A Monograph of the Lichen Genus Parmelina Hale - Smithsonian ...

SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 33 

A Monograph of the Lichen Genus 

Parmelina Hale (Parmeliaceae) 

Mason E. Hale, Jr. 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1976

ABSTRACT 

Hale, Mason E., Jr. A Monograph of the Lichen Genus Parmelina Hale (Par- 

meliaceae). Smithsonian Contyibutions to Botany, number 33, 60 pages, 21 

figures, 1976.-The 47 species of Parmelina are revised on the world level. Two 

sections are recognized: section Parmelzna with 30 species widely distributed in 

temperate to tropical montane regions and section Myelochroa with 17 terpene- 

containing species concentrated in eastern and southern Asia. The genus is 

most closely related to Parmotrema hlassalongo. Five new species, P. crassata 

Hale, P. degelii Hale P. indica Hale, P. rhytidodes Hale, and P. schindleri Hale, 

are described, and six new combinations proposed, P. amagiensis (Asahina) 

Hale, P. damaziana (Zahlbruckner) Hale, P. endoleuca (Taylor) Hale, P. irrugans 

(Nylander) Hale, P. jamesii (Hale) Hale, and P. pastillifera (Harmand) Hale. 

New combinations are also made for Hypotrachyna baguioensis (Hale) Hale and 

Parmotrema nylanderi (Lynge) Hale. 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded 

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

katsura tree Cercidiphyllum juponicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Hale, Mason E. 

A monograph of the lichen genus Parmelina Hale (Parmeliaceae) 

(Smithsonian contributions to botany ; no. 33) 

Bibliography: p. 

Includes index. 

Supt. of Docs. no.: SI 1.29:33 

1. Parmelina. I. Title. 11. Series: Smithsonian Institution. Smithsonian contributions to 

botany : no. 33. 

QKl.SZ747 no. 33 [QK585.P2] 581’.08s [589’.1] 76-608065

Contents 

Page 

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

Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 

Chemistry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 

Phytogeography and Speciation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 

Classification of Parmelina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 

Key to the Species of Parmelina . . . . . . . . . . . . . . . . . . . . . . . . . 16 

Species Treatment . . , . . . . . . . . . . . . . . 18 

Doubtful and Rejected Names . . . . . . . . . . . . . . . . . . . . . . ... 53 

Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55 

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 

iii

A Monograph of the Lichen Genus 

Parmelina Hale (Parmeliaceae) 

Introduction 

The genus Parmelina Hale is a segregate of Par- 

melia Acharius comprising 47 species widely dis- 

tributed in temperate and tropical regions. It is 

recogni7ed by the narrow, adnate lobes, marginal 

cilia, and absence of usnic acid in the cortex (Hale, 

1974:438). Some of the species included here were 

formerl) classified in the ill-defined (and invalid) 

genus Zmbitca? ia (Schreber) Michaux (Hale and 

Kurokawa, 1964: 130). This world level monograph 

is based on the study of all available type speci- 

mens, collections preserved in the major herbaria, 

and personal field work. The format follows my 

earlier treatments of Bulbothrtx (Hale, 1976b), 

Hypotrachyna (Hale, 1975a), Pseudopawnella 

(Hale, 1976a), and Rellcina (Hale, 197513). 

I wish to thank curators of the following insti- 

tutions who sent specimens on loan so promptl) and 

often allowed extensions of loan periods: BM, BO, 

BP, COLO, DUKE, F, FH, FI, G, GLAM, H, KAN, 

KR, L, LD, Rl, LlICH, NSC, RlVAf, NSLV, NY, 

PC, PH, REX;, S, SI, TNS, TRH, TUR, UC, UPS, 

W, LVIS, WU, and ZT. Specimens in TNS were 

annotated during 1964-65 while I studied in Tokyo. 

The priyate herbaria of Dr. D. D. Awasthi, Dr. 

Gunnar Degelius, Dr S Nakanishi, Dr. Clyde Reed, 

Mason E Hale, Jr, Department of Botan), National Mu- 

seum of Aatiital Histor), Smithsonian Institution, Washang- 

ton, D. C. 20160. 

Mason E. Hale, Jr. 

I 

and Dr. R. Santesson were also geneiously placed 

at my disposal. The material in these various her- 

baria was chemically tested and annotated between 

1958 and 1975, but not all of it has been re- 

examined for possible name changes in the light of 

new synonymies, species concepts, or improved 

chemical techniques. 

Any monograph is bound to be more complete if 

the author has had an opportunity to observe and 

collect specimens in the field. I am especially grate- 

ful to the following colleagues for assistance in 

arranging excursions: blr, J. Anderson (Sarawak), 

Alrs. Sheila Collenette (Sabah), Dr. S. Kurokawa 

(Japan), Dr. 51. L6pez-Figueii-as (l’enezuela), Dr. 

W. Meijer (Sabah), Dr. hI. Nakanishi and Dr. S. 

Nakanishi (Japan), Dr. P. G. Patwardhan (India), 

Dr. Stella Thrower (Hong Kong), and Dr. Flora 

Uyenco (Philippines). I have also conducted field 

studies in Llexico and Central America, the Lesser 

Antilles, and Malaya. 

The scanning-electron photographs were pre- 

pared by the Smithsonian Scanning-Electron Rficro- 

scope Laboratory. Photographs of the specimens 

were taken by the Smithsonian Photographic Ser- 

vices. 

Grants in support of this ret,earch have been re- 

ceived from the National Science Foundation (Sys- 

tematic Biology and the U. S-Japan Cooperative 

Science Program), hforden-Smi thsonian Expedition 

to Dominica, National Geographic Society, and the 

Smithsonian Research Foundat ion.

2 

Morphology 

THALISIS STRLCTURE.-Pa?-melina basically has a 

closely appressed to adnate foliose thallus with narrow 

lobes (1-4 mm wide). Most species have more 

or less subirregular lobation and apically rotund 

lobes (Figures I&, 18b), although narrow lobed 

species are sublinear (Figures 12d, 16c,). There is, 

however, a wide range of variation in lobe configuration 

and width, much more than exists, for example, 

in Hypotmchynn (Hale, 1975a). 

The internal anatomy is similar to that of other 

epicorticate genera (Hale, 1973a). A thin, generally 

palisade plectenchymatous cortex is overlain by a 

pored epicortex (Figures 1 and 2). The medulla 

consists of loosely packed hyphae, often encrusted 

with lichen substances (Figure 2f). The lower cortex 

is p”p1ectenchymatous (Figure 20-d). 

The upper surface is strongly white maculate in 

Pwmelina con.son (Figure 3a), P. miielleii, and P. 

pilostr, less conspiciiously so in P. nzelanochaetn, P. 

pa.stillifei.a, P. qiiercina, and P. tiliacea, and faintly 

or not at all in the remaining species. 

The lower surface is black with only three exceptions. 

Pai.melina cnormis has a uniformly pale 

brown lower surface, and P. expallida and P. versiforrnis 

are darker brown tending toward black at 

the center. 

CILIA.-The marginal cilia which characterize 

Pnrrnelina are usually distinct but may be variably 

dispersed around the lobe margins. In many species 

they occur more or less regularly both on lobe tips 

and in the axils (Figures 3a, lia, 18b). In others 

cilia may be lacking at the lobe tips and confined 

to the axils (Figure 3b) or may be so sparse and 

inconspicuous in a few species, such as P. .rimplicioy, 

that they are overlooked. In these cases one might 

tend to classify them in the genus Pseutlopaimelia 

(Hale, 1976a), which is differentiated from Parmelina, 

among other ways, by the total absence of 

marginal cilia. Great care must be taken to establish 

the presence or absence of cilia in the lobe 

axils. Another example of this problem is Pa?-melin(i 

endoleum and P.rezidopnrmelin siibtiliacea (Nylander) 

Hale, two very similar species in Australia that 

differ in the production of cilia. Both have fatty 

acids and cannot be separated easily by a chemical 

test. 

Confusion may also arise with species of Hypot~a- 

SSIITHSONIAPi COSTRIBUTIONS TO BOTANY 

chyna that lack cilia but may have a very dense 

rhizinal mat below. The mat may assume a marginal 

position and resemble cilia. These “cilia,” 

however, will be dichotomously branched. Such a 

distinction fails in many specimens of H. rcvoluta 

(Floerke) Hale having rather sparsely branched 

rhizines and a few marginally positioned “cilia.” If 

incorrectly interpreted as having furcate rhizines 

and sparse marginal cilia, such a lichen might be 

identified as Parmelina cryptochlom, which has 

more powdery capitate soralia, or as pustulate P. 

spiimosn, which has a pale yellowish medulla. All 

of these species have gvrophoric acid. 

One further example of parallelism involving 

Pam e 1 in a rl is sec t a and Hypo t ru c h y n a n eod issec t a 

(Hale) Hale can be mentioned. The latter has 

clearly dichotomously branched rhizines and lacks 

cilia, yet both contain the same chemical, gyrophoric 

acid, and have virtually identical lobe configuration. 

It is difficult, if not impossible, to decide at 

this time whether intergeneric hybridization has 

occurred in these cases and resulted in “hybrid” 

species. 

RHIzIms.-?’he rhiLines of Parmelina are of two 

types: simple to sparsely furcate (Figure 3c) and 

squarrose. Squarrose rhizines, that is, those with a 

main axis and short lateral branches, are confined 

mostly to some species in section Myelochroa. This 

pattern is also known in a few species of Parmelia 

(e.g., P. ,szilcata Taylor) and Pai.motl-ema (the P. 

7.cticnlOtzLm group). 

ISIDIA AND LosuLEs.-Isidia are normally cylindrical 

and erect as in other parmelioid genera and 

occur in the following 13 species: P. antillensis, P. 

expallidla, P. di.csecta (Figure 34, P. horrescens, P. 

jamesii, P. indica, P. lindmanii, P. melanochaeta, 

P. obsessn, P. perisidians, P. tiliacea, P. usambarensis, 

and P. urnllichinna. Isidia are distinctly lobulate 

in P. degelii ant1 P. spatlziilata (Figure 44 and 

uniquely peltate in P. pastillifem (Figures 3e, 4b), a 

close relative of P. tiliacea, Apical cilia almost always 

occur in P. howescens and P. melanochaeta 

(Figure 4c) and resemble those in PaYmotTema crin i- 

tiinz (Acliarius) Choisy. 

Lobules not originating from isidia are found in 

P. schindle7.i and P. xantholepis (Figure 4e,f); they 

are mostly marginal. 

PvsTuLEs.-These are characteristic of four species, 

P. Iiaynchinensis, P. leucotylizn (Figure 5g), P.

NUMBER 33 

FIGURE 1.-Epicortical pores oti the surface of Parnze[irzn species: a,b, P. ettornzis (X 400 and 

1600) (Jeilicoe 150 in US); c,d, P. muelleri (X 400 and 1600) (Hale 42219); e,!, P. indica (X 400 

atid 1600) (Hale 43884). (Scatitiing-electron microphotographs.) 

3

4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 

2.-Internal structure of Parmelina; a, vertical cross section of P. degelii (Degelius in 

US) (x 800); b, upper cortex of P. degelii (same section as in a) (x 1600); c, vertical cross sec- 

tion of P. galbina (Hale 23415) (x 800); d, upper cortex of P. galbina (same section as d) (x 

1600); e, epicortex (arrow) of P. pastillifera (Schroppel 143) (x 5000); f, crystals of lecanoric 

acid on hyphae of P. pastillifera (Schroppel 143) (x 4000).

NUMBER 33 5 

FIGURE 3.-Morphological structures of Parmelina: a, white maculae and marginal cilia of P. 

consors (Malme 1282 in US) (x 10); b, axillary cilia of P. lindmanii (Nee and Mori 4258 in US) 

(x 10); c, rhizines of P. galbina (Hale 18930) (x 70 with scanning-electron microscope); d, 

isidia of P. dissecta (Montes 10121 pro parte) (x 90 with scanning-electron microscope): e, 

peltate isidia of P. pastillifera (Schroppel 143) (x 90 with scanning electron microscope): 

f, cross-section of a peltate isidium of P. pastillifera (Schroppel 143) (x 400 with scanning- 

electron microscope).


FIGURE 

4.-Morphological structures of Parwelina: a, isidia of P. dissecta (Hale 33375) (x 10); b, 

peltate isidia of P. pastillifera (Schroppel 143) (x 10); c, ciliate isidia of P. horrescens (Imshaug 

22174) (X 10); d, procumbetit isidia of P. spathulata (Koefler, in US) (x 10); e, lobules of P. 

schindleri (Schindler 4569 in US) (x 10); f, 1ol)ules of P. xantholepis (Hale 25993) (x 10).

NUMBER 33 7 

FIGURE 

5.-Morphological structures of Parrnelina: a, soralia of P. cc~irulenta (Weber anti McYean 

L-51280 in US) x 10); b, soralia of P. eryptochlora (Hale 35357) (x 10); c, pustules of P. su0- 

fatiTcens (isotjpe in US) (x 10); tl, coarse coralia of P. swinsco7~'ii (Swinscow K 31/13 in US) 

(x 10); e, pustules of P. spiiniosa (Hale 19897) (x 10): f, pustule of P. ~piiniosa (Hale 19897) 

(x 100 with scanning-electron microscope); g, pustules of P. leitcotyliza (Hale 29.544) (x 10).


spumosa (Figure 5e,f), and P. szibfatiscens (Figure 

5c). Soredia are not produced on these pustules. 

Parmelina denegans, on the other hand, is clearly 

pustulate initially but produces coarse soredia at 

maturity. 

SOREDIA.-SOralia are produced in seven species of 

Parmelina: P. auwlenta, P. cryptochlora, P. dene- 

guns, P. metawvoluta, P. muelleri, P. pilosa, and 

P. swinscowii (Figures 5a,b,d, 17b, 18b). They are 

orbicular and lamina1 or subterminal but never 

marginal and linear as in Parmotrema. 

The remaining 17 species of Parmelina produce 

no vegetative propagules, although the very 

wrinkled and often fragile upper cortex of P. 

entotheiochroa or P. rhytidodes might be mistaken 

for pustules (Figure 6). 

APoTHEcIA.-The apothecia are typically adnate 

or sessile and less than 5 mm in diameter, though 

Pal-nzelina irrugniis has discs up to 10 mm broad. 

The disc is occasionally perforate in P. co1zsors. No 

species has a coronate rim, and only two, P. qzier- 

cina and P. tiliacea, sometimes have retrorse 

rhizines on the lower part of the amphithecium. 

No fertile collections are known as yet for P. 

ci.yptochlora, P. degelii, P. hayachinensis, P. iradica, 

P. jamesii, P. spathulatcl, or P. ~winscowi~. 

Spore siLe is very uniform within a given species 

and not greatly different between unrelated species. 

For example, the species in the P. dissecta-P. hor- 

wscens group (Figure 9) have spores 8-12 X 12-18 

pm. The P. aurzilenta-P. subaurulenta group (sec- 

tion Myelochroa) has spores 5-10 X 7-15 pm. The 

largest spores have been measured in P. versiformis 

(10-14 X 18-28 pm), but no other species in the 

genus have spores greater than 19 pm long. The 

smallest spores (3-6 X 7-8 pm) occur in the two 

anamolous norstictic acid-containing species, P. 

antillensis and P. phlyctina. 

Chemistry 

The chemistry of Parmelina species was de- 

termined with microcrystal tests before 1965 (Hale 

and Kurokawa, 1964) and later with thin-layer 

chromatography (Merck F-254 silica gel pre-coated 

plates). Two solvent systems, hexane-ether-formic 

acid and benzene-dioxane-acetic acid, have been 

routinely employed using C. Culberson’s methods 

(1972). I have had the benefit of Dr. Culberson’s 

advice on the identification of many of the lichen 

substances, and she has very kindly communicated 

to me the results of her work on the “horrescens” 

unknown. It should be assumed, incidentally, that, 

unless otherwise stated, the components of each 

species in tlie taxonomic section refer to the liolo- 

type or lectotype specimens with additional com- 

ments on tlie results from other specimens. 

Before listing the lichen substances discovered 

so far in Pal-melinn, I might point out a most re- 

markable feature in its chemical profile. Orcinol 

FIGURE 

6,-MorpholoLgy of Parmelina: a, flaking cortex of P. entotheiochroa (Hale 29.534) (x 4); 

b, wrinkles of P. rhytidodes (Numnriri 156 in US) (x 10).

NUMBER 33 

depsides (except for gyrophoric and lecanoric acids), 

meta-depsides, orcinol depsidones (except for rare 

lobaric acid), and the p-orcinol depsides related to 

barbatic acid are completely lacking. Two other 

cortical substances, lichexanthone and usnic acid, 

are also missing. The general impression is that 

Parmelina has a relatively “primitive” assemblage 

of substances and would fall on a lower evolutionary 

scale than Hypotrachyna, Parmotrerna, and 

Pseudopamelia but significantly higher than Bzilbothrix, 

Painzelia, and Relicina (Hale, 1967). 

ALIPHATIC L\cIDs.-The identification of fatty 

acids presents many problems and no entirely satisfactory 

techniques have been worked out to deal 

with them. The most direct method is to spray the 

chromatographic plates ivith water and carefully 

watch for opaque, white, water-repellant spots as the 

plate dries. These are the fatty acids. Terpenes have 

a similar reaction but differ in turning bluish, purple, 

or I~roivn when sprayed with H,SO, and heated. 

Dr. AIyles Keogh of the Universitlatl de 10s Andes, 

hlerida, Venefuela, has also informed me (pers. 

comm.) that the fatty acids turn brownish when 

heated for 12 hours or more at 100°C (in contrast 

to the 10 minutes usually needed for development 

of phenolic substances). In any event, protolichesterinic 

acid has been identified in P. expallida 

(Kurokawa, 1968a), and unidentified fatty acids 

are the principle components in P. endolezica and 

P. !let ei.ochloti. 

hPsmEs.-These well-known substances are common 

in section Pamwlina (those species of Parmelina 

lacking terpenes) and occur in most of the 

pantemperate-montane pantropical species. 

Atranorin is the primary cortical substance found 

in all species of Paymelina. 

Gyrophoric acid, a tridepside, occurs in Paimelinn 

cryptochlora, P. dissecta, P. melanochaeta, P. spnthirhta, 

and P. spztrnosu. It is a probable minor component 

in P. tlninnziana, P. horl-escens, P. .schindlei~i, 

ancl P. subfritiscen.s along with the “horrescens” unknown. 

Gyrophoric acid is not easily chromatographed 

because of the tendency for streaking antl 

variable R, values that are highly influenced by 

solvent composition. In addition, one or more associated 

spots, still unidentified, may appear on 

the plates. 

“Horrescens” unknown is actually several substances 

presumed to be depsides related to lecanoric 

acid. The spots react with H,SO, the same as 

orcinol depsides and fall close to gyrophoric and 

lecanoric acids in the usual solvent systems. The 

best separation is achieved in benzene-dioxane 

where a single large spot, the main “horrescens” 

unknown, falls well above gyrophoric acid. This 

unknown has now been detected in P. damaziana, 

P. horrescens, P. schindleri, and P. subfatiscens. I 

have also recently discovered this substance in an 

undescribed species of Hypoti.achyna. 

The widespread medullary component lecanoric 

acid occurs in P. pastillifem, P. pnrinata, P. qiiercina, 

and P. tiliacea. It falls Close to gyrophoric 

acid on chromatographic plates antl must be carefully 

distinguished from it by running controls. 

?’he toluene-acetic acid solvent system should be 

used in addition to the other t\vo systems. 

DEPsmosiis.-?’liese substances occur in many 

lichen genera but are at best s~~oradically developed 

in Paim el ina. Pliy logene t icall y “primitive ” salazinic 

acid and its close acetate galbinic acid predominate 

in section Pai.melina. 

Constictic acid is an accessory substance with 

stictic acid ancl salazinic acid (see below). 

Fumarprotocetraric acid is extremely rare in the 

genus, occurring in only one species, the Australian-Sew 

Zealantl endemic P. jnmesii. It is accompanied 

by protocetraric acid. Succinprotocetraric 

acid is lacking. 

Galbinic acid is confined to a small group of 

very closely related species: P. grilbincr, P. haynchinensi,~, 

P. nic/awvolufa, arid P. ob.scsscr, accompanied 

by secalonic acid X and the ”s~~baui~i~le~ita” terpene 

series. It is easily identified as ;I deep H,SO, orange 

spot below norstictic acid in benzene-tlioxane. 

Lobaric acid is the only orcinol tlepsidone in 

Pni.melinci and occurs as an accessory substance 

with salwinic acid in P. su~inscou~ii. 

Korstictic acid is the main component in two 

closely related Caribbean species, P. antillensis and 

P. phlyctina. A lower spot of the, as yet, unidentified 

connorstictic acid may be present. 

Protocetraric acid occurs only in P. jnmesii with 

fumarprotocetraric acid (see above). 

Salazinic acid, a very common substance in the 

Parmeliaceae, occurs in P. enotmis, P. .rimplicior, 

P. ,\ ZL’ in .sc ow i i, P. 1 I sa ?n have n.c is, P. ve~s i f 07-rn is, ant1 

P. zuallichinna. It is a rare accessory with leucotylin 

in P. el-assata antl may be accompanied by constictic 

9

10 S\IITHSONIAS CONTRIBUTIONS TO BOTANY 

acid in P. metarevoluta. All of these species, ex- 

cepting P. versiformis, are distributed in the Old 

World following a pattern seen in Bulbothrix 

(Hale, 1976b). 

Stictic acid has been discovered in only one spe- 

cies, P. mnelleri, where it is accompanied by con- 

stictic acid and two unknown compounds falling 

between stictic and constictic acids in benezene- 

dioxane. 

TERPENEL-A closely related series of triterpenes 

with a liopane skeleton have been described for 

species in section Myelochroa. Their structures have 

been determined by Yosioka and his group in Ja- 

pan using nuclear magnetic resonance (NhIR) 

spectra, mass spectrometry, etc. The tliin-layer 

chromatography is not yet standardized or well un- 

derstood, and I have not tried to identify individual 

spots. Typical profiles for the terpene-containing 

species of Parmelina are illustrated in Figure 7. 

Leucotylic acid, a colorless compound closely 

related to both zeorin and leucotylin, was first elu- 

cidated by Yosioka (1966). It was isolated from Pa?.- 

melina leucotyliza, although my results indicate that 

leucotylin is, in fact, the main component of this 

species. I suspect that their material included a 

mixture of P. crassata, which does contain leuco- 

tylic acid. The species of Pnrmelina which contain 

leucotylic acid (or at least have the same terpenic 

profile as P. airrulentn, as illustrated in Figure 7) 

are P. aurzilenta, P. degelii, P. il-rugans, and P. 

rhytidodes. 

Leucotylin, the progenitor of leucotylic acid, is 

the main terpene component in P. amagiensis, P. 

crassata, P. denegans, P. entotheiochroa, P. galbina, 

P. hayachinensis, P. indica, P. leucotyliza, P. meta- 

revoluta, P. obsessa, P. perisidians, P. subaurulenta, 

and P. xantholepis. It forms a low major spot in 

benzene-dioxane, but I have generally identified it 

as a profile of several terpene spots as illustrated in 

Figure 7. 

Yosioka (Yosioka and Nakanislii, 1963; Yosioka 

and Nakanishi, 1966) has isolated at least five other 

compounds related to leucotylin in “Parmelia ento- 

theiochroa.” I presume that at least some of these 

make up the numerous spots resolved in hexane- 

ether (see Figure 7), but no attempt has been made 

to identify any of them on the plates. There seems 

to be great variation in the intensity of the spots, 

reflecting different concentrations in the thallus. It 

remains to be seen whether this variation has any 

taxonomic value. 

Zeorin is a well known lichen substance. The 

stereochemistry of its hopane skeleton was only recently 

studied by Yosioka et al. (1968b). It occurs in 

all species in section 1\4pelochroa and has been reported 

in Acroscyplirrs, Cladonin spp., Hppotrachpa 

majoris (Vainio) Hale, Lecanom, Sephroma, 

Peliigera, and the Physciaceae. It may be identified 

as the highest blue spot on the chromatographic 

plates in Iiotli solvent systems (Figure 7). 

Prc~iE.\’Ts.-.\Iedullary pigments are especially 

characteristic of species in section hfyelochroa. The 

chromatography of these is extremely difficult and 

~is~iallp unsuccessful. There is considerable streaking 

and individual components cannot be distinguished. 

Secalonic acid X was first ideniifietl by Yosioka 

et al. (1968a:2090), who established its identity with 

entothein. It also occurs in ergot (Shibata et al., 

1964). Yosioka found secalonic acid A in Parmelina 

nii~rilcntm, P. enfotheiochroa, P. pel-isiclians, and 

P. subauixlenta, and I presume it is the major 

pigment in all species of section ilfpelochroa with 

a yellowish orange medulla, as well as in P. immiscens 

and P. lintlmanii, both lacking terpenes. Unidentified 

reddish pigments accompany secalonic 

acid 4 in the lower medullary layer of P. amagien- 

,pis and P. dcnegans. 

