. cochleata, P. cunninghamii, P. fraudans (Figure 

9c), P. kerguelensis, P. meiophora, P. omphalodes, P. 

pseudoshinanoana, P. saxatilis, P. sulcata, P. squarrosa, P. 

testacea (Figure 9b)); cubic or parallelepiped shapes, also 

probably weddelite (P. cunninghamii, P. hygrophila (Figure 

94, P. protosulcata, and P. sulcata (Figure 98); and flat 

hexagons of whewellite (P. adaugescens, P. discordans, P. 

erumpens, P. fraudans (Figure 9e), P. isidioclada, P. 

kerguelensis, P. novae-zelandiae, P. pseudoshinanoana, P. 

saxatilis, F! tenuirima). There seems to be little if any 
NUMBER 66 3 

FIGURE 1.-Pseudocyphellae in Purmeliu: a, pore in upper cortex of P. uduugescens (hi 7809); b, 

pseudocyphellae of P. submututu (Poeit L-142); c, pseudocyphellae of P. murmriza (Kurokawa 63056); 4 

pseudocyphellae of P. murmurizu to show pored “roof ” (Kurokawu 63056); e, submarginal pseudocyphellae 

of P. niitukam (Hsu 1374); I: marginal pseudocyphellae of P. luevior (Kurokawu 58255). 
4 SMITHSONIAN CONTRIBUnONS TO BOTATTY 

FIGURE 2.4nentation of pseudocyphellae in Parmlia species: a, P. adaugescens (Asahina s.n.); b, P. cochleata 

(isolectotype in US); c, I? crambidiocarpa (Hale, 65541); 4 P. cunninghamii (Santesson 6792); e, P. discordans 

(Issin m.); 1: P. erumpens (Hale 43876). (All x8.5.) 
hWMBER 66 

FIGURE 3.4rientation of pseudocyphellae in Parmelia species: a, P. fertilis (Tagawa s.n.); b, P. fraudans 

(Hale 49870); c, F? hygrophila (Noble 6448); d I? isidioclada (Kurokawa 58010); e, F? kergueiensk (Harris 

5670); f f. iaevior (Culberson 11 104). (All x8.5.) 

5 
6 SMITHSONIAN CONTRIBUnONS TO BOTATTY 

FICLW 4.4rientation of pseudocyphellae in Parmelia species: a, P. marmariza (Kurokawa 58062); b, P: 

meiophora (Poelt L-151); c, P. neodircordanr (holotype in US); d, P. niitakiana (Hsu 1374); e, P. 

norcrambidiocarpa (holotype in US); f; P. novae-zelandiae (Hale 66232). (All x8.5.) 
NUMBER 66 

FIGURE 5.-Orientation of pseudocyphellae in Parmelia species: a, P. omphalodes (Santesson 12918); b, P. 

protosulcata (Santesson 2953); c, I? pseudolaevior (Kuroknwa 59205); d P. pseudoshinanoana (Kwoknwa 

59206); e, P. pseudotenuirima (Hale 58276); f; P. queenslandensis (holotype in US). (All x8.5.) 

7 
8 SMITHSONIAN CONTRIBUTIONS TO BOTAhY 

FICLRE 6.4rientation of pseudocyphellae in Parmelia species: a, P. ricarolioides (isotype of P. daliensis in 

US); 6, P. salcrambidiocarpa (holotype in US); c, P. saxutilis (Hale 48453); d, P. sectilis (holotype in US): e, 

P. shinattoam (isolectotype in US); f; P. signifera (Hale 66435). (All x8.5.) 
NUMBER 66 

FIGURE 7.4nentation of pseudocyphellae in Parmelia species: a, P. skultii (holotype in US); b, P. squarrosa 

(Hale 18170); c, P. submontam (VPzda 845); d F! submutata (Poelt L-142); e, P. subtestacea (Hale 58771): 

1: P. sulcata (Kjellmert s.n.). (All x8.5.) 

9 
10 SMITHSONIAN CONTRIBUnONS TO BOTANY 

FIGURE 8.4rientation of pseudocyphellae in Parmelia species: a, P. lenuirimu (James 488 pp.): b, P. tesfocea 

(Hale 65112). (All x8.5.) 

correlation with taxonomic features. For example, the crystal 

shapes of two specimens of P. fraudans (Figure 9c,e), are 

significantly different, whereas common P. sulcata shows three 

types among five specimens examined (three flat and one each 

pyramidal and cubic). It is obvious that many more specimens 

will have to be examined with SEM before any definite 

conclusion can be reached on the usefulness of crystal shape 

in taxonomy. 

The lower surface of all species of Parmelia is black and 

more or less continuously vested with rhizines. Broader-lobed 

species such as P. tenuirima may also have a rather 

conspicuous brown, bare to papillate marginal zone. The 

rhizines are black and shiny, 0.5-2 mm long, and either simple 

to sparsely furcate (Figure 10a,b), simple with a few weakly 

squarrosely branched (Figure lOc), or mostly strongly squarrosely 

branched (Figure 106). The following species have simple to 

furcate rhizines: P. adaugescens (Figure lOa), P. cochleata, 

P. discordans, P. fraudans, P. hygrophila, P. isidioclada, P. 

marmariza (Figure lob), P. neodiscordans, P. niitakana, P. 

omphalodes, P. pseudolaevior, P. saxatilis, P. sectilis, P. 

shinanoana, P. skultii, and P. submontana. The following 

species have simple rhizines and at least some (often densely) 

squarrosely branched: P. cunninghamii, P. erumpens, P. 

fertilis, P. kerguelensis, P. laevior, P. meiophora, P. norcrambidiocarpa, P. novae-zelandiae, P. protosulcata, P. pseudoshinanoana, P. pseudotenuirima, P. queenslandensis, P. 

ricasolioides, P. salcrambidiocarpa, P. signifera, P. squarrosa, P. submutata, P. sulcata (Figure lOc), P. subtestacea, 

P. tenuirima (Figure 10d ), and P. testacea. 

SoRmk-soredia appear to originate from pseudocyphellae in most species and become aggregated into orbicular or 

linear soralia. In P. cunninghamii they originate from marginal 

fissures not associated with pseudocyphellae, as well as from 

laminal pseudocyphellae. They are predominantly marginal in 

P. cunninghamii, P. fraudans (Figure lla), and P. protosulcata; laminal and marginal in P. sulcata (Figure 1 lb); and 

mostly laminal in P. submontana. Parmelia erumpens is unique 

in having mostly laminal pustular soralia. 

IsmA.-Although isidia are one of the most easily 

recognized vegetative propagules in lichens, their origin, 

development, and structure are not well known. The few SEM 

pictures published to date (see Hale, 1975, 1976b, for 

Hypotrachyna and 1976a for Parmelina) show in general a 

clearly defined radial structure. There is a more or less 

continuous polysaccharide layer over the surface, a rather 

loosely organized paraplectenchymatous cortical layer 20-30 

Fm thick, which is continuous with the thallus cortex at the 

base. The interior is filled with loose medullary hyphae, the 

algae occupying the periphery (Figure 118. 

Five species of Parmelia have typical shiny, corticate 

cylindrical isidia: P. kerguelensis (Figure 1 1 c,e), P. novaezelandiae, P. pseudotenuirima, P. saxatilis, and R squarrosa. 

The isidia of P. meiophora are rather short and basally 

constricted (Figure 10d ) in comparison with the other isidiate 

species. In P. isidioclada the isidia become densely coralloid 

branched and subsorediate. In P. hygrophila the isidia are 

initially corticate but appear to break down apically at maturity, 

becoming scurfy or subsorediate, an appearance that is 

enhanced by the dense production of pruina on the isidia. 

LOBuLES.--LobUkS in Parmelia are dorsiventral and usually 

originate along lobe margins; they are very narrow, suberect, 

and almost isidia-like in appearance. There are three lobulate 

species here in a strict definition: P. pseudolaevior, P. 

pseudoshinanoana, and P. sectilis. Parmelia pseudolaevior 

seems to be a well-matched morph of non-lobulate P. laevior; 

P. pseudoshinanoana differs from its presumptive parent P. 

shinanoana in lacking gyrophoric acid in the cortex. Parmelia 

sectilis has no extant parent morph. 

Parmelia discordans, P. neodiscordans, and P. omphalodes 

often produce lobulate morphotypes, although these lobules 
hTJMBER 66 

FICIJRE 9.Surface features in Parmlia: a, surface of P. squarrosa (Hale 18922, x425); b, pruina of P. testacea 

(Hale 58741); c, pruina of P. fraudam (Hale 36414); 4 pruina of P. hygrophila (Noble, 6448); e, pruina of P 

fraudam (Hale, s.n.); f; pruina of P. sulcata (Wetmore 38229). 

11 
12 SMITHSONIAN CONTRIBUTIONS TO BOTAh'Y 

FIGURE lO.-Rhizines of Parmelia: a, P. adaugescens (hi 7809); b, P. marmariza (Kurokawa 63056); c, P. 

sdcata (Hale 14415); 4 P. tenuirima (James 488 pp.). 

may be better defined as narrow secondary laciniae. Parmelia 

subtestacea and P. testacea often have roundish marginal 

lobulate secondary lobes. In these cases the lobules should be 

considered as secondary lobes. 

Reproductive Structures 

PYCsIDIA.-VobiS (1980) discussed the ontogeny and 

morphology of pycnidia in many different lichens, but he did 

not include in his study any species of Parmelia sensu stricto. 

However, he studied Melanelia acetabulum (Necker) Esslinger, which has Umbilicaria-type ontogeny and Type IV 

conidiophores, characteristic features of the family Parmeliaceae and apparently encompassing Parmelia. 

Pycnidia are typically immersed and lamina1 (Figure 4e), 

90-110 pm in diameter, with a tendency to group toward the 

lobe margins in some species. They have been observed in 

most species but not in P. hygrophila, P. isidioclada, P. 

kerguelensis, P. meiophora, P. novae-zelandiae, P. protosulcata, I? pseudolaevior, P. pseudoshinanoana, P. shinanoana, 

and P. submontana. Most of these species are isidiate or 

sorediate and often lack apothecia. 

The first lichenologist to examine pycnidia and conidia 

carefully was Lindsay (1859). By today's standards his 

drawings are rather crude and difficult to interpret. Even with 

better techniques, however, Hillmann (1936), Krog (1982), and 

other workers who have examined Parmelia species have not 

been able to describe the conidia of the genus with absolute 

clarity. Vague and often conflicting descriptions range from 

cylindrical and straight to somewhat bent or subbifusiform to 

bifusiform. 

It is indeed extremely difficult to see lichen conidia clearly 
NUMBER 66 

FIGLW 1 L-Soredia and isidia of Parmelia: a, soralia of P. fraudans (Hale 36414); b, soralia of P. sulcata 

(Hale 14415); c, isidia of P. kerguelensk (Brodo 11506); d isidia of P. meiophora (Togarhi s.n.); e, isidium of 

P. kerguelensis (Brodo 11506); 1: longitudinal cross section of isidium of P. saxatilk (Hale 36387). 

13 
14 SMITHSONIAN CONTRIBUnONS TO BOTANY 

with an ordinary light microscope. One should ideally use oil 

immersion, but even then some of the best photographs are 

rather fuzzy (Vobis, 1980). Enhancement with Nomarski lenses 

or phase contrast can improve clarity of images but the results 

are still far from perfect. In my own studies I have mounted 

freehand sections of pycnidia in glycerin-water and observed 

the conidia released under a Nomarski microscope. 

The conidia of Parmelia fall within a very narrow range of 

length and shape (Figure 12). They are either uniformly 

cylindrical, straight to slightly bent, slightly inflated at the 

center, or slighty bifusiform. The only species with strongly, 

unmistakably bifusiform conidia is P. signifera. Most conidia 

are between 5 and 7 p long, and only in P. adaugescens, P. 

laevior, P. marmariza, P. omphalodes, P. submutata, and P. 

sulcata do they consistently reach 8.0 pm. There is no 

dimorphism such as has been reported for Punctelia (Culberson 

and Culberson, 1980; Krog, 1982). 

APomw-Henssen (198 1) includes the Parmeliaceae 

among families in the Lecanorales with zeorine apothecia. I 

have not investigated the ontogeny of apothecia in Parmelia 

but assume it follows a pattern similar to that reported by 

Henssen for Melanelia exasperata (De Notaris) Esslinger. 

Mature apothecia are usually substipitate and may be rather 

large, up to 20 mm, as the disc flattens out and splits radially. 

The amphithecium is well developed, pseudocyphellate, and 

rugose or even sublobulate with age. The hymenium, which 

stains deep blue with IKI, is 50-90 pm high. The asci are 

poorly developed in a few species and lack spores. 

SmREs.-All species of Parmelia, except for P. neodiscordans, P. novae-zelandiae, P. skultii, and P. submontana, have 

been found with apothecia and spores. Spores fall into four 

size classes with relatively little overlap. One sharply defined 

group has large spores (range for the group 9-18 x 20-33 pm) 

with a thick episporium 2-3 pm wide. This group includes P. 

adaugescens, P. isidioclada, P. niitakana, P. ricasolioides, and 

P. sectilis, all Asian endemics. None of these species, it should 

be noted, are closely related. 

The remaining species have spores less than 21 p long 

with a thinner episporium 1-2 pm thick. They fall into three 

groups: one with very small spores, 3-6 x 6-9 pm (closely 

related F! meiophora and P. submutata); a second with large 

spores, 18-21 pn long (P. crambidiocarpa and P. kerguelensis); and the remainder with spores in the range 11-15 pm 

long. 

Chemistry 

When first studied systematically with microcrystal tests 

(Krog, 1951), Parmelia was thought to produce atranorin in 

the cortex and only two major medullary substances, salazinic 

acid and protocetraric acid, with accessory lobaric acid and 

protolichesterinic acid. Although these depsidones do indeed 

predominate as secondary metabolites, a number of additional 

products have been identifed with thin layer chromatography 

and more will probably be discovered. The genus has a distinct 

chemical profile, with predominantly p-orcinol depsidones, 

there being no representatives of the para- and meta-depsides 

(barbatic acid and lecanoric acid groups) in the medulla. 

The substances discovered so far are classified as follows. I 

have analyzed acetone extracts of all specimens in two solvent 

systems (toluene-dioxane-acetic acid and toluene-ethyl acetateformic acid), using Merck pre-coated, aluminum-backed TLC 

plates. 

Higher aliphatic acids: Protolichesterinic acid, unidentified acids 

Orcinol series: 

Tridepsides: Gyrophoric acid, 4-0-methylgyrophoric acid 

Depsidones: Lobaric acid 

Para-depsides: Atranorin and chloroatranorin 

Depsidones: Consalazinic acid, fumarprotocetraric acid, galbinic acid, 

p-Orcinol series: 

norstictic acid, protocetraric acid, salazinic acid 

Dibenzofurans: Usnic acid 

Anthraquinones: Skyrin 

Undetermined: 

Echinocarpic acid (and associated secondary metabolites) (probably related 

Parmelia testacea unknown #27 (probably a fatty acid according to C.F. 

to alectorialic acid according to J.A. Elk) 

Culberson) 

These substances occur in the following major combinations. Further details on their occurrence will be found in the 

species discussions. 

Cortical substances: 

Atranorin (probably always with chloroatranorin): All species in the genus 

Gyrophoric acid (with 4-0-methylgyrophoric acid): P. shinanwna 

Usnic acid: Soredia of P.fraudans 

Echinocarpic acid (k conechinocarpic acid): P. crambidiocarpo, P. 

Fumarprotocetraric acid: P. neodkcordanr, P. protosulcata 

Lobaric acid: P. dkcordans, P. kerguelenrk, P. omphalodes, P. protosulNorstictic acid: P. omphalodes, P. submutata 

Protocetraric acid: P. crambidiocarpa, P. dkcordanr, P. kerguelenrk, P. 

Protolichesterinic acid: P. dkcordanr, P. omphalodes 

Salazinic acid (often with consalazinic acid): P. adaugescenr, P. cochleata, 

P. cunninghamii, P. erumpenr, P. fertilk, P. fraudons, P. hygrophila, P. 

kidioclada, P. laevior, P. marmariza, P. meiophora, P. novae-zelandiae, 

P. niitakana, P. norcrambidiocarpa, P. omphalodes, P. pseudolaevior, 

P. pseudoshinanoana, P. pseudotenuirim, P. queenrlandenrk, P. 

ricasolioides, P. salcrambidiocarpa, P. saxatilis, P. sectilk, P. shinanoana, P. signifera, P. skultii, P. squarrosa, P. submontana, P. 

submutata, P. sulcata, P. tenuirimo, P. testacea 

Medullary substances: 

norcrambidiocarpa, P. protosulcata, P. subtestacea 

cata, P. pseudotenuirimn, P. saxatilk 

protosulcata, P. signifera 

Stictic acid: P. skultii 

Unknown fatty acid (unknown #27): P. testacea, P. subtestacea (chemotype) 

Chemotypes 

The question of “chemical species” arises in Parmelia as it 

does in most parmelioid groups. Lichenologists have still not 

agreed on the proper treatment of chemical variation, even 

though it is a fundamental species character in many other 

fungal groups. An overview by Hawksworth (1976) is the 

most recent summary of this topic. An objective solution to 
NUMBER 66 

FIGLRE 12.--Conidia of Parmelia species: a, P. cochleata (isolectotype in US); 6, P. cunninghamii (Santesson 

7814); C, P. dbcordanr (Kjellmerf m.); d P. tesfacea (James 561); e, P. erwnpenr (Hale 43657); J P. 

murmariza (h'akanbhi 2811); g, P. omphalodes (Hale 18851); h I? saxatilis (Hale 49288); i, P. sulcata (Hale 

49675a). (Scale in i = 10 pm.) 

15 
16 SMITHSONIAN CONTRIBUTIONS TO BOTATTY 

this controversial problem will probably not be reached until 

more experiments, such as those begun by C.F. Culberson 

(Culberson and Ahmadjian, 1980), are completed. 

Ultimately some “chemical species” will be found on closer 

study to have subtle morphological differences, obviating any 

decision as to their validity. Others will stand as good species, 

especially when strong correlating geographical or habitat 

differences are found. Finally some will be rejected as ever 

more refined analytical tests show them to be artifacts of 

earlier, imperfect tests, which were incapable of detecting 

minor metabolites (Elix, 1982). Another obstacle to an orderly 

solution is that very few “chemical species” have been 

adequately sampled and studied in the field. 

There are several species pairs in Parmelia that may be 

considered nearly identical except for chemistry and which I 

am recognizing at the species level in this monograph. The 

best known pair is P. omphalodes (salazinic acid) and P. 

discordans (protocetraric acid). Although these are broadly 

sympatric, Skult (1984) found that P. discordans is more 

strongly oceanic in distribution. A second pair is isidiate P. 

saxatilis (salazinic acid), which is mostly saxicolous and 

boreal, and P. kerguelensis (protocetraric acid), which is 

corticolous in North America and mainly saxicolous in South 

Africa. A third pair is P. norcrambidiocarpa (echinocarpic 

acid) and P. salcrambidiocarpa (salazinic acid), which are 

allopatric in Australia and New Zealand. Similar chemical 

variation is used in part to separate P. testacea (salazinic acid 

and unknown #27) and P. subtestacea (echinocarpic acid and 

unknown #27). On the other hand, I am not recognizing the 

protocetraric acid chemotype of Australian P. signifera 

(unnamed), because there are too few collections available to 

make a decision. 

Phytogeography 

Parmelia behaves as a typical boreal-temperate genus in 

North America and Europe. It is a small, well collected group 

of 9 species familiar to all lichenologists. Africa has a very 

small Parmelia flora with no endemic species. Eastern Asia 

has been the most important center of evolution for the genus. 

Japan alone has at least 12 species, and Asia as a whole 17, 

10 of them endemics. Three species, P. meiophora, P. 

ricasolioides, and P. submutata, seem to have originated in the 

mountains of southern China. 

Parmelia has also evolved extensively in Australasia, where 

14 species have been found. Twelve of these, P. crambidiocarpa, P. cunninghamii and P. protosulcata (both also in 

austral South America), P. norcrambidiocarpa, P. novaezelandiae, P. pseudotenuirima, P. queenslandensis, P. salcrambidiocarpa, P. signifera, P. subtestacea, P. tenuirima, and 

P. testacea, occur only in the southern hemisphere. The 

remaining two, P. kerguelensis and P. sulcata, occur in both 

hemispheres. 

The 38 species of Parmelia are distributed as follows in the 

major geopolitical units of the world: 

NORTH AMWCA 

Canada: P. fertilis, P.fraudans, P. hygrophila, P. kerguelensis, P. omphalodes, 

P. saxatilk, P. skultii, P. squarrosa, P. sulcala. 

USA: P. discordans, P. fraudans, P. hygrophila, P. kerguelensis, P. 

neodiscordans, P. omphalodes, P. saxatilis, P. skultii, P. squarrosa, P. 

sulcata. 

SOLTH AMWCA 

Argentina: P. cunninghamii, I? protosulcata, P. saxatilis, P. sulcata. 

Chile: P. cunninghamii, P. protosulcata, P. saxatilis, P. sulcata. 

Falkland (Malvinas) Islands: P. cunninghamii, P. smtilis, P. sulcata. 

EUROPE 

GreenlandAceland: P. fraudnns, P. omphalodes, P. saxatilis, P. sulcata. 

Europe (including Turkey): P. discordam, J? fiaudans, P. omphalodes, P. 

saxafilis, P. skultii, P. submontana, P. sulcata. 

AFRICA 

Tunisia: P. submontana. 

East Africa (Ethiopia, Kenya): P. sautilis, P. sulcata. 

South Africa: P. erumpens, P. kerguelensis. 

ASIA 

Japan: P. adougescens, P. cochleata, P. erwnpens, P. fertilis, P. isidioclada, 

P. laevior, P. marmariza, P. pseudolaevior, P. pseudoshinanwna, J? 

shinanoana, P. squarrosa, P. sulcata. 

Korea: P. cochleata, P. fertilis, P. squarrosa. 

Mongolia: P. cochleata, P. omphalodes. 

China: P. fertilis, P. laevior, P. marmariza, P. meiophora, P. ricasolioides, 

P. squarrosa, P. submutata, P. sulcata. 

Himalayan region (northern India, Nepal. Sikkim, Pakistan): P. adaugescens, 

P. marmariza, P. meiophora, P. omphalodes, P. ricasolioides, P. squarrosa, 

P. submutata, P. sulcata. 

Southern India: P. erwnpens. 

SO~HEAST AIA

Taiwan: P. adaugescens, P. erumpens, P. fertilis, P. laevior, P. marmariza, 

J? niitakana, P. submutata. 

Philippines: P. isidioclada, P. sectilis. 

Sabah: P. erumpens, P. sectilk. 

Indonesia: P. erumpens. 

AUSTRALASIA 

Australia: P. cunninghamii, P. erwnpens, P. norcrambidiocarpa, P. pseudotenuirima, P. queenslandensis, P. salcrambidiocarpa, P. signifera, P. 

sulcata, P. tenuirima, P. festacea. 

New Zealand: P. crambidiocarpa, P. cunninghamii, P. kerguelensis, P. 

norcrambidiocarpa, P. novae-zelandiae, J? protosulcata, P. salcrambidiocarpa, P. saxatilis, P. signifera, P. subtestacea, P. sulcata, P. tenuirima, P. 

festacea. 

Key to the Species of Parmelia 

1. Thallus with powdery, granular, pustulate or subisidiate soredia (under high magnification the isidia-like soredia of P.fraudans 

and P. isidioclada may be mistaken for true isidia). 

2. Pseudocyphellae punctiform, occurring only on lobe edges , . . . . . . . . . . . . . . . . . . . . . . . . . . R isidioclarla 

2. Pseudocyphellae predominantly effigurate, on lobe surface and/or margins. 
NUMBER 17 66 

3. Soredia pustular, mostly laminal; cortex deeply fissured ........................... l? erumpens 

4. Medulla K- (protocetraric acid) ...................................... f? Protosulcata 

6. Rhizines strongly squarrosely branched; pantemperate-panboreal ................... l? sulcata 

6. Rhizines simple; Mediterranean region .............................. Z? submontana 

7. Soredia yellowish; boreal regions ................................... l? fraudans 

7. Soredia greenish or whitish gray; austral regions ........................ P. cunninghamii 

9. Pseudocyphellae round, punctiform, only on lobe edges ......................... Z? pseudolaevior 

10. Pseudocyphellae conspicuous as a broad white rim; Japan ................... l? pseudoshinanoana 

10. Pseudocyphellae inconspicuous; southeastern Asia .............................. l? sectilis 

3. Soredia powdery, marginal or along ridges on surface; cortex not deeply cracked. 

4. Medulla K+ yellow turning red (salazinic acid). 

5. Soralia mostly laminal along ridges. 

5. Soralia mostly marginal. 

1. Thallus lacking soredia. 

8. Thallus with dense suberect, marginal (or in part laminal) dorsiventral lobules or isidiate lobules. 

9. Pseudocyphellae elongate, mostly marginal. 

8. Thallus without conspicuous suberect lobules (P. discordans, I? neodiscordans, P. omphalodes, P. subtestacea, and P. 

testacea may have dense appressed secondary lobules); if isidiate, the isidia mostly laminal. 

1 1. Thallus isidiate. 

12. Medulla K- (protocetraric acid) .................................... l? kerguelensis 

13. Isidia produced only along lobe margins .............................. I! isidioclada 

and Taiwan. ........................................... l? meiophora 

12. Medulla K+ yellow turning red (salazinic acid). 

13. Isidia produced mostly on lobe surface or along ridges on lobe surface. 

14. Upper surface finely white-maculate pseudocyphellate, the pseudocyphellae less than 0.5 mm long; China 

14. Upper surface with larger effigurate pseudocyphellae more than 0.5 mm long. 

15. Isidia dull, decomposing and becoming subsorediate; Pacific Northwest ......... P. hygrophila 

15. Isidia usually shiny, corticate, not becorning sorediate. 

16. Thallus small, 3-5 cm broad, closely attached; isidia very dense; Australia . . Z? pseudotenuirima 

16. Thallus larger, 5-20 cm broad, adnate to loosely attached; not occurring in Australia. 

17. Lobes broad and apically rotund, 4-10 mm wide .............. P. novae-zelandiue 

17. Lobes narrower, sublinear, 1-5 mm wide. 

18. Rhizines simple to furcate .............................. l? saxatilis 

18. Rhizines squarrosely branched .............. .......... P. squarrosa 

11. Thallus lacking isidia. 

19. Medulla K- (fumarprotocetraric acid, protocetraric acid or unknown #27). 

20. Thallus corticolous; New Zealand. 

21. Lobes subirregular, contiguous; pseudocyphellae mostly marginal ....... l? subtestacea (chemotype) 

21. Lobes sublinear, separate; pseudocyphellae mostly laminal ................ R crambidiocarpa 

22. Fumarprotocetraric acid present; North America ...................... l? neodiscordans 

22. Protoceuaric acid present; Europe ................................ Z? discordans 

23. Pseudocyphellae punctate, on lobe edges ................................ l? laevior 

23. Pseudocyphellae effigurate, laminal and/or marginal. 

20. Thallus saxicolous; Europe and North America. 

19. Medulla K+ yellow or yellow turning red (salazinic acid or echinocarpic acid). 

24. Pseudocyphellae marginal as a white rim around lobes; Japan and East Asia only. 

25. Saxicolous; cortex C+ rose (gyrophoric acid) ....................... l? shinanoana 

25. Corticolous; cortex C- (atranorin only). 

26. Lobes subirregular, apically rotund; Japan ........................ R cochleata 

26. Lobes sublinear, apically obtuse; Taiwan ........................ l? niifakana 

24. Pseudocyphellae laminal and marginal, not forming a conspicuous marginal rim (except in P. subtestacea 

and P. testacea from New Zealand). 

27. Rhizines squarrosely branched. 

28. Pseudocyphellae less than 0.3 mm long, appearing as fine white maculae; Taiwan and China 

.............................................. Z?submutata 
18 SMITHSONIAN CONTRIBUTIONS TO BOTATTY 

28. Pseudocyphellae distinct, effigurate, up to 1 mm long. 

29. Lobes broad and rotund, to 10 mm wide; pseudocyphellae separate, laminal ... P. tenuirima 

29. Lobes sublinear, 1-5 mm wide; pseudocyphellae marginal and/or laminal, often fusing. 

30. Lobes subirregular; pseudocyphellae mostly marginal. 

31. Salazinic acid present ............................. I! testacea 

31. Echinocarpic acid present ......................... Z? subtestacea 

32. Collected in Japan, East Asia and Canada ................... l? fertilis 

34. Spores more than 16 pm long ................ P. crambidiocarpa 

34. Spores less than 16 pm long ............... Z? norcrambidiocarpa 

and New Zealand ..................... l? salcrambidiocarpa 

crowded; Queensland and northern N.S.W., Australia ... I! queenslandensis 

37. Apothecia numerous, to 2 mm in diameter; Himalayan region ......... Z? ricasolioides 

39. Salazinic acid present ............................. I! testacea 

39. Echinocarpic acid present ......................... I! subtestacea 

40. Salazinic acid present (without norstictic acid) .............. l? omphalodes 

40. Salazinic and norstictic acids present ...................... P. skultii 

42. Pseudocyphellae laminal; upper surface becoming fissured; Australasia .... I! signifera 

......................................... l? omphalodes 

43. Pseudocyphellae separate, less than 0.5 mm long ............... I! marmarim 

30. Lobes sublinear; pseudocyphellae mostly laminal. 

32. Collected in Australasia. 

33. Echinocarpic acid present. 

33. Salazinic acid present. 

35. Rhizines forming a dense, projecting mat; lobes sublinear, separate; Tasmania 

35. Rhizines sparse to moderate, not projecting; lobes sublinear but short and 

27. Rhizines simple to furcate. 

36. Pseudocyphellae mostly marginal. 

37. Apothecia, if present, larger, to 15 mm in diameter; boreal areas or New Zealand. 

38. Thallus greenish mineral gray; Australasia. 

38. Thallus whitish to brownish gray or blackening; boreal regions. 

36. Pseudocyphellae mostly laminal. 

41. Collected on rocks or soil; thallus often turning brownish. 

42. Pseudocyphellae laminal and marginal; upper surface continuous; boreal regions 

41. Collected on trees; thallus greenish or whitish mineral gray. 

43. Pseudocyphellae separate or fusing into a network, to 1 mm long. 

44. Collected in Australasia (see 32). 

44. Collected in Japan and eastern Asia. 

45. Upper surface conspicuously cracked with age; pseudocyphellae mostly laminal 

45. Upper surface not cracking conspicuously; pseudocyphellae marginal, forming a 

.................................... I! adaugescens 

46. Spores about 30 pm long; Taiwan ................. I! niitakuna 

46. Spores about 15 pm long; Japan, East Asia ............ P. cochleata 

nearly continuous rim. 

Taxonomic Treatment 

The 38 species of Parmelia are arranged below in 

alphabetical order. Locations of specimens cited in synonymies 

and “Specimen Examined” sections are indicated by standard 

herbarium acronyms. 

Parmelia 

Parmelio Achanus, 1803: 153 [nomen conservanduml. 

Lichen L., 1753: 1140. [kctotype species: Lichen saxnrilis L.] 

Aspidelia Stirton, 1900231. [kctotype species Aspldeliu beckefrii Stirton 

(= Parmeliu tenuirima, see Culberson, 1966).] 

DESCmoN.-Thallus foliose, 4-60 cm broad, greenish to 

whitish mineral gray; lobes sublinear to more rarely subirregular, 1.5-10 mm wide; upper &face plane to foveolate, 

isidiate, sorediate, or lacking soredia and isidia, pseudocyphellate, the pseudocyphellae usually effigurate, rarely punctiform; 

lower surface black, moderately to densely rhizinate, the 

rhizines simple, furcate or squarrosely branched. Pycnidia 

common, immersed, laminal; conidia cylindrical to slightly 
NUMBER 66 19 

bifusiform, straight or slightly bent, 5.5-8.0 pm long. 

Apothecia adnate to substipitate, 1-20 mm in diameter, the 

disc pale brown, the amphithecium usually pseudocyphellate; 

spores 8/ascus, simple, colorless, 3-18 x 6-33 pm long, the 

episporium 14 pm thick. 

CHEMISTRY .-Atranorin and chloroatranorin (rarely gyrophoric acid) in the cortex: salazinic acid, consalazinic acid, 

protocetraric acid, fumarprotocetraric acid, echinocarpic acid, 

or unidentified substances and as accessory or trace substances 

conechinocarpic acid, galbinic acid, lobaric acid, norstictic 

acid, protolichesterinic or related fatty acids, or stictic acid in 

the medulla. 

Parrnelia adaugescens 

FIGURES 2a, 13a 

Parmelia adaugescens Nylander, 1890:28. [Type collection: Ichigome, Japan, 

Parmeiia pseudomarmariza Awasthi, 1976: 186. [Type collection: Mervakhola 

Alrnqubf (H, Nyl. herb. no. 34876, lectotype).] 

Valley, Nepal, Awasfhi 2305 (AWAS).] 

DEscmoN.-Thallus adnate on bark, rather firm, 8-15 

cm broad, greenish to whitish mineral gray (brownish in the 

herbarium); lobes sublinear, long, contiguous, 2-5 mm wide: 

upper surface shiny, plane, rugose-foveolate with age, transversely cracked in older parts, pseudocyphellae effigurate, 

conspicuous, 0.5-1.5 mm long, marginal and laminal, mostly 

separate or forming a loose network; lower surface black, 

densely rhizinate, the rhizines simple to furcate or dichotomously branched, 1-2 mm long. Pycnidia common but 

poorly developed: conidia cylindrical, straight, 7-8 pm long. 

Apothecia common, substipitate, urceolate and inrolled when 

young, 4-8 mm in diameter, the amphithecium rugose, 

pseudocyphellate; hymenium 65-70 pm: spores 13-15 x 

21-27 pm, the episporium distinct, 2-3 pm thick. 

CHEMISTRY.-Atranorin and salazinic acid (Asahina, 195 lb) 

and consalazinic acid, 

REMARKs.-This widespread Asian species is characterized 

by the large spores and large, uniformly dispersed pseudocyphellae. It is most common in Japan and Taiwan but extends 

as far west as Pakistan in Asia. It seems rather isolated from 

other large-spored species such as P. isidioclada and P. sectilis, 

which have different orientation of the pseudocyphellae. There 

is some intergradation with P. cochleata, which has small 

spores, as well as with P. fertilis, which also has small spores 

and richly branched rhizines. 

Parmelia pseudomarmariza, described from Nepal, has 

similarly large spores and slightly smaller but comparable 

pseudocyphellae. It does not seem sufficiently different from 

P. adaugescens to be kept as a distinct species. 

Specimens Examined 

Pakistan: Iqbal 742 (US). Sikkim: Hara ef al. s.n. (TNS, US). India: West 

Bengal, Hara ef al. s.n. (Th’S, US). China: Prov. Heiho, Takahashi 2928 

(TNS). Japan: Prov. Awa, Fujikawa (TNS); Prov. Inaba, Naknnbhi 24 WS); 

Prov. Ishikari, Asahina s.n. (TNS); Prov. Iwashiro, Kwokawa 58169 (TNS, 

US); Prov. Iyo, Nakanishi 56 (US); Prov. Kii, Numajiri s.n. (TNS); Prov. 

Kozuke, Asahina s.n. (TNS); Prov. Musashi, Asahina s.n. (TNS); Prov. Mutsu, 

Hale 29301,29302,29306,29312,29355 (US); Prov. Shimotsuke, Kwokawa 

56530A (TNS); Prov. Tosa, Fujikawa s.n. (TNS). Taiwan: Kwokawa 1209 

mS, US); Miaoli County, hi 6874,7795,7809,7813 (US); Nantou County, 

Suzuki 10901 (US); Taichung County, hi 6876,6875 (US). 

Parmelia cochleata 

FIGURES 26, 136 

Parmelia cochleafa Zahlbruckner, 1927b3350. [Type collection: Mt. Fuji, 

Japan, Asahina 27 (W, holotype; TNS, US, isotypes).] 

Parmelia marmariza var. physcioides Zahlbruckner, 1927b3352. [Type 

collection: Mt. Buko, Musashi, Japan, Asahina 24 (W, holotype, TNS, 

isotype).] 

Parmelia pseudosaxafilis Asahina, 195 1b:354. [Type collection: based on P. 

marmariza var. physcioides Zahlbruckner.] 

DEscmIoN.-Thallus loosely adnate on bark, firm, greenish mineral gray, 6-20 cm broad; lobes sublinear to 

subirregular, crowded, 1-5 mm wide, the margins lobulate, 

lobules rotund, 1-2 mm wide, more or less concave or 

hood-shaped, becoming suberect; upper surface plane to very 

weakly foveolate, continuous, shiny, becoming pruinose at the 

lobe tips, pseudocyphellae marginal, forming a nearly continuous narrow white rim, only sparsely developed on the lobe 

surface, separate: lower surface black, moderately to densely 

rhizinate, the rhizines simple to furcate, 1-2 mm long. Pycnidia 

numerous: conidia (Figure 12a) cylindrical to weakly bifusiform, slightly bent, 5.5-7.0 pm long. Apothecia numerous, 

substipitate, 3-8 mm in diameter, the rim somewhat inrolled, 

finely crenate, the amphithecium sparsely pseudocyphellate; 

hymenium 60-70 pm: spores poorly developed, about 6 x 10 

pm, the episporium 1 pm thick (Asahina reports 6-8 x 12-13 

pm in type description). 

CmmsmY.-Atranorin and salazinic acid (consalazinic 

acid lacking). Protolichesterinic acid also reported by Kurokawa and Nakanishi (1971) in 2 of 5 specimens tested. 

REMARKs.-Parmelia cochleata is widespread but not 

especially common in Japan. In the typical form it has strongly 

cochleate lobes but at other times the lobes are quite flat and 

appressed and the species lacks other distinguishing features 

except the generally marginal pseudocyphellae, similar to those 

in P, rudior and P. testacea from New Zealand. 

Specimens Examined 

Korea: Kimwa s.n. (TA’S). Japan: Prov. Buzen, Kwokawa 62481 (TNS); Prov. 

Inaba, Ibma 2179 (Th’S), Nakanishi 12167 (KOBE); Prov. Iyo, Ogala 159 

(TKS); Prov. Kii, Kurokawa 60235, Numajiri 24 (ThS); Prov. Musashi, 

Kwokawa 64288 (TNS); Prov. Shinano, Hirafsuka s.n. (TNS); Prov. Sumga, 

Asahina 27 WS); Prov. Totomi. Nakanbhi 51 (Th‘S). Golubkova (1981) 

reports the species from Mongolia. 
20 SMITHSONIAN CONTRIBUTIONS TO BOTATTY 

FIGURE 13.Species of Parmelia: a, P. adaugescens (lectotype in H); b, P. cochleata (type of P. marmariza 

var. physcioides Zahlbruckner in W); c, P. crambidiocarpa (lectotype in W); d, P. cunninghamii (Santesson 

6792); e, P. discordam (Laurila in Lichenes Fennici Exriccati 195); f P. erwnpens (Hale 59471). (Scales in 

mm.) 
NUMBER 66 21 

Parmelia crambidiocarpa 

FIGURES 2c, 13c, 14 

Parmelia crambidiocarpa Zahlbruckner. 1941 :109. [Type collection: Wilkes 

Pools, Mt. Egmont National Park, North Island, New Zealand, Cranwell 

ZA205 (W, lectotype; CHR, isolectotype).] 

DEScruPnoN.-ThallUS adnate to loosely adnate on bark, 

firm, whitish or pale greenish gray, 6-15 cm broad; lobes 

sublinear, often little branched, subdivaricate, browning at the 

tips, 14 mm wide; upper surface shiny, continuous or cracking 

transversely with age, plane, pseudocyphellae forming an 

irregular but nearly continuous rim around the lobe margins, 

0.1-0.3 mm wide, also laminal and effigurate, 0.2-0.8 mm 

long, separate, fissuring with age; lower surface black, 

moderately to densely rhizinate, the rhizines simple to furcate 

or squarrosely branched, 1-2 mm long and often projecting as 

a mat around the lobe margins. Pycnidia common, 90-1 10 pm 

in diameter; conidia cylindrical, rod-shaped, 5.54.0 pm long. 

Apothecia common, becoming stipitate and cupuliform, 

splitting radially with age, to 20 mm in diameter, the disc pale 

tan to very dark brown, the amphithecium rugose and 

effigurate-pseudoc yphellate; hymenium 80-90 pm high; spores 

10-15 x 17-21 pm, the episporium about 2 pm thick. 

CHEMISTRY.-Atranorin, chloroatronorin, and protocetraric 

acid or echinocarpic acid (and associated unknowns) or 

protocetraric and echinocarpic acids together in nearly equal 

concentration. 

REMARKS.-Parmelia crambidiocarpa is a member of the 

P. testacea complex, closely related to P. norcrambidiocarpa 

and P. salcrambidiocarpa. Although Galloway and Elix (1983) 

synonymized it with I? testacea, I have concluded from my 

own field studies in New Zealand that it is a distinct species, 

separated by a series of highly correlated morphological and 

chemical characters. 

The morphological characters are quite distinctive. The 

thallus is typically loosely adnate, rather large, and with 

narrow, sublinear lobes. The pseudocyphellae are both 

marginal and laminal and usually fissure with age. The rhizines 

form a dense mat below and often project out from the margins. 

The apothecia tend to be strongly stipitate and nearly urceolate, 

although very large apothecia become flattened and radially 

split. Disc color ranges from very pale tan (11% of 37 fertile 

specimens examined) to light brown (59%) or very dark brown 

(30%). 

The spores of P. crambidiocarpa are consistently large. The 

average maximum length is 18.4 pm (range 15-21 pm) and 

width 11.4 pm (9-12 pm) (29 measurements). The episporium 

is about 2 pm thick. For the other two externally similar species 

in this group, P. norcrambidiocarpa and P. salcrambidiocarpa, 

spores do not exceed 15 pm in length and the episporium is 

only 1-1.5 prn thick. 

The chemistry is also distinctive. The type specimen from 

the Dawson Falls area at Mt. Egmont contains a mixture of 

echinocarpic and protocetraric acids. I visited this locality and 

made a random sample of 69 specimens in the subalpine scrub 

zone at 1000-1200 m elevation. The sample consisted of 39% 

with echinocarpic and protocetraric acids, 45% with echinocarpic acid (and associated unknowns)-although traces of 

protocetraric acid might have been missed on the TLC 

plates-and 16% with protocetraric acid alone. Unknown #27, 

so typical of the P. testacea group, was not detected in any 

specimens. 

Taking the whole range of P. crambidiocarpa, one finds a 

complex pattern for combinations of the acids. At Tongariro 

National Park, another isolated volcanic peak about 150 km 

east of Mt. Egmont, specimens with protocetraric acid alone 

made up an overwhelming 98% of the population (54 

specimens tested), the remainder containing both echinocarpic 

and protocetraric acids. A similar proportion appears to hold 

for the Urewera area farther to the east although the sample 

size (5) is too small to be sure. On the South Island, where the 

species is far less abundant and largely replaced by P. 

norcrambidiocarpa, all specimens (2 1 tested) contained protocetraric acid except for one collection in beech forest near 

Tuatapere and one (sterile) from Lewis Pass with both 

echinocarpic and protocetraric acids present. 

Parmelia crambidiocarpa has strong ecological requirements. On North Island of New Zealand it is abundantly 

developed-and in fact the only member of the P. testacea 

group except for rare P. salcrambidiocarpa-at 1000-1300 m 

elevation in the subalpine zone on a variety of trees and shrubs 

(Nothofagus, Dracophyllum, Nothopanax, Pseudopanax, etc.). 

I did not collect it below 600 m elevation. On South Island it 

occurs above 600 m in the highest passes but also grows near 

sea level at high latitudes in the extreme south, where it occurs 

with P. norcrambidiocarpa (Figure 14). In random samples 

made at 71 localities in New Zealand, I collected 149 

specimens (23%) of P. crambidiocarpa out of a total of 715 

collections in the P. testacea complex. A similar percentage 

(17% or 14 of 88 collections) was found in the herbarium 

collections at CHR. 

Representative Specimens Examined 

(protocetraric acid) 

Kew Zealand: North Island, Elix 8195, 8208 (ASUC), Hale 65108, 65527, 

65572 (US), Hayward H112.70 (US); South Island, Elix 7702 (AKUC). 8527 

(CHR), Hale 65146, 65236, 65497, 65782 (US), Imshaug 47879, 55985 

(MSC), Molloy CHR266952 (CHR). 

Representative Specimens Examined 

(echinocarpic acid) 

New Zealand: North Island, Davey CHR160226 (CHR). Hale 65296, 65533 

(US). 
22 SMITHSONIAN CONTRIBUTIONS TO BOTAh'Y 

FIGURE 14.-Distribution of Parmelia crambidiocarpa in New Zealand. 
NUMBER 66 23 

Representative Specimens Examined 

(echinocarpic and protocetraric acids) 

New Zealand: North Island, Hale 65279, 65537, 65783 (US); South Island, 

Elk 7507 (CHR), Hale 65779 (US). 

Parmelia cunninghamii 

FIGURES 26 13d 

Parmelia cunninghamii Crombie, 1876:228. [Type collection: Island Harbour, 

Falkland Islands, 18 April 1868, Cunningham (BM, lectotype; H, isotype).] 

Parmelia brownii Dodge, 1970:449. [Type collection: Camp Hill, Macquarie 

Island, Brown 69 (FH-Dodge).] 

DEscRIPnoN.-Thallus adnate on bark, more rarely on rocks, 

very firm, pale greenish to brownish mineral gray, 8-12 cm 

broad; lobes subirregular, apically subrotund, contiguous, 

usually turning up at the margins, 3-7 mm wide; upper surface 

shiny, becoming white pruinose at the tips, plane to 

rugose-foveolate with age, reticulately fissured only in older 

parts, sorediate, the soralia linear along the margins, orbicular 

when laminal, pseudocyphellae effigurate, 0.3-1 mm long, 

rather sparsely developed to well developed, marginal and 

laminal, separate, sinuous; lower surface black, moderately to 

densely rhizinate, the rhizines simple to squarrosely branched, 

1-2 mm long. Pycnidia rare, marginal; conidia (Figure 12b) 

cylindrical, straight to slightly bent, 5.5-7.0 p long. 

Apothecia rare, substipitate, 4-15 mm in diameter, the 

amphithecium arugose, pseudocyphellate, the disc brown; 

hymenium 65-70 p; spores 8-9 x 10-12 p, the episporium 

1 p thick. 

CHEMISTRY .-Atranorin and chloroauanorin, salazinic acid 

(Galloway and Elix, 1983), consalazinic acid, and accessory 

lobaric acid. 

REms.-This widespread austral species can be recognized by the firm, shiny thallus and well-developed marginal 

soralia. It differs from closely related P. sulcata, with which 

it occurs in austral regions, in several respects. For example, 

the surface is not strongly foveolate-ridged, the soralia are 

chiefly marginal, and the rhizines are not so densely 

squarrosely branched. Parmelia protosulcata, a narrow-lobed 

species with protocetraric acid, has sparser, often orbicular 

soralia. When these are better developed they become linear 

and approach P. cunninghamii very closely. 

Parmelia cunninghamii is preeminently a species of the 

moist forests of austral South America and New Zealand. 

Farther west in Australia it occurs in temperate rain forest. 

Specimens Examined 

Argentina: hv. Chubut, Lumb 5860 (US); Cerro Mazo, James 1644 (BM); 

SW Patagonia, James 579 (BM); Tiena del Fuego, Sawesson 1145 (S, US). 

Falkland Islands: lmhaug 40162 (US). Chile: Prov. Chiloe. Redon 03575 

(US); Prov. Malleco. Eyerdm 10860 (COLO, US), Mahu 2537 (US); Ten. 

Magallanes. lmshaug 44566 (MSC), Sonfesson 7814 (US), 8012, 8188 (S); 

Tiena del Fuego, Sawlosson 5910,6013,6792,7385,73% (S, US), 5812,6005, 

7491 (S); Valdivia, Mahu 1699 (US). New Zealand: Antipodes Island, Du 

Rietz 2598:3 (UPS, US); South Island Fiord District, Du Riefz 1856a (UPS, 

US); South Island-Eastem Botany District, Du Rietz 1462:26 (UPS, US); 

Otago, James 611/2 (BM), Martin 1116 (BM), Murray 1273 (BM). James 

1623 (BM, US), Thomon 2.136 (CHR); Canterbury, Elk 8570 (CHR), 

Galloway s. n. (CHR), Tibell 9232, 9439, 9682 (UPS); Auckland Islands, 

Imhaug 56283 (MSC), James 1345 (BM), 1378 (FH, US). Australia: New 

South Wales, Hale 66605 (US), South Australia,Elix 4901 (ANUC); 

Tasmania, Elix 5593 (ANUC); Victoria, Bastow (US), Elk57 (AXUC), 

James Au2136, Au2138 (BM); Marion Island, Huntley 971 (US). 

Parmelia discordans 

FIGURES 2e, 13e 

Parmelia discordans Nylander in Brenner, 1886:40. [Type collection: Hogland, 

Finland, Brenner s.n. (H, Nyl. herb. no 34916. lectotype).] 

Parmelia omphalodes f. insensitiva Magnusson, 1919:89. [Type collection: 

Vestrogothia, Goteborg, Sweden, Magnumon s.n. (UPS, lectotype (not 

seen); FLAS, isolectotype).] 

Parmelia omphalodes var. discordans (Nylander) Magnusson, 1929:89. 

Parmelia insensitiva (Magnusson) Anders, 1928:152. 