Phytogeography and Speciation 

Pawnelinti occurs primarily on trees in secondary 

forests in lemperate zones and at higher elevations 

in the tropics. A number of the commoner species 

may also occur on rocks. There are, however, only 

four oliligately saxicolous species, P. enownis, P. 

indica, P. obsessa, and P. usambaiensi.r. The Pnr- 

melina floras of various geopolitical units are 

enumerated below. It is altogether obvious that 

many countries are underrepresented because they 

have not been visited by lichen collectors. 

XORTH AMERIC.~ 

United States: P. antillensis, P. aurulenta, P. dissecta, P. 

galbi17n, P. hoi 1 escciis, P. ~ne/ar~~volii/a, P. obsessa, and P. 

sfiu in osa. 

MEXICO ALD C~NTRAL AMERIC~ 

Mexico: P. antillensis, P. aurulenta, P. dissecta, P. hor-

NUMBER 33 11 

rescens, P. ivztniscens, P. Zindnlanii, P. niuelieri, and P. WEIT ISDIE s 

spumosa. 

Guatemala: P. horrescens. Cuba: P. antiliensis, P. dissecta, P. horrescens, and P. 

Panama: P. dissect(/, P. Izorre,ce,ls, and P. subfutiscens. pizlyctinn. 

FIGURE 

7.-Chromatographic profiles of Pnrnielinu species: profiles in the hexane-ether solvent 

system (top) and benzene-dioxaiie solvent system (bottom). (1 1 P. uicndenla, 2 = P. degelii, 

3 = P. irrugans, 4 = P. rhytidot/u.i, 5 = P. ci)nrrgie!i,iis, G = P. ci-nssata, 7 = P. denegnns, 8 = P. 

entotheiochroa, 9 = P. indlica, 10 = P. iettcotyliza, 11 = P. perisidians, 12 = P. subaurulentn, 

13 = P. santholepis, 14 = P. plbi!rcr, 1.5 = P. tnetarevoltrtrr, lG = P. ohsessa, Z = zeorin spot, 

G = galbinic acid, S = salarinic acid, and C = constictic acid: leucotylic acid is presumed to 

be the series of spots in 1 to 4 just below rerorin, and leucotylin is the lowest series in 5-10 

in benzene-dioxane; atranorin falls just above the point where the photographs were trimmed.)

Jamaica: P. antillensis, P. dissecta, P. horrescens, P. 

phlyctina, P. spumosa, and P. subfatiscens. 

Hispaniola: P. antillensis, P. dissecta, P. horrescens, and 

P. phlyctina. 

Puerto Rico: P. phlyctina. 

Lesser Antilles (including Trinidad): P. anlillensis, P. 

cryptochlora, P. dissecta, and P. subfatiscens. 

SOUTH AMERICA 

Colombia: P. lindmanii, P. melanochaeta, and P. spumosa. 

Venezuela: P. antillensis, P. dissecta, P. horrescens, P. 

lindmanii, P. muelleri, and P. spumosa. 

Ecuador: P. pilosa. 

Peru: P. muelleri. 

Brazil: P. antillensis, P. aurulenta, P. consom, P. damaziana, 

P. dissecta, P. heteroloba, P. lindmanii, P. melanochaeta, 

P. muelleri, P. schindleri, P. spumosa, and P. 

wersiformis. 

Uruguay: P. cons or,^, P. horrescens, P. lindmanii, and P. 

pi losa . 

Paraguay: P. consors, P. lindmanii, and P. melanochaeta. 

Argentina: P. consors, P. lindrnanii, P. muelleri, P. pilosa, 

and P. uersiformis. 

Chile: P. horrescens, P. pilosa, P. spumosa, and P. swinscowii. 

EUROPE 

Western Europe: P. dissecta, P. horrescens, P. pastillifera, 

P. quercina, and P. tiliacea. 

Russia: P. aurulenta, P. qziercina, and P. tiliacea. 

Israel: P. tiliacea. 

AFRIC4 

Morocco: P. tiliacea. 

Tunisia. P. tiliacea. 

Ivory Coast: P. usambarensis and P. wallichiana. 

Guinea: P. usambarensis and P. wallichiana. 

Uganda: P. usambarensis and P. wallichiana. 

Kenya: P. dissecta, P. pilosa, P. swinscowii, and P. wallichiana. 

Angola: P. wallichiana. 

Rhodesia: P. wallichiava. 

Zambia: P. enormis. 

Tanzania: P. aurulenta, P. dissecta, P. usambarensis, and 

P. wallichiana. 

Union of South Africa (including Swaziland): P. dissecta, 

P. horrescens, P. pilosa, P. spathulata, P. spumosa, P. subfatiscens, 

and P. wallichiana. 

Madagascar-RPunion: P. aurulenta, P. spumosa, and P. 

wallichiana. 

ASIA AND PACIFIC AREAS 

Pakistan: P. aurulenta, P. quercina, and P. tiliacea. 

India (including Nepal): P. aurulenta, P. cryptochlora, P. 

denegans, P. dissecta, P. expallida, P. horrescens, P. indica, 

P. perisidians, P. rhytidodes, P. simplicior, P. spumosa, P. 

SAIITHSONIAN CONTRIBUTIONS TO BOTANY 

subaurulenta, P. tiliacea, P. wallichiana, and P. xantholepis. 

Sri Lanka: P. denegans, P. dissecta, P. perisidians, and P. 

subaurulenta. 

Thailand: P. expallida, P. perisidians, P. usambarensis, P. 

wallichiana, and P. xantholepis. 

Indochina: P. aurulenta. 

China (including Hong Kong): P. aurulenta, P. irrugans, 

P. quercina, P. subaurulenta, and P. wallichiana. 

Korea: P. entotheiochoroa. 

Japan: P. amagiensis, P. aurulenta, P. crassata, P. dissecta, 

P. entotheiochroa, P. galbiria, P. hayachiizensis, P. horrescens, 

P. irrugans, P. leucotyliza, P. metareuoluta, P. perisidians, P. 

quercina, P. rhytidodes, P. spumosa, and P. wallichiana. 

Taiwan: P. azcrulenta, P. dissecta, P. expallida, P. horrescens, 

P. spirmosa, P. subaurulenta, and P. wallichiana. 

Philippines: P. aurulenta, P. dissecta, P. horrescens, P. 

perisidians, P. wallichiana, and P. xantholepis. 

Malaya: P. wallichiana. 

Indonesia: P. aurulenta, P. dissecta, P. horrescens, P. spu- 

mom, P. wallichiana, and P. xantholepis. 

Sabah: P. denegans, P. leucotyliza, and P. wallichiana. 

New Guinea: P. aurulenta. 

Australia (including New Zealand): P. degelii, P. endoleuca, 

P. horrescens, P. jamesii, P. pruinata, P. qziercina, and 

P. spumosa. 

Hawaiian Islands: P. aurulenta. 

To summarize, t.he New World has a total flora 

of 20 species, 12 of them endemic, 3 (P. pilosa, P. 

subfatiscens, and P. swinscowii) shared with Africa, 

and the remaining 5 (P. aurzilenta, P. dissecta, P. 

horrescens, P. quercina, and P. spumosa) essentially 

pantemperate or montane pantropical. 

Europe has a small Paimelina flora. Dahl and 

Krog (1973) list only P. tiliacea in Scandinavia. The 

checklist by James (1965) for Great Britain includes 

P. dissecta, P. horrescens, P. qziercina, and P. 

tiliacen. For the rest of Europe, Poelt (1969) adds 

only P. caipoirhizans (= P. qziercina) and P. pastilli- 

fera. 

Africa has only two endemic species, P. enormis 

and P. spnthdatn, and nine other species either 

shared with the Americas, Asia, or pantemperate in 

distribution. 

Asia is especially rich in Parmelina with 15 spe- 

cies on the Indian subcontinent and 16 in Japan 

and eastern Asia. Tropical Southeast Asia has nine 

species, none of them endemic and all occurring 

in the higher elevation cloud forests. The genus is 

not represented at all in the lowland dipterocarp 

rain forests from Sri Lanka to Indonesia. 

The Australia-New Zealand region has seven 

species, of which four, P. degelii, P. endoleuca, P. 

jamesii, and P. pruinata, are endemic. No Parme-

NUMBER 33 

FIGURE 8.-Number of species in section Myeloclztm in each major geographical division. 

linae have been collected on the Pacific Islands ex- 

cept for P. cl~irttlenta at higher elevations in Ha- 

waii. The total Old World flora (Africa, Asia, and 

Australia-New Zealantl) comprises 26 species. 

One distrihution pattern of considerable phyto- 

geographic interest stands out. The species of sec- 

tion Myeloch ma, a rather homogeneous group, are 

concentrated in India and eastern Asia (Figure 8) 

and only one of them, P. aurulentn, occurs in all 

geographic areas (Figure 10). The species in section 

Pasmelina, on the other hand, are not strongly 

Concentrated in any region. 

TVithin section Myelochl-oa the species related to 

P. galbina form a particularly interesting group. As 

pointed out by Kurokawa (1972), P. galbina, the 

fertile progenitor species, behaves as an Arcto- 

tertiary plant which migrated southward and has 

survived in Japan and eastern Korth America. A 

rare sorediate morph, P. metawvoluta, has the 

same range, whereas the isidiate morph, P. ob.tessa, 

is restricted to North America, and the pustulate 

morph, P. hayachinensis, to Japan. W. Culberson 

(1972) also regards P. aimilenla as a Tertiary relict. 

Pai.melina has several complex groups of species 

that have evidently resulted from combined chem- 

ical and morphological evolution. The species in 

section iMyelochroa, for example, all of which con- 

tain terpenes and almost always have secalonic 

acid A, a yellow pigment, have evolved from two 

chemically dissimilar ancestors, one, the smaller 

group, with leucotylic acid and resembling P. ir- 

g.iigans, and the other, a larger series with leuco- 

tylin, resembling the present-day P. subazirzilenfa. 

The numerous morplis derived from these presump- 

tive parents or their ancestors, P. nmagiensis, P. 

aiti.itlenta, P. crassata, P. clegelii, P. denegnns, P. 

entotheiochroa, P. leucotyliza, P. perisidians, P. 

Thytidodes, and P. xnntholcpis, have diversified 

mostly by production of soretlia, isidia, pustules, 

and wrinkles, variation in apotliecial diameter, etc. 

AIorph forniation (see Hale, 1975a:13) is not 

strongly delineated for the most part since the par- 

ents cannot be traced directly. \Ve are left to con- 

clude that section Alyelochron is an ancient group 

13


evolved over a long period. The P. galbina group, 

as mentioned above, forms the only exception to 

this rule. 

ILIorph formation has also been an important 

mode of speciation in section Parmelina. The fol- 

lowing parent morph-vegetative morph series are 

indisputable: P. consors-P. pilosa (sorediate), P. 

quercina-P. tiliacea (isidiate), and P. imrniscens-P. 

lindmanii (isidiate). Parmelina phlyctina and 

isidiate P. antillensis are extremely close but less 

clear-cut morphs. A large complex of species have 

been derived from now extinct parents in the P. 

dissecta-P. horrescens group, as illustrated in Figure 

9. Like the P. subaurulenta group of section Mye- 

lochroa, the parent morphs of these numerous 

vegetative morphs are either extinct or have yet to 

be discovered. 

The remaining species in the genus appear to 

have no discernible interrelationships or morphs. 

These include P. endoleuca, P. enormis, P. expal- 

lida, P. heteroloba (part of the P. darnaziana se- 

ries?), P. pruinata, P. sirnplicior, P. swinscowii, P. 

usambarensis, P. veisiformis, and P. wallichiana. 

Classification of Parmelina 

The taxon Lichen section Imbricaria was first 

proposed by Schreber (1791:767). He cited no spe- 

cies, only Squamaria Hoffmann, which is invalid as 

the later homonym of Squamah Ludwig (Phane- 

rogamae). Acharius (1794:250) adopted the name as 

a tribe of Lichen and included in it L. olivaceus L. 

and L. tiliaceus Hoffmann, as well as several other 

species now recognized as belonging to Hypogymnta, 

Parmeliopsis, Physcia, and Xanthoria. XIichaux 

(1803) raised the name to generic rank and cited 

one species, a lichen now called Anzia colpodes 

(Acharius) Stizenberger. Fries (1825:242) later trans- 

ferred it to Parmelia as Parmelia section Imbricaria, 

and Koerber (1855:68) used it at the generic rank, 

even though he realized that the name was a later 

homonym of Imbricaria Decandolle (Phaneroga- 

mae). Koerber and contemporary lichenologists 

placed many species that we now consider parme- 

lioid in Imbricaria, while at the same time using 

Parmelia for many species now placed in Physcia. 

After the delimitation of Parrnelia in a modern 

sense by authors such as Sluller and Nylander, 

Imbricaria was relegated to synonymy under it. 

Hale and Kurokawa (1964: 130) grouped the narrow- 

lobed, marginally ciliate Parrneliae in Parmelia 

subgenus Parmelia section Imbricaria (Schreber) E. 

Fries, and it is essentially this group that I have seg- 

regated as a distinct genus, Parmelina (Hale, 1974: 

482). 

Parmelina is a rather heterogeneous assemblage of 

47 species differing in lobe width, production of 

cilia, pigmentation, and rhizine branching. Two 

major groups can be recognized, one including 

those species lacking terpenes and one with terpene- 

containing species. They may be conveniently re- 

garded as sections. 

Section Parmelina 

Parmelia section Hypotrachyna subsection Myelolezica Asa- 

hina, 1952:74 [type-species: Purrnelina tiliacea (Hoffmann) 

Hale]. 

The species in this section are characterized by 

the lack of any triterpenes. Except for two species, 

P. immiscens and P. lindmanii, the medulla is white 

and unpigmented. Several internally homogeneous 

groups can be recognized. For example, the P. 

dissecta-P. horrescens group (Figure 9) includes 

seven species with very similar lobe configuration, 

abundant marginal cilia, and gyrophoric acid or 

the closely related “horrescens” unknown. Heavily 

white-niaculate P. consow, P. pilosa, and P. rnuelleri, 

having coarse, leathery thalli and thick, furcate 

marginal cilia, are obviously related. 

The predominantly European complex of spe- 

cies, P. qztcrcina-P. iiliacea-P. pastillifera, contains 

lecanoric acid and forms an easily recognized but 

isolated group. Parmelina antillensis and P. phlyc- 

tina, both unusual in containing norstictic acid, 

have no close relatives in the genus. Two other 

New World species, P. immiscens and P. lindmanii 

with a yellow medulla, form an isolated branch of 

this section. 

The species with salazinic acid, P. enormis, P. 

simplicior, P. swinscowii, P. usambarensis, P. versi- 

formis, and P. wallichiana, have little in common 

except their chemistry. Finally, the remaining spe- 

cies, P. endoleuca, P. expallida, P. heteioloba, P. 

jamesii, and P. pririnata, have no obvious common 

ancestry or affinities with other species in the sec- 

tion.

NUMBER 33 15 

Progenitor 

I- 

Progenitor Progenitor 

(gyrophoric acid) (“horrescens” unkn.) 

P. horrescens IP. schindleri P. sirbfutiscens 

Progenitor ,(white Progenitor rnedullaj (isidiate) (lobulate) (pustulate) 

(yellow medulla) 

I P. spir ntosu 

(pustulate) 

r-l-l 

P. dissei.tu P. sputhulatu P. cryptochloru 

(isidiate) (lobulate-isidiate) (sorediate) 

FIGURE 9.-Hypothetical derivation of species in the Parmelina dissecta-P. horrescens group. 

hrot shoivn are P. damaziana, a presumptile nonisidiate, nonsorediate derivative of the “hor- 

rescens” progenitor, and P. melanochaetn, a possible relative of P. dissecta but derived from a 

different progenitor. 

Section Myelochroa (Asahina) Hale, new status 

Parmelia section Hypotrachyna subsection Myelochroa Asa- 

hina, 1952374 [type-species: Parmelina aurdenta (Tucker- 

man) Hale]. 

All of the species in this section contain zeorin 

and either leucotylin or leucotylic acid and asso- 

ciated terpenes, and all, excepting P. indica, produce 

varying amounts of the yellow-orange pigment 

secalonic acid A and possibly other, as yet, unidenti- 

fied pigments. This group is highly restricted to 

Asia, and the 17 species have very close affinities, in 

sharp contrast to section Parmelina. 

Two apparently unrelated groups are recognira- 

ble in the section. One, the P. galbina group (P. gal- 

bina, P. hayachinensis, P. metarevoluta, and P. ob- 

sessa), is characterized by galbinic acid in addition 

to leucotylin and secalonic acid A. The other group 

includes two closely related subgroups, the P. auru- 

lenta subgroup (P. aul-ulenta, P. degelii, P. irru- 

guns, and P. rhytidodes) with leucotylic acid, and 

the P. subazirulenta subgroup (P. amagiensis, P. 

crassata, P. denegans, P. entotheiochroa, P. leuco- 

tyliza, P. perisidzans, P. subauiulenta, and P. xan- 

tholepis) with leucotylin. Parmelina indica can 

probably be classified in the P. szi baurulenta sub- 

group although it has a small saxicolous thallus 

and lacks secalonic acid A. 

The affinities of Parmelina lie primarily with 

Parmotrema Alassalongo, a broader lobed, loosely 

attached, often ciliate genus (Hale, 1965). Palme- 

Zina, however, has a far less evolved chemistry, nar- 

rower adnate lobes, consistently small spores, and 

adnate or sessile, usually imperforate apothecia. 

Where soredia are produced, they are lamina1 

rather than marginal in contrast to most Parmo- 

trema species. All in all, however, the apparent 

intergradation between broad-lobed Parmelinae and 

the smaller lobed species of Parmot?-emu, involving 

perhaps six or eight species, poses difficult problems 

in the correct generic identification of individual 

specimens and for the “lumper,” at least, raises 

questions on the distinctness of the two genera as 

circumscribed here. I do not expect that the prob- 

lem can be resolved to the satisfaction of all lichen- 

ologists, but the basic differences should become 

clearer as more collections are made in tropical


countries and our knowledge of soredial formation, species lacking isidia, soredia, and pustules; (2) 

significance of chemical variation, etc., broadens. those with isidia, lobulate isidia, or lobules; and 

The key below is divided into three sections: (1) (3) those with soredia or pustules. 

Key to the Species of Parmelina 

1. THALLUS LACKING ISIDIA, SOREDIA, AND PUSTULES 

(Cortex flaking off in Parmelina entothetochroa and P. phlyctina, and heavily rugose in P. 

crassata and P. rlzytidodes) 

1. Medulla more or less completely yellow to yellow-orange (pigmented mostly below the 

apothecia in P. galbina). 

2. Upper surface hea\ily rugose (Figure 6b) 35. P. rhytidodes 

2. Upper surface more or less plane. 

3. Upper cortex vei) fragile, flaking awa) over extensive areas (Figure 6a) . 

4. Medulla deep yellow 13. P. entotheiochroa 

4. Medulla white (turning reddish if impropeily curated) 3 1, P. phylctina 

3. Upper cortex entire and continuous. 

5. Lower half of medulla darker reddish orange 1. P. amagiensis 

5. Loner half of medulla not darkly pigmented. 

6. Lobes narrow and sublinear, 1-2 mm wide 15. P. galbina 

6. Lobes broader, subirregular, 2-4 mm wide. 

7. Collected in Mexico 

19. P. immiscens 

7. Collected in Asia. 

8. Apothecia 3 mm or less in diameter 

40. P. subaurulenta 

8. Apothecia 3-10 mm in diameter. 

9. Thallus coarse, about 250 pm thick: leucotylin present 5. P. crassoto 

9. Thallus tnembranous, about 150 pm thick: IeuLotylic acid present 

21. P. irrugans 

1. Medulla entirely white. 

10. Lower surface pale brown. 

11. Collected on rocks in Africa 12. P. enormis 

11. Collected in treer in South America 45. P. versiformis 

10. Lone1 surface black. 

12. Medulla K+ yellow turning red. 

13. Lobes sublinear, 1-2 mm wide 15. P. galbina 

13. Lobes broader, 2-5 mm wide. 

14. Upper cortex fragile, flaking off 

31. P. phlyctina 

14. Upper cortex firm, continuous. 

15. Collected in India 37. P. simplicior 

15. Collected in South America 45. P. versiformis 

12. Medulla K negatike. 

16. Medulla Cf red. 

17. Cortex distinctly white maculate 

34. P. quercina 

17. Cortex emaculate 33. P. pruinnta 

16. Medulla C-. 

18. Thallus coriaceous, heavily whlte-maculate; lobes 2-5 mm jvide 

4. P. consors 

18. Thallus thin, emaculate: lobes 1-3 mm i\ide. 

19. Medulla KCf rose; “horrescens” unknor\n present 7. P. damaziana 

19. Medulla KC-: fatty acids present. 

20. Collected in Australia 

11. P. endoleuca 

20. Collected in South America 

17. P. heteroloba 

11. THALLUS ISIDIATE, LOBULATE-ISIDIATE, OR LOBULATE 

I. Medulla yellow to pale orange-yellow.

NUMBER 33 

2. Thallu\ densely lobulate uithout formation of isidial initials (Figure 4f) 

47. P. xantholepis 

2. Thallus noimally isidiate or lobulate-isidlate. 

3. Isidia large, doisiientral and lobulate 

8. P. degelii 

3. Isidia normal, cylindrical and erect. 

4. Lobes subirregular and apicall) rotund, 3-6 mm wide 24. P. lindmanii 

4. Lobes sublinear, 1-2 mni Ttide. 

5. Plants saxicolous (rarel) coi ticolous) 111 eastetn Sorth America 28. P. obsessa 

5. Plant5 corticolous in Southeast A5ia 30. P. perisidians 

1. Medulla M hite. 

6. Thallus densel) lobulate I\ ithout isidial initlals (Fignre 4e) 

6. Thallus isidiate 01 lobulate-isidiate. 

7. lsidia in part ptocumbent and lobulate (Fignre 4c,rl). 

36. P. schindleri 

8. Medulla C+ rose 38. P. qpathulata 

8. Medulla C- 

18. P. hori escens 

7. lsitlia erect, cylintliical. 

9. Medulla Kf jellou turning red. 

10. Collected in tiopical Ameiica, norstictic acid piesent 

10. Collected in Asia and Aflica; salazinic acid present. 

2. P. antillensis 

11. Lobes subirregulal, cioitded, apicall) iotund 

46. P. urallichiana 

1 I . Lobes bublineai, looselp dii aricate 44. P. usambarensis 

9. Medulla I(- or faintly )elloit. 

12. Medulla Cf rose 01 led. 

13. Isidia apically flattened and peltate 

13. Isidia cylindiical, not peltate. 

29. P. pastillifera 

14. Medulla C+ intense red (lecanoric acid) 

14. Medulla C+ lose (g)iophoiic acid). 

43. P. tiliacea 

15. Lobes narrow, 1-2 mni nide; upper surface emaculate: isidia 

eciliate 10. P. dissecta 

15. Lobes broader, 2 4 mm itide; upper surface maculate; isidia 

ciliate 25. P. melanochaeta 

12 4iedulln C-. 

16. Medulla P+ red 

16. Medulla P-. 

22. P. jamesii 

17. Louei surface bronn to hlacLening in palt 

17. Lower suiface unifoimly black. 

14. P. expallida 

18. Isidia apically ciliate; medulla KC+ rose 18, P. hort escens 

18. Isidia eciliate; medulla KC- 20. P. indico 

111. THALLUS SOREDIATE, SOREDIATE-PUSTULATE, OR PU5TULATE 

1. Medulla entirely white. 

2. Lobes broad and apicall) rotund; coi tex con


10. Pustules forming in capitate masscs (Figuie 5g) 23. P. Zeucotyliza 

10. Pustules irregular, intermixed with wrinkles (Figure 6b) 35. P. rhytidodes 

9. Lobes narrow and sublinear, less than 2 mm wide. 

11. Medulla K+ reddish, P+ orange 16. P. hayachinensis 

11. Medulla K-, P-, C+ rose, pigmented bery faint yellow 39. P. spumosa 

8. Thallus sorediate or pustulate-sorediate at maturit). 

12. Medulla deeper reddish orange In the lower half 9. P. denegans 

12. Medulla uniforiiil) pigmented to nhite and pigmented only under the soralia. 

13. Lobes 2-4 mm wide; medulla P- 3. P. aurulenta 

13. Lobes less than 2 mm wide; medulla P+ orange 26. P. metarevoluta 

Species Treatment 

The 47 species of Parmelina are arranged below 

in alphabetical order. Location of specimens listed 

under “Specimens Examined” are indicated by the 

standard herbarium acronyms except for those col- 

lected by Hale, all of which are preserved in the 

Smithsonian Institution (US). 