Parmelia omphalodes subsp. discordans (Nylander) Skult, 1984: 138. 

DEscmrIoN.-Thallus adnate to loosely adnate, very fragile 

and brittle, dark to blackish brown, 4-8 cm broad; lobes 

sublinear, short and imbricate, 1-3 mm wide, usually 

developing small secondary lobes with age; upper surface plane 

to weakly foveolate, shiny, continuous, sparsely cracked with 

age, pseudocyphellae mostly marginal but a few also laminal 

on older lobes, small, 0.2-0.5 mm long, effigurate and 

coalescing at times to form a loose, mostly marginal reticulate 

network; lower surface black, sparsely to moderately rhizinate, 

the rhizines simple to furcate, 1-1.5 mm long. Pycnidia not 

common: conidia cylindrical, straight to slightly bent, 5.5-6.5 

pm long (Figure 12c). Apothecia rather rare (25% frequency 

in Finland (Skult, 1984)), adnate to substipitate, 2-5 mm in 

diameter, the amphithecium pseudocyphellate-reticulate, the 

disc plane; hymenium 55-60 pm; spores 8-10 x 14-16 pm, 

the episporium 1.5-2 pm thick. 

CmhlrsmY.-Atranorin, protoceuaric acid, and lobaric acid 

(Culberson, 1970) with accessory substances galbinic acid, 

fumarprotocetraric acid and unidentified fatty acids (Skult, 

1984). 

REmms.-Lichenologists are not in agreement on the 

status of Parmelia discordans, as it appears to be a “chemical 

species.” Many Europeans (e.g., Degelius, 1931; Krog et al., 

1980: Santesson, 1984) consider it to be at most a variety (var. 

discordans (Nylander) Magnusson) of P. omphalodes. Skult 

(1984) calls it a subspecies of P. omphalodes. Anders (1928), 

Culberson (1970), and Kurokawa (1976) used specific rank. 

Morphologically P. discordans seems to be consistently 

smaller, with slightly narrower lobes. Parmelia omphalodes 

itself, however, has a very great range of variation, overlapping 

this character and almost any other morphological character 

one might want to designate as unique to P. discordans. 

Nylander himself compared it originally to “I? prolixa var. 

panniform’s,” a brown Parmelia. The apothecial and pycnidial 

characters are essentially identical for the two species. 
24 SMITHSONIAN CONTRIBUTIONS TO BOTAhY 

Parmelia discordans appears to be a relatively rare species 

in Europe, at least in comparison with P. omphalodes. Degelius 

(1931, 1944), however, found it to be more common on the 

basis of a KOH test in Angermanland and Gotland, Sweden, 

and in Westland, Norway (1934). Skult (1984) collected it 

intensively in Finland, where it behaves as an oceanic species 

in the southwestern part of the country. 

Specimens Examined 

Europe. Great Britain: Scotland, Denison 251, Dixon, s.n. (US). Finland: 

Satakunta, Luurila in Lichenes Fennine Exriccati 195 (US). Sweden: Bohuslan, Magnusson in Lichenes Selecfi Exsiccuti 106 (FLAS) and Krypfogamae 

Exsiccatae 257 1 (US); Smaland, Zetterstedf s.n. (US;) Vastmanland, Kjellmert 

s.n. (US); Vestrogothia, Magnusson, s.n. (US). Belgium: Prov. Luxembourg, 

Lornbinon in Vezda, Lichenes Selecti Exsiccafi 916 (US). France: Sarthe, 

Monguillon in des Abbayes, Lichenes Gallici 39 (BM). Spain: Asturias, 

Vmquez s.n. (US). 

Parmelia erumpens 

FIGURES 2J 13J 15 

Parmelia erwnpens Kurokawa, 1969, no. 74. [Based on Parmelia tenuirima 

f. corallina Muller Argoviensis.] 

Parmelia fenuirima f. corollino Muller Argoviensis, 1883:46. [Type collection: 

Gippsland, Australia, Stirling s.n. (G, lectotype; UPS, US, isolectotypes).] 

DEscmoN.-Thallus adnate to loosely adnate on rocks 

and trees, fairly firm, pale greenish to whitish mineral gray, 

8-20 cm broad; lobes short, subirregular to apically rotund, 

imbricate, 2-8 mm wide; upper surface shiny, plane, soon 

becoming conspicuously reticulately cracked, pseudocyphellae 

effigurate, 0.2-1.0 mm long, somewhat raised, dense, fusing 

into a reticulate network over the whole surface, sorediate, the 

soredia coarse and isidioid, often bursting apically, forming 

dense marginal and laminal soralia; lower surface black, 

moderately rhizinate, the rhizines simple or squarrosely 

branched at maturity, 1-2 mm long. Pycnidia rather rare; 

conidia (Figure 12e) cylindrical to weakly bifusiform, straight, 

5.5-7.0 pm long. Apothecia rame, stipitate, the rim inrolled 

but the disc flattening and splitting radially at maturity, 4-15 

mm in diameter, the amphithecium reticulately cracked, 

pseudocyphellate, sorediate; hymenium 65-70 pm; spores 6-8 

x 1&12 pn, the episporium 1 pm thick. 

CmmsmY.-Atranorin and chloroatranorin, salazinic acid 

(Galloway and Elix, 1983), and consalazinic acid. 

REmRKs.-This is one of the more widespread AsianAfrican species (Figure 15), easily recognized by the deeply 

reticulately fissured cortex and abundant coarse pustular 

soredia. In Australia it is collected most commonly on 

sandstone outcrops in sclerophyll forests but in other areas it 

is usually corticolous. In India, for example, it occurs at 

1400-2200 m elevation in open montane forests. In New 

Zealand it grows in mature, moist Nothofagus forests, and in 

South Africa it is found in the wet coastal forests in the 

Tsitsikama area. It seems to lack any recognizable parent 

morph, although P. signifera would have to be considered very 

closely related. Parmelia signifera has larger spores (13-15 

pm) and greater chemical variation. Another parent morph 

proposed by Galloway and Elix (1983), P. tenuirima, is a 

much larger lichen with separate pseudocyphellae and larger 

spores (12-15 pm long). 

Specimens Examined 

Australia: Lord Howe Island, Wafts (US); Australian Capital Territory, 

Streimann 4452. 7893 (US); New South Wales, Cheel and Borman 1045 

(BM, US), Craigie L1692 0, Degelius A-77 (US), Du Riefz 60a, 546h 

(UPS, US), Flockton 725 (US), Hale 58450, 58835, 59011, 59257. 59262, 

59471 (US), Kuroknwa 5151,6542 (TNS, US), Sfaer L1720 (NSW), Sfreimann 

9518 (US), Weber andMcVean L47307, L49615. L49912 (US); Victoria, Hale 

58254, 28258 (US), Sfreimann 2335 (US); Western Australia, Weber L50111 

(COLO, US). New Zealand: North Island, Galloway s.n. (CHR), Hale 59503, 

59499 (US), James 111 (BM, US), Tibell 13203 (UPS); South Island, Hale 

66234 (US). South Africa: Cape Province, Hale 72001, 72002 (US). India: 

Tamil Nadu, Hale 40245, 40250, 43578, 43657, 43813, 43815, 43876 (US). 

Sabah: Hale 29017 (US). Kurokawa 0969) has published additional records 

from Japan, Taiwan, and Indonesia. 

Parmelia fertilis 

FIGURES 3a, 16a 

Parmelia ferfilis Muller Argoviensis, 1887:3 16. [Type collection: Siberia, 

Parmelia subdivaricafa Asahina, 1951b:356. [Type collection: Mt. Arisan, 

USSR, corn. Lohrn, no. 7 (G, lectotype).] 

Nimandaira, Taiwan, Asuhina F.71 (TNS, lectoptype).] 

DEscmoN.-Thallus adnate on bark, firm, pale greenish 

to whitish mineral gray, 6-12 cm broad; lobes sublinear, fairly 

long and divaricate to contiguous (shorter and more adnate in 

Canadian specimens), 1.5-5 mm wide; upper surface plane to 

weakly foveolate, transversely cracked with age, pseudocyphellae effigurate, conspicuous, 0.5-1 mm long, marginal 

and laminal, sometimes forming a discontinuous white rim, 

fusing with age to form a coarse reticulate network; lower 

surface black, densely rhizinate, the rhizines simple to densely 

squarrosely branched, 0.5-2 mm long. Pycnidia common; 

conidia cylindrical to weakly bifusiform, straight to slightly 

bent, 5.5-6.5 pm long. Apothecia common, substipitate, the 

rim inrolled, 2-7 mm in diameter, the disc dark brown, the 

amphithecium finely reticulate-pseudocyphellate; hymenium 

55-60 pm; spores usually lacking or very poorly developed, 

6-8 x 12-14 pm, the episporium 1 pm thick. 

CmmsmY.-Atranorin and salazinic acid (Asahina, 1951b, 

in type of P. subdivaricata) along with consalazinic acid. 

REmRKs.-Asahina (1951b) described this as a new 

species, P. subdivaricata, unaware that Miiller Argoviensis had 

earlier published P. fertilis on the basis of a single rather poor 

specimen from Siberia. He noted the squarrosely branched 

rhizines, an important diagnostic character for the species. 

Asahina found bifusiform conidia, but material that I have 

examined has mostly cylindrical ones. 

Parmelia fertilis is very common in Japan on maples, oaks, 

and other trees in open forests along with its probable isidiate 

morphotype P. squarrosa. The probable sorediate morphotype 
NUMBER 66 25 

FIGURE 15.-Distnbution of Punneliu erumpens. 

is P. sulcata, a very rare species in the range of P. fertilis, 

which differs in having larger microconidia (6-8 pm long). 

Sharon Gowan and colleagues recently discovered the 

species in New Brunswick and Nova Scotia, where it is rare. 

The specimens are more crowded and adnate than the Japanese 

material but spore size and rhizine branching are identical. 

Specimens Examined 

Canada: New Brunswick, Gowan 3332-4 (CAN, US). Korea: Chosen, Asahina 

s.n. (TKS), Fujikuwu WS). Japan: Prov. Awa, Fujikuwa (TNS); Prov. Bingo, 

Safo 60 (TNS); Prov. Bungo, Kuroknwu 62356 (TNS), 63194 in Lichenes 

Critici el Selecti 27 (US); Prov. Buzen, Asuhim s.n. (TNS); Prov. Etchu, 

Nirhijimu 18 (TSS); Prov. Hoki, Asahina s.n. WS); Prov. Inaba, Nukanishi 

12076 (KOBE); Prov. Ishikari, Endo 216 (TNS); Prov. Iwaki. Kuroknwu 58083 

(TXS), 78009 in Lichenes Selecti ef Critici 426 (US); Prov. Iyo, Nukunishi 90 

(KOBE), Yunugiruwu 1484 mS), Kurokawa 60050 in Lichenes Rariores 

Critici Ersiccufi 85 (as P, subdivaricufa) (US); Prov. Kii, Nwnajiri s.n. (TNS); 

Prov. Mutsu, Asuhinu 46 (TNS), Hale 29333, 29342 (US), Kurokuwu 56218 

(TSS); Prov. Rikuchu, Nukunishi B92 (KOBE); Prov. Sagami, Kurokawu 

58050 (US); Prov. Shimotsuke, Hale 63138 (US), Ogafu 1815 (mS); 

Saghalin, Asuhinu s.n. (n'S); Prov. Yamato, Tugawu s.n. (US). Taiwan: 

Chia-yi County, Koponen 17314 (US); Taichung County, hi 6866 (US); 

Taitung Kwokuwu 2586 (US). Zhao et al. (1982) reported it from Anhui, 

Chekiang, and Kiangsi provinces in China (as P. subdivaricata). 

Parmelia fraudans 

FIGURES 36, 16b 

Parmelia saxutilis (L.) Acharius *frauduns Nylander, 1861:lOO. [Type 

collection: Savolaxia, Finland, Nylander (H, Nyl. Herb. no. 34869, lectotype. 

A syntype (Malmgren: Kajana) was not located at H).] 

Parmeliafraudans (Nylander) Nylander, 1890:28. 

DEscwrIox.-Thallus adnate on rock, rather brittle, greenish to brownish mineral gray with a yellowish cast, 8-16 cm 

broad; lobes sublinear, short, contiguous to imbricate, 1-4 

mm wide; upper surface shiny or dull and white pruinose, plane 

to rugose-foveolate, transversely cracked with age, pseudocyphellae effigurate, small, to 0.5 mm long, mostly marginal 

and soon becoming densely sorediate, the soralia sinuous along 

lobe margins with a few isolated orbicular laminal soralia, the 

soredia coarse to almost subisidiose at maturity; lower surface 

dark brown and shiny at the margin, black at the center, densely 

rhizinate, the rhizines simple to furcate, 1-1.5 mm long. 

Pycnidia rather rare and often poorly developed; conidia 

iciylindrical to weakly bitusiform, straight to slightly bent, 

5.5-6.5 pm long. Apothecia rare, adnate, 2-3 mm in diameter; 

hymenium 45 pm; spores 5-6 x 10-12 pm, the episporium 1 

pm thick. 
26 

SMITHSONIAN CONTRIBUTIONS TO BOTATTY 

FIGW 16.4pecies of Parmelib: a, P. fertilir (Hale 29333); b, P. fr0udan.v (Hale 49870); c, P. hygrophila 

(Noble 6448); d, P. iridioclada (Kurokawa 58010); e, P. kerguelenris (lectotype in BM); J P. krguelernis 

(Brodo 11506). (Scales in mm.) 
SL‘MBER 66 21 

CHEhlrsmY.-Atranorin, usnic acid (in the soralia), salazinic 

acid, and protolichesterinic acid (Dahl, 1952; Krog, 1968). 

REMRKs.-Parmelia fraudans is a well-known boreal 

European species that also occurs in North- America. The 

presence of small quantities of usnic acid in the soralia is 

unique in the genus. The orientation of soralia is similar to 

that in P. sulcata, which often occurs in the same habitat but 

has densely squarrosely branched rhizines and lacks usnic acid. 

In North America P. fraudans is collected most often on 

rocks just above the high water mark in large lakes in the 

boreal zones but has also been found as far south as Arizona. 

Thomson (1984:302) has plotted the distribution of this species 

in arctic America. 

Representative Specimens Examined 

Canada: Northwest Territories, Ahti 14441 (US), Scotter 1953,2588, Thornson 

11873 (WIS), 14047 (US, WIS); Manitoba, Scotter 3007 (WIS). Ontario: Cain 

26998. Hale, 36414, 36510, 36665, 49852 (US). USA: Arizona, Nash 22636 

(ASU); Michigan, Hale 33890,34998 (US); Minnesota, Hale 33325, Wetmore 

34882 (US); Colorado, Anderson 2416, Shwhan 1139 (COLO, US); Montana, 

Standley 18567 (US). Greenland: Gelling in Lichenes Groenlandici Exsiccati 

121 (US). Norway: Ericbon 355. Hasselrot s.n., Lynge 5147 (US). Sweden: 

Hasselrot s.n.. Osterlind s.n., Santesson 13415 (US). Finland: Ahlner s.n. 

(US), Ldng in Kryptogamae exsiccatae 1970 (US), Rasanen in Lichenes 

Fenniae Exsiccati 45 (US). 

Parmelia hygrop h ila 

FIGURES 3c, 16c 

Parmelia hygrophila Goward and Ahti, 1983:9. [Type collections: 17 km E 

of Nelson, Kokanee Creek Park, BC, Canada, Coward 81-1601 (UBC, 

holotype; US, isotype).] 

DEscmIox-Thallus adnate on bark, whitish gray, often 

becoming densely pruinose, 4-10 cm in diameter; lobes 

sublinear, 3-5 mrn wide: surface plane to weakly ridged, 

pseudocyphellae effigurate, 0.1-0.3 mrn long, forming a 

network in the marginal area, erupting at maturity with the 

formation of coarse, granular, subsorediate isidia, the isidia 

very weakly corticate with pale tips, densely clumped and 

branched with age, 0.14.15 mm in diameter and 0.5-1.5 mm 

high; lower surface jet black, moderately rhizinate, the rhizines 

simple to sparsely furcate. Pycnidia not seen. Apothecia rare, 

substipitate, about 2 mrn in diameter; hymenium 50-60 pm; 

spores 8, simple, 9-12 x 14-16 pm (from original description). 

CIJEMSTRY .-Atranorin and sdazinic acid. 

REMARKs.-This species was only recently discovered in the 

Pacific Northwest. It is close to P, saxatilis, differing primarily 

in the “ecorticate soredioid isidia.” Another difference is that 

P. hygrophila is prirnarly a corticolous lichen in humid, oceanic 

forests, whereas P. saxatilis is saxicolous. 

Specimens Examined 

Canada: British Columbia, Noble 6448 (US). Other records are given in 

Goward and Ahti (1983). 

Parmelia isidioclada 

FIGURES 34 1M 

Parmelia isidioclada Vainio, 1921:48. [Type collection: Prov. Mimasaka, 

Parmelia yasudae Rasanen. 1940:84. [Type collection: Prov. Iyo, Japan, 

Parmelia psoromoides Rasanen, 1949:79. [Type collection: Mt. Panai, Prov. 

Japan, Yasuda 210 (TUR, lectotype).] 

Yasuda 656 (H, lectotype).] 

Benguet, Luzon, Philippines, Copeland 1378 (H, lectotype).] 

DEscmoN.-Thallus loosely adnate on bark or rock, 

rather firm, greenish mineral gray (turning brownish in the 

herbarium), 8-12 cm broad: lobes sublinear, elongate, contiguous, 2-4 mrn wide, the margins rolled upward, punctatepseudocyphellate along the edges, the pseudocyphellae soon 

becoming granular and giving rise to dense, branched, 

cylindrical to somewhat lobulate, eventually coralloid-isidioid 

growths along the length of the lobes, to 1 mm high, apically 

crumbling and subsorediate with age; upper surface plane, 

shiny, continuous; lower surface brown to whitish in a narrow 

marginal zone, black at the center, densely rhizinate, the 

rhizines simple to furcate, 1-1.5 mm long. Pycnidia not seen. 

Apothecia rare, substipitate, less than 5 mm wide; hyrneniurn 

about 60 pm high; spores 13-16 x 23-30 pm (from description 

of P. psoromoides). 

CHEmsmY.-Atranorin and salazinic acid (Asahina, 195 la; 

Krog, 1968) and consalazinic acid. 

REmws.-This rare, montane-tropical, temperate species 

appears to be relatooed too P. laevior because of the lateral 

punctate pseudocyphellae. It is unusual, however, in having 

very large spores, as well as marginal granular isidia. 

Specimens Examined 

Japan: Prov. Idzu, Kurokawa 58010 (TNS, US); Prov. Ohmi, Hale 29467a 

(US). China: Kiangsi, Hamet-Ahti 3322 (H). Krog (1968) tentatively identified 

a specimen from Alaska. 

Parmelia kerguelensis 

FIGLRES 3e, 16e,f 

Parmelia kerguelensir Wilson, 1900:87. [Type collection: Royal Sound, 

Parmelia pseudosulcata Gyelnik, 1934:282. [Type collection: Mary’s Peak, 

Kerguelen Island, Robert Hall s.n. (.MEL, lectotype).] 

Oregon, Sipe 684 (BP, holotype: iWCH, isotype).] 

DEscRIPrIox.-Thallus adnate to loosely adnate on bark or 

rock, fairly firm, greenish to light brownish mineral gray, 6-10 

cm broad; lobes sublinear, contiguous, sometimes short and 

more crowded, 1-2.5 mm wide, brownish towards the tips, at 

times white pruinose; upper surface dull, plane to weakly 

foveolate, continuous but becoming fissured with age, 

pseudocyphellae irregularly effigurate, 0.2-0.5 mrn long, 

sparse and inconspicuous to fairly well developed, marginal 

and laminal, isidiate, the isidia developing mostly on the 

surface, scattered initially but at maturity clumped along faint 

ridges, cylindrical, little branched, 0.04-0.06 x 0.1-0.3 mm; 

lower surface black, moderately rhizinate, the rhizines simple 
28 SMITHSOhTAN CONTRIBUTIONS TO BOTAKY 

to furcate, rarely weakly squarrosely branched, 0.5-1.5 mm 

long. Pycnidia not seen. Apothecia rare, substipitate, 10-15 

mm in diameter, flat with a radially split disc at maturity; 

hymenium 65-70 pm; spores 9-12 x 14-18 pm, the episporium 1-2 pm wide. 

CmmTRY.-Atranorin, protocetraric acid, and lobaric acid. 

REwims.-Parmelia kerguelensis has the same northern 

hemispheresouthern hemisphere distribution pattern as P. 

suxatilis and P. sulcata but is much less common. It is the only 

Parmelia species in South Africa. In the northern hemisphere 

it is known only from the moist coastal forests of the Pacific 

Northwest, where it was first recognized as P. pseudosulcata 

by Gyelnik. Although the specific epithet implies a close 

relationship with sorediate P. sulcata, this species is actually 

very close to P. saxatilis, as both have mostly simple rhizines 

and isidia. Without careful examination, in fact, P. kerguelensis 

would be identified as P. saxatilis. Aside from the clear 

chemical distinction (K- in the medulla), it lacks the 

conspicuous reticulate network of pseudocyphellae on the lobe 

surfaces and has a tendency for squarrose rhizines. In Canada 

at least P. kerguelensis is found on tree bark, P. saxatilis on 

rocks, a basic habitat difference. In South Africa, however, 

where trees are rare, it is found on sheltered sandstone outcrops 

as well as on trees in forested areas. 

Specimens Examined 

Canada: British Columbia, Ohlsson 1638, 2410, 2445 (CAh?, Brodo 11506 

(CAN). USA: California, Hale 57877 (US). Republic of South Africa: Cape 

Province, Almborn 5090, Hale 72142 (US), Maas Geesteranu 6820 (LD, 

US). New Zealand: South Island, Hale 66266 (US); Campbell Island, Harris 

4358, 4488, 5644,5670 (US). 

Parmelia laevior 

FIGURES 3f; 170 

Parmelia laevior Nylander, 1890:28. [Type collection: Ichigome, Japan, 

Parmelia laevior f. denigrata Hue, 1899:166. [Type collection: Onikobe, 

Parmelia pefrophila Vainio, 1921:48. [Type collection: Prov. Kii. Japan, 

Parmelia hakonensis Zahlbruckner, 1927b3348. [Type collection: Hakone, 

Parmelia laevior f. hakonensis (Zahlbruckner) Asahina, 1952:112. 

Parmelia onfakensir Asahina, 1954:323. [Type collection: Mt. Ontake, 

Hidaguchi 6-gome, Prov. Hida, Japan, Asahina 54819 (TSS, lectotype; US, 

isolectotype).] 

Almquist s.n. (H, Nyl. herb. no. 34857, lectotype; S. isolectotype).] 

Japan, Faurie 305 (PC, lectotype).] 

Yasuda 193 (TUR. Vain. herb. no. 2933, lectotype).] 

Prov. Sagami, Japan, Yamashifa 13 (TNS, isolectotype).] 

DEscmrroh'.-Thallus loosely adnate on bark, firm, pale 

greenish to brownish mineral gray, 6-15 cm broad; lobes 

sublinear, contiguous, 1 .54 mm wide, somewhat lobulate; 

upper surface shiny, plane to weakly rugose, sparsely 

transversely cracked with age, pseudocyphellae on lateral 

edges of the lobes, round to elongate, about 0.2 mm long; lower 

surface black moderately rhizinate, the rhizines simple to 

furcate to sparsely squarrosely branched at maturity, 1-2 mm 

long. Pycnidia common; conidia cylindrical, straight to slightly 

bent, 6-8 pm long. Apothecia common stipitate and urceolate, 

flaring at maturity with an open, radially split disc, 5-10 mm 

in diameter, the rim and amphithecium warty pseudocyphellate; hymenium 60-65 pm; spores 8-9 x 14-16 pm, the 

episporium distinct, about 1 pm thick. 

CHEMISTRY.-Atranorin, salazinic acid (Asahina, 195 la), 

and consalazinic acid. 

REmxs.-Parmelia laevior is a representative of a small 

isolated group of Parmeliae with lateral, punctate pseudocyphellae. This group includes P. pseudolaevior, a very close 

lobulate morphotype, and P. isidioclada, a large-spored 

temperate-tropical species. It is rather common in Japan on 

conifers and hardwoods: there are few collections elsewhere. 

Specimens Examined 

Japan: Prov. Aki, Hale 29551 (US); Prov. Bungo, Kurokawa 63192 (TNS); 

Prov. Buzen, Kurokawa 62479 (TNS); Prov. Hida, Kurokawa 64072 VS); 

Prov. Higo, Kurokawa 63121 (TNS); Prov. Hitachi, Asahina 12b (TNS); Prov. 

Hizen, Kurokawa 52599 (TNS); Prov. Hoki. Yasudn s.n. (TNS); Prov. Hyuga, 

Hale 29640, 29688 (US); Prov. Inaba, Yasuuia s.n. (TNS); Prov. Ishikari, 

Asahina s.n. (TKS); Prov. Iyo, Kurokawa 550149 (US); Prov. Kii, Numajiri, 

s.n. (TNS); Prov. Oita. Omura 707 (US); Prov. Osumi, Kurokawa 63055 

WS); Prov. ,Musashi, Shibuichi 4546 (US); Prov. Shimotsuke, Culberson 

10641 (US), Hale 63134 (GS), Ogata 8Oc (TNS); Prov. Shinano, Asahina 

547, 2627 (TNS), Kurokawa 58200, 58255 (US); Prov. Sumga, Culberson 

10691,11104. Iishiba 64,69,72 (US). Taiwan: Nantou, hi 2355 (US). USSR: 

Primorsk, Vasileve 10 (US). Zhao et al. (1982) repon it from Chekiang 

Province in China. 

Parmelia marmariza 

FIGURES 4a, 176 

Parmelia marmarim Nylander, 1890:28. [Type collection: Ichigome, Japan, 

Parmelia marmariza f. angustifolio Asahina, 1951b:353. [Type collection: 

Parmelia submarmariza Asahina, 1953:65. [Type collection: Mt. Kenzan, 

Almquist, s.n. (H, Kyl. herb. no. 34879, lectotype).] 

Shimoda, Prov. Idzu, Japan, Asuhino s.n. (TNS, lectotype).] 

Prov. Awa, Shikoku, Japan, Fujiknwu, s.n. (TNS, lectotype).] 

DEscmroN.-Thallus adnate on bark, quite firm, whitish 

to greenish mineral gray, 8-12 cm broad; lobes subirregular 

to sublinear, rather short, contiguous, 2-8 mm wide: upper 

surface shiny, plane to weakly foveolate, continuous, blackrimmed, coarsely lobulate with age, pseudocyphellae effigurate, raised, small, 0.2-0.5 mm long, numerous and conspicuous, marginal and laminal, in part fusing into a weak reticulate 

network; lower surface black, densely rhizinate, the rhizines 

simple to furcate, 1-2 mm long. Pycnidia common; conidia 

(Figure 12f) cylindrical, straight to slightly bent, 6-8 pm long. 

Apothecia common, substipitate, the rim remaining inrolled 

but the disc splitting with age, 5-12 mm in diameter, the 

amphithecium coarsely rugose with pseudocyphellae on the 

ridges; hymenium 55-60 pm; spores 8-9 x 12-14 pm, the 

episporium 1 pm thick. 

C€lEMIsTRY.-Atranorin, chloroatranorin, and salazinic acid 

(Asahina, 195 lb), and consalazinic acid. 

REmxs.-The most important feature of this Asian species 
hTJMBER 66 

FIGURE 17.Species of Parmelia: a, P. laevior (Hale 29357); b, P. marmariza (lectotype in H); c, P. meiophora 

(Poelt L-151); 4 P. modkcorduns (holotype in US); e, P. niitakana (Hsu 1374); f P. norcrombidiocarpa 

(holotype in US). (Scales in mm.) 

29 
30 SMITHSONIAN CONTRIBUTIONS TO BOTATTY 

is the distinct and numerous but comparatively small and 

crowded pseudocyphellae. This is in contrast to the larger, less 

numerous pseudocyphellae of P. adaugescens, another common 

Asian species, with mostly simple rhizines but much larger 

spores. 

Asahina described P. submarmariza to accommodate some 

specimens with smaller spores (8-1 1 pm long)-in my opinion 

not significantly smaller-and a less strongly rugose amphithecium. 

Specimens Examined 

Japan: Prov. Awa, Fujikawa s.n. (TNS); Prov. Higo, Kurokawa 63120 (TNS); 

Hiroshima Pref., M. Nakanishi 2811 (US); Prov. Hizen, Kurokawa 62597 

WS); Prov. Inaba, Nakanishi 12157 (KOBE); Prov. Iyo, Ogata, s.n. (TNS), 

Nakanishi 72 (KOBE). Yoshimura 1675 (US); Prov. Izu, Asahina 30825 

(TNS), Kwokawa 57038 (US); Prov. Kii. Kurokawa 71099 in Lichenes Critici 

Selecti Exriccati 234 (US); Prov. Kozuke, Kwokawa 55480 (US); Prov. 

Ohsumi, Kurokawa 63056 in Lichenes Critici Selecti Exsiccati 29 (TNS. US); 

Prov. Sagami, Kurokawa 58062 (US); Prov. SuPruga, Asaahaaina 538 (US); 

Yakushoima, Fujikawa, s.n. (TNS); Prov. Yamato, Nakanishi 50 (KOBE). 

Taiwan: Asahina, s.n. (US). India: Awasthi 644 (AWAS), Watt 5391 (BM). 

Nepal: Lmgalelah, Watf 7087 (BM). Zhao et al. (1982) report it from Anhui 

and Shensi provinces in China. 

Parmelia meiophora 

FIGURES 4b, 17c 

Parmelia meiophora Iiylander, 1889:45. [Type collection: Song-pin, Yunnan, 

China, Delavay s.n. (H, Nyl. Herb. no. 35201, lectotype; US, W, 

isolecto-types.] 

Parmelia meiophora var. isidkta Chao [Zhao], 1964:156. [Type collection: 

Yunnan, China, Chao 1002 (not seen).] 

DEscmIoN.-Thallus loosely adnate on bark, firm, greenish mineral gray (turning brownish in the herbarium), 8-15 

cm broad; lobes subirregular, contiguous to imbricate, 3-5 

mm wide; upper surface shiny, plane, continuous to finely 

cracked with age, pseudocyphellae numerous and small, 

0.2-0.3 mm long, separate, isidiate, the isidia bullate, short, 

about 0.1 mm thick and 0.1-0.2 mm high, unbranched, the tips 

often darkening; lower surface black, densely rhizinate, the 

rhizines richly squmosely branched, 1-2 mm long. Pycnidia 

not found, Apothecia rare, adnate, 3-5 mm in diameter; 

hymenium 55-60 pm: spores 5-6 x 7-8 pm, the episporium 

1 pm thick. 

CHEMISTRY.-AtranOrin and salazinic acid (Misra et al., 

1976) and consalazinic acid. 

REmRKs.-Parmelia meiophora is unquestionably the 

isidiate morphotype of P. submutata. Together they make up 

an anomalous element in the genus, because of the numerous 

small pseudocyphellae, too small to be seen with the naked 

eye. In his original description Nylander did not mention the 

presence of pseudocyphellae at all. 

Very little is known of the habitat and ecology of the species, 

except to note that it occurs at high elevation (3000-4000 m) 

in conifer-Rhododendron cloud forests. 

Specimens Examined 

China: Prov. Setschwan, Handel-Mazzetti 2856 (US. W). Nepal Gola, Togashi 

s.n. (US); Khumbu, Poelf 151 (M, US). Zhao et al. (1982) report specimens 

from Yunnan. 

Parmelia neodiscordans Hale, new species 

FIGURES 4c, 17d 

DEscmox-Thallus P. discordantis sirnilis sed colore 

albicanti et substantiis chimicis aliis differt. 

Thallus adnate to loosely attached on rock, whitish gray but 

sometimes brownish with age, 5-8 cm broad; lobes sublinear, 

crowded and imbricate, 1-2 mm wide; surface plane, 

marginally and in part laminally lobulate, the lobules 1-2 mm 

wide and 0.5-3.0 mm long, suberect; pseudocyphellae 

inconspicuous along the lobe margins as a whitish raised rim; 

lower surface jet black, sparsely to moderately rhizinate, the 

rhizines coarse, simple, up to 1 mm long. Pycnidia rather rare, 

about 90 pm in diameter; conidia cylindrical, straight to 

slightly bent, 6-7 pn long. Apothecia not seen. 

CmwsmY.-Atranorin and fumarprotocetraric acid, with 

accessory unidentified fatty acids. 

HoLo?-YPE.-Deer Mountain, Acadia National Park, Washington Co., Maine, USA, M.E. Hale 37496 (US). 

REmRKs.-The few specimens of this unusual species were 

first identified as P. discordans on the basis of a mistaken 

chemical test for protocetraric acid. Dr. Richard Harris brought 

to my attention the correct chemistry. Although similar to P. 

discordans, it has a whitish cast and never turns so dark brown. 

It differs from P. omphalodes, which is also whitish in the 

Appalachian region, in the smaller size and dense, uniform 

lobules, Parmelia neodiscordans is a strictly Appalachian 

species occurring at high elevations (more than 1500 m) in the 

southern part of its range but near sea level in New England. 

Specimens Examined 

USA: New York, Anderson s.n. (NY), Brodb 2901 (US), Harris 16579 (US); 

West Virginia: Hale 101 12, 14957 (US). 

Parmelia niitakana 

FIGURES 44 17e 

Parmelia niilakana Asahina, 1951a:332. [Type collection: Sokei, Mt. Niitaka, 

Parmelia shinanoana f. calvescens Zahlbruckner, 193456. [Type collection: 

Taiwan, Kwnada 23712 (TM, lectotype).] 

Mt. Morrison, Taiwan, Sasaki s.n. (W, lectotype; "S, isolectotype).] 

DEscmIoN.-ThalluS loosely adnate on bark, firm, pale 

greenish mineral gray, 6-10 cm broad: lobes sublinear, 

contiguous, 1.5-3 mm wide; upper surface shiny, plane, 

continuous, pseudocyphellae effigurate, marginal, forming a 

continous white rim 0.1-0.2 mm wide around the lobes; lower 

surface black, densely rhizinate, the rhizines simple to furcate, 

1-2 mm long. Pycnidia common; microconidia cylindrical to 

a few vaguely bifusiform, 5.5-6.5 pm long. Apothecia 
NUMBER 66 31 

common, substipitate, urceolate, 24 mm in diameter, the 

amphithecium rugose-sublobulate with pseudocyphellae on the 

ridges; hymenium 65-70 pm; spores 13-15 x 27-30 pm, the 

episporium distinct, 3 pm thick. 

CmmsmY.-Atranorin and salazinic acid (Asahina, 195 la) 

and consalazinic acid. 

REms.-This Taiwan endemic has lobes with a conspicuous white marginal rim formed by the continuous linear 

pseudocyphellae. It is also characterized by the large spores 

and the curiously sublobulate ridging on the amphithecium so 

well depicted by Asahina. It is most closely related to the 

Philippine species P. sectilis because of the very large spores. 

It occurs at high elevations in Taiwan, probably on most major 

mountains above 3000 m high. 

Specimens Examined 

Taiwan: Hattsukan, Suzuki s.n. WS), Chiayi County, Hsu 1374 (US), Mt. 

Nan-Fu-Ta-San, Kurokowa 121 1 (US). 

Parmelia norcrambidwearpa Hale, new species 

FIGURES 4e, 17J 18 

DESCRIPTION.-T~I~US ut in P. crambidiocarpa sed sporis 

parvis (6-8 x 11-13 um) differt. 

Thallus adnate to loosely attached on bark, fairly firm, 

whitish gray, 6-12 cm broad; lobes subirregular to sublinear, 

little branched, becoming divaricate, browning at the tips, 14 

mm wide; upper surface shiny, continuous to transversely 

cracked with age, plane, pseudocyphellae forming a very 

narrow, nearly continuous marginal rim, 0.1-0.2 mm wide, 

also laminal and effigurate, 0.2-0.6 mm long, separate, 

fissuring with age; lower surface densely rhizinate, the rhizines 

simple to strongly squarrosely branched, 0.5-2 mm long, 

usually projecting as a mat around the lobe margins. Pycnidia 

common, 90-110 pm in diameter; conidia cylindrical, rodshaped, 5.5-6.0 pm long. Apothecia common, substipitate and 

cupuliform, splitting radially with age, to 20 mm in diameter, 

the disc brown to dark brown, the amphithecium rugose, 

effigurate-pseudocyphellate; hymenium 60-70 pm high; spores 

7-10 x 10-15 pm, the episporium 1 pm thick. 

CmmsmY.-Atranorin, chloroatranorin, and echinocarpic 

acid (and associated unknowns). 

HoLoTYPE.-St. James Walkway, Tarn Nature Trail, Lewis 

Pass road, South Island, New Zealand, elev. 850 m, M.E. 

Hale 65352 (US; isotypes in ANUC, BM, CHR, UPS). 

REmws.-This new species is a member of the P. testacea 

complex. It is closely related to P. crambidiocarpa and P. 

salcrambidiocarpa, the thallus of these three species being 

essentially identical with sublinear lobes, marginal and laminal 

pseudocyphellae splitting open with age, and a dense mat of 

rhizines below. However, there are significant differences in 

chemistry, distribution, and, in part, spores. 

The apothecia of P. norcrambidiocarpa are typical for the 

group except that the disc is even darker than in P. 

crambidiocarpa. Of 34 specimens examined, none had a pale 

tan disc, 56% were light to medium brown, and 44% were dark 

brown. Spore differences are more significant: average 

maximum length is 13.4 pm (25 measurements) with a range 

of 12-15 pm, and average width 8.8 pm (range 7-10 pm). 

There is no overlap with the larger spores of P. crambidiocarpa 

and in fact a highly statistically significant value of P = .005 

was calculated for spore length differences between the two 

species. Parmelia salcrambidiocarpa has identical small 

spores. 

Chemistry is extremely uniform: atranorin and echinocarpic 

acid and associated unknowns. Some specimens of P. 

crambidiocarpa on North Island also contain only echinocarpic 

acid (and were identified by spore size); the only two from 

South Island with echinocarpic acid also contained protocetraric acid. Parmelia salcrambidiocarpa contains only 

salazinic acid. 

Parmelia norcrambidiocarpa is by far the most common 

foliose lichen in the subalpine scrub zone on the major 

mountain chains running the length of the South Island (Figure 

18). I collected 161 specimens at 32 localities, the bulk (85%) 

between 300 m and 910 m elevation. It comprised 23% of the 

715 specimens I collected at random in New Zealand and 30% 

of the 88 collections in CHR. It has yet to be collected on 

North Island or in Tasmania. 

Representative Specimens Examined 

New Zealand: South Island, Allan CHR160255 (CHR), Child 1556 (CHR), 

Elk 8675 (ANUC), Hale 65118,65325, 65391,65472.65659,65670,65719, 

65789,65841 (US). 

Parmelia novae-zelandiae Hale, new species 

FIGURES 4J19a 

DEscmoN.-Thallus ut in P. tenuirima sed isidiis munitis 

differt. 

Thallus loosely adnate on bark, rather fragile, pale greenish 

straw colored, 6-15 cm broad; lobes broad and rotund, 4-10 

mm wide; upper surface plane, the pseudocyphellae mostly 

laminal, uniformly dispersed, angular, 0.3-1.0 mm long, 

simple or branched, splitting open with the formation of coarse 

isidia, the isidia somewhat inflated, branching with age; lower 

surface black and shiny, the rhizines coarse, sparsely 

developed, 1-2 mm long. Pycnidia and apothecia not seen. 

CmmsmY.-Atranorin and salazinic acid. 

HOLOTYPE.-~ Mile stream trail, Rimutaka State Forest, 

E of Lower Hutt, North Island, New Zealand, M.E. Hale 58809 

REms.-This New Zealand endemic was identified as P. 

pseudotenuirima by Galloway and Elix (1983), although they 

subsequently recognized the variation in the New Zealand 

collections (Galloway and Elix, 1984). It is amply distinct 

because of the much larger thallus, large separate pseudocyphellae (as in P. tendrima), and coarse, irregularly inflated 

(US). 
32 SMITHSOhTAN COhTRIBUTIONS TO BOTANY 

FIGURE 18.-Distribution of Parmelia norcrambidiocarpa in New Zealand. 
NUMBER 66 

FIGURE 19.Species of Parmelia: a, P. novae-zelandiae (holotype in US); b, P. omphalodes (Santesson 23706); 

c, R omphalodes @innatif& form) (Vezda in Lichenes Selecti Ersiccati 1735); d P. protosulcata (holotype in 

US); e, R pseudolaevior (Kurokawa 59205); f; P. pseudoshinanoana (Nakanishi 5172). (Scales in mm.) 

33 
34 SMITHSONIAN CONTRIBUnONS TO BOTANY 

isidia. Parmelia pseudotenuirima can therefore be considered 

as a strict Australian endemic and P. novae-zelandiae a New 

Zealand one, although, of course, it may yet be found in 

Tasmania. In New Zealand it occurs rather rarely in moist 

Nothofagus forests or even on large trees in pastures at low 

elevation. 

Specimens Examined 

New Zealand: North Island, Hale 58809 (US); South Island, Hale 65443,66232 

(US). Three additional records are listed under P. tenuirima in Galloway and 

Elk (1983), of which I have checked Eartlett s.n. from Pandora. 

Parmelia omphalodes 

FIGLRES Sa, 196,~ 

Parmelia omphalodes (L.) Acharius, 1803:204. 

Lichen omphalodes L., 1753:1143. [Type collection: specimens and Dillenius, 

1742, pl. 24: fig. 80~ (OXF, lectotype).] 

Parmelia omphalodes p. panniforrnis Acharius, 1803:‘204. [Type collection: 

Helvetia, Schleicher 257 (H-Ach, lectotype).] 

Parmelia omphalodes var. panniformir f. subconcentrica Crombie, 1872:3M. 

[Type collection: Braemar. Marrone, Aberdeen, Great Britain, Crombie s.n. 

(BM, lectotype).] 

Parmelia sulcala var. laevis f. hirsufa Crombie, 1875:140. [Type collection: 

Ben Lawers, Perth, Great Britain, Crombie s.n. (BM, lectotype).] 

Parmelia omphalodes var. herminica Tavares, 1945:120. [Type collection: 

Serra da Estrela, Tavares s.n. (LISU) (not seen).] 

Parmelia pimnnatijkia Kurokawa, 1976:378. [Based on Parmelia omphalodes 

p. panniformis Acharius.] 

Parmelia omphalodes subsp. pinnutipdo (Kurokawa) Skult, 1984:138. 

[Additional varieties and forms, which I consider to be synonyms, will be 

found in Hillmann (1936) and Skult (1984).] 

DEscmox-Thallus adnate to loosely adnate on rocks, 

firm to somewhat brittle, extremely variable in color, whitish 

mineral gray to dark chestnut brown or even black, 6-20 cm 

broad; lobes sublinear, short, becoming crowded and imbricate, 

often becoming laciniate with dense marginal secondary lobes, 

14 mm wide; upper surface shiny, plane to weakly foveolate, 

continuous or becoming cracked along older pseudocyphellae, 

pseudocyphellae effigurate, variable, to 0.5 mm long, mostly 

marginal as a nearly continuous rim (especially in “I? 

pinnatifida”), becoming more numerous on the lobe surface 

with age and forming a distinct network; lower surface black 

and shiny, moderately rhizinate, the rhizines simple to furcate, 

1-1.5 mm long. Pycnidia common; conidia cylindrical, straight 

or a few bent or very weakly bifusiform, 5.5-6.5 pm long 

(Figure 12g) (Hillmann (1936) reports 1 x 5-6 pm). Apothecia 

not common, substipitate and urceolate but at maturity open 

the disc splitting somewhat, 3-12 mm in diameter, the 

amphithecium rugose; hymenium 60-80 pm; spores often 

poorly developed, 7-9 x 10-15 pm, the episporium 1-2 pm 

thick. 

CmmsmY.-Atranorin, lobaric acid (lacking in “P. pinnatifida”), salazinic acid (Culberson, 1970; Krog, 1968; Kurokawa, 

1976; Thomson, 1979), consalazinic acid, protolichesterinic 

acid (Skult, 1984), rarely also with traces of accessory galbinic 

acid (Culberson in Vezda, 1980), fumarprotocetraric acid, or 

protocetraric acid (Skult, 1984) and several unidentified fatty 

acids @ey, 1978; Kurokawa, 1976; Skult, 1984). 

REmRKs.-The Swedish specimen (Flora Suecica 947) 

cited by Linnaeus is not in the Linnean Herbarium. I have 

selected the Dillenian sheet at OXF as the lectotype (see 

Crombie, 1880). 

As might be expected for a widespread boreal-arctic lichen, 

P. omphalodes exhibits a great range of variation in thallus 

color (from whitish mineral gray to blackish brown), 

development of pseudocyphellae (from sparse and marginal 

to well developed as a lamina1 network) and secondary lobation 

(some specimens without secondary lobes, others heavily 

lobulate), in large part variations that are modified by the harsh, 

exposed habitats it frequently occupies. Poelt (1974) does not 

recognize any of the numerous varieties and forms based on 

habitat modifications described from Europe. 

Skult (1984) has made an especially thorough study of the 

species in Finland, recognizing three taxa: P. omphalodes 

subsp. discordans, P. omphalodes subsp. omphalodes, and P. 

omphalodes subsp. pinnatifida. Subspecies discordans was 

separated by the presence of protocetraric acid and subsp. 

pinnatifida by the absence of lobaric acid in salazinic 

acid-containing specimens. All specimens with lobaric and 

salazinic acids were called subsp. omphalodes. Relatively few 

intermediates were found in Fennoscandia. 

Parmelia pinnatifida, long recognized by European lichenologists as P. omphalodes var. panniformis, is also differentiated from typical P. omphalodes by a number of subtle, 

intergrading morphological characters. Kurokawa (1976) emphasized the predominance of marginal pseudocyphellae. It also 

has somewhat narrower lobes (Skult, 1984), which are 

repeatedly branched, and a congested, nearly pulvinate thallus. 

The populations of P. omphalodes in North America, 

however, although almost always lacking lobaric acid (96% 

of 84 specimens tested), are not like the European ones. Their 

morphology is much closer to typical European P. omphalodes, 

and after studying a small sample Skult (1984) considered them 

to be intermediate between his subsp. omphalodes and subsp. 

pinnatifida. I am inclined not to make any taxonomic divisions 

in the group at this time. 

An unnamed chemotype first recognized by Skult is 

distinguished by seven1 subtle morphological characters and 

by the presence of norstictic acid in equal concentration with 

salazinic acid. I am calling this a new species, P. zkulfii (see 

below). 

In the United States Parmelia omphalodes is locally 

common on rocks at the highest elevations in the Appalachian 

Mountains (Dey, 1978; Hale 1959) but is very rare in the 

western states. Imshaug (19573249) found it only on Mt. 

Rainier in Washington in his intensive study of alpine lichens, 

and I have collected it only in Montana. It is, however, a very 

common tundra lichen in the arctic-boreal regions of Canada, 

Alaska (Krog, 1968), and Europe (see map in Thomson, 

1984:308). 
NUMBER 66 35 

Representative Specimens Examined 

(salazinic acid with or without lobaric acid). 

Canada: Northwest Territories, Ahti 144%, Hale 236, Thomon and Larsen 

5859, 11066 (US); Quebec, GaNo 3055, Morton 11922 (US); Ontario, Brodo 

5974, Cain 26407, Garton 5451, Hale 36547. Wetmore 28431 (US); British 

Columbia, Macoun 243 (US). USA: Maine, Parlin 11699 (US); New York, 

Hale 23570, Shurhan 3479 (US); West Virginia, Hale 14293 (US); Virginia, 

Egan 3947, Hale 18851, Lultrell 3318 (US); Noh Carolina, Esslinger 3489 

in Lichenes Selecti Exsiccati 1093 (US), Hale 18047 (US); Tennessee, Hale 

37594; Michigan, Imhaug 4249 (MSC), Thomon 17191 (US); Montana, 

Hale 49973 (US), Imshaug 8249 (MSC); Alaska, Croasdale 89b, Llano 396g, 

511b, Nash 13119, Palmer 1342, Shurhan and Maher 10, Thomon 5463, 

Thomon and Ahfi 18428, Weber and Viereck 7060 (US). Greenland: Andersen 

and Hansen in Lichenes Groenlandici 291 (US). Iceland: Kristinsson 9114, 

16489 (US). Great Britain: Caemarvon, Holl s.n. (BM); Comwall, Culberson 

and Culberson 11738 (US); Devon, Davk s.n. (BM); Devonshire, Hebden 

1942 (US); Kincardine, Crombie s.n. (BM); Scotland, Vickery s.n. (US). 

Norway: Norman 5156 (US). Sweden: Kjellmert s.n., Santesson 23706 (US), 

Vrang in Cryptogamae Exsiccatae 4846 (US). Finland: HaMinen s.n., Rasanen 

in Lichenes Fenniae Exsiccati 706 (US). Denmark: Christiansen 1121 (US). 

Austria: Tirol, Steiner in Cryptogamae Exsiccatae 4731, Herre 32 (US). 