1. Parmelina amagiensis, new combination 

FIGURE Ila 

Parnzelia arriagiensis Asahina, 1951a:228 [type collection: Mt. 

Amagi, Prov. Izu, Japan, Asahina 95 (TNS, lectotype)]. 

DEScRIPTIoN.-Thallus loosely adnate on bark or 

mosses over bark, buff mineral gray, 4-8 cm broad; 

lobes more or less subirregular, imbricate, 2-3 mm 

wide, the marginal cilia numerous, simple, up to 

1 mm long; upper surface plane, white-maculate, 

becoming heavily pycnidiate, isidia and soredia 

lacking; medulla deep salmon orange; lower sur- 

face black, densely rhizinate, the rhizines black, 

simple or sparsely squarrosely branched, 0.5-1 .O mm 

long. Apothecia sessile to subpedicillate, 2-4 mm 

in diameter; spores 8, 6-7 X 9-1 1 pm. 

CHEMISTRY.-cOrteX K + yellow (atranorin); rne- 

dulla more intensively yellow with color tests 

(zeorin, leucotylin and related “subaurulenta” ter- 

penes, secalonic acid A, and unidentified pigments). 

DIsTRIBuTIoN.-Japan. 

REMARKS.-ThiS rare Japanese species is very 

close to P. denegans, a pustulate or sorediate- 

pustulate species in tropical Asia. Both have a 

more deeply pigmented medulla, especially in the 

lower half, than other members of section Myelo- 

chroa. I have not been able to resolve these pig 

ments with the usual chromatographic solvent 

systems. Parmelina amagiensis appears to be some- 

what more robust than P. denegans, judging from 

the few specimens available, and does not seem to 

be a parent morph for the latter. 

SPPCIMENS Ex \MIsFD.-Japan: Prov. Ettchu, Yanagisawa 

599a (TNS); Prov. Hyuga, Kurokawa 50058 (TNS, US). 

2. Parmelina antillensis 

FIGURE 11 b 

Parmelirin antillensis (Nylander) Hale, 1974:482. 

Parmelia antillensis Nylander, 1868:264 [type collection: 

Matouba, Guadeloupe, Husnot 445 (H. Nylander herbar- 

ium number 35119, lectotype; isolectotypes in G, P)]. 

Parinelia blastica ’I’ainio, 1896:32 [type collection: Shawford 

Estate, Dominica, Elliott 899 (TUR, lectotype; isolectotype 

in BM)]. 

DEscRIPTIoN.-Thallus adnate on bark, fragile 

and membranous, up to 10 cm across, greenish min- 

eral gray, often more or less white-pruinose at the 

lobe tips; lobes irregularly branched, subimbricate 

to convoluted, the marginal cilia sparse, mostly in 

lobe axils, to 0.5 mm long; upper surface shiny, 

plane, cortex continuous or cracked with age; 

densely isidiate, the isidia fine, to 0.4 mm high, 

sparingly branched or simple; lower surface black, 

brown and shiny in a narrow zone along the mar- 

gins, densely rhizinate, the rhizines simple. Apo- 

thecia adnate, 3-5 mm in diameter, the amphi- 

thecium isidiate; spores 8, 5-6 X 7-8 pm. 

CHEMISTRY.-cOrteX K + yellow (atranorin); rne- 

dulla K+ yellow turning red, C-, KC-, P+ 

orange (norstictic and connorstictic acids). 

DIsTRIsuTIoN.-Caribbean region. 

REMARKs.-Parmelia antillensis and its close Ca- 

ribbean relative P. phlyctina are the only two spe- 

cies in the genus containing norstictic acid. This 

trait, along with the rather broad, apically rotund 

lobes, might tempt one to place them in Parmo- 

trema Massalongo, where, in fact, I had originally 

suggested they might belong (Hale, 1959). The

NUMBER 33 

lower surface, however, has at most a narrow bare 

or papillate zone, the lobes are generally adnate at 

the tips, not ascending, and the cilia are mostly in 

the axils of the lobes. Both P. antillensis and P. 

phlyctinn share traits intermediate between Parme- 

Zina and Parmotrema and have evolved in a very 

restricted region, the rain forests of the Caribbean 

basin (excepting the unusual record of P. until- 

1en.yi.r. from the southeastern United States discussed 

in Hale, 1971b:46). Other parmelioid groups have 

evolved here, eg., Pseudoparmelia martinicnnn 

(Nylander) Hale and P. raunkiaeri (Vainio) Hale 

and perhaps also Relicinn eximbricnta (Gyelnik) 

Hale (1975b, 1976a), but these are all lowland spe- 

cies in dry, disturbed habitats. Although Parmelina 

ontillensis is often collected on citrus trees at 600- 

700 m elevation, its primary habitat is the canopy 

branches of trees in virgin rain forest. 

SrECI\rEss Ex4\rrXED.-L?nited States: Tennessee, Yoshi- 

viiira et al. 660872 (US). Mexico: Chiapas, Hale 20238.’ 

Cuba: Oriente, Iinsliairg 250.51 (MSC). Jamaica: Cztlberson 

13648, 13777, 13787 (DUKE), Znishaug 14088, 14627, 14943, 

1.5321 (MSC), 14593, 15295 (MSC, US), Orcult 3551b, 5626b 

(US), Plitt (US). Haiti: Oueit, Inishaug 22642 (MSC). Puerto 

Rico: Stinisoti 1440 (DUKE). Guadeloupe: Culberson 14511 

(DUKE), Oitestal 1916 (US). Martinique: Citlberson 14682 

(DUKE), Uegeliuc (Degelius herbarium). Dominica: Irnshaug 

32726 (MSC), Hale in Lichenes Selecti Exsiccati 942 (H, LD, 

US) (for additional records see Hale 1971a:lO). St. Lucia: 

Inishaug 29712 (MSC). St. Vincent: Znishaug 30504 (MSC). 

Grenada: Hale 38298, Inishaug 16092, 16335 (MSC). Trini- 

dad: Hale 38291, Inisliaug 32042 (MSC). Barbados: Elliott 

(TUR). \’enezuela: MC-rida, Hale 42751, 42941. Brazil: Minas 

Gerais, T’ninio in Lichenes Brasilienses Exsiccati 675 (TUR) 

(di7rributed as Parnielia a~tiazonica). 

3. Parmelina aurulenta 

Parmelina aurulenta (‘Tuckerman) Hale, 1974:482. 

Parmelia auruletzta Tuckerman, 18583424 [type collection: 

Harpers Ferry, Virginia, Tuckernia?i (FH-Tuck, lectotype)]. 

Parmelia tiliacea var. eporescens Miiller Argoviensis, 

1887:316 [type collection: Siberia, Russia, Lahm 5 and 6 

(G, lectotype: isolectotype in W)]. 

Pnrnzelia albido-strarninea Hue, 1899:161 [type collection: 

Sanctum Dionysium, Reunion, Rodriguez (PC, lectotype)]. 

Parnielia subrevoluta Harmand, 1928:326 [type collection: 

Cha Pon, Indochina, Demnnge (PC, lectotype)]. 

Pornielia hitnunensis Zahlbruckner, 1930: 187 [type collection: 

’ Entries without herharium designations of specimens 

collected by Hale are in US. 

Tschangscha, Hunan, China, Hnndel-Mazzetti 11454 (WU, 

lectotyie)]. 

Parnielia sihestris Degelius, 1940:47 [type collection: Togue 

Pond near Mt. Katahdin, Maine, Degelius (Degelius her- 

barium [not seen]: isolectotype in US)]. 

Pnrmelia aurulenta var. silrwstris (Degelius) Degelius, 1941 :58. 

DEscRIPrIoN.-Thall~is adnate on hark or rock, 

pale greenish mineral gray, 4-10 cni broad; lobes 

sublinear to subirregular, apically subrotund, 2-4 

mni wide, the marginal cilia irregularly dispersed 

but mostly in the lobe axils, to 0.8 mni long; upper 

surface shiny, plane to rugulose, soon sorediate to 

i)iistLilate-sorecliate, the soralia up to 1 mm in diameter, 

coalescing into large subcapitate clumps, the 

soretlia coarse (Figure 5a); medulla white and becoming 

orange sulfur yellow only beneath soralia 

antl near exposed cracks in the upper cortex or 

entirely sulfur yellow; lower surface black and 

moderately to densely rhizinate, the rhizines black, 

simple or becoming sparsely f urcate or squarrosely 

branched. Apothecia rare, adnate to substipitate, 

the amphithecium sorediate-pListulate, 2-5 mm in 

diameter; spores poorly developecl, 7 x 12 pm. 

CmxmmY.-Cortex K + yellow (atranorin); medulla 

negative with color reagents or if pigmented 

areas tested becoming more intensely yellow (zeorin, 

leucotylic acid and related terpenes, antl secalonic 

acid A). 

DIsTRIsvi-Io~.-Panteni~)er~~e antl montane pantropical 

(Figure 10). 

REMARKs.-Parmelina auritlenta is one of the most 

widespread species in the genus and, at least in the 

United States, is one of the most commonly collected 

foliose lichens. It is also unusual in being the 

only species of the subaurulenta complex that occurs 

in the New World, perhaps suggesting merely 

that efficient dispersal by soredia is the prime reason 

for the pantemperate distribution. 

Soredial formation is extremely variable. The 

lectotype, for example, is densely pustulate with 

well developed soredia that obscure most of the 

pustular initials (Figure 5a). At other times, the 

pustules become only sparsely and coarsely sorediate, 

this being especially true of saxicolous forms. 

In no case, however, do the pustules remain intact 

as in P. leucotyliza, a Japanese species. In Japan the 

two species may also be separated by chemistry 

since P. leztcotyliza produces the “subaurulenta” 

terpenes. There is a very minor amount of variability 

in chromatograms of P. aul-zdenta from different 

19


FIGURE 10.-Distribution of Parmefina aurzrlenta hased on all available herharium specimens. 

geographic areas but it does not seem significant or 

consistent. I did not, incidentally, recheck the 

terpene profiles of Pawnelia tiliacea var. efiol-esrens, 

P. albido-strarninea, P. subrevoluta, or P. hunanen- 

sis, but assumed that they have the “aurulenta” ter- 

penes since they are sorediate and fall outside the 

range of P. leucotyliza. Should any of them be 

found to contain “subaurulenta” terpenes, the no- 

menclature of P. aurulenta would not change. 

The parent morph of Pal-melina awulenta might 

well be P. irl-ugans, which is identical in chemistry. 

It would be desirable, however, to examine more 

specimens of P. irrugans in relation to P. aurulenta 

in Japan before making a final decision. 

The ecology of P. aurulenta has been intensively 

studied in Wisconsin, an area of typical deciduous 

and conifer forests well within its range. For ex- 

ample, I found in 1955 that it occurred on 30y0 of 

a 2800 tree sample at the 1.3 m level on the trunk 

and on 21y0 at base level in mesic deciduous forests. 

It was most common in closed oak-hickory stands 

and much rarer in both open, savanna-like oak 

stands and in heavily shaded maple (Acer saccha- 

l-urn) climax forests. Parmelina aul-ulenta occurs 

frequently with Parmelia ruderta Acharius and 

Pseudopal-rnelia capel-ata (L.) Hale. 

In northern Wisconsin, an area of hardwood- 

conifer forests more comparable in climate to 

northern Europe, P. aurulenta had an overall fre- 

quency of only Il.670 at the 1.3 m level and 1.3% 

at the base. It avoided Pinus spp. and grew almost 

exclusively on Acel- and Querczis (W. Culberson, 

1955). This avoidance of Pinus was demonstrated 

even more graphically by W. Culberson (1958) in 

an ecological survey exclusively of Pinus in North 

Carolina. Although otherwise common in this state, 

not a single collection was made on conifers. 

SPECI~IESS 

Ex.\xIINED.-United States and Canada: See Hale 

(1958384) for a map of distribution based on specimens in 

various American herbaria. Mexico: Michoacin, Wirth 329 

(US). Brazil: Minas Gerais, Eiten 6932 (US). Europe: Russia, 

Vasiljeva 9 (US). Tanzania: Tanga Province, Santesson 23358, 

23359 (Santesson herliarium, US). Union of South Africa: 

Satal, Alniboria 8071, 8079 (LD); Transvaal, Alrnborn 7225 

(LD). Madagascar: des .4 bboyes (TUR), Lernaitre (H). Paki-

NUMBER 33 

FIGURE 11.-Species of Parmelina: a, P. nmagiensis (Kurokawa 550058 in US); b, P. antillensis 

(Hale 38291); c, P. aurulentn (Hale 18941); d, P. consors (Reit- and Klein 15973 in US); e, P. 

crassutn (Kurofmwu 550446 in US); 1, P. cryfitochlora (Hale 37782). (Scale in mm.) 

21

22 SMITHSOSIAN CONTRIBUTIONS TO BOTANY 

Stan: Ahmad 211 (L), Iqbnl 831, 839, 843 (US). Nepal: Noikett 

9368 (BM). India: Almora District, Arvasthi 3970 (Aivasthi 

herbarium, US); Kangra District, Hoeg 1492, 1497 (Awasthi 

herbarium); Punjab: Schelpe 3260 (BM); Tamil Natlu: Hale 

43664, 43812, 43816; Uttar Pradesh, Awasthi 3374 (Awasthi 

herbarium); !Vest Bengal, Togashi (TNS). Sri Lanka: Degel- 

ius As407 (Degelius herbarium), Thwaites 32 (BM). China: 

Kiangsi Province, Tai (BM). Taiwan: Kurokawa 1359, 1390, 

2694 (TNS). Korea: Den (TNS). Hong Kong: without collec- 

tor, 1897 (BM, US); Hale 46393. Japan: Prov. Aki, Yan- 

agisatca 746-1 (TNS); Prov. Awa, Inobe 23 (TNS); Prov. 

Higo, Mayebnra 169 (TNS) Togashi in Lichenes Japoniae 

Exsiccati 177 (US); Prov. Hizen, Kurokawa 6261.5 (TNS): 

Prov. Kii, Kurokawa 59129 (TNS, US), Sasnki 481 (TNS); 

Prov. Settsu, Hale 29417, 29428; Prov. Shimotsuke, Kiirokaroa 

56509 (TNS). Philippines: Benguet, Degeliiis As-888 (Degelius 

herbarium). Indonesia: Java, Groenhart 5872 (L, US). New 

Guinea: IMorobe District, Weber and McVean L-51280 

(COLO, US). Hawaiian Islands: Maui, Faurie 514 (LD), 866 

(BM, PC), Hnle 31543; Kauai, Hale 31747, 31755. 

4. Parmelina consors 

FIGUKE lld 

Pnrmelinci consors (N) lander) Hale, 1974:482. 

Pnrnielia co?isors Sylander, 1885:613 [type collection: Minas 

Gerais, Brazil, Weddell (H, Nylander herbarium number 

35277, lectotype)]. 

Parmelia balansae Miiller Argoviensis, 1888a: 1 [type collection: 

AsunciOn, Paraguay, Balansn 8 (G, lectotype)]. 

Parmelin sampaiana Hue, 1899: 170 [type collection: S%o 

Paulo, Brazil, Sampaio (PC, lectotype)]. 

Parmelia continentalis Lynge, 1914:lll [type collection: 

Corumba, Mato Grosso, Brazil, hfalme 48 (S, lectotype)]. 

DEsCRIPTIoN.-Thallus adnate on bark, coria- 

ceous greenish mineral gray and turning deep olive- 

buff in the herbarium, 5-10 cm in diameter; lobes 

irregularly branched, sublinear-elongate to subir- 

regular, apically rotund, often imbricate, 1-4 mm 

wide, the margins crenate, moderately to densely 

ciliate, the cilia stout, blackish brown to black, 

becoming furcate, 0.2-1 .O mm long; upper surface 

shiny, becoming heavily maculate, sometimes 

pruinose towards the tips, isidia and soredia 

lacking; medulla white; lower surface black, mod- 

erately rhizinate, the rhizines black, thick and thin 

intermixed, simple or squarrose. Apothecia substi- 

pitate, 2-8 mm in diameter, amphithecium smooth, 

the disc cinnamon brown, rarely perforate; spores 

8,S-lZ X 14-19 pm. 

CHEMISTRY.- Cortex K+ yellow (atranorin); 

medulla negative with color tests (no substances 

demonstrated). 

DIsTRIBuTroN.-South America. 

REhfARKs.-This robust, coriaceous lichen is char- 

acterized by its heavily white-maculate cortex (Fig- 

ure 3a), thick, furcate cilia (Figure 3a), and lack of 

medullary chemistry. It is also the only species as- 

signed to Parmelina with perforate apothecia, a 

trait usually associated with Parmotrema hfassa- 

longo. The dense rhizine mat below to the margin 

and the generally close adnation, however, do not 

conform to Parmotrema. Paymelina consors is evi- 

dently a common lichen in dry, scrubby forests and 

on trees along roads. The presumptive sorediate 

morph, P. pilosa, has a broader distribution in 

South America and also occurs in Africa. 

SPECIMENS Ex4MIsEo.-Brazil: WawrU 669 (M); Mato Groqso, 

Monte\ 10146 (DUKE, LD, US), Minaq Gerais, hfalme 201B 

(S), I’ainio in Lichenes Brasilienses Exsiccati 398 (BM, M, 

TUR); Rio de Janeiro, Burchell 2303 (BM), Gardner 37 

(BM), Glnziou 1821, 1836 (M), 1840 (M, S), Hemmendorff 

5386 (S, UPS), Milne (BM); Rio Grande do Sul, Malme 1282 

(LD, S, UC, UPS, US); Santa Catarina, Reitz and Klein 

12901, 15973, 16032a (US): SZo Paulo, Eiteti 5731 (US), Gehlt 

(US), 5924 (MICH), Puiggari (G). Paraguay: Itapua, Malme 

1462 (S, UPS, US). Uiuguay: Latellaja, Lamb (H). Argentina: 

Buenos Aires, Kuhnemann 22 (S), Santesson 76, 77a (S); 

Misiones, Montes 36 (US), 3353 (WIS). 

5. Parmelina crassata, new species 

FIGURE lle 

DEscRIPTIoN.-Thallus laxe adnatus, corticola, 

crassatus coriaceusque, cinereo-albidus vel obscurascens, 

usque ad 15 cm latus, lobis sublinearibus 

vel subirregularibus, contiguis vel congestis, 3-6 

mm latis, margine ciliatis, ciliis irregulariter dispersis, 

nigris, 0.3-0.8 mm longis, superne planus vel 

aetate rugulosus, nitidus, continuus, isidiis sorediisque 

destitutus, cortex superior ca. 12 pm crassus, 

epicorticatus, stratum gonidale 16-18 pm crassum, 

medulla omnino sulphureo-salmonea, 160-2 10 pm 

crassa, cortex inferior fuscus, 14-16 pm crassus, 

subtus niger, dense rhizinosus, rhizinis nigris, nitidis, 

1-2 mm longis, simplicibus vel pro parte 

furcatis vel squarroso-ramosis. Apothecia numerosa, 

subpedicillata, ainphithecio rugoso, usque ad 10 

mm diamentro, sporis octonis, 6-8 X 14-15 pm. 

CHEnmTR\r.-Cortex K + yellow (atranorin); 

medulla more intensively yellow with color tests 

(zeorin, leucotylin and associated terpenes, and 

secalonic acid A).

NUMBER 33 

HoLOTYPE.--;\ft. Akagi, Japan, S. Kzirokazun 

550466, 23 August 1955 (US; isotype in TNS). 

DIsTRrBLJTr0N.-Japan. 

REivARKs.-This is another member of the P. 

sitbaic~iilenfa group so abundantly developed in 

Japan. It has the same chemistry as P. subazirulenta 

but is well separated by the larger, leathery thallus, 

averaging 218 pm in thickness (sample of 10 speci- 

mens), as opposed to 150 pm for P. subaurzilenta. 

It also has significantly larger, substipitate apothe- 

cia. Parmelia in-ugans, which has large apothecia, 

can be distinguished by chemistry (the “aurulenta” 

terpenes) and thinner thallus. 

This species previously had been identified 

(along with P. irrugans) as “Parmelia homogenes” 

in Japan, but, as explained below, P. homogenes 

was misinterpreted and is in reality the same as 

P. .siibnicrulentn, Puvmelia cmssatn is a common 

species in Japan, but since I have not had an 

opportunity to re-examine tlie chemistry of the rich 

collections in TNS, I have cited below only those 

specimens also represented in US. 

SIV CIMFX Ex \~is~o.--Japaii: Prov. Kozuke, Kurokarc~a 

.58583 (TNS, US) ; Prov. Sagami, Kurokarca 58039 (TNS, US) : 

I’rov. Suruga, ‘4saliina 539, 2G27 (US). 

6. Parmelina cryptochlora 

FIGURE llf 

PorrridiTia crjptochloro (Vainio) Hale, 19743482. 

Pnrrnelin crjptoclilorn 1-ainio, 1896:34 [type collection: Lau- 

dat, Dominica. Elliott 912 (BM, lectot!pe)]. 

DEscRIPTIos.-Thallus closely adnate on bark, 

whitish mineral gray, 1-3 cm broad; lobes sublinear, 

separate to crowded, 1-2 mm wide, the marginal 

cilia sparsely developed, 0.1-0.3 mm long; 

upper surface plane to convex, sorediate toward the 

tips, tlie soralia capitate, up to 1 mm in diameter 

(Figure 5b), the sorediate lobe tips becoming 

revolute: lower surface black, sparsely rhizinate, tlie 

rliizines black, simple. Apotliecia not seen. 

CHEhIISTRY.-COrteX K + yellow (atranorin): 

medulla K-, C+ rose, KC+ red, P- (gyroplioric 

acid). 

DISTRIBUTIOS.-~YeSt Indies and India (?). 

REMARKS.--AS I explained in my study of the 

Pawneline of Dominica (Hale, 1971a: 12), Vainio’s 

type collection is so poor that I had not been able 

to interpret it until I collected additional speci- 

mens from the type-locality in Dominica. It is 

rather common at one locality (base of Morne 

Anglais) on trees in a pasture at 600-800 m elevation, 

One collection that I made in South India 

also appears to lie this species. It is so small and 

inconspicuous that it has undoubtedly been overlooked 

elsewhere or, iC collected, has been set aside 

as unidentifiable. It has the same chemistry as 

P. dissecta, a common species with which it is 

clearly allied as the sorediate morph of the extinct 

or still undiscovered parent morpli. It also has the 

same chemistry as sorediate Hypotl-achynn l-evoliita 

(Floerke) Hale, which woulcl normally be amply 

distinguished by a larger tliallus, more ascending 

lobes, dichotomously branched rhizines, and 

broader soretliate pustules. A poorly developed 

specimen of H. iavolicta, however, and a large specimen 

of P. c~pfochlol-a would have to be separated 

with extreme care. 

QI’ECIMFNS ExA\rINl.o.-~ominic;i: Hale 3535i, 37782. India: 

Tamil Nadu, Hale 40203. 

7. Parmelina damaziana, new combination 

FIGURE 120 

Partrielin danzaziatia Zahlbruckner, 1905:.541 [type collection: 

Mt. Ituculumi, Brazil, Daniazio 137j (W, lectotype; isolectotype 

in G)]. 

Partnelin brachpconidia Zahlbruckner, 1908:465 [type collection: 

\‘ellozo, Serra (lo Our0 Preto, Braiil, Daniazio 1741 

(LV, lectot!pe)]. 

Pnririelin brnchyrovidin var. cliloroccirpn Zahlbruckner, 

1908:466 [type collection: Cachoeira do Campo, Brazil, 

Dnmnzio I740 (W, lectotype)]. 

Pnrtnelici crystallorum Lynge, 1914: 128 [t\pe collection: Corcovado, 

Rio de ,janeiro, Brazil, Mnlme 59* (S, lectotype)]. 

D~sc~~~l~o~.-Thallub 

closely adnate on twigs, 

whitish ashy mineral gray, 3-6 cm broad; lobes 

short, sublinear-elongate to subirregular, 1-3 mm 

wide, tlie marginal cilia evenly dispersed, about 0.5 

mm long; upper surface plane, shiny; lower surface 

Black, moderately rhizinate, the rhizines simple, 

black. Apothecia common, sessile to substipitate, to 

12 mm in diameter, the disc flat, often radially split; 

spores 8,s-12 X 12-18 pm. 

CHEmsTRY.-Cortex K + yellow (atranorin); 

medulla K-, C-, KCf rose, P- (“horrescens” 

unknown). 