Switzerland: Schleicher 86 (US). Czechoslovakia: Moravia occ., Crnohorsky 

1245 (PRC), Vezda in Lichenes Selecfi Exsiccati 23 (US); Slovakia, Vezda in 

Lichenes Bohemoslovakiae Exsiccati 49 (US). Hungary: Magas Tatra, Forks 

3280 (US). Bulgaria: Szatala s.n. (US). France: Finistkre, Culberson 10483 in 

Vizda, Lichenes Selecti Exsiccati 1740 (US). Portugal: Beira Aha, Tavares 

4640 (US); Braga, Sampaio in Lichenes de Portugal 260 (US). Faroes: 

Torshavnd, Cernohorsy 1909 (PRC). USSR: Siberia, Elias, ef al. L-68228, 

Laurow 2301 (US). Mongolia: Schubert M600 (US). Nepal: Khumbu, Poelf 

130, 133 (M). 

Parmelia protosulcata 

Fxcms 56, 19d 

Parmelia protosulcata Hale, 1982: 162. [Type collection: Lago Fagnano, Tierra 

del Fugw. Argentina, Santesson 7955 (S, holotype; US, isotype).] 

DEscmoN.-Thallus adnate to loosely adnate on bark, 

pale greenish to brownish mineral gray, 2-6 cm broad; lobes 

sublinear, short, contiguous, 2-3 mm wide; upper surface plane 

to weakly foveolate, becoming white pruinose at the tips, 

deeply reticulately fissured in older parts, pseudocyphellae 

effigurate, small and rather sparsely developed, mostly on the 

lobe tips or margins, sorediate, the soralia developing on lobe 

tips and margins, usually orbicular but fusing and becoming 

linear or densely aggregated with age, the soredia coarse; lower 

surface black, moderately rhizinate, the rhizines simple to 

sparsely furcate or squarrosely branched, 0.5-1 mm long. 

Pycnidia not seen. Apothecia rare, substipitate, 4-5 mm in 

diameter, the amphithecium rugose, pseudocyphellate; hymenium 60-65 pm; spores 7-9 x 11-13 pm (spore data from 

Imshaug and Ohlsson 45613). 

CHEMISTRY.-A~~~~OM, chloroatranorin, protocetraric acid, 

accessory lobaric acid, echinocqic acid, and conechinocarpic 

acid (Galloway and Elk, 1984) or rarely fumarprotocetraric 

acid. 

REms.-This austral species occurs in South America 

and on some of the subantarctic islands, as well as in Australia 

and New Zealand (Galloway and Elix, 1984). It may be 

accompanied by P. sulcata, which would be distinguished by 

the different chemistry, a more foveolate surface, lamina1 

soralia, large fissurine pseudocyphellae, and densely squarrosely 

branched rhizines. It is most closely related to another austral 

species, P. cunninghamii, which has broader lobes, linear 

soralia, and salazinic acid. 

Specimens Examined 

New Zealand: Campbell Island, Harris 5455 (MSC, NY). Falkland Islands: 

Howkins 2964 (FH). Chile: Prov. Magallanes. lmshaug and Ohlsson 43842, 

43896. 44401, 44568, 45155, 45613 (MSC), 45613 (MSC, US), Sanfesson 

1843 (S, US); Prov. Osomo, Imhaug 42961 (MSC); Brunswick Peninsula, 

Imhaug and Harris 39095, 39260,39328,39329,39404 (MSC). 

Parmelia pseudolaevior 

FIGURES 5c, 19e 

Parmelia pseudolaevior Asahina, 1951a:331. [Type collection: Sengenjunja. 

Prov. Suruga, Japan, Asahina s.n. (TNS, lectotype; US, isolectotype).] 

DEscmoN.-Thallus adnate on bark or rock, rather brittle, 

pale greenish to brownish mineral gray, 8-12 cm broad; lobes 

sublinear, contiguous to imbricate, 1-3 mm wide, the margins 

becoming densely lobulate, the lobules suberect to erect, 

0.1-0.3 mm wide, 1-2 mm long, the lower surface whitish; 

upper surface shiny, plane to weakly rugose-foveolate, 

continuous, pseudocyphellae punctate, round, lateral on lobe 

edges, rarely submarginal, 0.14.2 mm long; lower surface 

black, shiny moderately rhizinate, the rhizines simple to 

sparsely furcate, 0.5-1 mm long. Pycnidia not seen. Apothecia 

rare, substipitate, 2-10 mm in diameter, at first urceolate, then 

expanded, flat and radially split, the amphithecium sparsely 

punctate-pseudocyphellate; hymenium 60-65 pm; spores 6-9 

x 10-15 pm, the episporium 1 pm thick. 

CmmmY.-Atranorin, salazinic acid (Asahina, 1951a), 

and consalazinic acid. 

REui\RKs.-Parme/ia pseudolaevior is clearly the lobulate 

morphotype of P. laevior, both having lateral punctate 

pseudocyphellae. It is rather common in Japan and apparently restricted there. A synonym cited by Asahina, P. laevior 

f. microphyllina Hue, is actually better identified with P. 

pseudoshinanoana. 

Specimens Examined 

Japan: Prov. Aki, Hale 29363, 29377, 29505 (US); Prov. Bungo, Kurokawa 

63193 (TNS); Prov. Izu, Asahina, s.n. mS); Prov. Kii, Kwokawa 56061, 

57281 (US), 60262 (TNS); Prov. Ishikari, Yoshimura 12235b in Lichenes 

Japonici Exriccati 41 (US); Prov. Rikutyu. Kurokawa 59277 (US); Prov. 

Shimotsuke, Hale 63127 (US); Prov. Shinano, Kwokawa 59205 in Lichenes 

Critici Selecti Exriccafi 34 (US); Prov. Suruga, Asahina 92 (TNS). 

Parmelia pseudoshinanoana 

FIGURES 54 19f 

Parmelia pseudoshinanoana Asahina, 1951a:334. [Type collection: Omiyaguchi, 2-gome, Mt. Fuji, Prov. Suruga, Japan, Asahina 52 (TNS, lectotype; 

US, isolectotype).] 
36 SMITHSONIAN CONTRIBUTIONS TO BOTAh’Y 

Parmelia hevior f. t?u.crophyllina Hue, 1899:166. [Type collection: Japan, 

Fawie 518 (PC, lectotype).] 

DEscmoN.-Thallus adnate on bark, fragile, pale greenish 

to brownish mineral gray, 6-12 cm broad; lobes sublinear, 

contiguous, 1.5-3 mm wide, the margins becoming densely 

lobulate, the lobules 0.24.3 mm wide, 1-2 mm long, suberect; 

upper surface plane, shiny, white pruinose at the tips, 

continuous, pseudocyphellae marginal, appearing as a more 

or less continuous white rim 0.5-1 mm wide; lower surface 

black, densely rhizinate, the rhizines shiny, becoming densely 

squarrosely branched, 1-3 mm long. Pycnidia not seen. 

Apothecia not common, barely substipitate, 2-4 mm in 

diameter, the rim finely crenate, pseudocyphellate; hymenium 

55-60 p; spores 6 x 10-12 p, the episporium 1 pm thick. 

CmmmY.-Atranorin and salazinic acid (Asahina, 1951a). 

REMARKs.-Asahina considered this Japanese species to be 

a lobulate relative of I? shinanoana. Although they share 

continuous rim-forming pseudocyphellae, P. shinanoana has 

mostly simple rhizines and produces gyrophoric acid in the 

cortex. Parmelia pseudoshinanoana is widespread in Japan 

but not especially common. 

Specimens Examined 

Japan: Rov. Iyo. Kurokawa 60048 WS); Rov. Kii, Koizwni (TNS); Prov. 

Musashi, Kurokawa 550574,59153,70156 in Lichenes Critici Selecti Exriccati 

129 (Us); Saitama Pref., Omura 332 (Us); Prov. Shinano, Asahina 33 (TNS), 

Kwokawa 520639, 58554, 59206 in Lichenes Critici Selecti Exriccati 35, 

59207 (US); Prov. Shimotsuke. Kurokawa 64059 (Th’S); Shizuoka Pref., M. 

Nakanishi 5172 (US); Prov. Suruga, Fawie 512 WS). 

Parmelia pseudotenuirima 

FIGURES 5e, 20a 

Parmelia pseudotenuirima Gyelnik, 1931:289. [Type collection: based on P. 

fenuirima f. isidiosa Miiller Argoviensis.] 

Parmelia tenuirimr f. isidiosa Muller Argoviensis, 1896:90. [Type collection: 

Victoria, Australia, Wilson 83 (G, lectotype). Collector on the label is 

Knight1 

DESC~~PI?ON.-T~~~~US closely adnate on bark, firm, pale 

greenish to whitish mineral gray (brownish in the herbarium), 

3-5 cm broad; lobes sublinear to subirregular, short, contiguous, 1-2 mm wide; upper surface shiny, finely foveolate, 

reticulately cracked with age, pseudocyphellae small, 0.24.3 

mm long, effigurate in outline, numerous, mostly laminal, 

isidiate, the isidia cylindrical, 0.05-0.07 x 0.14.3 mm, simple 

to branched, very dense on older parts of the thallus and 

obscuring the lobes; lower surface black, moderately rhizinate, 

the rhizines simple to squarrosely branched, 0.5-1 mm long. 

Pycnidia rare, about 100 pm in diameter; conidia cylindrical, 

rod-shaped, 5.5-6.0 p long. Apothecia very rare, substipitate, 

3-6 mm in diameter, the amphithecium warty, isidiate, the disc 

brown; hymenium 45-55 pm; spores poorly developed, 

7-9 x 11-13 pm, the episporium 1 pm thick. 

Cmmsmy.-Atranorin and chloroatranorin, lobaric acid 

and salazinic acid (Galloway and Elix, 1983). 

REmms.-This small lichen is known only from Australia 

(see distribution map in Galloway and Elix, 1983:403). It is 

not related to any other species in the genus. Literature reports 

of P. savatilis from Australia probably represent this species. 

New Zealand records cited by Galloway and Elix (1983) are 

all P. novae-zelandiae. A typical habitat is the base and lower 

trunks of Callitris endlicheri and Casuarina in open eucalypt 

forest and especially on old burned stumps. 

Specimens Examined 

Australia: New South Wales, Hale 58276, Weber 365 in Lichenes Ersiccati 

365 (US), Weber and McVean L-49416. L-50060 (US); Victoria, James 

Au2139, Streimann HS3223 (US). Filson (1982555) gives additional records 

for Australian Capital Territory and South Australia. 

Parmelia queenslandensis Hale, new species 

FIGURES Q. 20b 

DEsmmIoN.-Thallus similis P. signifera sed thallo 

persistenti albo-cinereo, corticola, et sporis minoribus differt. 

Thallus corticolous, adnate to closely adnate, leathery, 

whitish to pale greenish mineral gray, 5-8 cm broad; lobes 

broadly sublinear, rather short and becoming imbricate, 2-5 

mm wide; upper surface shiny, smooth, the pseudocyphellae 

effigurate, dense and small in a marginal zone but larger and 

more or less separate toward the center, 0.2-1.0 mm long, 

becoming fissured with age; lower surface sparsely to 

moderately rhizinate, the rhizines rather coarse, simple to 

weakly squarrosely branched with age. Pycnidia common, 

90-110 pm in diameter; conidia weakly bifusiform, 6-7 pm. 

Apothecia common, substipitate, radially split with the rim 

folded upward with age, 5-10 mm in diameter, the disc light 

brown; hymenium 50-60 pm; spores 6-9 x 11-12 pm, the 

episporium 1 pm thick. 

CmmsmY.-Atranorin, chloroatranorin, and salazinic acid. 

HoLoTYPE.-Near the north exit, Mt. Bunya National Park, 

Queensland, Australia, elev. -600 m, M.E. Hale 65970 (US; 

isotypes in BM, MEL). 

REMARKS.-I first identified this lichen as P. signifera in 

spite of the unusual substrate, canopy branches in rain forest. 

Parmelia queenslandensis is strictly corticolous, persistently 

whitish or greenish gray and occurs at elevations of 400 to 

1100 m from extreme northern New South Wales to the 

Atherton Tablelands in northern Queensland, well north of the 

range of P. signifera. Parmelia signifera is almost always 

saxicolous in rather exposed habitats and usually turns pale to 

dark brown. In Australia it occurs from northern New South 

Wales to Tasmania with no reports yet from Queensland 

(Galloway and Elix, 1983). Spore differences, which I have 

not tested statistically, are minor but consistent: 11-12 pm 
NUMBER 66 

FIGURE 20.4pecies of Parmelia: a, P. pseudotenuirima (Hale 58276); b, P. queemlandenris (holotype in US); 

C, P. ricasolioides lectotype in H); 4 P. salcrambidiocarp (holotype in US); e, P. samtilis (Arnold in Lichenes 

Universalis 9); f; P. sectilk (Hale 26536). (Scales in mm.) 

37 
38 SMITHSONIAN CONTRIBUTIONS TO BOTATTY 

long in P. queenslandensis and 13-15 pm in P. signifera. 

Another related corticolous species, P. tenuirima, also has 

larger spores (12-15 pn long), brown rotund lobes (4-10 

mm wide), and rather small, separate pseudocyphellae. It is 

common in New Zealand, much less so in Australia and 

represented by only one record as far north as Queensland. 

Specimens Examined 

Australia: Queensland, Hale 59480,64001, 64128,65971, 65972 (US); New 

South Wales, Hale 66686 (US). 

Parmelia ricasolioides 

FIGURES 6a 2Oc 

Parmelia ricasolioides Nylander, 1887:135. [Type collection: Yunnan, China, 

S.C. (H, Nyl. Herb. no. 35283, lectotype; PC, TUR, isolectotypes).] 

farmelk daliemis Zahlbruckner, 1930:183. [Type collection: between Dali 

and Hodjing, Yunnan, China, Handel-MarzetZti S658t,2 (W, lectotype; 

TNS, US, isolectotypes).] 

Parmelia doliensir f. tardiva Zahlbruckner, 1930:184. [Type collection: Muli, 

Yunnan, China, Handel-Mazzetti 7368 (W, lectotype; WU, isolectotype).] 

DESCI~P~ON.-T~~I~LIS adnate to loosely adnate on bark, 

brittle, brownish mineral gray in the herbarium, 6-8 cm broad; 

lobes subirregular, short and crowded, contiguous, 2-3 mm 

wide; upper surface shiny, plane to weakly rugose, continuous, 

pseudocyphellae inconspicuous, entirely marginal, elongate; 

lower surface black, densely rhizinate, the rhizines simple to 

furcate, 0.5-1 mm long. Pycnidia common; conidia cylindrical, 

straight 5-6 pm long (Nylander, 1887). Apothecia numerous, 

substipitate, 1-2 mm in diameter, the rim inrolled, crenate; 

hymenium 55 pm; spores 9-13 x 20-24 pm, the episporium 

about 2 pm thick. 

CmmsmY.-Atranorin and salazinic acid. 

REms.-This species occupies an isolated position in the 

genus. Nylander likened it to both “P. perlata” and P. 

tenuirima. The pseudocyphellae are very weakly developed 

along the lobe margins and the surface is continuous. It occurs 

at high elevations (to 3700 m), probably in Rhododendron 

forests. 

Specimens Examined 

Nepal: Okhaldunga, Poeft L715 (US). India: NEFA, Panigrahi 16079A 

(AWAS). Zhao et al. (1982) reported it from Yunnan Province in China. 

Parmelia salcrambidiocarpa Hale, new species 

FIGURES 6b, 204 21 

DEsmmIoN.-Similis P. crambidiocarpae sed sporis 

parvis (7-10 x 12-15 pm) et thallo acidum salazinicum 

continenti differt. 

Thallus corticolous, adnate to loosely attached, nearly 

pulvinate with age, firm, whitish mineral gray, 4-12 cm broad; 

lobes sublinear, little branched, becoming divaricately branched, 

1.W mm wide; upper surface shiny, plane, smooth to rugose, 

continuous or transversely cracked with age, the pseudocyphellae forming a nearly continuous narrow rim around the 

margins, 0.1-0.2 mm wide, also lamina1 and effigurate, 0.2-1 

mm long, separate, fissuring with age; lower surface moderately to densely rhizinate, the rhizines simple to moderately 

squarrosely branched, 1-3 mm long, usually projecting as a 

mat around the lobe margins. Pycnidia numerous, 90-1 10 pm 

in diameter; conidia cylindrical to weakly bifusiform, 5-6 pm 

long. Apothecia common, stipitate, 5-18 mm in diameter, the 

disc sometimes flat and radially splitting, light to dark brown; 

hymenium 55-65 pm; spores 7-10 x 12-15 pm, the 

episporium 1.0-1.5 pm thick. 

CmmsmY.-Atranorin, chloroatranorin, and salazinic acid 

with or without consalazinic acid (one collection, Kanfvilas 

and James 633181, also contains lobaric acid). 

HoLoruPE.-Whakapapanui Walk Trail Head near the 

Chateau, Tongariro National Park, North Island, New Zealand, elev. 1180 m, M.E. Hale 65558 (US). 

REmxs.-This species is in the P. testacea complex, 

closely related to both P. crambidiocarpa and P. norcrambidiocarpa (see discussions under these species). It is easily 

distinguished from P. crambidiocarpa by the smaller spores 

and chemistry. The average maximum spore size, 8.9 x 13.4 

pm (range 7-10 x 12-15 pn for 17 collections), is the same 

as in P. norcrambidiocarpa. As in P. crambidiocarpa the 

apothecial disc ranges from very pale brown (29%) to brown 

(23%) or dark brown (47%) (17 collections examined). 

Parmelia salcrambidiocarpa differs from morphologically 

identical P. norcrambidiocarpa in containing salazinic acid 

rather than echinocarpic acid. In addition the two species are 

allopatric, P. salcrambidiocarpa occurring in Tasmania and 

the Tongariro-Urewera region of North Island of New Zealand 

(Figure 21). Parmelia norcrambidiocarpa occurs only on 

South Island of New Zealand. I collected P. salcrambidiocarpa 

at 8 localities in Tasmania, a total of 28 specimens out of 186, 

the remainder being P. testacea. 

Ecologically P. salcrambidiocarpa behaves much as the 

other species in this group. It is usually collected on 

Nothofagus cunninghamii in Tasmania at 600-1200 m 

elevation and in New Zealand on Dracophyllum and other 

subalpine shrubs up to 1300 m elevation. 

Specimens Examined 

Australia: Tasmania, Hale 68654, 68656, 68695-698, 68821 -828, 68690, 

68768-772, 68785, 68786 (US). Kanfvilas 192/82 (AhLJC). Kantvilas and 

James 633/81 (ANUC). New Zealand: North Island, Hale 65114-1 16,65372, 

65551,65556,65557,65567-569 (US). 

Parmelia saxatilis 

FIGLXES 6c, 20e 

Parmelia saxatilis (L.) Acharius, 1803:204. 

Lichen saxatifis L., 1753:1142. [Type collection: Sweden, sheet 1273.61 

(LINN, lectotype).] 
?\STAMBER 66 39 

i BOTANY DIVISION 0 XI im MILES 

).'..'.: ' ' , 

0 XI iM KILOMETRES 

FIGCRE 21 .-Distribution of Partnelb saicrambidiocarpa in Sew Zealand. 
40 SMITHSOiWV COh'UTIONS TO BOTANY 

Parmelia saxatilk var. laevis Nylander, 1860:389. [Type collectim: Pyrenees 

Parmelia saxatilis f. rubricosa Steiner, 1904:399. [Type collection: Cumbre 

Parmelia acervata Hue, 1915:43. [Type collection: Cape Tuxen, Graham Land 

Parmelia saxatilis f. squumigera Gyelnik, 193 la:152. [Type collection: 

Parmelia saxatilis f. corailicola Gyelnik, 1931a:152. [Type collection Domos. 

Parmelia saxatilk f. prk Gyelnik, 1931b3284. [Type collection: Magas Tatra, 

Parmelia saxatilk var. imbricatoi&s Gyelnik, 1932450. [Type collection: 

Parmelia saxatilk var. pseudoviridir Gyelnik, 1932:448. [Type collection: St. 

Parmelia saxatitis f. acervata (Hue) Lamb, 1948:241. 

[See HiUmann (1936) for additional subspecific taxa, all of which appear to 

cent, Nylander s.n. (H, Nyl. herb. no. 34863, lectotype).] 

Nueva, La Palma. Canary Islands, Bornmuller 3256 (WLJ, lectotype).] 

S.C. 190 (PC, lectotype; see Lamb, 1948:241).] 

Helsinki, Finland, Gyelnik s.n. (BP, lectotype).] 

Mt. Vadallokovek, Pest, Hungary, Gyelnik s.n. (BP, lectotype).] 

Kqmva Vdey, Hungary,Timko s.n. (BP, lectotype).] 

Artukainen, Finland, Linkula s.n. (BP, lectotype).] 

Gilgen, Austria, Gyelnik s.n. (BP, lectotype).] 

be synmyms of P. sutilir.] 

DEscmoN.-Thallus adnate to loosely adnate on rock, 

more rarely on bark, fairly firm, greenish mineral gray, turning 

brownish in exposed habitats, sometimes whitish with varying 

amounts of pruina, 8-20 cm broad; lobes sublinear, contiguous 

to crowded, 24 mm wide; upper surface shiny, continuous or 

cracking with age along the pseudocyphellae, reticulatefoveolate, the pseudocyphellae effigurate, large, 0.3-1 .O mm 

long, marginal and laminal, often fusing into a recitulate 

network and soon becoming isidiate, the isidia cylindrical, 

simple to sparsely branched, darkening at the apices, 0.06-0.15 

mm in diameter, up to 0.5 mm high, becoming very dense on 

older lobes; lower surface black, densely rhizinate, the rhizines 

simple to furcate or rarely subsquanosely branched, 1-2 mm 

long. Pycnidia uncommon, about 100 pm in diameter; conidia 

cylindrical, straight to slightly bent, 6-7 pm long (Hillmann 

(1936) reported 1 x 5-7 pm). Apothecia occasional, substipitate, 2-8 mm in diameter, cupuliform, the amphithecium and 

rim coarsely rugulose, isidiate; hymenium 80-100 p; spores 

9-1 1 x 16-18 p, the episporium 2-3 p thick. 

CHEMISTRY .-Atranorin, chloroatranorin, and salazinic acid 

(Galloway and Elix, 1983), accessory lobaric acid (Krog, 1951) 

with or without protolichesterinic acid (Kurokawa and 

Nakanishi, 1971) or unidentified fatty acids (Dey, 1978). 

REms.-This is the most widespread species in the 

genus, occurring in both hemispheres and known to all 

lichenologists. For such a common species it has very little 

morphological and chemical variation. The presence or absence 

of the accessory substance lobaric acid is loosely correlated 

with geography. In Europe about a third of the collections (50 

tested) have only salazinic acid, the rest having salazinic and 

lobaric acids. In Alaska and Canada (see map in Thomson, 

1984:310) these two chemotypes are nearly equally common, 

but in western United States and South America most 

specimens contain only salazinic acid (91% of 75 tested). All 

specimens in the Appalachian region of eastern North America, 

where the species occurs on rocks at the highest elevations, 

seem to have lobaric acid constantly (Dey, 1978). Parmelia 

kerguelensis appears to be a protocetraric acid-containing 

chemotype. 

Most corticolous specimens from Japan, eastern North 

America, and the west coast of North America, usually 

identified as P. saxatilis in herbaria, have proved to be P. 

squarrosa, a closely related species with densely squarrosely 

branched rhizines. This probably includes the reports of Zhao 

et al. (1982) for P. savatilis in China. 

Representative Specimens Examined 

Canada: Northwest Territories, Inskip 11 (BM, US); Newfoundland, Fernald 

2210 (US); Quebec, LeGallo 2943, Samuel 62 (US); Ontario, Hale 36240. 

Imhaug 4132, Wefmore 29481 (US); British Columbia, Brodo 10982, Noble 

6336, Norris 2325D (US). USA: Maine, Egan 61 19, Hale 37477, Merrill in 

Lichenes Exsiccati 152, Pike 4341 (US); New Hampshire, Rathburn s.n. (US); 

New York, Harris s.n., Hermann 14693 (US); West Virginia, Hale 15463 

(US); North Carolina, Hale 33628 (US); Michigan, Egan 3522 (US); Colorado, 

Anderson 2843, Kiener 4523, Shushan 5033, Weber and Klockenbrink in 

LichenesExsiccati 538 (US); Montana, Hale 49319 (US); Idaho, Hale 48533 

(USA); Washington, Imhaug 1184, Thomon 14717 (US); Oregon, Brown 

857, Hale 49288, Pike 379 (US); California, Hale 51588. 56220, Herre 702 

(US); Alaska, R. Hale 159, Hermann 21857, Llano 416c, Schaack 250, 

Thomon and Shushan 6208, Viereck 2292 (US). Chile: Aysen, Santesson 

4820 (S, US); Cautin, Redon 3576 (US); Magallanes, Santesson 2158 (S, US); 

Malleco, Mahu 2466 (US); Tierra del Fuego, Sanfesson 1246 (S, US); Juan 

Femandez, Imhaug 37035,37410 (MSC). Argentina: Tierra del Fuego, Dusen 

262 (US), Sanfesson 346, 7532, 8224 (S). Falkland Islands: Hawkins 2962 

(FH), Lechler 76 (BM). Greenland: Alsfrup in Lichenes Groenlandici Exsiccafi 

195. Hansen in Lichenes Groenlandici Exsiccafi 19, Lynge s.n. (US). Iceland: 

Reykjavik, dernohorskj 1924 (PRC), Kristinsson 18087, 22174 (US); 

Reydarvatn, Smarda 1610 (PRC). Great Britain: Comwall, Curnow s.n. (BM); 

Merioneth, James 2077 (BM). Ireland: W. Galway, Larbalestier 21 1 (BM). 

Norway: Lynge 5165 (US). Sweden: Magnusson 8562 (US). Finland: Lai 

10911, Rdsanen in Lichenes Fennicae hiccati 856 (US). Denmark: 

Christianren 9857 (US). Netherlands: Maas Geesteranus 7005 (US). Germany: 

Erichen 667, Hillmann in Kryptogamae Ersiccatae 2862 (US). Austria: Herre 

32 (US). Czechoslovakia: Cernohorsyy 589 (PRC), Matousek in Lichenes 

Slovakiae Ersiccati 72 (US). Hungary: Foriss 1885, Timko in Flora Hungarica 

Emiccata 319 (US). France: Beraud s.n., Ilfis 3122 (US). Spain: Vuzquez s.n. 

(US). Pottugal: Sampaio in Lichenes de Portugal 259 (US). Teneriffe: Imhaug 

33860 (MSC). New Zealand: North Island, Galloway s.n. (CHR); South Island. 

Hale 63216 (US), James 564 (BM), Martin A-167 (CHR). There are also 

repotts by Krog (in litt.) from Kenya, by Golubkova (1981) from Mongolia, 

by Kurokawa (1966) from Nepal, and by Lindsay (1973b) from Adelaide Island 

in Antarctica. 

Parmelia sectilis 

FIGURES 6d 2of 

Parmelia sectilis Hale, 1968:326. [Type collection: summit of Mt. Data, 

Mountain Prov., Philippines, Hale 26536 (US, holotype; TNS, isotype).] 

DEscmoN.-Thallus adnate to closely adnate on bark, 

brittle, light greenish mineral gray, 8-12 cm broad; lobes 

sublinear, contiguous, 1.5-2 mm wide, marginally dissected 

and densely isidiate-lobulate, the lobules erect, branched, 

0.1-0.2 mm wide, to 1 mm long, the tips cylindrical; upper 

surface shiny, plane to weakly rugose, continuous, pseudocyphellae effigurate, elongate. mostly marginal, forming an 
NUMBER 66 41 

interrupted white rim around the lobes; lower surface black, 

densely rhizinate, the rhizines simple to furcate, 1-1.5 mm 

long. Pycnidia rare; conidia cylindrical, straight, 5-6 pm long. 

Apothecia not common, substipitate, 24 mm in diameter, the 

rim and amphithecium deeply rugose with numerous lobules 

or isidia-like projections, the disc brown; hymenium 90-100 

pm high; spores 15-18 x 28-33 pm, the episporium distinct, 

34 pm. 

CHEmsmY.-Atranorin, salazinic acid, and consalazinic 

acid. 

REMRKs.-Parmelia sectilis occurs at higher elevations 

(1600-2300 m) in the pine forests of the Philippines and in 

oak forests in Sabah. It is the only species of Parmelia that 

seems to have evolved and remained in the tropics. It is 

distantly related to P. niitakana, P. pseudoshinanoana, and P. 

shinanoana, which have similar though more strongly 

developed rim-forming pseudocyphellae. It has the largest 

spores in the genus. 

Specimens Examined 

Philippines: Mountain Prov.. Hale 26104, 26194, 26219, 26397, 26536 (US). 

Sabah: Kinabalu National Park, Hale 28378, 28241, 28507, 28630, 28786, 

28935,29050,29090 (US). 

Parmelia shinanoana 

FIGURES 6e. 22a 

Parrnelia shinanoana Zahlbruckner, 1927b:349. [Type collection: Mt. 

Yatsu-gadake, Shinano, Japan, Asahina 548 (w, lectotype; US, isolectotype).] 

DEscmox-Thallus adnate on rock, fragile, pale greenish 

to distinctly brownish mineral gray, 8-20 cm broad; lobes 

sublinear, contiguous, 1.5-3 mm wide, sparsely lobulate in 

older parts; upper surface shiny, plane, continuous, pseudocyphellae marginal, linear, forming a conspicuous white rim 

about 2 mm wide around the lobes; lower surface black, 

sparsely to moderately rhizinate, the rhizines simple to furcate, 

1-2 mm long. Pycnidia not seen. Apothecia rare, adnate and 

inconspicuous, 1-2 mm in diameter, the rim crenate, pseudocyphellae; hymenium 60-65 pm; spores poorly developed, 5-6 

x 9-11 pm, the episporium 1 pm thick (after Asahina, 

1952:114). 

CmMIsmY.-Atranorin, gyrophoric acid, and 4-0- 

methylgyrophoric acid in the cortex, salazinic acid in the 

medulla (Kurokawa and Takahashi, 1970). 

REmms.-Parmelia shinanoana has very conspicuous 

rim-forming pseudocyphellae, similar to P. niitakana and P. 

pseudoshinanoana and, as the main chemical feature, the 

production of gyrophoric acid in the cortex, unique in the entire 

family Parmeliaceae. It is known only from Japan, where it is 

not commonly collected. 

Specimens Examined 

Japan: Prov. Awa, Fujikawa s.n. (TNS); Prov. Buzen, Kurokawa 62477 ("3); 

Prov. Iyo, Kwokawa 60047 (TNS), Ochi s.n. (TNS): Prov. Kozuke, Kurokawa 

58588 (TNS); Prov. Musashi, Kurokawa 56140 (US), 64289 (TANS); Saghalin, 

Asahina s.n. (TNS); Prov. Shinano, Kwokawa 58500 (US); Prov. Yamato, S. 

Nakanishi 41 (KOBE). 

Parmelia signifera 

FIGURES 61: 22b 

Parmelia sign;fera Nylander, 1888:25. [Type collection: Sew Zealand, Knight 

Parmelia saxatilis var. sign;fera (Nylander) Muller Argoviensis, 1892:30. 

s.n. (H, Nyl. Herb. no. 34828, lectotype; BM, isolectotype).] 

DEscmIoN.-Thallus adnate to loosely adnate on rock 

(rarely on bark), sometimes forming dense, pulvinate mats, 

rather firm to brittle, darkish green to brownish mineral gray, 

the tips usually distinctly brownish, 8-20 cm or more broad; 

lobes sublinear, crowded and imbricate, 2-8 mm wide, more 

or less lobulate with age in the center, the lobules marginal, 

suberect; upper surface shiny, plane to rugose or foveolate, 

extensively fissured with age along pseudocyphellae, the 

pseudocyphellae effigurate, 0.3-1 mm long, separate or in part 

fusing into a loose white network, appearing raised; lower 

surface black, the margin shiny and dark brown, densely 

rhizinate, the rhizines rather dense, about 1 mm long, simple 

to sparsely squarrosely branched. Pycnidia common; conidia 

bifusiform, 5.0-56.5 pm long. Apothecia common, adnate to 

substipitate, 6-12 mm in diameter, the disc brown, flattened, 

often radially split, the amphithecium heavily effiguratepseudocyphellate; hymenium 55-60 pm; spores 6-8 x 13-15 

pm, the episporium distinct, about 1 pm thick. 

CHEMISTRY.-Atranorin and chloroatranorin, salazinic acid 

(Galloway and Elix, 1983) and consalazinic acid, and accessory 

lobaric acid; or rarely atranorin and protocetraric acid. 

REbfAms.-This is by far the most commonly collected 

saxicolous Parmelia in Australia and New Zealand (see 

distribution map in Galloway and Elix, 1983:408). In Australia, 

in particular, it forms large, pulvinate, exfoliating colonies on 

granite and sandstone outcrops in open sclerophyll forests. In 

New Zealand the typical habitat is more exposed ridges in 

open mountainous areas. In a sense it is the austral counterpart 

of P. omphalodes, which has much less conspicuous pseudocyphellae and is, of course, a northern arctic-alpine lichen. A 

few specimens (Hale 58374, Melville 2009) containing 

protocetraric acid instead of salazinic acid are morphologically 

indistinguishable. 

When corticolous, there is rarely confusion with I? 

tenuirima, a large greenish mineral gray lichen that has less 

dense, separate pseudocyphellae and cylindrical to vaguely 

bifusiform conidia. However, a few narrow-lobed, usually 

corticolous specimens may be difficult to name as P. signifera 

or P. lenuirima (see also Hillmann, 1939). Filson (1982:565) 

feels that the two species may prove to be identical, but both 

morphological and conidial differences seem to preclude this 

unlikely possibility. 
42 SMITHSONIAN CONTRIBUTIONS TO BOTAh'Y 

FIGURE 22.-Species of Parmelia: a# P. shinanoana (Kurokawa 58588); b, P. signifera (Hale 58289); c, P. 

shltii (holotype in US); 4 I? squnrrosa (holotype in US); e, I? submantanu (lectotype of P. contorta Bory in 

F'C); f; I? submufata (Poelt L-142). (Scales in mm.) 
NUMBER 66 43 

Specimens Examined 

Australia: New South Wales, Du Riefz 704, 792a (UPS, US). Flockfon 1443 

(US), Hale 58289,58301,58927,59106,59286,59296,59443,66122,66639 

(US), Streimann 2496, 4017, 7414, 7473, 9138, 9286 (US), Verdon 1364, 

1428, 1513, 1587, 1832, 7415 (US); Australian Capital Territory, Hoogland 

7775, 7779 (US), Weber 251 (US); Victoria, Hale 58256, 58330, 58326, 

58374, 58474, 58977, 59159, 59414, 59541 (US), Melville 2009 (BM. US), 

Streimann 2820, 3267, 3410, 3414 (US); Macquarie Island, Isaird s.n. (US). 

New Zealand: South Island, Du Rietz 1802a (UPS, US), Hale 66177, 66435 

(US), James 482, 1419, 1471 (BM, US), Galloway s.n. (CHR), Martin s.n. 

(BM, US), Mason 134 (BM). 

Parmelia skultii Hale, new species 

FIGURES 7a, 22c 

Parmelia omphalodes subsp. glacialis Skult, 1985201. [Type collection: 

Mould Bay, Prince Patrick Island, SWT, Canada, MacDonald s.n. (CAN,, 

holotype).] 

DESCmIoN.-SubSimilis Parmelia omphalodi sed thallo 

acidum salazinicum continenti differt. 

Thallus usually growing on soil among mosses or on rocks, 

fragile, brown to nearly black at maturity (or whitish with 

pruina), 6-10 cm broad; lobes sublinear, rather short, 24 mm 

wide, crowded and imbricate with secondary laciniae; upper 

surface plane, smooth to minutely rugulose, often becoming 

white pruinose with age, the margins pseudocyphellate, the 

pseudocyphellae more or less continuous, narrow and inconspicuous, larninal pseudocyphellae rarely seen; lower surface 

black and shiny, sparsely to moderately rhizinate, the rhizines 

simple to fucate, 0.5-2 mm long. Pycnidia rare, 90-110 pm 

in diameter; conidia cylindrical, rod-shaped, 6-8 pm long. 

Apothecia not seen. 

CH?%LSTRY.-AtranOrh, salazinic acid, norstictic acid (equal 

or exceeding the concentration of salazinic acid), consalazinic acid (k), protolichesterinic acid (k), fumarprotocetraric 

acid (k), stictic acid (k), and several unknowns (Skult, 1984). 

HoLoTYPE.-West wall of pass, on soil over rocks, 

Anaktuvuk Pass, 151-52OW, 68"20'N, Alaska, USA, G.A. 

Llano 299a (US). 

REmRKs.-Skult (1984) first pointed out the existence of 

a norstictic acid-containing population of P. omphalodes in his 

broader study of the P. omphalodes complex in eastern 

Fennoscandia. He noted the rather broad, marginally pseudocyphellate lobes and frequent occurrence of pruina, the two 

most important morphological features that separate it from 

typical P. omphalodes. I believe this combination of characters 

to recognize it as a distinct species. It appears to be restricted 

to high latitudes, especially near the ocean, from the Aleutian 

Islands to Novaya Zemlya. 

Specimens Examined 

USA. Alaska: St. Matthew Island, Trelease 1165 (US); Port Clarence, Trelease 

1129 (US); Cape Lisbume, Bean s.n. (US); Diomede Island, Palmer 801 (US); 

Point Barrow, Thomon et al. 25 (US, WIS), Scholander s.n. (US); Attu Island, 

van Schaak 205 (US). Canada. Northwest Territories: Bylot Island, Scoffer 

68129 (WIS); Axel Heiberg Island, Scotfer 46165 (WIS); Kaminceriak Lake, 

Scofter 4316 (WIS). Novaya Zemlya: Lynge s.n. (US). Skult (1985) also reports 

a specimen from Spitsbergen. 

Parrnelia squarrosa 

FIGURES 76, 226 

Parmelia squarrosa Hale, 1971:29. [Type collection: Shenandoah National 

Park, Madison County, Virginia, USA, ME. Hale 36494 (no. 142 in 

Lichenes Americani Exsiccati) (US, holotype).] 

Parmelia saxof ilis *divaricafa Delise ex Nylander, 1890:27. [Type collection: 

Itchigomi, Japan, Almquirf in 1879 (H, Nyl. herb. no. 34853, lectotype).] 

DEsCmoN.-ThalluS adnate to loosely adnate on bark, 

less commonly on rocks, fairly firm, pale greenish to whitish 

mineral gray, 4-15 cm broad; lobes sublinear, becoming 

imbricate, 1-5 mm wide; upper surface plane to foveolate, 

shiny or white pruinose with age, continuous but cracking 

along pseudocyphellae with age, pseudocypellae angular, 0.5-1 

mm long, abundant, marginal and laminal, often forming a 

reticulate network, isidiate, coarse isidia forming along ridges 

and pseudocyphellae, cylindrical and barely constricted at the 

base, 0.10-0.15 x 0.1-0.5 mm high, clustered; lower surface 

black and shiny, densely rhizinate, the rhizines squarrosely 

branched, 1-1.5 mm long. Pycnidia rare; conidia cylindrical, 

straight to slightly bent, 5.5-6.5 pm long. Apothecia rare, 

substipitate, poorly developed, 1-3 mm in diameter, the 

amphithecium densely isidiate; hymenium 55-60 pm; spores 

poorly developed, 8-9 x 13-15 pm, the episporium 2 pm thick. 

CmmsmY.-Atranorin, salazinic acid, and consalazinic 

acid. 

REhuRKs.-Parmeh squarrosa is one of the commoner 

corticolous lichens in both North America (see map in 

Thomson, 1984:313) and Japan, growing on oak and other 

deciduous trees in open forests. The sparse population 

extending from Alaska to California may in fact be a 

continuation of the Japanese one. It is rare in the Alps in Europe 

(Hyvonen, 1985). The overlap in range with P. sauatilis, with 

which it was lumped in the past, is extremely limited. In eastern 

United States, for example, P. saxatilis occurs in exposed 

habitats on the highest mountains (more than 1500 m), above 

the habitats where P. squarrosa normally occurs (Dey, 1978). 

It does grow with P. saxatilis in Maine and along the eastern 

shore of Lake Superior, but even in these areas where they 

appear to be sympatric, the two species occupy different 

habitats, P. saxatilis growing on exposed rocks, P. squarrosa 

on less-exposed rocks and trees back from the shorelines. 

Representative Specimens Examined 

Canada: Nova Scotia, MMSS 73 (US); Quebec, Fabiur 7836, GaNo 3095 (US); 

Ontario, Hale 33556, 34818, 34945. 36556, 49836 (US), Sharp 440 (US); 

British Columbia, Brodo 14257 (US). USA: Maine, Egan 5833, Hale 37534, 

Merrill in Lichenes Exsiccafi 35 (US); New Hampshire, Tuckerman in 

Reliquiae Tuckermaniarue 72; Massachusetts, Hermann 14087, Willey s.n. 

(US); Connecticut, Hale 33, 14915, 15406 (US); New York, Brodo 1213, Hale 

16661 (US); New Jersey, Hale 15288 (US); Pennsylvania, Hale 16046, 17217 
44 SMITHSONIAN CONTRIBUnONS TO BOTANY 

(US); Maryland, Hale 14489,Pliff s.n. (US); West Virginia, Hale 12501, 14966 

(US); Kentucky, Hale 13739 (US); Indiana, Hale 14191 (US); Missouri, Hale 

4241 (US); Michigan, Hale 34126, 33995 (US); Minnesota, Fink 57, Hale 

23045 (US); Iowa, Fink s.n. (US); Virginia, Hale 15170, 18170 (US); 

Tennessee, Hale, 18033, Moore 274 (US); South Carolina, Hale 7706 (US); 

Alabama, Hale 7072, McCullough 535 (US); Georgia, Hale 30922 (US); 

Arkansas, Hale 3518 (US); California, Becking 610602, Hale 49579 (US); 

Oregon, Imshaug 17653b (US); Alaska, Palmer 786 (US). China: Manchuria, 

Asahina s.n. (TNS). Nepal: Khumbu, Poeli 126, 155, 170 @I); Langtang 

Himal, Sfainfon 4104 (BM). Sikkim: Togashi s.n. (TNS). Korea: Fujikawa 

s.n. (TNS). USSR: Kurile Islands: Okaah s.n. (TNS). Saghalien: Fujikawa s.n. 

(TNS). Japan: Hokkaido, M. Nakanishi 3493 (US); Prov. Honshu. Kurokawa 

80003 in Lichenes Rariores Crifici Exsiccafi 483 (US); Prov. Ishikari, Asahina 

s.n. (TNS); Prov. Izu, Asahina s.n. (TNS); Prov. Mutsu, Hale 29293 (US); 

Prov. Kai, Jinzenji 64016 (TNS); Prov. Omi, Asahina s.n. (TNS); Prov. 

Shimotsuke, Harhimoio s.n. (TNS); Prov. Shinano, Kurokawa 59242 (US); 

Prov. Tanba, Asahina 71 (TNS); Prov. Tango, Togashi s.n. (TNS). 

Parmelia submontana, new name 

FIGURES 7c. 22.9 

Parmelia conforfa Bory, 1832:305. [Type collection: Atlas Cedres, Taygefe 

1420 (PC, lectotype). Not Parmelia contoria (Hoffmann) Sprengel, 

1827:298 (= Aspicitia conforfa (Hoffmann) Krempelhuber).] 

Parmelia sarafilir var. conforfa (Bory) Zahlbruckner, 1907:68. 

Parmeha sdcaia f. contorioides Zahlbruckner, 1927a:97. [Type collection: 

Hohenberg, Brennalpe, Austria, Suza s.n. and Neuwalde. St. Agyd, Austria, 

Suza s.n. (W, syntypes; not seen).] 

Parmelia bohemica Nbdvornik, 1951:244.[Type collection: Planavy, Hlinsko, 

Bohemia, Nddvornik s.n. in 1931 (PRM, lectoty-pe; US, isolectotype). 

Not Parmelia bohemica Gyelnik, 1932x21 8 (= Xanthoparmelia conspersa 

(Acharius) Hale).] 

Parmelia bohemica f. conforfoides (Zahlbruckner) h'&dvomik, 1951 :244. 

Parmelia submontana Nbdvornik, 1957:72. [Nomen illeg. Basionym not cited 

in description. Type collection: Based on P. bohemica Nadvornik.] 

DESCRKTION.-T~~~~US loosely attached to bark, trailing in 

well-developed specimens, firm, greenish mineral gray, 10-15 

cm broad; lobes elongate, linear, up to 30 mm long, little 

branched, divaricate, sometimes weakly canaliculate, 2-5 mm 

wide; upper surface shiny, plane to rugulose, continuous, 

pseudocyphellae small, round to effigurate, laminal and 

marginal, 0.3-1 mm long, soon becoming sorediate, the soralia 

orbicular to linear with an eroded center with age, the soredia 

granular to nearly isidioid; lower surface black, sparsely 

rhizinate, the rhizines simple to sparsely furcate, 1-1.5 mm 

long. Apothecia and pycnidia not seen. 

Cmmsmy.-Atranorin, salazinic acid, and consalazinic 

acid. 

REms.-This relatively rare European species has not 

been well understood. Nylander (1860:389), who must have 

examined the type specimen in Paris, considered it an elongate 

form of "l? suxatilis var. sulcata." He said it also occurred in 

North America but cited no specimens. Zahlbruckner (1907) 

gave the first modern review (as R saxatifis var. conrorta), 

including an excellent figure of the species, stressing the long 

separate lobes. Recent collections in Austria and Corsica have 

confirmed in my mind the distinctiveness of the species: long, 

little branched lobes, numerous orbicular soralia, small 

pseudocyphellae, and sparse, mostly simple rhizines, as 

characterized by Poelt (1974:447). Schindler (1975) has written 

an up-to-date summary of the distribution and ecology of the 

species in Europe. It is a Mediterranean-South European 

montane element. 

Specimens Examined 

France: Corsica, Lambinon, Rondon and Vezda in Lichenes Selecti Exsiccaii 

845 (US). Austria: Linz, Schauer s.n. (M). Czechoslovakia: Bohemia, 

Nddvornik s.n. (FH); Orlicke, Nadvornik s.n. (US). Schindler (1975) cites 

further collections from Germany, Poland, Russia, Yugoslavia, Italy, Greece, 

Turkey, and Tunisia. 

Parmelia submutata 

FIGURES 74 22f 

Parmelia submuiaia Hue, 1899:172. [Trpe collection: San-tchang-kiou. 

Yunnan, China, Delavay s.n. (PC, lectotype).] 

Parmelia rhododendri Zahlbruckner, 1930: 187. [Type collection: Hsiangschuiho, Yunnan, China, Handel-Mazzetii 6518 (w, lectotype; TNS, US, 

isolectotypes).] 

of Kopati, Djago, and Muli, China, Rock s.n. (w, lectotype).] 

Parmelia leiocarpodes Zahlbruckner, 1934b:207. [Type collection: mountains 

DEScmIoN.-Thallus adnate to closely adnate, firm, 

greenish mineral gray (brownish in the herbarium), 8-12 cm 

broad; lobes subirregular to sublinear, imbricate, 2-5 mm wide; 

upper surface shiny, plane, becoming finely reticulately 

cracked, pseudocyphellae numerous and small, 0.1-0.2 rnm 

long, laminal; lower surface black, densely rhizinate, the 

rhizines simple to densely squarrosely branched, 1-1.5 mm 

long. Pycnidia poorly developed; conidia cylindrical, straight, 

6-8 pm long (after Zahlbruckner for the type of P. 

rhododendri). Apothecia common, adnate to substipitate, 3-5 

mm in diameter, the amphithecium smooth, the disc brown; 

hymenium 45-50 pm; spores 3-5 x 6-9 pm, the episporium 

1 pm thick. 

CHEMISTRY.-Atranorin, salazinic acid, and consalazinic 

acid with or without norstictic acid. 

REmms.-This is the parent morph of l? meiophora. In 

his descriptions Zahlbruckner recognized the dendroid branched 

rhizines but not the fine reticulations on the surface 

representing pseudocyphellae. It is more widesp read than i? 

meiophora but seems to occur in similar habitats, high 

elevation pine-Rhododendron forests. About a third of the 

specimens tested from Taiwan have relatively high concentrations of norstictic acid. 

Representative Specimens Examined 

Taiwan: Prov. Ilan, Kurokawa 929, 931, 1038 in Lichenes Crifici Selecfi 

Exsiccafi 38 (US); Mt. Chien-San, Kurokawa 932 (US); Mt. Arisan, Ogaia 

s.n. (TNS); Taichung County, Chun s.n. (US). China: Yunnan, Delavay s.n. 

(PC). It has also been reported by Kurokawa (l966:610) from Nepal. 
NUMBER 66 45 

FIGURE 23.-Species of Parmelia: a, P. subtestacea (holotype in US); b, P. sulcaia (Kjellmert s.n.); c, P. 

fenuirima (James 488 pp.); d P. rudior (type of P. erimis in H). (Scales in mm.) 

Parmelia subtestacea Hale, new species 

FIGURES le, 23a, 24 

DEscmoN.-Simi~is I? testacea sed thallo acidum echinocarpicum continenti differt. 