DIsrRIBuTIo,v.-So~itli !imerica. 

RE~r,~RIts.-AlthoUgll clearly distinct because of 

23

24 SSIITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 

12.4pecies of Parmelina: a, P. damaziana (Dainazio 1375 in W); b, P. degelii (Degelius 

A-I82 in Degelius herbarium); c, P. de?iegans (Hale 43855); d, P. dissecta (Hale 14999); e, P. 

endoleuca (Weber in Lichenes Exsiccati 244 in US): f, P. enorinis (Jellicoe in US). (Scale in mm.)

NUMBER 33 

the chemical constitutents, P. damaziana is a rare 

and not well comprehended species. The lectotype 

was collected on tree branches whereas Lynge’s 

species was apparently saxicolous. Burchell 1105-06 

is tentatively placed here since the chemistry, while 

not clear, is closest to the “horrescens” type. Parme- 

lina damaziana has rather large spores, just as the 

other members of the P. horrescens group. It could 

be regarded theoretically as the parent morph for 

P. howescens, P. schindleria, and P. subfatiscens, but 

all of these vegetative morphs have smaller, more 

fragile thalli. 

SPECIXIENS ExAalrNED.-BraLil: Burchell 1105-06 (BM, US). 

8. Parmelina degelii, new species 

FIGURE 12h 

DEscRIPrroN.-Thallus adnatus, corticola, fragilis, 

pallide viridi-albicans, 6-8 cm latus, lobis plus 

minusve subirreqularibus, 2-3 mm latis, margine 

aetate lobulato-dissectis, ciliatis, ciliis usque ad 0.8 

mm longis, superne planus vel aetate rugulosus, 

nitidus, sparse vel modice isidiatus, isidiis primum 

cylindricis, mox procumbentibus, dorsiventralibus, 

expansis, ad 0.6 mm longis, cortex superior 12 pm 

crassus, epicorticatus, stratum gonidiale 12-14 pm 

crassum, medulla pallide sulfurea, 60-90 pm 

crassa, cortex inferior paraplectenchymatus, brunneus, 

12 p,m crassus, subtus niger, dense rhizinosus, 

rhizinis nigris, simplicibus, 0.5-09 mm longis. 

Apothecia ignota. 

CHEMIsTRY.-Cortex K + yellow (atranorin); 


(zeorin, leucotylic acid, apparently secalonic acid 

A, and traces of other pigments and terpenes). 

HoLoTYPE.-Rain forest, Cairns Cove, Queensland, 

Australia, elevation 600 m, G. Degelius A-I82 

(Degelius herbarium; isotype in US). 

DI~TRIRUTI~N.-A~S traha. 

REMARKS.-This is the only species of section 

Myelochroa that has been discovered in Australia. 

It is remarkable that such a conspicuous lichen 

should be collected so recently in Queensland. The 

terpene chemistry is close to P. aurulenta except 

for the absence of one spot (Figure 7). Regardless 

of chemistry, however, it is distinct from all other 

species in the P. subaurulenta complex because of 

the production of large lobulate isidia. Parmelina 

degelii is named in honor of Dr. Gunnar Degelius, 

who has enriched our knowledge of lichen distribu- 

tions through his careful collecting efforts in the 

tropics. 

9. Parmelina denegans 

FILLRF 12C 

Parmelzna denegans (Nylander) Hale, 1974,482. 

Parnelta denegans Njlander, 1900 6 [tjpe collection: Ram- 

podde, Ceylon, Almquzst (H, Njlander herbarium number 

35129, lectotype; isolectotype in S)]. 

DE~CRIPTI~X .-Thallus adnate to closely adnate 

on bark, light greenish mineral gray, 5-9 cm broad; 

lobes irregularly branched, sublinear, imbricate, 1-3 

mm wide, the marginal cilia irregularly dispersed, 

0.4-0.7 mm long; upper surface shiny, weakly maculate, 

plane but soon developing small pustulate 

areas, the pustules eventually breaking open 

apically and forming coarse soredia; medulla pale 

orange-salmon, in part K+ purple; lower surface 

black, densely rhizinate, the rhizines black, simple 

to sparsely branched. Apothecia sessile, 2-3 mm in 

diameter; spores 8, 5-7 X 7-9 p.m. 

CHEhmTRY.-Cortex K + yellow (atranorin); 

medulla more intensively yellow-orange with color 

tests (zeorin, leucotylin and associated terpenes, 

secalonic acid A, and possibly anthraquinones). 

DIsTRIsuTIoN.-India, Sri Lanka, and Sabah. 

REMARKS.-Except for the darker reddish orange 

pigment in the lower part of the medulla, this species 

is the pustulate morph of P. subaurulenta. 

They have similar lobe configuration and habitats, 

occurring at about 2000 m elevation in open forests. 

The lectotype specimen is fertile and has few 

pustules. The specimens that I collected are heavily 

pustulate-sorediate by contrast. 

SPECIMENS ExAM1xFD.-India: Tamil Nadu, Foreau 4128 

(Awa3thi herbarium, US), Hale 40239, 43636, 43762, 43855. 

Sri Lanka: Thruaites (UPS). Sabah: Hale 29020. 

10. Parmelina dissecta 

FIGCJRE 12d. 13 

Parmelina dissecta (Nylander) Hale, 1974:482. 

Parmelia dissecta h’ylander, 18823451 [type collection: Fon- 

tainebleau, France, A’ylander (H, Nylander herbarium 

number 35131, lectotype; isolectotype in PC)]. 

Parinelia laeuigata var. gracilis f. furfuracea Miiller Argovien- 

25


sis, 1888c:529 [type collection: Faxina, Brazil, Puiggari 47 

(G, lectotype)]. 

Parmelia minarum Vainio, 1890:48. 

Parmelia amazonica var. hwnotii Hue, 1899:158. 

Parmelia puiggari Gyelnik, 1931:288 [based on Parmelia 

lamigata var. gracilis f. furfuracea Muller Argoviensis]. 

Parmelia camtschadalis var. epiphylla Cengia Sambo, 

1938:379. 

Parmelia hubrichtii Berry, 1941:102. 

[For full citation of synonymy, see Hale, 1971a:6.] 

DEScRIPTIoN.-Tha1Ius adnate on bark or rock, 

yellowish glaucous to pale greenish mineral gray, 

3-7 cm broad: lobes sublinear-elongate, contiguous, 

1-3 mm wide, the marginal cilia irregularly dis- 

persed, mostly simple, to 0.7 mm long: upper sur- 

face shiny, emaculate, plane to convex, moderately 

to densely isidiate, the isidia cylindrical, erect, 

often branched, less than 0.5 mm high; medulla 

white; lower surface black, moderately rhizinate, 

the rhizines black, shiny, simple or sparsely 

branched. Apothecia adnate, the rim crenate, the 

amphithecium isidiate: 1-4 mm in diameter: spores 

8,8-10 X 12-17 pm. 

CHEMIsmY.-Cortex K+ yellow (atranorin); 

medulla K-, C+ rose, KC+ red, P- gyrophoric 

acid with or without associated unknown sub- 

stances). 

DIsTRIBuTI0N.-Pantemperate and montane pan- 

tropical (Figure 13). 

REMAws.-This is one of the most widespread 

and commonly collected species in the genus, 

especially in temperate zones, occurring on a wide 

variety of substrates in open, secondary forests. In 

tropical regions it is strictly montane, usually being 

most abundant in cloud forests up to 2300 m eleva- 

tion. Morphological variation is wide, even though 

the basic characters, isidia, ciliate lobes, and gyro- 

phoric acid, are constant. The lobes, for example, 

are narrow, sublinear, and separate in the lectotype 

specimen. At the other extreme are plants such as 

those collected in Dominica (Hale, 1971a) with 

broader, contiguous to subimbricate lobes. 

FIGURE 13.-Distribution of Parmelina dissecta based on all available herbarium specimens.

NUMBER 33 

Parmelina dissecta has been correctly identified 

by European lichenologists both on the Continent 

and in tropical regions, Degelius (1941:60) was the 

first to identify the species correctly in North 

America. He noted two populations differing in C 

test (C+ rose or C-) and presence or absence of 

cilia on the isidia. Hale and Kurokawa (1962) 

determined that the C- ciliate population is P. 

horrescens. No parent morph has been discovered 

for P. dissecta, but it forms a convenient nucleus 

of the “dissecta” group, including isidiate but sig- 

nificantly larger P. melanochaeta, sorediate P. cryp- 

tochlora, and isidiate-lobulate P. subfatiscens (Fig- 

ure 9). 

SPECIMENI EXAMINED (selected).-United States: Pennsylvania, 

Hale 17140; Marjland, Hale 14367; West Virginia, 

Hale 11470: Ohio, Hale 13910, 15735; Kentucky, Hale 13770; 

I’irginia. Hale 38419, Liittrell 3799 (US); Tennessee, Hale 

31124, 37038; North Carolina, Culberson 7099 (DUKE), Hale 

30645; South Carolina, Culberson 7459 (DUKE), Hale 16601; 

Alabama, Hale 33937, Skorepa 4547 (US); Georgia, Culberson 

7249 (DUKE), Hale 7509; Florida, Nelson (US) (see 

Moore, 1968:220 for additional records in Florida); Mississippi, 

Hale 7977, Hubricht 1532 (US): Texas, Hale 5242. 

Mexico: Chiapas, Hale 20304; \‘era Cruz, Hale 21231. Panama: 

Chiriqui, Hale 38879. Cuba: Oriente, Imshaug 24720 

(MSC). Jamaica: Inishaug 14172 (MSC). Guadeloupe: Dim 

1030 (TUR). Dominica: Hale 35569 (see Hale, 1971a:13, for 

additional records). Martinique: Degeliits (Degelius herbarium). 

St. T’incent: Guilding 46 (BM). Trinidad: Imshaug 

32103 (MSC). Colombia: Antioquia, Nee and Mori 4258 (US). 

l’enezuela: Distrito Federal, Dennis 1568 (BM); Mkrida, Hale 

42543; TAchira, Hale 42540, 45112. Brazil: Minas Gerais, 

Eiten 6867 (US); Parana, Montes 10121 (hIVM). Europe: 

Portugal, Persson (UPS), Sampaio in Lichenes de Portugal 

252 (LD, US), Tauares 97 (H, LD, WIS), 632 (US); Spain, 

Degelius (S), Santesson 19428 (UPS), Schauer in Lichenes 

Selecti Exsiccati 265 (LD, US); France, des Abbayes in 

Lichenes Armoricani Spectabiles Exsiccati 88 (LD, W), Hasselrot 

(S); Italy, Gresino (S), Sbarbaro (US). Kenya: Rift 

Valley Province, Maas Geesteraniis 6067 (L). Tanzania: Tanga 

Prov., Santesson 23137 (UPS). Union of South Africa, Cape 

Province, Almborn 301, 670, 3201 (LD); Natal, Almborn 

9192, 9525 (LD), Hoeg (TRH). Swaziland: Almborn 7904 

(LD). India: Tamil Nadu, Degelius As-238 (Degelius herbarium), 

Hale 40222, 43715, 43852; West Bengal, Degelius As- 

21 1 (Degelius herbarium). Sri Lanka: Degelius As-466 (Degelius 

herbarium, US). Malay: Pahang, Hale 30137, 30483. 

Taiwan: Kurokawa 2400 (TNS). Japan: Prov. Aki, Hale 

29368: Prov. Higo, Mayebara 1.58 (TNS); Prov. Kazusa, Kurokawa 

64096 (TNS); Prov. Ohmi, Hale 29484; Prov. Owari, 

Asahina 146 (TNS); Prov. Settsu, Hale 29411. Philippines: 

Mountain Province, Degelius As-% (Degelius herbarium), 

Hale 26040. Indonesia: Java, Groenhart 6022 (L), 8483 

(BOR); Sumatra, Groenhart 967 (L). Hawaii: Hale 31340, 

32933. 

11. Parmelina endoleuca, new combination 

FIGURE 12e 

Parmelia endole.uca TaFlor, 1847: 167 [type collection: Swan 

Rixer, Australia, Drurnmond (FH-Tajl, lectotjpe)]. 

DEscRIP’rIos.-Thallus closely adnate on bark, 

whitish mineral gray, 2-4 cm broad; lobes sublinear 

to subirregular, short and crowded, 1-2 mm wide, 

the marginal cilia sparse, less than 0.3 mm long; 

upper surface shiny, plane to minutely rugulose, 

becoming more strongly rugose and lobulate toward 

the center, heavily pycnidiate, soredia and isidia 

lacking; medulla white: lower surface dark brown 

at the margins and black toward the center, mod- 

erately rhizinate, the rhizines simple, brown but 

blackening at maturity. Apothecia very numerous, 

sessile, 1-2.5 mm in diameter, the disc dark brown, 

plane; spores 8, 6-7 X 11-12 pm. 

CHExmrRY.--Cortex K + yellow (atranorin); 

medulla negative with color tests (an unidentified 

fatty acid). 

DIsTRIsuTIoN.-Austra~ia. 

REalARKs.-In my earlier work on Parmelia I 

placed this species in synomymy under Parmelina 

galbina, a Japanese-North American species. Since 

the Australian locality seemed at variance with the 

Arcto-Tertiary distribution pattern of P. galbina 

and since my first chemical tests had been done 

with crystal techniques, I have checked the type of 

Parmelia endoleiica again and found that it con- 

tains neither galbinic acid nor any of the associated 

terpenes that characterize P. galbina. I have con- 

cluded, therefore, that Parmelia endoleuca is a good 

species in spite of the very close external resem- 

blance to P. galbina. The fatty acid falls higher on 

the chromatographic plates than either caperatic 

acid or protolichesterinic acid. The species is 

endemic to the dry scrub forests of eastern 

Australia. 

SPECIVENS ExA\rIN~.o.-Australia: Australian Capital Terri- 

tory, Weber in Lichenes Exsiccati 244 (COLO, US). 

12. Parmelina enormis 

FIGURE 12f 

Parmelina enormis (Hale) Hale, 1974:482. 

Parmelia enormis Hale, 1972c:344 [type collection: Nyaka 

Plateau, Zambia, Jellicoe, September 1968 (BM, holotype; 

isotype in US)]. 

27


DESCRIPTIoN.-Thahs expanded, loosely attached 

on rock, coriaceous, whitish to ivory mineral gray, 

10-30 cm broad; lobes sublinear, dichotomously 

branched, imbricate and crowded toward the cen- 

ter, 5-8 mm wide, the margins entire, ciliate mostly 

in the lobe axils, the cilia to 2.0 mm long; upper 

surface plane to convex, continuous, lacking soredia 

and isidia; medulla white; lower surface pale 

brown, densely rhizinate, the rhizines pale brown, 

simple. Apothecia numerous, subpedicillate, 3-8 

mm in diameter; spores 8, 6-7 x 8-11 

CHEMIsTRY.-Cortex K + yellow (atranorin); 

medulla I

NUMBER 33 

FIGURI: 14.--Species of Parmeliiza: a, P. entotheiochroa (Hale 29456a); b, P. expollida (Kurokair-a 

2930 in TKS); c, P. gnlbina (Hole 2341.5); tl, P. hoyachitzensis (Kurokawa 67081 in TNS): e, 

P. hetemloba (Schiflner in \\'); f, P. liorrescens (Hnle 36999). 

29


CHEMIs-rRY.-Cortex K + yellow (atranorin); 

medulla negative with color tests (protolichesterinic 

acid and an unidentified fatty acid). 

DIsmInuTIoN.-India, Thailand, and Taiwan. 

As pointed out by Kurokawa (1968a:191), an 

unusual feature of this species is the brown lower 

surface, indicating some relationship with P. versi- 

forrnis, a South American species. It is also one of 

the few Parrnelinae to produce a fatty acid. 

SrEcIarENs EXAMINF‘.D.-See Kurokawa (1968a:193) for records 

from India, Taiwan, and Japan. 

15. Parmelina galbina 

FIGURES 14c, 15 

Parmelina galbina (Acharius) Hale, 1974:482. 

Parmelia galbina Acharius, 1814: 195 [type collection: North 

America (?Pennsylvania), Muhlenberg (H-Ach, lectotype; 

isolectotype in PH)]. 

Parmelia tiliacea var. minor Muller Argoviensis, 1877:X 

[type collection: near Dallas, Texas, Boll (G, lectotype; 

isolectotypes in FH, M, W)]. 

Parmelia tiliacea var. sulphurosa Tuckerman, 1882:57 [type 

collection: Illinois, Hall (FH-Tuck, lectotype)]. 

Parmelia deminuta Hue, 1899:156 [type collection: Texas, 

Boll (PC, lectotype)]. 

Parmelia subquercifolia Hue, 1899: 157 [type collection: Ohio, 

Sullivant (PC, lectotype)]. 

Parmelia subquercifolia var. rugosa Hue, 1899: 175 [type col- 

lection: Oyama, Japan, Faurie (not seen)]. 

Parmelia tiliacea subquercifolia (Hue) Merrill and Burnham 

in Burnham, 1922:75 [rank not designated]. 

Parmelia quercina var. sulphurosa (Tuckerman) Zahlbruck- 

ner, 1929:192. 

Parmelia sulphurosa (Tuckerman) Fink, 1935:328. 

Parmelia laeuigata ssp. extremi-orientalis f. rugosa (Hue) 

Asahina, 1951c:291. 

Parmelia subquercifolia f. subradiata Asahina, 1952:98 [type 

collection: base of Mt. Fuji, Japan, Yamamoto (TNS, lecto- 

type) (nomen nudum in Asahina, 1951c:291)]. 

Parmelia galbina var. rugosa (Hue) Asahina, 1963:225. 

Parmelia galbina var. subradiata (Asahina) Asahina, 1963:225. 

DEscRIPTIoN.-Thallus closely adnate on bark, 

yellowish-glaucous to greenish mineral gray, 3-10 

cm in diameter; lobes short, sublinear, contiguous, 

1-2 mm wide, marginal cilia mostly in axils, some- 

times only sparsely developed, less than 0.5 mm 

long; upper surface plane to rugulose, continuous, 

pycnidia usually numerous; medulla white or pale 

yellow to orange, especially beneath the apothecia; 

lower surface densely rhizinate. the rhizines black, 

simple. Apothecia very common, adnate, 2-5 mm in 

diameter, the disc cinnamon brown; spores 8, 7-9 X 

10-13 pm. 

CHEMIsrRY.-Cortex K + yellow (atranorin); 

medulla K+ yellow, C- or C+ yellow, KC-, 

P + pale orange (galbinic acid, zeorin, secalonic 

acid A, trace of salazinic acid, and leucotylin and 

associated terpenes). 

DIsTRIsvTIo;v.-Japan and eastern North America. 

REMARKs.-During a visit to the Academy of 

Natural Sciences in Philadelphia in 1958 I found a 

specimen in the hfuhlenberg herbarium without a 

name but with a penciled number “15.2.” I later 

found the same plant in the Acharian collection 

(H) also numbered 15.2 and described by Acharius 

as Parrnelia galbina. This discovery clarified the 

status of one of the very common corticolous lichens 

in eastern North America. W. Culberson (1961: 173) 

subsequently studied the problem and lectotypified 

P. subquercifolia Hue as synonymous with P. gal- 

bina. The second syntype of Hue’s species was 

determined by Culberson to be Parmelia livida 

(= Hypotrachyna livida (Taylor) Hale), another 

very common corticolous lichen in eastern North 

America distinguished by more or less distinctly 

dichotomously branched rhizines and the lividic 

acid complex (Hale, 1975a:45). As a rule, H. livida 

has a whiter cast than P. galbina, is more robust, 

and lacks pigments in the medulla or under the 

apothecia. Still, care must be taken when identi- 

fying these two species. One other distinguishing 

character is the unique moniliform cells in the 

medulla of P. galbina and its morphs, P. hayachin- 

ensis, P. metarevoluta, and P. obsessa, first described 

by Asahina (1951~). The cells, however, are not 

easy to find and their exact significance is unknown 

at this time. 

SPECIMENS EXAMINED.-United States: See W. Culberson 

(1961:lil) for a discussion of the distribution and map of 

localities in North America based on specimens in DUKE 

and US (Figure 15). Japan: Prov. Aki, Kurokawa 64431 

(TXS); Prov. Bungo, Kurokawa 63191 (TNS); Prov. Hizen, 

Kurokawa 62588 (TNS); Prov. Hoki, Yasuda (TNS); Prov. 

Hyogo, Tagawa L15 (TNS); Prov. Kozuke, Degelius As-1072 

(Degelius herbarium); Prov. Mikawa, Takaki 353 (TNS): 

Prov. Musashi, Kurokawa 64279 (TNS); Prov. Mutsu, Kuro- 

kawa 550385 (TNS); Prov. Shimotsuke, Kurokawa 64057 

(TNS); Prov. Shinano, Kurokawa 51758 (TNS, US), 59179 

(TNS); Prov. Totomi, h’akanishi 27 (Kobe University); Prov. 

Ugo, Suzuki 272 (TNS).

NUMBER 33 31 

FIGURE 15.-Distribution of Parrnelina galbina in Korth 

America (taken froin W. Culberson, 1961). 

16. Parmelina hayachinensis 

FIGURE 14d 

Parrnelina hayaclzinensis (Kurokait-a) Hale, 1971:482. 

Prrrnlelia ha)achinensis Kurokawa, 1968b:350 [type collection: 

hit. Hayachine, Proy. Rikuchu, Japan, Kurokawa 67081 

(TSS, holotype)]. 

DEscRIPnoN.-Thallus adnate on bark, whitish to 

greenish mineral gray, 5-8 cm broad; lobes sub- 

linear, 1.5-3 mm wide, marginal cilia simple, about 

0.2 mm long; upper surface shiny, faintly maculate, 

becoming densely pustulate, the pustules becoming 

aggregated, erupting but not forming soredia; 

medulla white; lower surface black, densely rhizin- 

ate, the rhizines black, simple. Apothecia not seen. 

CHE~IsTRY.-Cortex K + yellow (atranorin); 

medulla K+ yellow turning red, C-, KC-, P+ 

orange (galbinic acid, leucotylin, and zeorin). 

DxsmIsuTIoN.-Japan. 

REMARKS.-AS pointed out by Kurokawa (1968b: 

350), this species is a member of the P. galbina 

complex, representing the pustulate morph. It is 

known only from the type collection growing on 

Cryptomeria in Japan. 

17. Parmelina heteroloba 

FIGURE 14e 

Parrnelina heteroloba (Zahlbruckner) Hale, 1974:482. 

Parrnelia heteroloba Zahlhruckner, 1909: 171 [tlpe collection: 

Mt. Itatiaya, Rio de Janeiro, Brazil, Schiflner (W, lecto- 

type)l. 

DEscRIPTIoN.-Thallus adnate on bark, light buff 

mineral gray in the herbarium, 5-8 cm broad; lobes 

more or less sublinear, short and imbricate, be- 

coming lobulate toward the center, 2-3 mm wide, 

the marginal cilia mostly in the lobe axils, to 0.4 

mm long; upper surface shiny, plane to rugulose, 

becoming somewhat lobulate; lower surface black 

except for a narrow brown zone at the tips, densely 

rhizinate, the rhizines black, simple or sparsely 

furcate. Apothecia numerous, substipitate, the disc 

splitting, 2-9 mm in diameter; spores 8, 8 X 12 pm. 

CHEMIsTRY.-Cortex K + yellow (atranorin); me- 

dulla negative with color tests (an unidentified fatty 

acid and a faint unknown spot, perhaps related to 

one of the “horrescens” unknown). 

DIsmIsuTIoN.-Brazil. 

REMARKS.-siXICe this species is represented only 

by the single type collection, it is difficult to gen- 

eralize on its affinities to other Parmelinae. It would 

probably be mistaken for P. damaziana, which has 

a different chemistry (“horrescens” unknown) and 

smaller spores. It is only one of three species in 

Paimelina with fatty acids, the others being P. 

endolezica and P. expallida. Parmelilza heteroloba 

has not been recollected at Alt. Itatiaya, a well 

known site that has been visited by several experi- 

enced lichen collectors, and it may be an extinct 

species. 

18. Parmelina horrescens 

FIGURE 14f 

Paririelina horresce?is (.laylor) Hale, 1974:482. 

Pnr,,relin korrescens Taylor in Mackay, 1836:144 [t!Iie collec- 

tion: Dunkerron Mountains, Kerry, Ireland, Taylor (FH- 

Tayl, lectotype)]. 