Thallus adnate on bark, usually thin and rather brittle, pale 

greenish mineral gray, 5-12 cm broad; lobes sublinear to 

mostly subirregular, short, imbricate, 1-5 mm wide, usually 

with marginal secondary lobes 1-2 mm wide, flaring to 

rounded, developing with age; upper surface shiny, plane 

to distinctly rugose-foveolate, continuous, pseudocyphellae 

forming a nearly continuous white rim 0.1-0.2 mm wide 

around the main and secondary lobes, also laminal on the main 

or older lobes, separate, sparse, 0.2-0.4 mm long, usually not 

fissuring with age; lower surface black, sparsely to moderately 

rhizinate but with a distinct bare to papillate brown zone around 

the margins, the rhizines simple to furcate to sparsely 

squarrosely branched, 0.5-1.5 mm long. Pycnidia numerous, 

90-100 pm in diameter; conidia cylindrical, rod-shaped to 

weakly bifusiform, 5.5-6.5 pm long. Apothecia common, 

substipitate, generally remaining flat but sometimes cupuliform, 4-20 mm in diameter, the disc very pale brown, rarely 

darkening, splitting radially at maturity, the amphithecium 

rugose and effigurate-pseudocyphellate; hymenium 60-65 pm; 

spores 7-10 x 11-15 pm, the episporium 1.0-1.5 pm thick. 

Cmmmy.-Atranorin, chloroauanorin, echinocarpic acid 

(and associated unknowns), with or without unknown #27 or 

rarely with unknown #27 alone. A fatty acid near protolichesterinic acid was detected in several specimens tested by Dr. 

C.F. Culberson). 

HOLOTYPE.-silver beech forest with understory grazing, 

near entrance to Mt. Aspiring National Park, Routeburn Road, 
46 SMITHSONIAN CONTRIBUTIONS TO BOTATTY 

/1 BOTANY DIVISION 3 50 IM MILES 

0 50 iw I(lL0MEIRES 

FIGURE 24.-Distribution of Parmelio subtestacea in New Zealand. (Circles = unknown #27 chernotype; solid 

dots = echinocarpic acid and unknown #27 chemotype.) 
NUMBER 66 47 

elev. 350 m, South Island, New Zealand, M.E. Hale 65402, 

30 Jan 1984 (US; isotypes in CHR, BM). 

REMARKS.4alloway and Elix (1983) recently summarized 

their knowledge of the intergrading morphologies and diverse 

chemical patterns that characterize the P. testacea complex, 

corticolous lichens especially common in New Zealand and 

Tasmania. They concluded from a study of herbarium 

specimens that this complex represented a single variable 

species, P. testacea, but also stressed the need for more detailed 

field studies. 

Hoping to elucidate this species more fully myself for this 

monograph, I conducted field studies of several weeks duration 

in New Zealand and Tasmania in 1984 and 1985, randomly 

collecting 902 specimens at 86 localities. All of the material 

was chromatographed and examined for spores and morphology. I concluded that the P. testacea complex can be divided 

into two distinct major groups, the P. crambidiocarpa group 

(R crambidiocarpa, P. norcrambidiocarpa and P. salcrambidiocarpa) and the P. testacea group (P. subtestacea and P. 

testacea). 

In terms of gross morphology, the P. testacea group is 

distinguished by the adnate to closely adnate thallus, irregularly 

lobed thallus with frequent production of roundish to flaring 

marginal secondary lobes, by the relatively sparse, mostly 

marginal pseudocyphellae, and by sparse to moderate rhizines. 

In contrast the P. crambidiocarpa group has sublinear lobes, 

infrequent production of secondary lobes, marginal and laminal 

pseudocyphellae, and dense rhizines (see P. crambidiocarpa 

for fuller discussions of the differences). 

The Parmelia testacea group has three different lichen 

substances produced in four different combinations: echinocarpic acid-unknown #27, salazinic acid, salazinic acid-unknown 

#27, and unknown #27 alone. Of 381 random collections on 

North and South islands of New Zealand, the echinocarpic 

acid-unknown #27 chemotype made up 56% (215 specimens) 

and unknown #27 alone 7% (26 specimens). These two 

chemotypes define P. subtestacea. The other two combinations 

with salazinic acid (salazinic acid-unknown #27 in 99 

specimens (26%) and salazinic acid alone in 41 (11%)) 

constitute P. testacea. 

The chemotypes have strong geographical correlations. 

Parmelia subtestacea with echinocarpic acid-unknown #27 

does not occur on North Island at all but predominates on 

South Island (Figure 24), where it makes up 71% of the group 

(215 of 302 collections). It is virtually the only member of the 

group found south of the Mt. Cook region, and north of Mt. 

Cook it occurs chiefly in the Arthurs Pass and Lewis Pass 

regions. Salazinic acid-containing P. testacea is essentially 

allopatric, occumng mostly at low elevations north of Lewis 

Pass, eastern Canterbury, and on North Island. All specimens 

of the group in Tasmania are P. testacea. 

The apothecia of I? subtestacea are adnate to substipitate, 

most often with an open pale tan disc (69% of 32 fertile 

specimens examined) or light to medium brown (28%) or rarely 

dark brown (3%). Spores are small and thin-walled: Average 

maximum dimension 8.7 x 12.6 pm (range 7-10 x 11-15 pm 

for 22 specimens examined). Eight fertile collections of the 

unknown #27 chemotype have virtually identical apothecia and 

spore dimensions but 75% of the specimens have a brown or 

dark brown apothecial disc with only 25% pale tan. 

Parmelia subtestacea is especially common on roadside 

Nothofagus and other trees (e.g., Griselina lucida, Leptospermum ericoides, Metrosideros tomentosa) in most forested 

areas of South Island. It is, however, rare at base level in 

mature, dense forests, where it occurs only on canopy branches. 

The upper limit of elevatioun is about 600 m, above which the 

P. crambidiocarpa group occurs. 

The chemotype unknown #27 alone is considered to be the 

“acid-free” chemotype of P. subtestacea. It is rare and I 

collected only 27 specimens out of 241 in this group. It 

occurred with the echinocarpic acid-unknown #27 chemotype 

at 4 localities, intermixed on the same trees, and alone at 4 

other localities. On North Island, where the echinocarpic 

acid-unknown #27 chemotype is lacking, I collected 13 

specimens with unknown #27 at just two localities, 12 of them 

in a mass sample of a fallen tree near the Tongariro National 

Park headquarters on Okahune road. No specimens have been 

collected at the same sites as P. testacea, and this is the main 

justification for recognizing this chemotype as the acid-free 

chemotype of P. subtestacea. 

Representative Specimens Examined 

(echinocarpic acid and unknown #27) 

New Zealand: South Island, Allan CHR160348 (CHR), Galloway CHR342966 

(CHR), Hale 63246, 65220, 65466. 65585, 65643. 65758, 65820 (US), 

Tihomson CHR342997 (CHR). 

Representative Specimens Examined 

(unknown #27 alone) 

New Zealand: North Islandn, Hale 65552 (US); South Island, Elk 6989,7085. 

7138 (ANUC), Hale 65505,65777 (US), Martin ZAlll (CHR). 

Parmelia sulcata 

FIGURES 7j 23b 

Parmelia sulcata Taylor, 1836: 145. [Type collection: Dunkerron, Keny, 

Parmelia saxatilk var. rosiformis (“rosaefomk”) Acharius, 1810:470. [Type 

Parmelia saxatilk var. sulcata (Taylor) Nylander, 1860:389. 

Parmelia cruenta Darbishire. 1912:13. [Type collection: Navarino 

Island, Tierra del Fuego, Chile, Skottsberg s.n. (S, lectotype; BM. 

isolectotype).] 

Ireland, Taylor s.n. (FH-Tayl, lectotype; BM, US, isolectotypes).] 

collection: Sweden, S.C. (H-Ach., lectotype).] 

Parmelia rosiformis (Acharius) Gyelnik. 1928592. 

Parmelia sulcata f. aberrans Zahlbruckner, 193 1:84. [Type collection: 

Cerro El Roble do Libun, Chile, Espinosa s.n. (W, lectotype).] 

Parmelia rosiformk f. cretaceosorediosa Gyelnik. 193 1 b:289. [Type collection: between Monor and Pilis, Hungary, Timko s.n. (BP, lectotype).] 
48 SMITHSONIAN CONTRIBUnONS TO BOTANY 

Parmelia rosiformis f. pomazensis Gyelnik, 1931b:289. [Type collection: near Porn&, Pest, Hundary, Gyelnik s.n. (BP, lectotype).] 

near PiLissanto, Pest, Hungary, Kummerli et al. s.n. (BP, lectotype).] 

Parmelia rosiformis f. rufescenfisorediosa Gyelnik, 193 lb:290. [Type 

collection: Zlatn6, Hungary, Va‘ngel s.n. (BP, lectotype).] 

Parmelia rosiformis f. subnullisorediosa Gyelnik, 193 lb:290. [Type collection: Mt. Paulmauer, St. Aegyd, Austria, Swa s.n. (BP, lectotype).] 

Parmelia rosiformis f. furocensis Gyelnik, 1931b:290. [Type collection: near 

Stubnyafurdo, Turoc, Hungary, Margittai s.n. (BP, lectotype).] 

Parmelia rosifomis f. viridisorediosa Gyelnik, 1931b:289. [Type collection: 

Nagyszenas Mountains near Budapest, Hungary, Gyelnik s.n. (BP, 

lectotype).] 

Parmelia rosiformis var. pseudoviridis Gyelnik, 1932b:448. [Type collection: 

St. Gilgen, Austria, Gyelnik s.n. (BP, lectotype).] 

Parmelia rosiformis f. fagicola Gyelnik, 1932b3450. [Type collection: near 

Kudsir, Hungary, Gyelnik s.n. (BP, lectotype).] 

[See Hillmann (1936) for additional subspecific epithets that I consider to be 

synonyms.] 

DESCRIPI?ON.-T~~~~US adnate to loosely adnate on bark and 

rock, rather firm, greenish to whitish mineral gray, 8-20 cm 

broad; lobes sublinear but rather short, crowded to imbricate, 

2-5 mm wide; upper surface shiny, becoming strongly 

foveolate, continuous to irregularly cracked with age, pseudocyphellae effigurate, 0.3-1 mm long, laminal and marginal, 

separate, soon becoming sorediate, the soralia marginal and 

laminal along the ridges; lower surface black, densely rhizinate, 

the rhizines becoming densely squarrosely branched. Pycnidia 

poorly developed; conidia (Figure 12i) cylindrical, straight to 

slightly bent, 68 pm long (Hillmann (1939) reports 5-8 pm). 

Apothecia rare, substipitate, 2-6 mm in diameter, the 

amphithecium smooth, the rim sorediate, disc dark brown; 

hymenium 55-60 pm; spores usually poorly developed, 6-8 

x 11-14 pm, the episporium 1 pn thick. 

CHEMISTRY.-A~~~~OM and chloroatranorin, salazinic acid 

(Galloway and Elix, 1983), and consalazinic acid. Dey (1978) 

also reports accessory lobaric acid. 

REms.-Parmelia sulcata is easily typified by material 

in the Taylor herbarium. Gyelnik (1930) chose the name 

“rosaeform’s” because it is older (but of a different rank!) 

than sulcata. Although Magnusson (1933) disapproved of this 

name change for such a long-established species, he need not 

have worried, because this epithet was never actually used in 

species rank. Zahlbruckner’s (1929:216) reference to “P. 

rosaeformis Rtihl.” is erroneous. Rohling merely cited “P. 

rosaeformis” as a variety of P. saratilis. 

The most important diagnostic characters of P. sulcata are 

the well-developed laminal soralia and the richly squarrosely 

branched rhizines. Morphological variation in lobe configuration is slight. Most of the numerous varieties and forms have 

been described on the basis of soredial characters, such as 

coarseness and color. The chemistry is very uniform. No 

accessory substances were detected among approximately 100 

specimens tested with thin layer chromatography, although 

Dey (1978) reported specimens from the Appalachian Mountains with lobaric acid. 

Parmelia sulcata is very closely related to P: fertilis, its 

probable parent morph, which is less foveolate and has less 

dense pseudocyphellae and slightly smaller conidia. The two 

species are largely allopatric. The companion isidiate morph, 

P. squarrosa, is more closely related to P. fertilis. 

This is a very widespread, even weedy species (see map of 

the American arctic distribution in Thomson, 1984:318, and 

of the Australasian distribution in Galloway and Elix, 

1983:412), which is more common than P. saratilis in 

temperate and subboreal regions. The rarity of the species in 

Japan, however, is remarkable (Kurokawa, 1968). In austral 

regions it has been collected as far south as Georgia (59”S), 

where it may in fact have been introduced from whaling ships 

(Lindsay, 1973a). 

Representative Specimens Examined 

Canada: Newfoundland, Waghorne 116 (US); Quebec, Bigelow s.n., Fabius 

696, 7830 (US); Ontario, Coin 26889 (TRT, US), Hale 34806, 34912 (US); 

Alberta, Hale 38944, Turner 10174 (US); British Columbia, Macoun 150 

(US); Northwest Territories, Hale 329 (US). USA: Maine, Hale 37626, Harris 

and Bailey 5855 (US); Vermont, DrLlshel 10581 (US); Massachusetts. Gates 

2244, Smifh 12388 (US); Connecticut, Hale 158, 14768 (US); New York, 

Hale 16664, Hermann 14441, Ogden 54109 (US); New Jersey, Hale 15315, 

Leonurd 6428 (US); Pennsylvania, Hale 16058, 19107, Olday 356 (US); 

Maryland, Hale 14488, 14506, Norden 36 (US); West Virginia, Allard 10858, 

Hale 12608,16647 (US); Michigan, Hale 11081 (US), Imhaug 25423 (MSC, 

US); Minnesota. Fink 57, Hale 23148 (US), Wetmore 21759, 22774 (MIN, 

US); North Dakota, TraM and Disrud 71038 (US); South Dakota, Hale 49737 

(US), Wetmore 10352 WIN, US); Montana, Hale 48612,49201 (US); Idaho, 

Hale 49374 (US); Wyoming, Hale 49178 (US); Colorado, Weber andShwhan 

29132, Shushan 5205 (COLO, US); New Mexico, Hale 48924, Imhaug 10008, 

Sfandley 13985 (US); Arizona, Darrow 1853 (US); Washington, Ireland 5997 

(US); Oregon, Hale 21562, 48499 (US); California, Brown 737, Hale 51762, 

Herre 450, Lindroy 258, Tucker 6219 (US). Argentina: Tierra del Fuego, 

Imhaug 54304, 54445 (MSC), Santesson 432 (S, US), 410, 611, 1098,7878 

(S). Chile: Magallanes, Imhaug 38699,39310 (MSC), Santesson 1844, 6384, 

7999, 8189 (S). Greenland: Hansen in Lichenes Groeniandici Exsiccati 12. 

Ireland: W. Mayo, Wattam s.n. (BM). Great Britain: Cumberland, Johnson 69 

(BM). Finland: Fagerstrom in Lichenotheca Fennica 685 (US), hi 10954 

(US). Sweden: Almborn and Hale 33458, Kjellmert s.n. (US). Denmark: 

Chistiansen s.n. (US). Germany: Erichsen 159 (US). France: Crozals s.n. 

(US). Hungary: Fbriss in Lichenotheca Hwtgarica 527. Spain: Vasquez and 

Rico s.n. (US). Portugal: Sampaio in Lichenes de Portugal 258, Tavares in 

Lichenes Lwitaniae Selecti Exsiccati 169 (US). USSR: Elias et al. L-67970 

(COLO, US). Kenya: Hedberg 1846b (UPS, US). Nepal: Khumbu, Poelf 124, 

137. 201 (M). India: Kashmir, Awusthi 2628, 993 (AWAS). Sew Zealand: 

South Island, Du Rietz 1713:7 (UPS, US), Hale 65095, 66429 (US), Martin 

5373 (BM, US). Australia: Sew South Wales, Elk s.n. (AKUC). 

Parmelia ten uirima 

FIGURES 8a, 23c 

Parmelia fenuirima Hooker and Taylor, 1844:645. [Type collection: Tasmania, 

“van Diemens Land,” Gum s.n. (FH-Tayl. herb. sheet 1131. lectotype; BM, 

isolectotype).] 

Parmelia tenuircypha Taylor, 1847:175. [Type collection: Maquany River, 

Australia, Ball herb. (FH-Tayl, lectotype). See Miiller Argoviensis, 

1888:203 .] 

Aspidelia beckettii Stirton. 19003 1. [Type collection: New Zealand. Beckeft 
NUMBER 66 49 

s.n. (BM, lectotype (not seen); see Culberson, 1966:114).] 

Mt. Cargill, Dunedin, New Zealand, Thornon ZA56 (W, lectotype).] 

Parmelia tenuirima var. platynu Zahlbruckner. 1941:108. [Type collection: 

DEscmoN.-Thallus adnate to loosely adnate on bark, 

rather firm, pale to dark greenish mineral gray, 8-60 cm broad; 

lobes broad and subirregular, 4-10 mm wide, often clearly 

black rimmed; upper surface shiny, sometimes coarsely white 

maculate, plane to foveolate in older parts, continuous but 

cracking along old pseudocyphellae, the pseudocyphellae 

effigurate, fairly small but conspicuous, 0.5-1 mm long, 

marginal and laminal, uniformly scattered, separate; lower 

surface black except for a brown marginal bare zone, sparsely 

to moderately rhizinate, the rhizines simple to thickly 

squarrosely branched, 1-1.5 mm long. Pycnidia common; 

conidia cylindrical to slightly bent, a few vaguely bifusiform, 

5.5-7.0 pm long. Apothecia common, stipitate, 5-15 mm in 

diameter, the amphithecium rugose, pseudocyphellate, the disc 

brown, radially split and sometimes perforate with age; 

hymenium 65-70 pm; spores 8-10 x 12-15 pm, the 

episporium about 1 pm thick. 

CmmsmY.-Atranorin and chloroatranorin, salazinic acid 

(Culberson, 1966; Galloway and Elix, 1983), and consalazinic 

acid. 

REms.-This distinctive species has the broadest lobes 

of any in the genus, with conspicuous, separate pseudocyphellae. It often has a faintly greenish cast. Closely related P. 

signifera, largely a saxicolous species, has denser, net-forming 

pseudocyphellae and a brownish cast. Parmelia tenuirima is 

common at higher (and wetter) elevations in southeastern 

mainland Australia, but reaches its best development at higher 

latitudes in New Zealand and Tasmania (see distribution map 

in Galloway and Elix, 1983:415), occurring most often on 

lower trunks of large Nothofagus trees in open mossy forests, 

where colonies attain enormous size. 

Hillmann (1939) examined this species rather carefully and 

disposed of most of the published subspecific names. He 

regarded f. corallina Muller Argoviensis as a probable distinct 

species, at least a variety (it is now recognized as R erumpens 

Kurokawa). A f. sorediata Miiller Argoviensis (1894:258), 

based on a Holst specimen from Usambara, is Parmotrema 

reticulatum (Taylor) Choisy. A variety described by Wilson 

(1893), vat. multifida, was not found by Hillmann or me. 

Finally, f. isidiosa Muller Argoviensis had already been raised 

to species level by Gyelnik as P. pseudotenuirima over 

Hillmann’s objections (“Von der Anschauung ausgehend, dass 

jede ‘planta isidiosa’ eine eigene Species darstellt hat Gyelnik 

die f. isidiosa Miill. Arg. . . . zur Art erhoben”). 

Specimens Examined 

New Zealand: North Island, Hale 58886, 66468 (US); Soulh Island, Du Rietz 

1760:1, 1851:b (UPS, US), Galloway s.n. (CHR), Hale 65086, 65755 (US), 

James 488, 1531 (BM, US), Marfin 557 (BM), Thomson T522 (CHR), Wade 

96 (BM). Australia: Australian Capital Territory, Hale 59395 (US); New South 

Wales, Hale 59357,59370 (US), Weber andMcVean L-49328 (US); Tasmania, 

Brown s.n. (BM), James Au2112, Au2116 (BM), Ofdfiefd s.n. (US), Pearcey 

s.n. (BM); Victoria, Degeliw A-218 (US), Filson 6508, 7038 (US), James 

Au2115 (BM). Lorn 7509b (US). 

Parrnelia testacea 

FIGURES 8b, 234 25 

Parmelia testacea Stirton, 1877-1878:203. [Type collection: near Wellington, North Island, New Zealand, Buchanan s.n. (BM, lectotype; CHR, 

solectotype).] 

Parmelia tenuirima *P. rudior Nylander. 1888:25. [Type csollection: New 

Zealand, Knight 27 (H, Nyl. herb. no. 35289, lectotype).] 

Parmelia tenuirima Hooker and Taylor var. erimis Nylander, 1888:25. [Type 

collection: New Zealand, Knight 55 (H, Nyl. herb. no. 35286, lectotype).] 

Parmelia rudior (Nylander) hhlbruckner, 1929: 198. 

Parmelia erimis mylander) HiUmann, 1939:259. 

Parmelia signifera f. pallidior Zahlbruckner, 1941 :107. [Type collection: 

Dunedin, South Island, New Zealand, Thomson ZA249 (W, lectotype). The 

syntype (Otago Heads, Thornson ZA211 in W) is Parmotrema reticulatum 

(Taylor) Choisy.] 

DESCmIoN.-Thallus adnate to closely adnate on bark, 

thin and rather brittle, pale greenish gray, 5-15 cm broad; lobes 

sublinear to mostly subirregular, short and imbricate, 1-5 mm 

wide, usually with marginal secondary lobules 1-2 mm wide, 

flaring to rounded, becoming dense with age; upper surface 

shiny, plane to rugose-foveolate, continuous, the pseudocyphellae forming a nearly continuous white rim 0.1-0.2 mm 

wide around the main and secondary lobes, also sparsely 

laminal on the main or older lobes, separate, 0.2-0.4 mm long, 

usually not fissuring with age; lower surface black, sparsely 

to moderately rhizinate but with a distinct narrow bare to 

papillate brown zone around the margins, the rhizines simple 

to furcate to sparsely squarrosely branched, 0.5-1.5 mm long. 

Pycnidia numerous, 90-100 pm in diameter; conidia cylindrical, rod-shaped to weakly bifusiform, 5.5-6.5 pm long 

(Hillmann (1939) reported them as bifusiform, 6.0-6.5 pn 

long for the type of P. erimis). Apothecia common, adnate to 

substipitate, generally remaining flat but sometimes cupuliform, 4-20 mm in diameter, the disc very pale tan (sometimes 

concolorous with the thallus) to medium brown, rarely 

darkening, splitting radially at maturity, the amphithecium 

rugose and effigurate-pseudocyphellate; hymenium 60-65 p; 

spores 7-10 x 11-15 pm, the episporium 1-1.5 pm thick. 

CmmsmY.-Atranorin, chloroatranorin, salazinic acid, and 

usually consalazinic acid, with or without unknown #27 (all 

types contain unknown #27). 

REMARKs.-Parmelia testacea (see under P. subtestacea 

for additional discussions of the P. testacea complex) is 

characterized by the adnate, irregularly lobed, marginally 

pseudocyphellate thallus with sparse to moderate rhizines, 

small spores, and the presence of salazinic acid. Externally it 

is indistinguishable from P. subtestacea. Adnation and lobe 

configuration are the same, as are apothecial characters. The 

apothecia have an average maximum diameter of 8.4 mm and 

a range of 4-20 mm (29 specimens examined). The disc tends 

to be pale tan and rarely darkens to brown. Of 29 specimens 

scored for color, 52% were classified as pale tan, 45% as light 
50 SMITHSONIAN CONTRIBUnONS TO BOTANY 

FIGURE 25.-Distribution of Parmelia testacea in New Zealand. (Circles = salazinic acid and unknown #27 

chemotype; solid dots = salazinic acid chemotype.) 
NUMBER 66 51 

brown, and only 3% as dark brown. Spores are small, the 

average dimensions being 8.5 x 12.7 pm (range: 7-10 x 11-15 

pm for 21 specimens examined), virtually identical with those 

of P. subtestacea. 

Parmelia testacea is not chemically uniform. Although the 

main component is salazinic acid, most specimens (71% or 

99 of 140 randomly collected specimens) also contain 

unknown #27. Specimens containing salazinic acid-unknown 

#27 were collected at 6 localities, specimens with only salazinic 

acid at 3 localities, and specimens of both chemotypes at 8 

localities, often intermixed on the same trees. In Tasmania, 

where P. testacea was the commonest member of the P. 

testacea complex, all specimens (156 tested) contained 

salazinic acid and unknown #27. 

Parmelia testacea occupies habitats very similar to those of 

P. subtestacea, although the two species are almost completely 

allopatric, occurring together at only three localities (Tuatapere, Mt. Cook, and Tongariro) (Figure 25). It grows very 

frequently on Nothofagus species in New Zealand and 

Tasmania, especially on trunks of trees along highways or river 

banks. In dense forest it grows on canopy branches and is 

difficult to collect. 

Representative Specimens Examined 

(salazinic acid) 

New Zealand: North Island, Allan CHR160212 (CHR), Elk 8030 (ANUC), 

Hale 58837,65194,65763 (US); South Island, Elk 7807 (AhTJC, CHR), 8171 

(ANUC), Galloway CHR3483 12 (CHR), Hale 65302,65377,65839 (US). 

Representative Specimens Examined 

(salazinic acid and unknown #27) 

New Zealand: North Island, Barton CHR342984 (CHR), Elk 8003 (ANUC), 

Hale 65112,65196,65385,65740,65775 (US); South Island, Elix 6677,7116 

(ANUC), Gnlloway CHR240705 (CHR), Hale 65304, 65335, 65769 (US), 

Taylor 8a (CHR). Australia: Tasmania, Kantvilas 125/82 (ANUC), Hale 

68613,68636642,68671,68688 (US). 
Literature Cited 

Acharius, E. 