Dk:scRrp?.xos.-Thallus closely adnate to adnate on 

bark, rocks, or mosses over rocks, whitish to green- 

ish mineral gray, 2-5 cm broad; lobes more or less 

dichotomously branched, sublinear, often crowded 

and imbricate, 0.5-2.0 mm wide, the margins cre-

nate, often becoming lobulate, ciliate, the cilia more 

or less evenly dispersed, black, simple, 0.3-0.8 mm 

long; upper surface shiny, emaculate, densely isidi- 

ate, the isidia cylindrical, often branched and 

apically spinulate or short-ciliate, in part becoming 

procumbent; medulla white; lower surface black, 

moderately rhizinate, the rhizines black, simple. 

Apothecia rare, sessile, 2-4 mm in diameter, the 

amphithecium isidiate, the disc splitting at ma- 

turity; spores 8, 10-12 X 16-18 pm. 

CHExmTRY.-Cortex K + yellow (atranorin); me- 

dulla K -, C - or C + faint rose, KC + rose or red, 

P - (trace of gyrophoric acid, “horrescens” un- 

known falling above gyrophoric acid on chroma- 

tographic plates, and one or two other unidentified 

s PO t S) . 

DIsTRIsuTIozr.-Pantemperate and montane pan- 

tropical. 

REMARK\.-,& discussed under P. dissecta, this 

species only recently was differentiated correctly 

from P. dissecta (Hale and Kurokawa, 1962:2), an 

isidiate lichen with gyrophoric acid. Parmelina 

horrescens is characterized by dense fine isidia with 

greater or lesser development of short apical cilia 

(Figure 4c). The isidia sometimes become procum- 

bent and lobulate. On the whole, the lobes are 

somewhat narrower and more appressed than in 

P. dissecta. Parinelina horrescens has essentially the 

same geographic range as P. dissecta and occurs 

most abundantly in the temperate deciduous 

forests of the eastern United States and Japan. It is 

montane in the tropics, occurring as high as 3000 m 

in the paramos of Venezuela. 

The chemistry of P. horrcscens is now being 

studied by several lichen chemists. A number of 

unidentified spots appear in both hexane and ben- 

zene solvent systems, one of which is probably a 

trace of gyrophoric acid. The other spots fall above 

and below gyrophoric acid and seem to represent 

closely related depsides. 

The parent morph of P. horrescens is probably 

extinct. Chemically identical P. damaziana, a non- 

isidiate Brazilian species, is larger and more robust 

although obviously from the same stock as the 

parent of P. horrescens. Among the parallel morphs, 

both pustulate-sorediate P. subfatisceiis and lobulate 

P. schindzeri are very close in lobe configuration, 

adnation, and thallus texture. 

SPECIMENS EX&MINED.-United States: Illinois, Skorepa 4627 

SMITHSONIAN CONTRIBUTIONS TO BOTANY 

(US); Kentucky, Hale 13743a; West Virginia, Hale 10612, 

11772, 11878; Virginia, Hale 18388, 33153, Reed 9091 (Reed 

herbarium), Roller 400 (US); h‘orth Carolina, Culberson 

5114, 5706, 5744, 7134 (DUKE), Hale 18042, 15058, Imshaug 

22174, 22178, 22353 (MSC, US); Tennessee, Hale 31106, 36921, 

36959, 36970, Moore 284 (US), Phillips 358, 377 (US), Skorepa 

5527 (US); South Carolina, Hale 7723; Georgia, Hale 7405, 

7537, 16761, 16776, 30882, 30885; Alabama, Hale 7186, 7216, 

31164, 33779, 33918, 34121, 34174, McCullough 2195 (US); 

Florida, Hale 21685 (for additional records see Moore, 

1968:220). Mexico: Chiapas, Hale 20204, 20223, 20400a, 20414, 

20549, 21086. Guatemala: Baja Vera Paz, Hale 45828. Pan- 

ama: Darien, >Vori and Gentry 4309 (US); Panami, Hale 

38451. Cuba: Oriente, Imshaug 24737, 24810, 24932 (MSC). 

Jamaica: Zmshnug 14216 (MSC). Dominican Republic: Cordil- 

lera Central, Imshaug 23516 (MSC), Wetmore 3739 (MSC); La 

T’ega, Allnrrl 17695a (US). Haiti: Ouest, Fahius 2-4 (US), 

Ini.rhaug 22768, 22837, 22860 (MSC, US), Wetmore 3222 

(MSC); Sud, Imshaug 23233 (MSC, US). Venezuela: Distrito 

Federal, Deiinis 2394 (BM), Santesson 6679 (S); Wrida, Hale 

42038, 42067, 42952, 45121, 45201. Urugna): Trienta y Tres, 

Osorio 5931 (MT‘M). France: Harmand (DUKE). Spain: Ponte- 

vedra, Schnurr (XI); Tenerife, Zmshazcg 34476, 35677 (MSC). 

Union of South Africa: Cape Province, Alniborn 1442 (LD). 

India: Tamil Nadn, Hale 43784. Philippines: Mountain Pror- 

ince, Hale 26531. Indonesia: Java, Groenhart 2903 (L, US), 

6279 (L), Kurokau,a 2071 (TNS). Taiwan: Kurokawa 693 

(TSS). Japan: Proy. Buzen, Kiirokawa 62468, 63166 (TNS); 

Prov. Kii, Kurokawa 64127 (TNS); Prov. Ohmi, Hale 29465. 

Australia: Sew South \Vales, Cizeel L1708 (NSW), Craigie 

(NSW). Sew Zealand: Wade 85 (BM, US). 

19. Parmelina immiscens 

FIGURE 16a 

Pnrmelina irnmiscens (Nylander) Hale, 1974:482. 

Parmelia immiscens Nylander, 1885:606 [type collection: 

Orizaba, Mexico, Galeotti 6879 (PC, 1ectot)pe; isolectotype 

in H, Xilander herbarium number 35674)l. 

Pnrmelia michoacanensis Body de Lesdain, 1914:7 [type 

collection: Jeslis del Monte, Morelia, Michoacan, Mexico, 

Arsene 4456 (US, lectotype; isolectotypes in COLO, DUKE, 

G, LE, and UPS)]. 

DEscRIPTIoN.-Thallus adnate on bark, pale tur- 

tle green, 5-10 cm in diameter; lobes subirregular 

and apically rotund, 2-6 mm wide, the marginal 

cilia mostly in the axils; upper surface plane, con- 

tinuous, often pruinose near the tips; medulla 

sulphur yellow; lower surface densely rhizinate, 

short rhizinate and pale brown along the margins, 

the rhizines simple or squarrosely branched. Apoth- 

ecia numerous, adnate, the disc pale, 2-7 mm in 

diameter; spores 8, 4-6 X 7-12 pm. 

CmmsmY.-Cortex K + yellow (atranorin);

NUMBER 33 

FIGURE 16.4pecies of Parrnelina: a, P. irnrniscens (Andne 4456 in US); b, P. indica (Hale 

43884); c, P. irrugans (Kurokauu 55342); ti, P. jarnesii (Du Rieti 3117:3 in US); e, P. leucotyliza 

(Hale 29402); f, P. lindrnnnii (Maline 450 in S). (Scale in mm.) 

33


medulla K+, C+, KC+ more intensely yellow, 

P - (unidentified yellow pigments). 

DIsTRIsuT1oN.-Mexico. 

REMARKs.-Parmelina immiscens is the only spe- 

cies in the genus endemic to Mexico. It occurs 

rather rarely on trees in open oak-pine forests in 

the arid highlands (2000-2400 m elevation). Al- 

though the medulla is distinctly yellow, no terpenes 

are produced. The pigment seems to be secalonic 

acid A but chromatographic tests are inconclusive. 

This species and its presumptive isidiate morph 

P. Zindmanii, therefore, are unrelated to P. sub- 

aurulenta and similar Asian species with a yellow 

medulla. 

SPECIMENS EXAMINED.-MeXicO: Durango, Cramer 1998 

(KAN, US); Jalisco, Pringle 10706 (US); Oaxaca, Hale 20825. 

20. Parmelina indica, new species 

FIGURE 166 

DEscRIPTIoN.-Thallus arcte adnatus, saxicola, 

obscure albo-cinereus, 2-3 cm latus, lobis sub- 

linearibus, contiguis, 0.8-1.2 mm latis, ciliis mar- 

gine irregulariter dispersis, simplicibus, 0.1-0.3 mm 

longis, superne planus, continuus vel rimosus, 

nitidus, modice vel dense isidiatus, isidiis cylin- 

dricis vel leviter inflatis, praecipue simplicibus, 

usque ad 0.3 mm altis; cortex superior 12 pm 

crassus, epicorticatus, stratum gonidiale 15-18 pm 

crassum, medulla alba, 100-120 pm crassa, cortex 

inferior paraplectenchymatus, 12-14 pm crassus; 

subtus niger, dense rhizinosus, rhizinis nigris, sim- 

plicibus, 0.4 mm longis. Apothecia ignota. 

CHEMIsmY.-Cortex K + yellow (atranorin); 

medulla negative with color tests zeorin, leucotylin 

and traces of associated “subaurulenta” terpenes). 

HOL0TYPE.-on rocks in river below Silver Cas- 

cade, Kodaikanal, Tamil Nadu, India, elevation 

about 1800 m, M. E. Hale 43884, 24 January 1975 

(US: isotype in Poona). 

DIsmIsuTIoN.-India. 

REMARKS.-This small saxicolous lichen was col- 

lected on a large rock outcropping in an undis- 

turbed stream bed. Externally it is close to the 

American P. obsessa but lacks both yellow pig- 

ments and galbinic acid. It is the only terpene- 

containing Parmelina without pigments. 

21. Parmelia irrugans, new combination 

FIGURE 16c 

Parmelia irrugans Nylander, 1890:26 [type collection: Simon- 

oseki, Japan, A lmquist (H, Nylander herbarium number 

35551, lectotype; isolectotype in S)]. 

Parmelia insinuata Hue, 1899: 158 [type collection: Ta-Pin- 

Tze, Yunnan, China, Delavay 3008 (PC, lectotype); not P. 

insinuata ”).lander, 1856: 3243. 

Parmelia xanthocarpa Hue, 1899:178 [type collection: Ta- 

Long-Tan near Ta-Pin-Tze, Yunnan, China, Delavay 23 

(PC, lectotype)]. 

Parmelia insinuatula Zahlbruckner, 1929: 169 [based on P. 

insinuata Hue]. 

DEsCRIPTIoN.-Thallus adnate to loosely adnate, 

corticolous, firm, 4-10 cm broad; lobes sublinear 

to subirregular with more or less rotund apices, 

contiguous, 2-5 mm wide, the marginal cilia dis- 

tinct, mostly in the axils, to 0.8 mm long; medulla 

pale orange or yellow; lower surface black, densely 

rhizinate, the rhizines simple or becoming squar- 

rosely branched. Apothecia very common, adnate 

to sessile, 3-10 mm in diameter; spores 8, 8-10 X 

12-15 pm, often poorly developed. 

CHEMIsmY.--Cortex K + yellow (atranorin); 


(zeorin, leucotylic acid and associated terpenes, and 

secalonic acid A). 

DIsmIsuTIoN.-China and Japan. 

REMARKS.-This species has not been recognized 

by contemporary lichenologists or included in lists 

of Asian Parmelinae. It is characterized by the 

presence of the “aurulenta” terpene series, the thin, 

expanded thallus, and large apothecia (3-10 mm in 

diameter), invariably larger than those of P. sub- 

aurulenta (3 mm or less). The spores are slightly 

but not significantly larger than those in P. SUB- 

aurulenta. As a rule, the size of apothecia will 

separate the two species. Two of Nylander’s syn- 

types of P. subaurulenta (see “Specimens Exam- 

ined” below) are actually P. irrugans. 

Parmelina irrugans has a more restricted range 

than P. subaurulenta. It does not occur in India 

where P. subaurulenta is so common. It is, how- 

ever, more frequent in Japan than one might 

expect from the specimens examined because I did 

not borrow the specimens at TNS for redetermina- 

tion. Many annotated by me in 1964 in Tokyo as 

P. homogenes are probably this species. 

SPECIMENS EXAMINED.-China: Montiguy (H. Nylander her-

NUMBER 93 35 

barium number 35668, a syntype of Parmelia subaurulenta 

Kllander); Hunan, Handel-Mazzetti 2469 (US, \V); Yunnan, 

Delaruy (H, K)lander herbarium number 35666). Japan: 

illaries (H, Sylander herbarium number 35667, probable syn- 

type of Pormelia subaurulenta Nylander); Prov. Kii, Kuro- 

kawn 59064 (TNS, US); Prov. Kikuchu, Kzirokazua 59322 

(TNS, US); Prov. Kozuhe, Kurokazua 58581 (TNS, US); Prov. 

Mutsu, Kurokawa 550341, 55342, 56162, 58582 (TNS, US); 

Prov. Ohmi, Hale 29456, 29470; Prov. Rikuchu, Kurokawa 

59322 (TNS, US); Prov. Shimane. Omura 23 (US); Prov. 

Suruga, Culberson 10738 (DUKE, US). 

22. Parmelina jamesii, new combination 

FIGURE 16d 

Parmelia jamesii Hale, 1972b:179 [type collection: Welling- 

ton, New Zealand, James SZ2118 (US, holotype; isotypes in 

BM, TNS)]. 

DEscRIPTros.-Thallus adnate on bark, rather 

fragile, whitish mineral gray, 5-10 cm broad; lobes 

sublinear, contiguous, 1.5-3.0 mm wide, the mar- 

gins sparsely ciliate, the cilia irregularly dispersed, 

long; upper surface plane but becoming rugulose 

with age, shiny or becoming opaque and white- 

pruinose at the tips, sometimes faintly white- 

reticulate, moderately isidiate, the isidia cylindrical, 

thin, erect, up to 0.3 mm high; lower surface black, 

moderately rhizinate, the rhizines simple or sparsely 

squarrosely branched. Apothecia unknown. 

CHEmsTRY.-Cortex K+ yellow (atranorin); 

medulla K-, C-, KC-, P+ red (fumarproto- 

cetraric acid and a trace of protocetraric acid). 

DIsTRIBuTI0N.-Eastern Australia and New 

Zealand, 

REMARKs.-Parmelina jamesii is the only species 

producing ,B-orcinol depsidone fumarprotocetraric 

acid. It is also one of the few species endemic to 

the Australian region. It might be mistaken super- 

ficially for a Hypotrachyna except that the lobe 

margins are distinctly ciliate and the rhizines 

simple. 

23. Parmelina leucotyliza 

FIGURE 16e 

Parmelina leucotylizn (Sylander) Hale, 1974:482. 

Parmelia leucotyliza Nylander, 1890:27 [type collection: Rock- 

osan, Japan, Almquist (H, Nylander herbarium number 

35196, lectotype: isolectotype in S)]. 

Parmelia fmudans ssp. subfraudans Zahlbruckner, 1927:352 

[type collection: Inokashira, near Tokyo, Japan, Asahina 

23a (W, lectotype)]. 

Parmelia leucotyliza f. rugulosa Asahina, 1952:94 [tvpe col- 

lection: Mt. Higane, Prov. Suruga, Japan, Yamashita 17 

(TNS, lectotype)]. 

Parmelia leucotyliza f. sziblaevis Asahina, 1952:95 [type collec- 

tion: Kadoike, Pro\.. Suruga, Japan, Asahina (TSS, lecto- 

type)]. 

DEscRIPTIo;u.-Thallus adnate to loosely attached 

on bark or rocks, light greenish mineral gray, 8-12 

cm broad; lobes sublinear to subirregular, con- 

tiguous, 2-4 mm wide, the marginal cilia mostly 

in lobe axils, about 0.5 mm long; upper surface 

shiny or becoming dull white pruinose at the tips, 

plane but soon becoming pustulate, the pustules 

breaking open and coalescing into large clumps 

without formation of soredia; medulla very pale 

salmon colored to white; lower surface black ex- 

cept for a dark brown zone at the tips, densely 

rhizinate, the rhizines black and shiny, simple or 

sparsely squarrosely branched. Apothecia rare, sub- 

stitpitate, the amphithecium pustulate, 1-3 mm in 

diameter; spores 8, 6 X 11-12 pm. 

CHEmsTRY.-Cortex K + yellow (atranorin); 


(zeorin, leucotylin and related “subaurulenta” ter- 

penes, secalonic acid A, and traces of unidentified 

substances). 

DIsTRIBuTIoN.-Japan and hlalaysia (Sabah). 

REMARKS.-Originally when studying the Jap 

anese Paymeliae, I synonymized P. leucotyliza under 

P. aurulenta. These two species, however, are differ- 

ent in several important respects. First, they have 

different terpene chemistry. Second, the pustules of 

Paymelina leucotyliza do not become sorediate, 

whereas those in P. aurulenta are often densely 

sorediate. Lastly, P. leucotyliza is restricted to Japan 

except for a tentatively identified specimen from 

Sabah. It seems to be much more common than 

P. aurulenta in Japan, occurring on rocks and trees 

in open forests, trees along roads, and rocks in rice 

fields. 

SPECIXIE~S 

ExAaIisED.-Japan: Prov. Aki, Hale 29535, 29445, 

29544; Prov. Hyuga, Hale 29677; Prov. Iwaki, Kurokarua 

58079 (TNS, US); Prov. Izu, Asahina in Lichenes Japoniae 

Exsiccati 139 (US), Kurokawa 58004, 58600 (TNS, US); Prov. 

Kazusa, Kurokawa 59002 (TSS, L’S); Prov. Kii, Kurokawa 

59087, 59131, 59132 (TNS, US); Prov. Musashi, Kurokawa 

51040 (TNS, US); Prov. Settsu, Hale 29402, 29433; Shizuoka 

Prefecture, Omura 157 (US). Malaysia: Sabah, Hale 29102.


24. Pamtelina lindmanii 

FIGURE 16f 

Parmelina lindmanii (Lynge) Hale, 1974:483. 

Parrnelia lindrnanii Lynge, 1914:74 [type collection: Porto 

Alegre, Rio Grande do Sul, Brazil, Maime 450 (S, lecto- 

type)l. 

Parmelia tiliacea var. sulphurosa f. asperata Miiller Argoviensis, 

1883:46 [type collection: Argentina, Balansa 5 (G, 

lectotype)]. 

DESCRIPTION.- Thallus adnate on bark, pale buff 

to yellowish mineral gray, 5-8 cm broad; lobes subirregular, 

apically rotund, 3-5 mm wide, the marginal 

cilia mostly in lobe axils, less than 0.5 mm 

long; upper surface plane to rugose with age, shiny, 

moderately to densely isidiate, the isidia cylindrical, 

unbranched, to 0.3 mm high; medulla uniformly 

pale sulfur-yellow tinged with orange; lower surface 

black except for a narrow marginal brown 

zone, moderately rhizinate, the rhizines black, simple. 

Apothecia rare, sessile, 2-5 mm in diameter; 

spores 8,6-10 X 9-14 pm. 

CHExmTRY.-Cortex K + yellow (atranorin); 

medulla more intensely yellow with K and C, P- 

(probably secalonic acid A and unidentified 

pigments). 

DISTRIBUTION.-RleXiCO and South America. 

REhlARKS.-This corticolous lichen is character- 

ized by the pale yellow-orange medulla and the 

isidia. It is probably the isidiate morph of the 

Mexican endemic P. immiscens. When I first began 

study of Pal-nzelia, I confused Pal-melina lindmanii 

and Pal-melia endosulphur-ea ( = Parmotl-ema endo- 

sulphur-eum (Hillmann) Hale) (Hale, 1960:20). Both 

species have a yellow-orange medulla and isidia, but 

P. lindmanii has more adnate lobes, cilia in the 

axils, and no gyrophoric acid. Both species lack 

terpenes. 

SPECIMENS Ex.4JrINEo.-Mexico: Queretaro, Nakanishi 257 

(US); Tamaulipas, Nakanishi 106 (US): \'era Cruz, Hale 

19389, 19685. Colombia: Antioquia, Nee and Mori 4238 (US). 

Venezuela: Merida, Rodrigues 168 (US). Brazil: Rio Grande 

do Sul, Lima 3 (US). Paragua:; Caacupe, Balansa (G, H). 

Uruguay: Durazno, Osorio 2772 (DUKE, UPS): Lavalleja, 

Osorio 3822 (MVM, US): Paysandu, Lamb (BM, H); Soriano, 

Rosengurtt (H); Trinidad, Osorio 3603 (LD). Argentina: 

James (BM): Buenos Aires, Santesson 58, 100, 446 (S). 

25. Parmelina melanochaeta 

FIGURE 17a 

Parmelina nielanochaeta (Kurokawa) Hale, 1974:483. 

Parmelia inelanochaeta Kurokaira in Hale and Kurokawa, 

1964:133 [type collection: Santa Anna de Chapada, Mato 

Grosso, Brazil, Malme 2243 (S, lectot) pe: isolectotypes in 

UC, US)]. 

DEscRIrTIos.-Thallus adnate on bark, turning 

olive-buff to cream-buff in the herbarium, 4-7 cm 

in diameter; lobes irregularly branched, sometimes 

sublinear-elongate, 2-6 mm wide, the margins more 

or less crenate, ciliate, the cilia black, mostly simple, 

1-2 mm long; upper surface shiny, maculate, mod- 

erately to densely isidiate, isidia thin, cylindrical, 

usually branched, often with black spinules or short 

cilia; medulla white; lower surface black, dark to 

pale brown in a rather wide zone near the tips, 

moderately rhizinate, rhizines black to blackish 

brown, simple. Apothecia adnate to substipitate, 

1-3 mm in diameter, the amphithecium isidiate, 

spinulate, the disc Vandyke-brown; spores 8, 8-10 x 

13-15 pm. 

CHEiwsTRY.-Cortex K + yellow (atranorin); me- 

dulla K-, C+, KC+ rose, P- (gyrophoric acid). 

DIsTRIBuTIoN.-South America. 

REMARKS.-This species is very close to P. dissecfa 

in general morphology and has the same chemistry. 

The lobes, however, are much broader, the isidia 

ciliate, the marginal cilia long and distinct, and 

the upper cortex strongly white-maculate. It occurs 

in a rather restricted area from Brazil into Para- 

guay with one specimen recorded from Colombia. 

SPECIMF\S ExA\IINED.-Colombia: Cundinamarca, Flenniken 

1745 (US). Brazil: Mato Grosso, Malme 2397B (S). Paraguay: 

Asuncih, Schinini 17333 (M\'M): Gran Chaco, hlalme (S). 

Additional records from Brazil and Paraguai are listed in 

Hale and Kurokana, 1964:133. 

26. Parmelina metarevoluta 

FIGURE 17b 

Par??7e[i?7a rnetaraloluta (Asahina) Hale, 1974:483. 

Pnrmelia tnetarevolirtn Asahina, 1960:97 [type collection: 

AzusaJama, Prov. Shinano, Japan, h'uno and Kurokawa 

59243 (TSS, lectot) pe; isolectotype in US)]. 

DEscRIPTIo;u.-Thallus adnate on bark or rock, 

pale olive-buff, 2-6 cm in diameter; lobes dicho- 

tomously branched, sublinear-elongate, more or 

less ascending at the tips, 1-4 mm wide, lobules 

sometimes present on older lobes, the margins more 

or less crenate, narrowly black rimmed, ciliate, the 

cilia black, shiny, simple, 0.2-0.5 mm long; upper

NUMBER 33 

FIGURE 17.4pecies of Parinelina: a, P. melanochaeta (21ialnie 2243 in S); b, P. inetarevoluta 

(Kurokawa 59180 in US); c, P. niiielleri (Hale 42277): d, P. obsessa (Hale 19186); e, P. pnsr~lltfera 

(specimen in US); f, P. perisidrans (Togashi in US). (Scale in mm.) 

37


surface smooth and shiny, not maculate, more or 

less rugose on older lobes, sorediate, soralia subterminal, 

capitate, pale orange below; lower surface 

black, densely rhizinate; the rhizines black, shiny, 

simple, about 0.5 mm long. Apothecia very rare, 

adnate, 1-5 in diameter, the rim more or less 

crenate, sorediate, the disc dark brown; spores 8, 

7 X 10-12pm. 

CmMIsmY.-Cortex K + yellow (atranorin); medulla 

K + reddish, C -, KC -, P + orange (galbinic 

acid, rarely constictic acid, trace of salazinic acid, 

zeorin, leucotylin and associated terpenes, and 


DIsTR1suTIoN.-Eastern United States, Japan, and 

China. 

REMARKL-AS pointed out by Kurokawa (1968b: 

351) this is the sorediate morph of P. galbina. It 

was first collected and correctly identified in the 

United States by Dr. Satoshi Nakanishi in 1971. 

Two additional collections have since been identified, 

one of them (Plitt 236) filed under Physcia 

orbicularis, which is very superficially similar in 

lobe configuration and capitate soralia. It has the 

same Arcto-Tertiary distribution pattern as P. galbina 

but is extremely rare in North America. 

SPECIMENS ExAMmEu.-United States: Ohio, Wetmore 18051 

(MI", US); Maryland, Plitt 236 (US); Tennessee, Nakanishi 

238 (US). Japan: Prov. Bungo, Kurokawa 62329 (TR'S); Prov. 

Ise, Asahina (TNS); Prov. Omi, Asahina (TNS); Prov. 

Shinano, Asahina (TNS). Kurokawa 58326 (TNS); 59180 

(TNS, US) 59243, 59244 (TNS); Prov. Suruga, Asahina (TR'S); 

Prov. Totomi, Asahina 160 (TNS); Prov. Yamashiro, Asahina 

(TNS). China: Manchuria, Asahina 29 (TNS). 

27. Parmelina muelleri 

FIGURE 17c 

Parmelina muelleri (Vainio) Hale, 1974:483. 

Parmelia muelleri Vainio, 1890:49 [type collection: Sitio, 

Minas Gerais, Brazil, Vainio in Lichenes Brasilienses 

Exsiccati 948 (TUR, Vainio herbarium number 2677, lecto- 

type; isolectotypes in BM, FH, M, UPS)]. 

DEScRIPTIoN.-Tha11us closely adnate on bark, 

lichen green, turning honey yellow in the her- 

barium, 5 cm or more in diameter; lobes irregularly 

branched, rounded at the tips, 2-3 mm wide, the 

margins evenly ciliate, the cilia simple or rarely 

branched, black, shiny, up to 1.5 mm long; upper 

surface strongly white-maculate, irregularly cracked 

on older lobes, sorediate, soredia laminal, more or 

less granular, the soralia round, separate; lower 

surface uniformly black, moderately rhizinate, the 

rhizines simple, black. Apothecia rare, sessile, the 

amphithecium sorediate, disk brick red, 2-5 mm in 

diameter; spores 8,7-I0 X 12-15 pm. 


dulla K+ persistent yellow, C-, KC-, P+ pale 

orange (stictic acid with or without constictic acid). 

DISTRIRUTI0N.-Mexico and South America. 

REMARKS.--When 1 first collected this species on 

shade trees in a coffee plantation in Venezuela, I 

identified it tentatively as P. pilosa because of the 

laminal soralia and distinct white maculae. Chem- 

ical tests, however, showed that it contained stictic 

acid and would have to be identified with P. 

muelleri. After comparing the two species in the 

herbarium, I concluded that P. pilosa is larger and 

more robust on the average, but the two are very 

closely related. Vainio (1890) had in fact compared 

P. muelleii with Parmelia balansa f. sorediata (= 

Parmelina pilosa) which he distinguished with a 

negative K test. They have different geographic 

ranges, P. muelleri being more common in northern 

South America and possibly occurring at a higher 

elevation than P. pilosa. There is no esorediate, 

stictic acid-containing parent morph comparable to 

the P. consors-P. pilosa species pair. 

SPECIMENS ExAMINED.-Mexico: Michoackn, Wirth 329 (US); 

Vera Cruz, Hale 21124. Venezuela: MCrida, Hale 42219, 

42230, 42256, 42277, Oberroinkler and Poelt 7529 (M, US); 

Thchira, Hale 45716. Peru: Cuzco, lltis 3214 (WIS). Argentina: 

TucumPn, Culberson 14909 (DUKE, US). 

28. Parmelina obsessa 

FIGURE 17c 

Parmelina obsessa (Acharius) Hale, 1974:483. 

Parmelia obsessa Acharius, 1814:195. [type collection: North 

America (?Pennsylvania), Muhlenberg (H-Ach, lectotype; 

isolectotype in PH)]. 

Parmelia finkii Zahlbruckner in Hedrick, 1934: 162 [type 

collection: Williamsville, Wayne County, Missouri, Russell 

(W, lectotype; isolectotype in MICH)]. 

DEScRIPTIoN.-Tha11us closely adnate on rock 

(rarely on bark), pale smoke gray to light mineral 

gray, 3-6 cm broad; lobes irregularly branched, 

sublinear, contiguous, 1-2 mm wide, the margins 

dissected, sparsely ciliate, especially in axils, the 

cilia black, simple, to 0.3 mm long; upper surface 

densely isidiate, the isidia cylindrical, sometimes

NUMBER 88 39 

branched; medulla white to pale dull green-yellow; 

lower surface black, sparsely to moderately rhi- 

zinate, dark brown and short rhizinate near the tips, 

the rhizines simple. Apothecia adnate, 1-3 mm in 

diameter, the disc brown, the amphithecium isidi- 

ate; spores 8,4-5 X 6-10 pm. 

CHEmsTRY.-Cortex K + yellow (atranorin); me- 


(zeorin, galbinic acid, trace of salazinic acid, leuco- 

tylin and associated terpenes, and secalonic acid A). 

DIsTRIsuTI0N.-Eastern United States. 

REMARKs.-Parme~ina obsessa is another Acharian 

species that had been lost from the literature since 

1814. Not only were extremely few collections 

available, but also many were incorrectly identified. 

One in US was listed as “Parmelia prolixa,” and as 

recently as 1958 P. obses,ra was identified as Parme- 

lia finkii (Hale, 1958:82). Recent collecting by 

American lichenologists, however, has shown that 

it is extremely common on sandstone outcrops in 

closed oak forests and can tolerate considerable 

shade. As a closely appressed saxicolous lichen it is 

not easy to collect. Kurokawa (1968b) correctly 

recognizes it as the isidiate morph of P. galbina. No 

comparable isidiate population has evolved in 

Japan. 

SPECIMENS EXAMINED (Selected).-New Hampshire, Willey 

(US): Pennsylvania, Hale 16288, 17486; Maryland, Hale 

14422; West Virginia, Hale 10311, 10660, 12408, 15003; Kentucky, 

Hale 13866; Indiana, Hale 14182; Illinois, Hale 13935, 

14022; Michigan, Hale 33889; Wisconsin, Hale 23529: Minnesota, 

Hale 23602; Virginia, Hale 12018, 18803, Luttrell 

3864 (US), Tucker 6588 (US); North Carolina, Culberson 

6491, 10418 (DUKE, US), Hale 16358; Tennessee, Hale 37083; 

South Carolina, Hale 7698; Georgia, Hale 7404, 30605; Alabama, 

Hale 7098; Louisiana, Hale 33862; Arkansas, Hale 

2972, 4030; Kansas, Hale 4749; Oklahoma, Hale 4886, 5054, 

Keck 1493 (US); Texas, Hale 5452. 

29. Parmelina pastillifera, new combination 

FIGURE 17e 

Parmelia scortea var. pastillifera Harmand, 19103558 [type 

collection: Bussang, Vosges, France, Claude1 and Harmand 

in Lichenes Gallici Exsiccati 491 (H, lectotype; isolectotypes 

in BM, BPI)]. 

Parmelia pastillifera (Harmand) Schubert and Klement, 

1966:58. 

DEscRIPTIoN.-Thallus largely as in P. tiliacea 

(p. 48) except isidia dark tipped, peltate, and 

apically flattened (Figures 3e,f, 4b). 

CHEMISTRY.-cOrteX K + yellow (atranorin); me- 

dulla K-, C+, KC+ red, P- (lecanoric acid). 

DIsTRrsuTIoN.-Great Britain and Europe. 

REMARKS.-while this species has long been 

recognized as a distinct variety (Poelt, 1961:194), it 

was only recently raised to species level. Dobson 

and Hawksworth (1976) have compared it in detail 

with the normally isidiate P. tiliacea, finding that 

P. pastillifera has, on the average, somewhat nar- 

rower lobes (2-6 mm) than P. tiliacea (3-8 mm) and 

denser rhizines.2 Parmelina pastillifera also has a 

somewhat smaller, slightly more bluish thallus. The 

ecological differences are pronounced, for P. pastil- 

Zifera occurred in England west of the 813 mm 

rainfall isopleth and the 2OC January mean iso- 

therm. This agrees with Schauer’s (1965:80) defini- 

tion of P. pastillifera as an oceanic species in 

central Europe. 

SPECIMFNS EXAMINED.-EUrOpe: France, Lambinon in 

Lichenes Selecti Exsiccati 363 (LD, US); Germany, Schroppel 

in Lichenes Alpium 143 (US); Austria, Strasser in Crypto- 

gamae Exsiccatae Vindobonensi 3062b (US); Yugoslavia, 

Vtzda in Lichenes Selecti Exsiccati 762 (US). 

30. Parmelina perisidians 

FIGURE 17f 

Parmelia perisidians (Nylander) Hale, 1974:483. 

Parmelia perisidians (h’jlander, 1900:6 [type collection: Ram- 

podde, Ceylon, Almquist (S, 1ectot)pe; isolectotype in H, 

Nylander herbarium number 35673)l. 

Parmelia subsulphuuata Asahina, 1951a:228 [t)pe collection; 

Higashi-Shirakawa, Prov. Mino, Japan, Yasue (TNS, lecto- 

type)l. 

DEscRIPTIoN.-Thallus adnate on bark, greenish 

mineral gray, 3-6 cm in diameter; lobes sublinear, 

0.5-2 mm wide, the marginal cilia mostly in the 

axils; upper surface plane, continuous, densely 

isidiate, the isidia simple or branched; medulla 

sea-foam yellow; lower surface densely rhizinate, 

the rhizines black, simple or squarrosely branched. 

Apothecia adnate, 2-5 mm in diameter, the am- 

phithecium isidiate; spores 8,5-7 X 7-1 1 pm. 

CHEMISTRY.-cOrteX K + yellow (a tranorin); me- 

dulla K+, C+ KC+ more intensely yellow, P-, 

(zeorin, leucotylin and related terpenes, and 


*I wish to thank Dr. Hawksworth for allowing me to see 

his manuscript before publication.


DIsTRIBuTI0N.-Southeast Asia from India to 

Japan. 

REMARKS.-This densely isidiate species is closely 

related to P. amagiensis and may represent its isidi- 

ate morph, although differences in pigments, so far 

unresolved, may exclude this possibility. Both 

species have the typical “subaurulenta” terpene 

series. Parmelina perisidians has a broad distribu- 

tion in the higher elevation forests of tropical Asia 

and the temperate forests of Japan. 

SPECIMENS ExAMIKED.-India: Karnataka, Patwardhan 

74.3283 (Poona, US): Tamil Nadu, Hale 40101, 40.102, Togashi 

(TNS, US). Sri Lanka: see Kurokawa and Mineta (1973:74) 

for records in Ceylon. Thailand: Kurokawa 1818 (TNS, US). 

Philippines: Rlountain Province, Hale 26166a. Japan: Prov. 

Kozuke, Asahina (TNS); Proc. Ifusashi, Asahina (TNS); 

Kurokawa 550009 (TKS, LS), 64284, 64285 (TNS); Prov. 

Settsu, Togashi (TNS, US). 

3 1. Parmelina phlyctina 

FIGURE 18a 

Parmelina phlyctina (Hale), Hale, 1974.483. 

Parmelia phlyctina Hale, 1959:129 [tjpe collection, Blue 

Mountains, Jamaica, lnzshaicg 14908 (MSC, holotjpe; iso- 

qpe in US)]. 

DEscRIPTIoN.-Thallus adnate to loosely adnate 

on bark, membranous and fragile, light mineral 

gray but turning pink in the herbarium if improp- 

erly dried, 5-15 cm broad; lobes subirregular, 

apically rotund, 5-10 mm wide, the margins entire 

to lobulate, sparsely ciliate in the lobe axils, the 

cilia up to 0.5 mm long; upper surface plane, 

smooth to rugose in older parts, upper cortex 

fragile and flaking off in small pieces, isidia and 

soredia lacking; lower surface black except for a 

narrow brown zone at the tips, sparcely rhiAnate, 

the rhizines simple. Apothecia rare, adnate, 3-8 mm 

in diameter; spores 8, 3-5 X 7-8 pm. 

CHEMIsmY.-Cortex K + yellow (atranorin); me- 


orange (norstictic and connorstictic acids). 

DISTRIBUTION.-hkXiCO and the West Indies. 

REMARKS.-sinCe my discription of this species in 

1959, additional collections have been made in 

Mexico, Cuba, and other islands in the West Indies. 

It occurs in wet upland rain forests or secondary 

mist forests. The main distinguishing features are 

the fragile thallus with a flaking cortex, presence 

of norstictic acid, and small spores. It is closely re- 

lated to isidiate P. antillensis but is not the direct 

parent morph since P. antillensis has a continuous 

cortex. It is not, as implied in my original descrip- 

tion, a member of Amphigymnia (= Parmotrema) 

because the thallus is adnate overall, cilia are pro- 

duced in the axils, and the bare zone at the 

margins below is very narrow. Both P. phlyctina 

and P. antzllensis, however, constitute an anomolous 

element in Parmelina, not only because of the 

somewhat amphigymnioid lobation but also be- 

cause of the production of norstictic acid. 

SPECIW ?s Exs~i~\k~.-hfexico: Chiapas, Hale 20202 (S, 

US), Cuba: Oriente, Imshnug 27174 (MSC, US), 25146, 25148 

(MSC), ,\lorton 9394 (US). Jamaica: Imshaug 14614, 14714, 

15211, 15362, 15375, 15458, 15502, 15524, 15536 (MSC), 13179, 

14727, 15146, 15453, 15513 (MSC, US). Haiti: Sud, Wetmore 

3373 (MSC). Dominican Republic: Cordillera Central, Wetmore 

3728 (LD, MSC), 3738 (MSC, US). Puerto Rico: Imshaug 

29562 (MSC). 

32. Parmelina pilosa 

FIGURE 18b 

Parmelina pilosa (Stizenlxrger) Hale, 1974:483. 

Parmeha pilosa Stizenberger, 180: 165 [type collection: 

Rhenoster Ricer, Taaibosch Kranz Mountains, Orange 

Free State, Union of South Africa, Rehmann (ZT, lectotype; 

isolectotype in H)]. 

Parrnelin halansae var. sorediata Muller Argoviensis, 1888a:Z 

[tjpe collection: Montevideo, Uruguaj, Arechavaleta (G, 

lectotype)]. 

Parnielia subbalansae Gyelnik, 1931:288 (based on Parmelia 

balansae ~ar. sorediata Miiller Argoviensis. 

DEscRIPTIoN.-Thallus adnate to loosely attached 

on bark, coriaceous, light buff mineral gray, 5-12 

cm broad; lobes subirregular, apically subrotund, 

the marginal cilia coarse, becoming furcate, 0.5-1 .O 

mm long; upper surface shiny and heavily white- 

maculate, plane to minutely pitted, sorediate, the 

soralia orbicular, about 0.5 mm in diameter, sepa- 

rate or coalescing into extensive lamina1 sorediate 

areas; medulla white; lower surface black, densely 

rhizinate, coarse and fine rhizines intermixed, 

simple to sparcely furcate. Apothecia rare, sub- 

stipitate; spores not seen. 

CHE;MIsTRY.-Cortex K + yellow (atranorin); me- 

dulla negative with color reagents (no substances 

demonstrated). 

DISTRIRLTIoN.-~outh America and southern 

Africa.

NUMBER 33 

FIGURE 18.-Species of Parmelina: a, P. phlyctina (Imshaug 13179 in US); b, P. pilosa (McOwan 

in US); c, P. pruinata (Kurokawa in Lichenes Selecti Exsiccati 81 in US); (1, P. quercina (“carporrhizans” 

population) (Schroppel and Poelt in Lichenes Alpiurn 8 in US); e, P. quercina 

(Tavares 1082 in US); f, P. rhytidodes (Kurokawa 58601 in US). (Scale in mm.) 

41


REMARKs.-Parmelina pilosa is the sorediate 

morph of P. consors. It occurs on a variety of trees 

(Celtis, Erythz’na, Scutia), even on fence posts, and 

in open or secondary habitats up to 2000 m eleva- 

tion. It must be distinguished carefully from P. 

muelleri, which rarely occurs in this range and con- 

tains stictic acid. 

SPECIMENS EXAMINED.-Ecuador: Pichincha, Wiggins 38945 

(COLO). Uruguay: Canelones, Felippone 434 (G); Durazno, 

Osorio 2979 (DUKE); Flores, Osorio 3519 (MVM, US); 

Florida, Osorio 1299 (F); Minas, Osorio 2049, 2056 (MVM, 

US); Soriano, Rosengurth (H). Argentina: Buenos Aires, 

Grassi 719 (US), Santesson 75 (S), Schnyder (G), Venturi 

2853 (SI); Cordoba, Giardelli 1025 (SI), Petersen 23 (S); Entre 

Rios, Santesson 126 (S, US); Salta, Fries 21 (S), James (BM), 

Lorentz (M). Chile: Valparaiso, Santesson 3084 (S). Kenya: 

Nyanza Province, Maas Geesteranus 4957 (L, US). Rhodesia: 

Hoeg (TRH). Union of South Africa: Basutoland, KofEer 

(LD); Cape Province, Hoeg (LD, TRH), Kofler (LD), Mc- 

Owan (BM, US); Natal, Hoeg (TRH). 

33. Parmelina pruinata 

FIGURE 18c 

Parmelina pruinata (Muller Argoviensis) Hale, 1974:483. 

Parmelia pruinata Muller Argoviensis, 1883:46 [type collec- 

tion: Yorke Peninsula, Australia, Tepper (G, lectotype)]. 

Parmelia tiliacea var. afixa Stirton, 1899:485 [type collection: 

Queensland, Australia, Bailey (?) 376 (BM, lectotype)]. 

DEsCRIPTIoN.-ThaIIus closely adnate on twigs, 

ashy white, 1-5 cm broad; lobes sublinear-elongate, 

variable, 1-3 mm wide, marginal cilia sparse, mostly 

in lobe axils, up to 0.5 mm long; upper surface 

plane to convex, rugulose, more or less pruinose; 

medulla white; lower surface black, sparsely rhi- 

zinate, the rhizines black, simple. Apothecia con- 

spicuous, adnate, 2 4 mm in diameter, the disc 

often pruinose; spores 8,7-8 X 10-12 pm. 

CHEMIsTRY.-Cortex K+ yellow (atranorin); me- 

dulla K -, C +, KC + red, P - (lecanoric acid). 

DISTRIBUTION.-Australia and New Zealand. 

REMARKS.-This species is rather variable in lobe 

configuration and width but typically grows closely 

adnate on small twigs and branches of trees in arid 

scrub land. It has no apparent relation with P. 

quercina, the only other nonsorediate, lecanoric 

acid-containing species in the genus, characterized 

by much coarser lobes and white maculae in the 

cortex. Muller mentions “subtus alba” in his origi- 

nal description, but he was actually describing a 

Physcia species growing intermingled with the type. 

The marginal cilia are sparse and sometimes diffi- 

cult to determine. It could, in fact, be misidentified 

as a Pseudoparmelia but no corticolous nonisidiate 

species of that genus with lecanoric acid are pres- 

ently known. 

SPECIMENS ExAMINED.-Australia: Australian Capital Territory, 

Kurokawa in Lichenes Rariores Selecti Exsiccati 81 

(US), Leuthner (US); New South Wales, Boorman 1387, 

L1389 (NSW), Doing (L, US), McVean 6358 (COLO), Weber 

and McVean L-47913, L-49886 (COLO, US). Wilson 96 

(NSW); Queensland, Gwytherl (BM); South Australia, Rogers 

886 (US); Tasmania, Bratt 1380b (US), James 2117 2125 

(BM, US), Weymouth 65 (BM, US), 432 (BM); Victoria, Filson 

6639, 7004 (US), James 223 (BM, US). New Zealand: 

Mason 90 (BM). 

34. Parmelina quercina 

FIGURE 18d,e 

Parmelina quercina (Willdenow) Hale, 1974:483. 

Lichen quercinw Willdenow, 1787:353 [type collection: Thier- 

garten and near Tegel, Germany (not seen; illustrated in 

pl. 7: fig. l3)]. 

Imbricaria quercina (Willdenow) de Lamarck and de Can- 

dolle, 1805:389. 

Parmelia carporrhizans Taylor, 1847:163 [type collection: 

Canary Islands, Lemann (FH-Tayl, lectotype)]. 

Parmelia sinuosa var. hypothrix Nylander, 1856:301 [type 

collection: Pyrenees, France, Nylander (H, lectotype)]. 

Parmelia atricha Nylander, 1873:271 [type collection: La 

Preste, Pyrenees, France, Nylander 16 (H, lectotype (not 

seen); isolectotypes in FH, BM)]. 

Parmelia tiliacea var. hypothrix (Nylander) Muller Argovien- 

sis, 1888b:196. 

Parmelia revoluta var. grantilata Harmand, 1897:221 [type 

collection: Docelles, Vosges, France, Harmand 284 (DUKE, 

lecto type)]. 

Parmelia quercina (Willdenow) Vainio, 1899:279. 

Parmelia budapestinensis Gyelnik, 1932b3212 [type collection: 

Budapest, Hungary, Timkd (BP, holotype)]. 

Parmelia carporrhizans f. malicola Gyelnik, 1932b:212 [type 

collection: St. George, Hungary (Austria?), Zahlbruckner 

(BP. holotype)]. 

Parmelia yalungana Zahlbruckner, 1934b:206 [type collec- 

tion: between the Litang and Yalung rivers between Muli 

Gomba, Baurong, and Wa-Erh-Dje, Szechuan, China, Rock 

16720 (W, lectotype)]. 

DEscRIPTIoN.-Tha1Ius closely adnate on bark, 

rarely on rock, rather coriaceous, light olive-gray, 

sometimes turning olive-buff in the herbarium, 

5-10 cm broad; lobes dichotomously or irregularly 

branched, sublinear, separate or becoming im- 

bricate, 1.5-4.5 mm wide, the margins smooth to 

more or less crenate, often narrowly black rimmed,

NUMBER 33 43 

moderately to densely ciliate, the cilia black, simple, 

0.2-1 mm long; upper surface smooth and shiny, 

faintly to strongly white-maculate, irregularly 

rugose and cracked on older lobes, sometimes partly 

pruinose, isidia and soredia lacking; lower surface 

black, densely rhizinate, the rhizines black, shiny, 

simple or more rarely squarrose, 1-3 mm long. 

Apothecia numerous, substipitate, 1.5-5 mm in 

diameter, the amphithecium smooth or retrorse- 

rhizinaee, the disc burnt umber; spores 8, 5-7 x 

6-10 pm. 

CHEmsmY.--Cortex K + yellow (atranorin); me- 

dulla K-, C+ KC+ red, P- (lecanoric acid). 

DISTRIBUTION.-califOrnia, Europe, Pakistan, Ne- 

pal, eastern Asia, and Australia. 

REMARKS.-AS a common member of the Euro- 

pean foliose lichen flora this was one of the first 

species of Parmelia to be described. The records 

from eastern North America previously identified as 

Parmelia quercina are now recognized as Parmelina 

galbina or Hypotrachyna Zivida (Taylor) Hale. The 

interesting distribution pattern, centered in western 

North America and Europe, is typical of the Medi- 

terranean type, although the species occurs less 

commonly in eastern Asia and Australia. Still this 

is a remarkably broader range than its isidiate 

morph P. filiacea. 

The development of rhizines on the lower side of 

the apothecia has been used to justify a separate 

population, called Parmelia carporrhizans. As W. 

Culberson (1961: 168) points out, however, virtually 

all specimens in California have these rhizines, and 

many European specimens identified as P. quercina 

are often found to have them, sometimes only very 

sparsely developed (Dobson and Hawksworth, 1976). 

This trait does not seem to warrant specific status, 

although Schauer (1965:71) found this population 

to have a strongly oceanic distribution pattern in 

western Europe. The problem will have to be solved 

through careful field studies in Europe. 

SPECIMFNS EXAMINED (selected).-Without Retrorse Rhizines: 

Europe: Denmark, Rostrup (LD); Germany, Schroppel and 

Poelt in Lichenes Alpiurn 7 (H, LD, US); Switzerland, 

Mereschkovsky (US), Seifriz (US); Austria, Eggerth in Flora 

Exsiccata Austro-Hztngarica 1542 (US); Czechoslovakia, Vdzda 

in Lichenes Bohernoslouakiae Exsiccati 201 (LD, US); Hungary, 

Tirnkd in Lichenotheca 135 (H, LD); Spain, Schindler 

4303 (KR, US); Portugal, Tavares 1082 (US). Pakistan: Iqbal 

180 (US). Nepal: Stainton 4098 (BM, US). China: Manchuria, 

Asahina 28 (TNS); Shensi Prov., Hugh (BM). Japan: Prov. 

Hyuga, Hale 29650, Kurokawa 65065 (TNS); Prov. Tango, 

Asahina (TNS); Prov. Tosa, Kurokawa 64091 (TNS), 64094 

(TNS, US). Australia: New South Wales, Du Rietz 577b 

(UPS, US), Flockton 884 (US). 