1803. Methodur qua oms detectos lichenes. 394 pages. Stockholm. 

18 10. Lichenographia universalis. 696 pages. Gottingen. 

Anden, J. 

1928. Die Strauch- und Laubflechten Mifteleuropas: Anleifung zwn 

Bestimmen der in Mitfeleuropa vorkomnden Strauch- und 

Laubfechten. 217 pages. Jena: Gustav Fischer. 

The Upper Cortex of Parmelia saxatilk and Other Lichen Thalli. 

Lichenologist, 14:29-38. 

Anglesea, D., C. Veldkamp, and G.N. Greenhalgh 

1982. 

Asahina, Y. 

1951a. Lichenes Japiae novae vel minus cognitae (7). JOIUM~ of 

Japanese Botany, 26:329-334. 

1951b. Lichenes Japniae novae vel minus cognitae (8). Journal of 

Japanese Bofany, 26:353-357. 

1952. Lichens of Japan, II: Genus Parmelia. 162 pages. Tokyo. 

1953. Lichenes Japniae novae vel minus cognitae (10). Journal of 

Japanese Botany, 28:6548. 

1954. Lichenologische Notizm (1 10-1 11). Journal of Japanese Botany, 

29:321-324. 

Awasthi, D.D. 

1976. Lichen Genus Parmelia in India, I: Subgenera Parmelia and 

Amphigymnia. Biological Memoirs, series 1 (Lichenology), 1:155- 

229. 

Expidition scientifique de Moree: Section des Sciences physiques. 

Volume III. part 2 (Botanique), 367 pages. Paris. 

Bidrag till kannedom af Finska Vikens Ovegetation, IV: Hoglands 

Lafvar. Medielanden af Societas pro Fauna et Flora Fennica, 

133-143. 

On a New Erratic British Parmelia. Journal of Botany, British and 

Foreign, 10:306,307. 

Recent Additions to the British Lichen-flora. Journal of Botany, 

British and Foreign, 13:140-142. 

On the Lichens Collected by Prof. R.O. Cunningham. JOIUM~ of the 

Linnean Society of London, 15:222-234. 

On the Lichens of Dillenius’s “Historia Muscorum.” JOWM~ of the 

Linnean Society of London, 17:553-581. 

Artificial Reestablishment of Lichens, II: Secondary Products of 

Resynthesized Cladonia cristatella and Lecanora chrysoleuca. 

Mycologia, 72:90-109. 

The Lichen Genus Aspidelia Stirt. Bryologist, 69:113,ll4. 

Parmelia discordans, lichen peu connu d’Europe. Revue Bryologique et Lichenologique, 37:183-186. 

Microconidid Dimorphism in the Lichen Genus Pannelia. MycoBory de Saint-Vincent, J.B. 

1832. 

Brenner, M. 

1886. 

Crombie, J.M. 

1872. 

1875. 

1876. 

1880. 

Culberson, C.F., and V. Ahmadjian 

1980. 

Culbenon, W.L. 

1966. 

1970. 

Culberson, W.L., and C.F. Culberson 

1980. 

Dahl, E. 

1952. 

108 ia, 72 27-135. 

On the Use of Lichen Chemistry in Lichen Systematics. Revue 

Bryologique et Lichdnologique, 21:119-134. 

The Lichens of the Swedish Antarctic Expedition. In N. Nordenskjold, editor, Wissenschafrliche Ergebnisse der Schwedischen 

Sudpolar Expedition 1901-1903 mer Leifung von Dr. Otto 

Nordemkjold 4(11):1-73.3 plates. Stockholm. 

Darbishire, O.V. 

1912. 

Degelius, G. 

1931. Zur Flechtenflora von Angermanland. Arkiv for Botanik, 24A(3):1- 

122. 

1934. Flechten aus Nordfjord und Sunnfjord (Norwegen). Bergens 

MurewnArbok, 3:3-31. 

1944. Nya Bidrag till Kannedomen om Lavfloran pa bark, lignum och 

urbergsblock pa Gotland. Svensk Botanisk T&kri$t, 38:2743. 

Dey, J.P. 

1978. Fruticose and Foliose Lichens of High-mountain Areas of the 

Southern Appalachians. Bryologist, 8 1 : 1-93. 

Historia Murcor um.... 576 pages, 85 plates. Oxford. 

Lichenological Xotes on the Flora of the Antarctic Continent and 

the Subantarctic Islands, E-XI. Nova Hedwigia, 19:439-502. 

Peculiarities of the Australasian Lichen Flora: Accessory Metabolites, Chemical and Hybrid Strains. Journal of the Hattori Botanical 

Laboratory, 52:407415. 

Esslinger, TL. 

1978. A New Status for the Brown Parmeliae. Mycotaxon, 7:45-54. 

Filson, R.B. 

1982. A Contribution on the Genus Parmelia (Lichens) in Southem 

Dillenius, J.J. 

Dodge, C.W. 

1742. 

1970. 

Elk, J.A. 

1982. 

Australia. Aurtralian JOUrMl of Botany, 30:511-582. 

Fries, T.M. 

1871-1874. Lichenographia Scandinavica sive dispositio lichenm in 

Danica, Suecia, Norvegia, Fennia, Lnpponia Rossica hactenus 

collectorum. 639 pages. Uppsala. 

The Lichen Genera Parmelia and Punctelia Krog in Australasia. 

New Zealand Journal of Botany, 21:397420. 

Additional Notes on Parmelia and Punctelia (Lichenised Ascomycotina) in Australasia. New Zealand Journal of Bofany, 22:441445. 

Konspekt Flory lishaynikov Mongol’skoy Narcdnoy Respubliki. In 

Biologicheskie Resursy i Prirodnye Usloviya Mongol’skoi Narodnoy 

Respubliki, 16: 1-200. Leningrad. 

Ahfiana, a New Lichen Genus in the Pameliaceae. Bryologkt, 

Galloway, D., and J. Elk 

1983. 

1984. 

Golubkova, N.S. 

1981. 

Goward, T 

1985. 

88~367-371. 

Goward, T. and T Ahti 

1983. 

Gyelnik, V. 

Parmelia hygrophila, a New Lichen Species from the Pacific 

Northwest of North America. Annales Botanici Fennici, 20:9-13. 

1928. 

1930. 

1931a. 

1931b. 

1932a. 

1932b. 

1934. 

Adatok Magyaronzag Zumo Vegetatiojahoz, 2. Folia Cryptogamica, 1 :571-6O4. 

Lichenologiai Kozlemenyek, 8-19. Magyar Botanikai Lopok, 

28:57-65. 

Additamenta ad cognitionem Parmeliarurn. Fedde Repertorium 

Specierum Novarum Regni Vegefabilis, 29:149-157. 

Additamenta ad cognitionem Parmeliarum, lI. Fedde Repertorium 

Specierwn Novarum Regni Vegetabilis, 29:273-291. 

Additamenta ad cognitionem Parmeliarum, III. Fedde Repertorim 

Specierum Novarum Regni Vegetabilis, 30:209-226. 

Enumeratio lichenum europaeonun novarum rariorumque. Annales 

Mycologici, 30:442-455. 

Lichenes Sipeani ex Oregon. Annales Musei Historico-Naturalis 

Hwrgarici, 28:278-284. 

52 
NUMBER 66 

Hale, M.E., Jr. 

1959. New or Interesting Species of Parmelia in North America. 

Bryologist, 62:16-24. 

1968. New Parmeliae, from Southeast Asia. Journal of Japanese Botany, 

43:324-327. 

1971. Parmelia squarrosa, a New Species in Section Parmelia. Phytologin, 22:29. 

1973. Fine Structure of the Cortex in the Lichen Family Parmeliaceae 

Viewed with the Scanning-electron Microscope. Smithsonian 

Contribuiionr to Botany, 10: 92 pages. 

A Revision of the Lichen Genus Hypotrachyna (Parmeliaceae) in 

Tropical America. Smithonian Contribuiionr to Botany, 25: 73 

pages. 

1976a. A Monograph of the Lichen Genus Parmelinu Hale (Parmeliaceae). 

Smithsonian Contribufionr to Botany, 33: 60 pages. 

1976b. Lichen Structure Viewed with the Scanning Electron Microscope. 

In D.H. Brown, D.L. Hawksworth, and R.H. Bailey, editors, 

Lichenology: Progress and Problem, pages 1-15. New York: 

Academic Press. 

Taxonomy and Distribution of the Parmelia Javentior Group 

(Lxhens: Parmeliaceae). Journal of the Hattori Botanical LaboraPseudocyphellae and Pored Epicortex in the Parmeliaceae: Their 

Delimitation and Evolutionary Significance. Lichenologist, 13:l-10. 

A New Species of Parmelia (Lichenes) with Protocetraric Acid. 

Mycotaxon, 16: 162-164. 

Flavopunctelia, a New Genus in the Parmeliaceae (Ascomycotina) 

Mycotaxon, 20:68 1-682. 

Lichen Chemotaxonomy. In D.H. Brown, D.L. Hawksworth, and 

R.H. Bailey, editors, Lichenology: Progress and Problem, pages 

139-184. New York: Academic Press. 

The Lecanoralean Centrum. In D.R. Reynolds, editor, Ascomycete 

Systematics: The Luttrellian Concept, pages 138-234. New York: 

Springer-Verlag. 

Familie Parmeliaceae. In Rabenhorst’s Kryptogamenfora, 9(3)1:1- 

309. Leipzig. 

Bemerkungen uber einige Arten der Flechtengattung Parmelia, I. 

Hedwigia 783249-267. 

Lichenes Antarctici. Hooker’s London Journal of Botany, 3:634- 

658. 

1975. 

1980. 

tory, 47:75-84. 

1981. 

1982. 

1984. 

Hawksworth, D.L. 

1976. 

Henssen, A. 

1981. 

Hillmann, J. 

1936. 

1939. 

Hooker, J.D.. and T. Taylor 

1844. 

Hue, AM. 

1899. Lichenes extra-Europaei a pluribus collectoribus ad Museum 

parisiense missi et ab A.M. Hue elaborati. Nouvelles Archives de 

MusewnParir, series 3, 1:l-250. 

Lichens. In DeuxiemeExpedition Antarctique Francaise (1908- 

1910) commandde par le Dr. Jean Charcot, 202 pages. 

1915. 

Hyvonen, S. 

1985. Parmelia squarrosa, a Lichen New to Europe. Lichenologist, 

17:3 1 1-3 14. 

Imshaug, H.A. 

1957. Alpine Lichens of Westem North America and Adjacent Canada, I: 

The Macrolichens. Bryologist, M):177-272. 

An SEM Study of Lichen Pruina Crystal Morphology. Scanning 

Electron Microscopy, 3 :279-284. 

Microchemical Studies on Parmelia. Nytt Magasin for Naturvidenskapene, 8857-85. 

The Macrolichens of Alaska. Norsk Polarinstitutt Sbifler, 144:l180. 

Punctelia. a New Lichen Genus in the Parmeliaceae. Nordic Journal 

of Botany, 2:287-292. 

Jackson, D.W. 

1981. 

Krog, H. 

1951. 

1968. 

1982. 

53 

Krog, H., H. Osthagen, and T. Tonsberg 

1980. LavJora: Norske bwk- og bladlav. 312 pages. Oslo: Universitetsforlaget. 

Anaptychiae and Parmeliae. In H. Hara, The Flora of the Eustern 

Himalaya, pages 605-610. Tokyo. 

New or Noteworthy Species of Parmelia of Japan. Journal of 

Japanese Botany, 43:349-353. 

A Note on Some Rare Lichens of Japan. Journal of Japanese 

Botany, 44:225-229. 

A Note on Parmelia omphalodes and Its Related Species. Journal 

of Japanese Botany, 51:377-380. 

Lichens of the Hidaka Mountains. Hokkaido. Memoires of the 

National Science Museum (Tokyo), 459-70. 

Gyrophoric Acid as a Chemical Constituent in the Cortex of Lichen 

Thallus. Journal of Japanese Botany, 45:230-232. 

New Rare or Interesting Lichens from the Southern Hemisphere, 1. 

Lilloa, 14:203-251. 

Kurokawa, S. 

1966. 

1968. 

1969. 

1976. 

Kurokawa, S., and S. Nakanishi 

1971. 

Kurokawa, S., and R. Takahashi 

1970. 

Lamb, LM. 

1948. 

Lindsay, D.C. 

1973a. Probable Introductions of Lichens to South Georgia. British 

Antarctic Survey Bulletin, 33 and 34:169-172. 

1973b. Notes on Antarctic Lichens, VII: The Genera Cetraria Hoffm., 

Hypogymnia (Nyl.) Nyl., Menegazzia Massal.. Parmelia Ach. and 

Platismatia Culb. & Culb. British Antarctic Survey Bulletin, 

36:105-114. 

Lindsay, W.L. 

1859. Memoir on the Spermogones and Pycnides of Filamentous, 

Fruticulose, and Foliaceous Lichens. Transactions of the Royal 

Sociefy of Edinburgh 22:lOl-303. 

Species plantarum. 1200 pages. Stockholm. 

Linnaeus, C. 

Magnusson, A.H. 

1753. 

19 19. Material till Vastkustens lavflora. Svensk Botanisk Ttihbifi, 

1929. Flora over Skandinaviens bwk- och bladlavar. 127 pages. 

Stockholm. 

1933. Gedanken uber Flechtensystematik und ihre Methoden. Meddelanden fran Goteborgs Botaniska Tradgard, 8:49-76. 

Mitteilungen uber Flechten-inhaltsstoffe, CVm: Die Flechtenstoffe 

einiger indischer Parmeliaceen. Philippia, 3:20-23. 

13173-92. 

Misra, G., S. Huneck, and M.E. Hale 

1976. 

Miiller Argoviensis, J. 

1879. Lichenes Japonici. Flora, 62:481-487. 

1883. 

1887. 

1888. 

1892. 

Lichenologische Beitrage, XVII. Flora, 66:4548. 

Lichenologische Beitrage, XXVI. Flora, 70:3 16-320. 

Lichenologische Beitrage, XXTX. Flora, 71:195-208. 

Lichenes Knightiani, in Nova Zelandia lecti additis nonnulis aliis 

ejusdem regionis, quos exponit. Compte-Rendu Societe Royale 

Belgique, 3 1 :22-42. 

Lichenes usambarensis. Botanische Jahrbucher, 20:238-298. 

Analecta australienses. Bulletin Herbier Bossier, 4:87-96. 

New or Interesting Lichens. Studio Botanica Czechoslovaca, 12:244. 

Ein Beitrag mr Kenntnis der Flechtenflora des Tatranationalparkes. 

In Sammlung von Studien uber den Tatranationalpark. Sbornik Prac 

o Tatr. Narodnom Parku, 1 :67-72. 

Synopsis methcdica lichenwn. Volume 1, part 2. Paris. 

Lichenes Scandinaviae. Nother ur Sallskapets pro Fauna et Flora 

Fennica Forhandlingar, 5:l-312. 

Addenda nova ad lichenographiam Europaeam: Continuatio quadragesima septima. Flora, 70:129-136. 

Lichenes Novae Zelandiae. 156 pages. Paris. 

1894. 

1896. 

1951. 

1957. 

Nddvomk, J. 

Nylander. W. 

1860. 

1861. 

1887. 

1888. 
54 NUMBER 66 

1889. Lichenes Insularum Guineensium. 54 pages. Paris. 

1890. Lichenes Japoniae. 122 pages. Pans. 

1973. New and Interesting Macrolichens of British Columbia. Bryologist, 

76:366-387. 

Poelt, J. 

1974. Bestimmwtgsschlussel Europaischer Flechten. 757 pages. Vaduz: 

J. Cramer. 

Rasanen, V. 

1940. Lichenes ab A. Yasuda et aliis in Japonia collecti (I). Journal of 

Japanese Botany, 16:82-98. 

1949. Lichenes novi, IV. Suomalaisen elain-ja Kasvitieteellisen seuran 

Vanamon Tiedonannot ja Poytakirjat, 3:78-89. 

The Lichens of Sweden and Norway. 333 pages. Stockholm and 

Uppsala. 

1975. Uber die Flechte Parmelia contorta Boy und ihre bisher bekannte 

Verbreitung. Herzogia, 3:347-364. 

Skult, H. 

1984. The Parmelia omphalodes (Ascomycetes) Complex in Eastem 

Fennoscandia. Annales Botanici Fennici, 21:117-142. 

1985. A New Subspecies of Parmelia omphalodes (Ascomycetes) 

Described from the Arctic. Annales Botanici Fennici, 22:201-205. 

Caroli Linnaei, System Vegetobiliwn. Edition 16, volume 4, part 

1,592 pages. Gottingen. 

Ohlsson, K.E. 

Santesson, R. 

1984. 

Schindler, H. 

Sprengel, C. 

1827. 

Steiner, J. 

1904. Flechten, auf Madeira und den Kanaren gesammelt von J. 

Bommiiller in den Jahren 1900 und 1901. Oesterreich Botanische 

Zeitschrifi, 54:399409. 

Stinon, I. 

1877-1878. On Certain Lichens Belonging to the Genus Parmelia. 

Scottish Natural ist, 4: 20&203,252-25 4,29 8-299. 

1899 [1900]. On New Lichens from Australia and New Zealand. 

Transactions of the New Zealand Institute, 32:7&82. [Date on title 

page is 1899; actually published 1900.1 

Contribuicao para o estudo das Parmeliaceas Portuguesas. Portugaliae Acta Biologica, 1B:1-204. 

Lichens. In J.T. Mackay, Flora Hibernica, 279 pages. Dublin. 

New Lichens, Principally from the Herbarium of W.J. Hooker. 

Hooker’s London Journal of Botany, 6148-197. 

Lichens of the Alaskan Arctic Slope. 3 14 pages. Toronto: University 

Tavares, C. 

1945. 

Taylor, T 

1836. 

1847. 

Thomson, J.W. 

1979. 

of Toronto Press. 

American Arctic Lichens, 1: The Macrolichens. 504 pages. New 

York: Columbia University Press. 

Lichenes ab A. Yasuda in Japonia collecti, Continuatio I. Botanical 

Magazine (Tokyo), 35:45-62. 

Lichenes Selecti Exriccati. Fascicle 70. Pruhonice. 

Bau und Entwicklung der Flechten-Pycnidien und ihrer Conidien. 

Bibliotheca Lichenologica, 14: 1-1 41. 

Calcium Oxalate Hydrates on the Surface of Lichens. Liehenologist, 

1984. 

Vainio, E.A. 

1921. 

VEda, A. 

1980. 

Vobis, G. 

1980. 

Wadsten, T, and R. Moberg 

1985. 

17~239-245. 

Wilson, F.R.M. 

1893. 

1900. 

Tasmanian Lichens, Part 1. Papers and Proceedings of the Royal 

Society of Tasmania, 1892:133-178. 

Lichenes Kerguelenses: a Roberto Hall anno 1898 prope Royal 

Sound in Kerguelen insula lecti et in Herbario nationali Melbourniensi depositi at a Rev. F.R.M. Wilson determinati. Mimires de 

I’Herbier Bobsier, 1837-88. 

Glushinskite, a Naturally Occurring Magnesium Oxalate. Minerological Magazine, 43:837-840. 

Wilson, J., D. Jones, and J.D. Russell 

1980. 

Zahlbruckner, A. 

1907. Vorarbeiten zu einer Flechtenflora Dalmatiens, IV. Oesterreich 

1927a. Beitrage zur Flechtenflora Niederosterreichs. W. Verhandlwtgen 

1927b. Additamenta ad Lichenographiam Japoniae. Botanical Magazine 

1929. 

1930. 

1931. 

1934a. Flechten der Insel Formosa. Fedde Repertoriwn Specierwn Novorwn 

1934b. Nachtrage zur Flechtenflora Chinas. Hedwigia, 74:195-213. 

1941. 

Botanische Zeitschrifl, 57:65-73. 

der Zoologisch-Botanischen Gesellschaft in Wien, 76:76-101. 

(Tokyo), 41 :313-364. 

Catalogus lichenwn universalis. Volume 6, 618 pages. Leipzig. 

Lichenes. In H. Handel-Mazzetti, Symbolae Sinicae, 3:l-254. 

Neue Flechten. X. Annales Mycologici, 29:75-86. 

Regni Vegetabilis, 33:2248. 

Lichenes Novae Zelandiae a cl. H.H. Allan eiusque collaboratoribus 

lecti. Denkrchriflen der Akademie Wissenschafi Wien, Mathemtischnatiirliche Klasse, 104:l-132. 

A Preliminary Study on Chinese Parmelia. Acta Phytotamnomica 

Sinica, 9:139-166. [In Chinese.] 

Prodromus Lichenwn Sinicorum. 156 pages. Beijing. [In Chinese.] 

Zhao. J.-D. 

1964. 

Zhao, J.-D., L.-W. Xu, and Z.-M. Sun 

1982. 
Index 

(Synonyms in italics) 

Aspidelia 

Lichen 

beckettii, 18, 48 

omphalodes, 34 

saxatilis, 18.38 

acervata, 40 

adaugescens, 19 

bohemica f. contortoides, 44 

bohemica, 44 

brownii, 23 

cochleata, 19 

contorta, 44 

crambidiocarpa, 21 

cruenta, 41 

cunninghamii, 23 

daliensis f. tardiva, 38 

doliensis, 38 

discordans, 23 

erimis, 49 

erumpens, 24 

fertilis, 24 

fraudans, 25 

hakonensis, 28 

hygrophila, 27 

insensitiva, 23 

isidioclada, 27 

kerguelensis, 27 

laevior, 28 

laevior f. denigrata, 28 

laevior f. hakonensis, 28 

[aevior f. microphyllina, 36 

leiocarpodes, 44 

marmariza, 28 

marmariza f. angwtifolia, 28 

marmariza var. physcioides, 19 

meiophora. 30 

meiophora var. isidiota, 30 

neodiscordans, 30 

niitakana, 30 

norcrambidiocarpa. 3 1 

Parmelia 

novae-zelandiae, 3 1 

omphalodes, 34 

omphalodes p. panniformis, 34 

omphalodes f. insensitiva, 23 

omphalodes ssp. discordans, 23 

omphalodes ssp. glacialis, 43 

omphalodes ssp. pinnatifida, 34 

omphalodes var. discordans, 23 

omphalodes var. herminica, 34 

omphalodes var. panniformis f. concentrica, 

ontakensis, 28 

petrophila, 28 

pinnatifida, 34 

protosulcata, 35 

pseudolaevior, 35 

pseudomarmariza, 19 

pseudosaxatilk, 19 

pseudoshinanoana, 35 

pseudosulcata, 27 

pseudotenuirima, 36 

psoromo ides, 27 

queenslandensis, 36 

rhododendri, 44 

ricasolioides, 38 

rosiformis, 47 

rosiformis f. cretaceosorediosa, 47 

rosiformis f. domestica, 41 

rosiformis f. fagicola, 48 

rosiformis f. griseosorediosa, 47 

rosifowis f. pomazensis, 48 

rosiformis f. roseolosorediosa, 48 

rosiformis f. rufescentisorediosa, 48 

rosiformis f. subnullisorediosa, 48 

rosiformis f. turocensb, 48 

rosiformis f. viridisorediosa, 48 

rosiformis var. pseudoviridis, 48 

rudior, 49 

salcrambidiocarpa, 38 

saxatilis, 38 

saxatilis 'divaricata, 43 

34 

saxatilis *fraudans, 25 

saxatilis f. acervata, 40 

saxatilis f. corallicola, 40 

saxatilis f. prima, 40 

saxatilis f. rubricosa, 40 

saxatilis f. squamigera, 40 

saxatilis var. contorta, 44 

saxatilis var. dimorpha, 36 

saxatilis var. imbricatoides, 40 

saxatilk var. laevis, 40 

saxatilis var. pseudoviridis, 40 

saxatilis var. rosiformis, 47 

saxatilis var. signifera, 41 

saxatilis var. sulcata, 47 

sectilis, 40 

shinanoana, 41 

shinanoana f. calvescens, 30 

signifera, 41 

signifera f. pallidior, 49 

skultii, 43 

squarrosa, 43 

subdivaricata, 24 

submarmariza, 28 

submontana, 44 

submutata, 44 

subtestacea, 45 

sulcata, 47 

sulcata f. aberrans, 47 

sulcata f. contortoides, 44 

sulcata var. laevis f. hirsuta, 34 

tenuirima, 48 

tenuirima *rudior, 49 

tenuirima f. corallina, 24 

tenuirima f. isidiosa, 36 

tenuirima var. erimis, 49 

tenuirima var. platyna, 49 

tenuiscypha, 48 

testacea, 49 

yasudae, 27 

*US. GOVERNMENT PRINTING OFFICE: 1987-181-171/60002 

55