Mostly with Retrorse Rhizines on the Apothecia: United 

States: See W. Culberson (1961:172) for a list of specimens 

examined and a distribution map. Europe: England, Borrer 

(US), Crombie (BM, US), Holl 15 (BM), James 75 (BM): 

France, Crozals (US), Rondon in Lichenes Selecti Exsiccati 

268 (US); Spain, Bauza (LD); Portugal, Tavares in Lichenes 

Lusitaniae Selecti Exsiccati 217 (CS); Germany, Poelt in 

Lichenes Alpiurn 8; Switzerland, Barkman 4294a (UPS): Austria, 

Stezner in Crjptogamae Exsiccatae Vindobonensi 4134 

(F, NY); Italy, Alrnborn (LD), Baunigartner in Cryptogarnae 

Exsiccatae Vindobonensi 3165 (NY, US). Tunisia: Runeinaik 

(LD). 

35. Parmelina rhytidodes, new species 

FIGURE 18f 

DEscfuPTIo;u.-Thallus adnatus ve1 laxe adnatus, 

corticola et saxicola, pallide olivaceo-cinereus, 8-12 

cm latus, lobis subirregularibus, imbricatis con- 

gestisque, 2-4 mm latis, margine irregulariter 

ciliatis, ciliis nigris, 0.3-0.6 mm longis; superne 

nitidus vel apicem versus albo-pruinosus, primum 

planus sed mox dense et omnino rugosus (Figure 

6b), rugis non eruptentibus, cortice integro vel rare 

rumpente; cortex superior 12-14 pm crassus, epicor- 

ticatus, stratum gonidiale 14-16 pm crassum, me- 

dulla pallide salmonea, 110-150 pni crassa, cortex 

inferior brunneus, paraplectenchymatus, 15-17 pm 

crassus; subtus niger, dense rhizinosus, rhizinis nigris 

vel apice brunneis, simplicibus vel sparse squarroso- 

ramosis. Apothecia numerosa, sessilia vel substipi- 

tata, disco plano, 3-8 mm diametro; sporis 8, 

9-11 X 12-14,um. 

CHEMIsmY.-Cortex K + yellow (atranorin); me- 


(zeorin, leucotylic acid and related terpenes, seca- 

Ionic acid A, and traces of unidentified substances). 

HOLoTYPE.-Amagi Pass, Prov. Izu, Japan, S. 

Kurokawa 58601, 1 December 1958 (US, holotype; 

isotype in TNS). 

DIsTRIBuTIoN.-Japan and Nepal. 

REMARKS.-After I had examined specimens iden- 

tified as Pa? melina entotheiochron with thin-layer 

chromatography, there remained a small group with 

the “aurulenta” terpene profile. On further study 

I found that these are morphologically different 

from typical P. entotheiochroa in that the ridges 

which develop very densely do not burst open or

44 SSIITHSONIAX CONTRIBUTIONS TO BOTANY 

flake off. While P. rhytidodes can be recognized 

with practice from the external morphology alone, 

a chemical test is desirable for confirmation. The 

species is probably not as common in Japan as P. 

entotheiochroa, but I have not had the opportunity 

to re-examine the numerous specimens in TNS with 

chromatography. The collections made by Poelt in 

Nepal are tentatively placed here. They are very 

densely rugose and have crowded lobes, modified 

ecologically by the high exposed elevation where 

they were collected. Parmelina rhytidodes is appar- 

ently an offshoot with sorediate P. aurulenta from 

P. irrugans or a now extinct parent morph similar 

to it. 

SPECIMENS ExAMINED.-Japan: Prov. Awa, Kurokawa 56554 

(TSS, US); Prov. Hyuga, Hale 29602; Hiroshima Prefecture: 

Nakanishi and Oshio 5731 (US); Kanto, Nurnariri 156 (US). 

Nepal: Poelt L-747, L-754 (M, US). 

36. Parmelina schindleri, new species 

FIGURE 19a 

DEscRIPrIoN.-ThallUs adnatus vel appressus, 

fragilis, corticola, pallide albo-cinereus, 2-4 cm 

latus, lobis sublinearibus, brevibus, contiguis, ca. 

1-1.5 mm latis, margine ciliatis, ciliis 0.2-0.4 nim 

longis, simplicibus, margine et pro parte superficie 

lobulatis, lobulis congestis, suberectis, ramosis, 0.1- 

0.2 mm latis et usque ad 1 mm longis, margine 

breve ciliatis; superne nitidus, planus; cortex supe- 

rior 14-15 pm crassus, epicorticatus, stratum gonidi- 

ale 12 pm crassum, medulla alba, 50-65 pm crassa, 

cortex inferior paraplectenchymatus, 12 pm crassus; 

subtus niger, modice rhizinosus, rhizinis nigris, sim- 

plicibus vel sparse furcatis. Apothecia rara, sessilia, 

margine crenato, 3-4 mm diametro, sporis 8:nae, 

10 X 16-18pm. 

CHEMIsrRY.-Cortex K + yellow (atranorin); me- 

dulla K-, C-, KC+ rose, P- (traces of gyro- 

phoric acid and the “horrescens” unknown). 

HoLoTYPE.-Caraca, h has Gerais, Brazil, I/ainio 

in Lichenes Brasilienses Exsiccati 1284 (BM; isotypes 

in FH, RI, TUR, and UPS). 

DIsTRIBuTIoN.-Brazil. 

RExmKs.-The chemical constituents place this 

species in the “horrescens” group, where it is the 

lobulate morph of a parent species represented by 

P. damaziana or a now extinct progenitor similar 

to it. The dense lobules are easily recognized (Fig 

Lire 4e). They do not originate from isidia and for 

this reason P. schindleri cannot be considered as 

a modified form of P. horrescens, which may occa- 

sionally have cylindrical and flattened isidia inter- 

mixed. The species is named in honor of Dr. H. 

Schindler, the first lichenologist to collect the 

species since Vainio in 1885, although this region 

has been visited by many lichen collectors. Vainio 

had misidentified his exsiccate number 1284 as 

Parmelia coronata var. isidiosa Rluller Argoviensis 

(= Bztlbothrix fungicola (Lynge) Hale). 

SPFCIMENS ExAMIxED.-Brazil: Rio de Janeiro, Schindler 

4569,4577 (KR, US). 

37. Parmelina simplicior 

FIGURE 19b 

Parmelina simplicior (Hale) Hale, 1974:483. 

Parrnelia sirnplicior Hale, 1972a:99 [type collection: Panch- 

gani, \Vestern Ghats, India, D. D. Awasthi 4056 (US, holo- 

type; isotype in Awasthi herbarium)]. 

DEscRIPTIoN.-Thallus adnate on bark, buff min- 

eral gray, coriaceous, 8-10 cm broad; lobes elongate, 

more or less subirregular, contiguous and becoming 

imbricate, 3-5 mm wide, the axils sparsely ciliate, 

the cilia 0.5 mm long; upper surface plane, contin- 

LIOLIS, emaculate, isidia and soredia lacking; medulla 

white; lower surface black, sparsely to moderately 

rhizinate, the rhizines simple, black. Apothecia 

common, sessile, 3-6 mm in diameter; spores 8, 

4 X 6pm. 

CHEMIsmY.--Cortex K + yellow (atranorin); me- 


orange (salazinic acid). 

DISTRIBUTIoN.-~outh India. 

REMARKs.-Parmelina simphior has a rather 

leathery thallus and very sparsely developed axil- 

lary cilia. As with many other Indian lichens it 

does not exhibit clear-cut traits that enable one to 

place it immediately in a particular genus. I had 

mentioned a possible relationship to isidiate P. 

wallichiana, which has much larger spores and a 

more membranous thallus, but these two species are 

not at all related. Pal-melina simplicior grows on 

roadside trees in the Western Ghats region of 

India where an intense monsoon season from June 

to September alternates with a long period of almost 

total drought. Parmelina wallichiana, on the other

NUMBER 33 

FIGURE 19.-Species of Purmelina: a, P. schindleri (I’ainio 1284 in BM); 6, P. .rirnplicior 

(Arcnstlzi 4056 in US); c, P. spathulato (Alnzborn 931B in US); d, P. spurnma (Halt, 42934a); 

e, P. subaw-ulenta (Hale 43705); f, P. subfatitcens (Hale 35091). (Scale in mm.) 

45

46 

hand, occurs at higher elevations with less climatic 

stress. 

SPECIMENS ExAMIxED.-lndia: Maharashtra, Hale 40004, 

40007,40046, 40090,43972. 

38. Parmelina spathulata 

FIGURE 19c 

Parmelina spathulata (Kurokawa) Hale, 19743483. 

Parmelia spathulata Kurokawa in Hale and Kurokawa, 

1964:133 [type collection: Skeleton Gorge, Wynberg, Union 

of South Africa, Alrnbortz 305 (LD, holotype; isotype in 

Wl. 

DEsCRIPTIoN.-Thallus adnate on bark, pale 

whitish glaucous-green, fragile, 2-5 cm broad; lobes 

sublinear, crowded, 1-3 mm wide, the marginal 

cilia evenly dispersed, about 0.5 mm long; upper 

surface plane, continuous, moderately isidiate, the 

isidia initially cylindrical and erect but soon be- 

coming procumbent and flattened; lower surface 

black, moderately rhizinate, the rhizines black, 

simple to rarely furcate. Apothecia not seen. 

CmhiIsmY.-Cortex K + yellow (atranorin); me- 

dulla K-, C+, KC+ rose to red, P- (gyrophoric 

acid). 

DIsTRIBuTIoN.-South Africa. 

REMARKs.-This lobulate-isidiate species (Figure 

4d) is obviously a member of the P. dissecta group. 

It occurs only in southern Africa and is much less 

common than P. dissrcfn, which has smaller, normal 

is idia . 

SPrcihfEss Ex.iMlsED.-~nion of South Africa: Cape Pror- 

ince, Kofler (LD, US). Additional records from South Africa 

are given in Hale and Kurokaiva (1964:134). 

39. Parmelina spumosa 

FIGURE 19d 

Parmrlina spumosa (Asahina) Hale, 1974:483. 

Parinelia spz~mosa Asahina, 1951b:259 [type collection: 

Higashi-Murayama, Kita-Tatna-gun, Prov. Musashi, Japan, 

Asahina (TNS, lectotype)]. 

DEsCRIPTIoN.-Thallus closely adnate on bark, 

fragile, pale olive gray, 2-6 cm in diameter; lobes 

sublinear, 0.5-2 mm wide, the marginal cilia dis- 

tinct and evenly dispersed, about 0.5 mm long; 

upper surface plane, continuous, emaculate, densely 

pustulate-isidiate, the pustules bursting but not 

becoming sorediate; medulla faintly yellow; lower 

SMITHSONIAN COSTRIBUTIONS TO BOTANY 

surface moderately rhizinate, black, the rhizines 

simple or furcate. Apothecia rare, adnate, 1-3 mm 

in diameter, the amphithecium pustulate; spores 8, 

7-8 X 12-14,um. 

CHmiIsTRY.-Cortex K + yellow (atranorin); medulla 

K-, C+ rose, KC+ red, P-, (gyrophoric 

acid, an unidentified pigment, and frequently a 

white fluorescent spot). 

DIsTRInuTroN.-Pantropical at higher elevations. 

REMARKS.-T~~S widespread but rather rare 

species is characterized by the dense, erupting but 

nonsoretliate pustules (Figure 5e,f) and the pale 

yellowish medulla. The main constituent, gyrophoric 

acid, places it in the P. dissecta group, but 

it probably represents a different line of evolution 

from a now extinct parent morph (Figure 9). The 

chemistry is somewhat aberrant from P. dissecta and 

related species judging by the presence of a white 

UV-fluorescent spot in more than half of the specimens 

(including the lectotype) from both the New 

TVorltl and the Old World. The spot is not reactive 

with H2S0, and its identity is not known. 

Parmelina spurnosa has ecological requirements 

similar to those of P. dissecta and P. horrescens. 

Much less common at temperate latitudes than the 

latter two, it grows throughout the tropics on 

exposed trees (hardwoods and conifers) and more 

rarely on rocks in forests up to 2500 m elevation. 

SPEcIhfENs Ex.AarIxm.-Mexico: Chiapas, Hale 19897, 20110a, 

20150. Cuba: Hioram 6682 (US). Jamaica: Imshaug 14279, 

15646 (MSC, US). Colombia: Antioquia, Nee and Mori 

4258a (US). Venezuela: Merida, Hale 42957, 42934a, 43037a. 

Brazil: Sao Paulo, Eiten and Mimura 5738 (US). Chile: 

Chiloe, Santesson 2264 (S, US): Valdivia, Santesson 7066 (S). 

St. Helena: Loveridge (AM). Union of South Africa: Cape 

Province, Almborn 749, 2663 (LD), 3966 (LD, US), Degelius 

SA-403 (Degelius herbarium, US); Natal, Almborn 10069 

(LD). Madagascar: Lemaitre (H). Indonesia: Java, Groenhart 

2218 (L), Neemoort 3354 (BOR), Oka 4089 (L). India: Tamil 

Nadu, Hale 43829. Japan: Prov. Aki, Hale 29529, 29540; Prov. 

Hizen, Kurokawa 63159 (TNS): Prov. Izu, Asahina (TNS): 

Prov. Mino, Asahiim (TNS); Prov. Ohmi, Hale 29472, 29482; 

Prov. Sagami, Asahina 40 (TNS), Kurokawa 58038 (TNS, US); 

Prov. Shimofusa, Asahina (TNS); Prov. Survo, Asahina 125 

(TNS). Taiwan: Kurokawa 494 (TNS). Australia: Victoria, 

Johnson (BM). New Zealand: Knight (UPS), Martin 493, 566 

(AM). 

40. Parmelina subaurulenta 

FIGURE 19e 

Parmelina siibaurulenta (Nylander) Hale, I974:483.

NUMBER 33 47 

Parrnelia subaurulenta Nylander, 1885:606 [type collection: 

Narkanda, 5. W’. Himalayas, India, Skoliczka (H, Nylander 

herbarium number 35672, lectotype)]. 

Parmelia homogenes Nylander, 38853607 [type collection: 

India, Hooker and Thotnson 1942 (H, Nylander herbarium 

number 35664, lectotype)]. 

Parmelia conspicita Hue, 1899: 1/15 [type collection: Kiao Che- 

Tung, above Kiang-l’n, Yunnan, China, Delavay (PC, 

lectotype)]. 

Parrnelia homalotera Hue, 1899:159 [type collection: above 

Mo-So-Yn, Yunnan, China, Delavay (PC, lectotype)]. 

Parrnelia fecrrnrla Hue, 1899:169 [type collection: Ta-Pin-Tze, 

Yunnan, China, Delavay 5 (PC, lectotype)]. 

Parmelia srcbcrernea Zahlbruckner, 1934b:208 [type collection: 

Betiveen Muli Gomba, Baurong, and T\’a-Erh-Dje, Setch- 

Ivan, China, Rock 16720 pro parte (I%’, 

lectotype)]. 

Parmelia subaurulenta var. myriocarpa Asahina, 19.5la:227 

[type collection: Mt. Koya, Prov. Kii, Japan, Inuinaru 

1232 (TSS, lectotype)]. 

Parmelia homogenes f. mitior Asahina, 1952:78 [type collectioil: 

Yamanaka, Prov. Kai, Japan, Asahitla (TNS. 

lect o t ype)]. 

Parmelia rnyriocarpa (Asahina) Chao, 1964:149. 

Parnielina hotnoge?res (Sylander) Hale, 1974:482 

DE~CRIPTION.-T~~~~L~S 

adnate to closely adnate on 

bark, light greenish mineral gray, rather fragile, 

4-1 0 cm broad; lobes sublinear, subimbricate, 2-4 

mm wide, the marginal cilia quite dense in the 

axils, irregularly dispersed at the lobe tips, 0.4-0.7 

mm long; upper surface plane, faintly maculate, 

shiny, isidia and soredia lacking: medulla pale 

orange-yellow; lower surface black, densely rhizinate, 

the rhizines simple or becoming squarrosely 

branched. Apothecia very numerous, adnate, 1-3 

mm in diameter; spores 8, 6-10 X 8-14 pm, usually 

abundantly developed 

CHExmTRY.-Cortex K + yellow (atranorin); medulla 

more intensively yellow with color tests 

(zeorin, leucotylin and associated terpenes, secalonic 

acid A, and traces of unidentified substances). 

DIsTRIiivTIoN.-Eastern Asia from India to Japan. 

RE:hfARKs.-Parmelina su bazirirlentn is characterized 

by a thin thallus (average thickness for 20 specimens 

154 pm), numerous apothecia rarely exceeding 

3 mm in diameter, and well developed spores 6-9 x 

8-12 pm. Nylander (1885:60i) described Paimelin 

homogenes at the same time on the basis of a single 

specimen, citing as the main difference spore size 

(“similis siibazmilenta . . . sed sporis majoribus”). 

Nylander measured spores 8-10 X 14-16 pm but my 

own measurements on the type specimen gave 

7-8 X 12-13 pm, putting it well within the range 

of spore size for Pal-rnelina subaurulenta. Without 

a significant difference in spore size, P. homogenes 

must be regarded as a synonym of P. siibauixlenta. 

The species most often confused with P. sitbaum- 

lenfn, especially outside of India, is P. if-rugans, 

which has consistently larger apothecia (up to 10 

mm in diameter), somewhat larger spores, and the 

“aurulenta” terpene series. T’wo of Nylander’s 

syntypes for Parmelia sztbaurulenta are, in fact, 

P. imgans, as discussed above under that species. 

A11 specimens should be tested with chromatog- 

raphy, although diameter of the apothecia is often 

adequate to separate the species in this group. All 

type specimens listed in the synomymy above were 

tested with thin-layer chromatography except for 

Pnmelia coiispiczta Hue, which is placed here on 

the basis of the small apothecia. 

SPECIMEKS Ex.iMIs.ro.-Sepal: Poelt 156 (hf). India: Sikkim, 

Hnra et al. (TSS, US); Tamil Sadu, Degelius As352 

(Degelius herbarium), Hale 40214, 43705, 43761, 43834, 

Perinftet (H, Nylander herbarium number 35670, syntype of 

Parrnelia srcbaurdeiita), Watt 119 (BM): TTest Bengal, 

.4wnsthi 3919, 3921 (Aivasthi herbarium. US). Sri Lanka: 

Almquist (H, Nylander herbarium number 35665). China: 

Yunnan, Handel-Mazzrtti 89, 2453, 2453 (US, JV), Rock 11747 

(US). Taiivan: Kntrokawn 873, 1208 (TSS). Japan: Afoseley 

(H, Nylander herbarium number 35669, syntype of Parmelia 

s7tbaurulen/n); .4lrnquist 

36671). 

(H, Nylantler herbarium number 

41. Parmelina subfatiscens 

FIGURE 19f 

Parinelina subfatiscens (Kurolawa) Hale, 1974:483. 

Parmelfa siibfafiscens Kuroka\\a in Hale and Kurokawa, 

1964:134 [t)pe collection: Louis Trichartlt, Zoutpansberg, 

Transiaal, Union of South Ahica, Al/nborn 6443 (LD, 

holotype; isotype in US)]. 

DEscRIPTIos.-ThallLis closely adnate on bark, 

fragile, whitish mineral gray, 3-5 cm broad; lobes 

sublinear-elongate, separate to contiguous, 03-15 

mm wide, the marginal cilia distinct, simple, to 1.0 

mm long: upper surface plane, shiny, continuous, 

pustulate laminally and subterminally (Figure 5c), 

exposed medulla in center of pustules turning black, 

pustules remaining entire or producing very coarse 

sorediate or isidiate-sorediate masses, often short 

ciliate in African material but bery rarely so in the 

New 1Yorld; medulla white; lower surface black, 

densely rhizinate, the rhizines simple, black. 

Apothecia rare, adnate, 1.5-4.0 mm in diameter; 

spores 8,8-9 X 12-14 pm.



dulla K -, C -, KC + rose, P - (trace of gyrophoric 

acid and the “horrescens” unknown substances). 

DISTRIBUTIoN.-~outh Africa and the Caribbean 

region. 

REMARKS.-The pustules of P. subfatiscens (Fig- 

ure 5c) often erupt, leaving a central area of exposed 

medulla that blackens. Other pustules become 

coarsely sorediate with the soredia sometimes as- 

suming the form of tiny isidia-like masses. The 

specimens from Africa usually produce short cilia 

on these isidioid masses and on the surface of the 

pustules. The tropical American population very 

rarely has any cilia. The chemistry of the neotrop- 

ical and African populations is, as far as I can 

determine, within the limits of the solvent systems 

available, identical and equivalent to that in P. 

horrescens. Dey (1975:433) suggests that the neo- 

tropical specimens, along with some from the 

southern Appalachian Mountains in the United 

States, may represent a new species, based in part 

on the absence of cilia on the pustules. It is also 

true that the ecological requirements seem to be 

different. The neotropical specimens were all col- 

lected in upland virgin rain forest at or above 500 

m elevation, whereas the African localities are in 

much drier forests. I prefer to keep a broader spe- 

cies concept until more complete morphological and 

ecological data can establish the case one way or the 

other. 

SPECIMFUS EXA\IINID.-Panama. Code, Hale 43564. See Hale 

(1971a:24) for additional records from Dominica. 

42. Parmelina swinscowii 

FIGURE 20a 

Parmelina swinscowii (Hale) Hale, 1974:483. 

Parmelia swinscowii Hale, 1973b:1 [type collection: Mt. Kenja, 

Central Province, Kenya, T. D. V. Swinscow K 31/33 

(BM, holotype; isotype in US)]. 

DEscRIPTIoN.-Thallus growing on soil or over 

mosses on soil, loosely attached, whitish mineral 

gray or darkening, fragile, 3-5 cm broad; lobes 

sublinear-elongate, imbricate, 1.5-2.0 mm wide, the 

margins sparsely ciliate, the cilia black, simple, 

about 0.5 mm long; upper surface plane to convex, 

shiny, becoming rugose and cracked toward the 

lobe tips, the tips remaining entire or becoming 

minutely dissected, turning coarsely pustulate- 

sorediate; medulla white; lower surface black, 

densely rhizinate, the rhizines black, simple. 

Apothecia unknown. 


dulla K-, C-, KC+ rose, P- (lobaric acid and 

salazinic acid). 

REMARKS.-sinCe describing P. swinscowii, 1 have 

identified it among collections made by Rolf 

Santesson in southernmost Chile. The Chilean and 

African populations are identical in morphology 

and chemistry. In Kenya P. swinscowii grows in the 

alpine zone at over 3500 m elevation, and in Chile 

it occupies similar barren habitats on rock and soil 

near sea level. 

As mentioned in the original description, P. 

swinscowii does not fit well into any of the parmeli- 

oid groups. It superficially resembles an “evernii- 

form” Parmelia, such as sorediate, salazinic acid- 

containing Everniastwm sorocheilum (Vainio) Hale, 

but the rhizines are relatively short and the lobes 

not at all canaliculate. It intergrades to a certain 

extent with abnormally small forms of Parmotrema 

cetratum (Acharius) Hale or P. reticulatum (Taylor) 

Choisy, except that the upper surface is not reticu- 

late and the lobes are sublinear. The species is 

probably best left in Parmelina where it is 

anomolous largely because of the presence of lobaric 

acid. 

SPECIVEM EXAhZISED.-chile: Magallanes, Santesson 2159, 

6381a, 6381b, 6478 (S, US); Tierra del Fuego, Santesson 1244 

(S, US). Specimens from Kenya are listed in Hale, 1973b:4. 

43. Parmelina tiliacea 

FIGURE 20b 

Parmelina tiliacea (Hoffmann) Hale, 1974:483. 

Lichen tiliaceus Hoffmann, 1784:96, pl. 16: fig. 2 [type collection: 

Europe (not seen but illustrated in Hoffmann)]. 

Lichen scorteus Acharius, 1798:119 [type collection: Sweden 

(H-Ach, lectotype)]. 

Parmelin tiliacea (Hoffmann) Acharius, 1803:215. 

Zmbricaria tiliacea (Hoffmann) Koerber, 1855:70. 

Parmelia quercifolia var. scortea f. microphylla Massalongo, 

1856: 175 [type collection: Campofontana, Verona, Italy, 

Massalongo in Lichenes Ital. 329 (UPS, lectotype)]. 

Parmelia scortea var. papillosa Gylenik, 1931:284 [type collection: 

Ndgrad, Hungary, Foriss 599 (BP, holotype)]. 

Pnrmelia scortea var. ramificn Gyelnik, 1931 :285 [type collection: 

S6sko Mountains, Hungary, Timkd 4521 (BP, 

holotype)]. 

Parmelia scortea f. visegradensis Gyelnik, 1932a:444 [type 

collection: Visegrld, Pest, Hungary, Timkd (BP, holotype)].

NUMBER 33 

49


DEscRIPTION.-Tha1lus adnate on bark or rock, 

light mineral gray to mineral gray, turning olive- 

buff to deep olive-buff in the herbarium, 5-15 cm 

or more in diameter: lobes irregularly branched, 

sublinear-elongate, often imbricate, rounded at the 

apices, 2-6 mm wide, the margins more or less 

crenate and undulate, sometimes narrowly black 

rimmed, ciliate, the cilia black, coarse, shiny, simple, 

0.2-0.7 mm long; upper surface more or less shiny, 

white maculate, usually partly pruinose, irregularly 

cracked on older lobes, densely isidiate, the isidia 

cylindrical, short, rarely branched, usually black- 

ening at the tips; medulla white; lower surface 

black, moderately to densely rhizinate, rhizines 

black, simple, 1-2 mm long. Apothecia rare, adnate, 

sometimes retrosely rhizinate, to 4 mm in diameter; 

spores 8, 5-6 X 8-11 pm. 


dulla K-, C+, KC+ red, P- (lecanoric acid). 

DIsTRIBuTIoN.-Europe to western India. 

REMARKS.-ThiS classical European species was 

the first Parmelina to be described. Except for the 

nomenclatural confusion with Acharius’ Parmelia 

scortea, it has been correctly identified by all 

lichenologists, excluding, of course, those in Amer- 

ica, who used the name for Parmelina galbina. 

Hypotrachyna liuida (Taylor) Hale, and other 

narrow-lobed lichens. 

Parmelina tiliacea is the isidiate morph of P. 

q uercina, which, surprisingly, has a much broader 

geographic range, contrary to the situation for most 

species pairs. A close relative in Britain and Europe 

is P. pastillifera, which has peltate isidia and differ- 

ent ecological requirements (Dobson and Hawks- 

worth, 1976). Taken together, the three species 

form a distinct group in Parmelina characterized 

by the white maculae and presence of lecanoric 

acid. Only one other species in the genus, P. 

pruinata, contains this acid. 

As one of the commonest parmelioid lichens in 

Europe, P. tiliacea has been mentioned and stud- 

ied in detail by many workers. Sernander-Du Rietz 

(1926), for example, found that it tends to occur 

in ornithocoprous or dust-laden habitats in Scan- 

dinavia. She later conducted a careful study of 

apothecial formation, showing that growth of new 

apothecia could be correlated with heavy precipita- 

tion in warm periods of the summer (Sernander-Du 

Rietz, 1957). 

SxczcIhfEvs EXAMINED (selected) .-Europe: England, Crombie 

(BM), Davies (BM), Larbalestzer 292 (BM); Norway, 

Hoeg (US), Magnusson 9193 (US); Sweden, Almborn (LD), 

Asplund (US), Blomberg in Lichenes Sueciae Exsiccati 64 

(LD), Santesson 12711 (US), Vrnng in Kqptogamae Exsiccatae 

Vindobonensi 3718 (LD, US); Finland, Kari in Lichenes 

Fenniae Exsiccati 194, 1103 (H), Linkoln in Lichenes Fenniae 

Ewsiccati 392 (H, US); Poland, Glanc in Lichenotheca Polonica 

171 (UPS); Russia, in Lichenotheca Rossica 17 (WIS), Oxner 

(US): Bulgaria, Szatala (US): France, Crozals (US), Rondon 

in Lichenes Selecti Exsiccati 440 (US); Germany, Erichsen 

(US), Hiltmann in Kryptogamae Exsiccatae Vindobonensi 

3062 (US). Paul and Schroppel in Lichenes Alpium 9 (US); 

Czechosloi akia, Pisut in Lichenes Slovakiae Exsiccati 73 

(US): Spain, Culberson 11954 (DUKE, US); Portugal, Sampaio 

in Lichenes de Portugal 250 (US); Italy, Steiner in 

Ki yptogamae Exsiccatae Vindobonensi 4327 (US). Israel: 

Pharaii 11 (US). Tunisia: Runemark (LD). Morocco: Ernst 

2206 (US), hrewboula 198 (BM). Bahrain: Ahrnad L162 (L, 

US). Pakistan: Iqbal 180 (L), 511,540 (US). India: Kashmir, 

Kapoor 991 (Awasthi herbarium), Kaitl 4006 (Awasthi herbarium), 

Watt 117, 455 (BM). 

44. Parmelina usambarensis 

FIGURE 2Oc 

Parmelina usambarensis (Steiner and Zahlbruckner) Hale, 

1964:483. 

Parmelia usambarensis Steiner and Zahlbruckner in Zahl- 

bruckner, 1926:524 [type collection: Lutindi, Tanzania, 

Brunnthaler (W, lectotype)]. 

Parmelia lneuigatoides des Abbayes, 1951:970 [type collec- 

tion: Fouta-Djalon, Dalaba, Guinea, des Abbayes (RES, 

lectotype)]. 

DEscRIPTIox.-Thallus loosely attached on rock, 

whitish mineral gray, to 10 cm broad; lobes broadly 

sublinear, divaricate to contiguous, 2-5 mm wide, 

the marginal cilia irregularly dispersed but mostly 

in the lobe axils, up to 1 mm long; upper surface 

shiny, plane, continuous or cracking in older parts, 

sparsely to densely isidiate, the isidia cylindrical, 

simple to branched, to 0.4 mm high; medulla 

white; lower surface black and shiny except for a 

narrow dark brown zone at the tips, moderately to 

sparsely rhizinate, the rhizines black, simple, 1 mm 

or more long. Apothecia rare, substipitate, the 

amphithecium isidiate, 2-3 mm in diameter; spores 

8,6 X 12pm. 

REMARKS.-PaYmelina usambarensis is a rather 

large, loosely attached, saxicolous lichen. The isidia 

are often quite sparse. It does not conform perfectly 

to the concept of Paymelina as I define the genus, 

but the simple rhizines and coarse, long marginal

NUMBER 33 51 

cilia at least remove it from Hypotrachyna (Vainio) 

Hale. On the other hand, it cannot be accommo- 

dated in Parmotrema LIassalongo because of the 

sublinear subdivaricate lobes. There are no close 

relatives, except, very superficially, the African en- 

demic Parmelina enormis. 

SPECIMFSS Ex \vIsm.-Guinea: Baldruin 9849a (BM) . Ivory 

Coast: Man, tles .4bbnyes (LD, REN), Santesson 10632 (S, 

US). Uganda: Su’inscow 2U 15/6 (BM, US), Thornas 1678, 

3029 (BM). Thailand: Kuroknwa 1804, 1813, 1939, 1956 (TXS), 

1958 (TSS, US), Kurokawa in 1-icheria Rariores et Criiici 

Exsiccati 41 (US). 

45. Parmelina versiformis 

FIGURE 2Od 

Parmeliiza versiformis (Krempelhnber) Hale, 1964:483. 

Pnvr!ielin uersiformis Krempelhul)er, 1878:464 [type collection: 

Argentina, Lorentz crnd Hieronymuy (M, lectotype: 

isolectotypes in G, PC)]. 

Parmeiia inuta/a Vainio, 1890.3‘) [t)pe collection: Sitio, 

Minas Gerais, Brazil, Vainio in Lichenes Brasilienses 

Exsiccati 539 (TUR, 1ectot)pe: isolectotypes in Bhl, FH, 

UPS)]. 

Parmelia catliari?ie,!sis Miiller Argoviensis, 1891:239 [type 

colleclion: Near Santa Catarina, Brazil, Uie 1891 (G, lectotype; 

isolectotjpe in \V)]. 

Parmelia weltsteinii Zahlbrnckner, 1909:173 [type collection: 

Near Taipas, Sao l’aulo, Brazil, Schiffner and Wettsteiii 

(W, lectotype, isolectotype in G)]. 

DE~CRIPTION.-T~~~~US adnate to loosely adnate 

on bark, turning deep olive-buff in the herbarium, 

about 8 cm in diameter; lobes irregularly branched, 

sublinear-elongate, more or less imbricate, 2-8 mm 

wide, the margins crenate, ciliate mostly in lobe 

axils, the cilia often sparse, to 0.5 mm long; upper 

surface more or less shiny, not maculate, wrinkled 

and irregularly cracked on older lobes, soredia and 

isidia lacking; medulla white; lower surface dark 

brown or blackening, more or less wrinkled, sparsely 

rhizinate, the rhizines pale brown, simple, 0.1-0.3 

mm long. Xpothecia common, substipitate, 2-10 

mm in diameter, the rim crenate and undulate, the 

amphithecium rugose, the disc blackish brown; 

spores 8, 10-14 X 18-28 pm, the episporium about 

3 pm thick. 

CHEmsmY.-Cortex K + yellow (atranorin); medulla 

K+ yellow turning red, C-, KC-, P+ 


DISTRIBUTION.-~OUth America. 

REMARKS.--This is the only species of Parmelina 

in the New World with salazinic acid, excepting 

rare P. min.rcowii. It is also unusual in having very 

large spores with a thick episporium as well as a 

lower surface that is dark brown to black rather 

than entirely black. The closest relative appears to 

be the Asian P. expallida, an isidiate species with 

smaller spores (Kurokawa, 19G8a: 192). Although P. 

veisiformis occurs in a botanically well-known re- 

gion, it has not been collected since 1901. 

Si~ec~si~,ss Ex.iwAFD.-Brazil: hhas Gerais, r’C’flTr7lifig 283, 

289, 286 (M); Rio de Janeiro, B~lrclzell 2409 (BM), Glaziou 

1822, 1823 (M). 

46. Parmelina wallichiana 

FIGURES 20e, 21 

Parmelinn wallichiniia (Taylor) Hale, 1974:483. 

Parmelia wallichiana Taylor, 1847:l 76 [type collection: Sepal, 

Wnlliciz (FH-TayI, lectotype)]. 

Parinelia tiliacea rar. fximfa Steiner, 1888: 138 [type collection: 

Usambara, Tanzania, Meyer (G, lectotype)]. 

Parmelia juflodii Steiner, 19073640 [type collection: Sanatorium, 

Cape Proiince, Union of South Africa, Junod 978 

(G, lectot!pe)]. 

Parmelia nimandaira,la Zahlbrnckner, 1934a:55 [type collection: 

Mt. rlrisan, Taiivan, Asahimi 63 (W, lectotype; isolectotype 

in TSS)]. 

Parrnelia nimandairana f. sirblaeui.~ Asahina, 19523131) [type 

collection: hlt. Buko, Prov. hfusashi, Japan, Asahiiza 35 

(TNS, lectotype)]. 

DEscRIPTioN.-Thallus adnate to loosely adnate 

on bark or rock, yellowish glaucous to pale 

glaucous-green, 5-20 cm in diameter; lobes irregu- 

larly branched, apically rotund, 3-10 mm wide, the 

margins more or less crenate, lobulate with age, 

short ciliate, especially in the axils, the cilia about 

0.5 mm long; upper surface shiny, not maculate, 

irregularly cracked on older lobes, sparsely isidiate, 

isidia cylindrical, mostly Simply, less than 1 mm 

high; medulla white; lower surface black, mod- 

erately to sparsely short rhizinate, dark brown and 

naked or papillate in a rather wide Lone near the 

tips, the rhizines black, simple. Apothecia rare, 

adnate, 2-7 mm in diameter; spores 8, 8-10 x 

14-18 Fm. 

CHEnmTRu.-Cortex K + yellow (atranorin); me- 



DISTRIBuTIoN.-Africa and Asia (Figure 2 1). 

REMARKS.-This is the most widespread and com-


monly collected Paimelina in the Old World. It is 

easily recognized by the adnate, moderately isidiate 

thallus that grows in a variety of habitats in the 

highlands of Africa, India, and eastern Asia, rang- 

ing from temperate Japan to both higher elevations 

in the tropics and to rather low elevations near 

the equator in Guinea and the Ivory Coast. The 

parent morph is no longer extant nor are the 

parallel sorediate or pustulate morphs. 

SPECIMENS EXAMINED.-GUinea: N’Zerekore, Santesson 10568b 

(UPS, US). Ivory Coast: Man, Santesson 10650 (UPS). Uganda: 

Dummer 5064 (US), Swinscow 2U 24/20 (BM, US). Kenya: 

Nyanza Province, Maas Geesteratius 10935 (L, LD, US), 4950, 

11164 (L); Rift \’alley Province, hfaas Geesteranus 4832 (L). 

Angola: Cuanza Sul, Degelius (Degelius herbarium, US). 

Rhodesia: Hoeg (TRH). Tanzania: Arusha Province, Santesson 

21339 (UPS, US); Kiliuiarijaro Province, Santesson 

20961, 20989, 20990 (UPS, US); Moshi District, Burnet T122 

(BM, US). Swaziland: Almborn 7901 (LD), Kofler (LD). Union 

of South Africa: Natal, Almborn 8494 (LD), 8683, 9732, 9871 

(LD, US), Hoeg (TRH), Kofler (LD); Transvaal, Almborn 

6430 (LD). Madagascar: des Abbayes in Licheiies Madagascarienses 

et Borbonici Selecti Exsiccati 10 (UPS, US, dis- 

tributed as Parmelia isidiza). Nepal: Awasthi 2156, 2157, 

2198, 2203 (Arvasthi herbarium), i\Torkett 5533 (BM), Togashi 

(TNS, US). Sikkih: Bose 136 (Awasthi herbarium). India: 

Uttar Pradesh, Awasthi 3175, 3509, 3814, 3980, 3986 (Awasthi 

herbarium), Hara (TNS, US); Tamil Nadu, Foreau 43 

(Awasthi herbarium), Hale 40220, 40274, 43620, 43740, 43870. 

Thailand: Kerr L4 (BM). Malaya: Pahang: Hale 30094, 

30217, 30218, 30219. Taiwan: Kurokawa 29, 530, 626, 2926 

(TNS). China: Kwantung Province, Petig 12578 (US); Man- 

churia, Asahina 25 (TNS). Hong Kong: Thrower 1586 (US). 

Japan: Prov. Aki, Hale 29508; Prov. Chikuzen, Kurokawa 

62545 (TNS); Prov. Higo Kiirokawn 63076 (TNS); Prov. Hizen, 

Kurokawa 63157 (TXS); I’rov. Hyuga, Hale 29645; Prov. Iyo, 

Kurokau~a 60098 (TNS); Prov. Iru, Asahina 94 (Th’S); Prov. 

Kii, Asahina (TNS), Kzirokawa 60237 (TNS, US), Kurokawa 

in Lichenes Rariores et Critici Selecti 42 (US); Prov. Musashi, 

Asahina 11 (TNS), Ktirokawa 59155 (TNS, US); Prov. Ohmi, 

Hale 29460; Prov. Satsuma, Kurokawa 63043 (TNS); Tosa, 

Fujikawa (TNS). Yoshimirra 1157 (US). Philippines: Benguet, 

Degelius As-787 (Degelius herbarium), Hale 26760, Hale 

in Lichenes Selecti Exsiccati 672 (US); Mountain Province, 

Haze 26413. Sabah: Hale 28707, 28807, 29063, 29071. Indonesia: 

Java, Groenhart 3238 (L, US), 4358, 5881, 5913, 5917 (L), 5948 

(L, US), Schiflner 3294 (US), Voogd 2079 (L).

NUMBER 33 53 

47. Parmelina xantholepis 

FIGURE 20f 

Parmeliria sun tholepis (Montagne and van den Bosch) Hale, 

1974:438. 

Parrne/ia xantholepis hIontagne and van den Bosch, 

1855:428 [type collection: Gode, Java, Teydom (L, 

lectotype)]. 

Parrnelia hiformis T’ainio f. dataensis Vainio [= f. biformis], 

1909:660 [type collection: Mt. Data, Luzon, Philippines, 

Alerrill 4987 (TUR, T’ainio herbarium number 2607, 

lectotype)]. 

Parmelia biformis f. pauniensis T’ainio, 1909:660 [type collec- 

tion: Pauai, Benguet, Luzon, Philippines, Mearns 4434 

(TUR, T’ainio herbarium number 2605, lectotype)]. 

DEscRIP’rIos.-Thallus adnate on bark or rock, 

yellowish mineral gray, very fragile, 4-15 cm broad; 

lobes sublinear to subirregular, 1-3 mm wide, 

crowded, dissected, the marginal cilia coarse, 0.2-0.8 

mm long; upper surface plane, maculate, becoming 

densely lobulate toward the lobe margins, isidia 

and soredia lacking; medulla barium yellow; lower 

surface black, densely rhizinate, the rhizines black, 

long, simple or squarrose. Apothecia adnate, 1-3 

mm in diameter; spores 6-8 X 9-13 pm. 

CHEXIISTRY.-cOl.teX K + yellow (atranorin); me- 

dulla more intensively yellow with K and C, P- 

(zeorin, leucotylin and related terpenes, and seca- 

lonic acid A.) 

DISTRImTIoN.-hdia and Nepal to the Philip- 

pines. 

REM.mKs.-This species is easily identified by the 

fragile, lobulate thallus with a pale yellow-orange 

medulla. It is closely related to the P. amagiensis-P. 

clenegans series, definitely in chemistry but less so in 

thallus texture. It is usually collected at the base of 

trees or on rocks, often growing among mosses, at 

higher elevations (1000-2300 m) in evergreen hard- 

wood cloud forests. 

SPECIMESS Ex \MIsIm-h’epaI: Poelt 101, 111 (hI) , Norkett 

9227 (BM). India: Tamil Nadu, Foreau 4118 (ALvasthi herbarium, 

US), Hale 43862; Uttar Pradesh, Awasthi 3839 

(Awasthi herbarium). Thailand: Kziroknwa 1669 (TNS, US). 

Philippines: Mountain Province, Hale 25993, 26096, 26187, 

26344, 26554. Indonesia: Java, Groenhart 2841, 5939 (L), 

5941, 6004 (L, US). 

Doubtful and Rejected Names 

Parmelia coilocarpa 

Parriwlia coilocnrpn Stirton, 1877-78:202 [t)pe collection: 

Feinando Po, West Africa, G. Thornson (Bhf, lectotype; 

CLAM, isolectotype)]. 

The type material is too fragmentary for ade- 

quate study. It is a nonsorediate, nonisidiate, sub- 

irregularly lobed species with a black lower surface, 

simple rhizines, and numerous marginal cilia. The 

spores are large, 15 X 28 pm. Atranorin and sala- 

zinic acid are present. It is almost certainly a 

Pnrmelina, and its status will only be clarified as 

more collections are made in TYest Africa. 

ParmeIin orchidophlia 

Pnrnielin orcliidoljhila Dodge, 1953:374 [type collection: 

Nyinabitsa, TVestern Pro\-ince, Uganda, Omastin 118.1 

(Bhf, lectot! pe)] 

The type collection is too Eragmentary for study. 

Salazinic acid was proved niicrochemically. It may 

be Parmelina zunllichiann (Taylor) Hale. 

Parmelia tiliacea var. lezicina 

Parmelin tilicea Tar. leucintr Miillrr Argoviensis, 1880:267 

[type collection: Sear Petropolis, Brazil, Deventer (G, 

lec totype)]. 

This variety is equal to Hypotrachyna dactylifera 

(Vainio) Hale (Hale, 1975a:30). 

Pal-melia tilacea var. rimzilosa 

Pnrrrielin tilicea tar, rriricilosa Muller Aigoiiensis, 1882:458 

[tJpe collection: Socotra, h’nlfoirr (C, lectotipe)]. 

No type specimen could be located at Geneva. 

One of two specimens so labeled by hluller is Par- 

motrema reticiilatum (Taylor) Choisy, and the 

other is Pseudoparmelia carneopruinata (Zahl- 

bruckner) Hale. 

Pal-melina bagziioensis 

Panneliizu bnguioensis (Hale) Hale, 1974:482 [type collection: 

Baguio, Mountain Province, Philippines, Ha/e 26768 (US, 

holotype)]. 

Closer examination of this sorediate species con- 

vinced me that it is a member of the genus Hypo- 

tl-achyna (H. bagtrioensis (Hale) Hale, new combi- 

nation). The rhizines are rather sparse but clearly


dichotomously branched at maturity. In any event, 

the chemistry-fumarprotocetraric acid-was ano- 

malous for Parmelina, and this was the first clue to 

its correct generic status. 

Parmelina carporrhizans 

Parmelina carporrhizans (Taylor) Hale, 1974:482. 

I now consider this to be a synonym of Parme- 

lina quercina. 

Parmelina crystallorum 

Parmelina crystallorum (Lynge) Hale, 1974:482 

This species is synonymized under Parmelina 

damaziana. 

Parme 1 ina h om ogen es 

Parmelina homogenes (Nylander) Hale, 1974:482. 

I have placed this species in synonymy under 

Pa rm e 1 in a szi bau ru 1 en t a. 

Parmelina nylanderi 

Parmelina nylanderi (Lynge) Hale, 1974:483. 

Parmelia nylanderi Lynge, 1914:82 [type collection: Sear $450 

Jeronyno, Serra da Chapada, Mato Grosso, Brazil, Malme 

2747 (S, lectotype)]. 

I originally considered this loosely attached saxi- 

colous lichen to be a member of Parmelina, but 

after examining a second collection (Eiten 3429 

(US) from S5o Paulo, Brazil), I now feel that the 

chemistry (usnic acid in the cortex and salazinic 

and gyroplioric acid in the medulla) and general 

habit, in spite of the narrow ciliate lobes, place it 

more appropriately in Parmotrema (P. nylanderi 

(Lynge) Hale, new combination). It is related to 

the much broader lobed P. delicatulzim (Vainio) 

Hale group so common on sandstone in South 

America.

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3:1-234. 74: 193-21 3.

lmbricaria quercina, 42 

tiliacea, 48 

Lichen quercinus, 42 

tiliaceus, 48 

Par in elia a 1 b ido -s tra min ea, 19 

amagiensis, 18 

ainazonica var. hirsnotii, 26 

antillensis, 18 

atricha, 42 

aurulenta, 19 

halansae, 22 

balansae \ar. sorediata, 40 

biformis f. dataensis, 53 

biforinis f. pnuaiensis, 53 

blastica, 18 

brachpconidia, 23 

brachsconidia \ ar. chlorocarpa, 23 

budapestiiiensis, 42 

ca m t rchadalis i'ar. epiph y lla , 26 

carporrhizans, 42 

carporrhizans f. inalicola, 42 

catharinensis, 51 

coilocarpa, 53 

consors, 22 

conspiciia, 47 

con t in en talis, 22 

cryptochlora, 23 

crystalloruin, 23 

damaziana, 23 

deminuta, 30 

denegans, 25 

dissecta, 25 

endoleuca, 27 

enormis, 27 

entotheiochroa, 28 

expallida, 28 

fecunda, 47 

finkii, 38 

fraudans ssp. subfraudans, 35 

galbina, 30 

galbina var. rugosa, 30 

galbina 1-ar. sitbradiata, 30 

hayachinensis, 31 

heteroloba, 31 

homalotera, 47 

homogenes, 47 

homogenes f. minor, 47 

horrescens, 3 1 

Index 

(Synonyms in italics) 

hubrichtii, 26 

hrinanensis, 19 

immiscens, 32 

insinuata, 34 

insin itat ula, 34 

irrugans, 34 

jarnesii, 35 

junodii, 51 

laruignta ssp. estremi-orienialis, 30 

iaeuigata var. gracilis f. furfuracea, 25 

laevigatoides, 50 

lericotylita, 35 

leucotqlix f. rugulosa, 35 

leiico/ylizn f. siiblaevis, 35 

lin d ilia nii, 36 

mrlaiiochneta, 36 

nietnrevoiiita, 36 

inichoncnnensis, 32 

ni inn JIL m, 26 

riii/el/eri, 38 

uiiitata, 51 

myriocnrpa, 47 

11 ima nda i ra n a, 5 1 

iiimaiidairnna f. sublaevis, 51 

nylanderi, 54 

obsessa, 36 

orchidophila, 53 

pastillifera, 39 

perisidians, 39 

phlyctina, 40 

pilosa, 40 

pruinata, 42 

piiiggari, 26 

quercifolia var. scrotea f. inicrophylla, 

48 

quercina, 42 

querrina var. stclphurosa, 30 

revolnta i.ar. granulata, 42 

sampniana, 22 

.scortea var. papillosa, 48 

scortra var. pastillifera, 39 

scortea iar. ramifica, 48 

scortea var. visegradensis, 48 

silvestris, 19 

simplicior, 44 

sinuosa var. hypothrix, 42 

spathulata, 46 

sp umosa, 46 

59 

subaurulenta, 47 

siibaurulenta var. inyriocarpa, 47 

subbalansae, 40 

subcremea, 47 

sribfatisceiis, 47 

subquercifolia, 30 

subqiiercifolia f. Aiibradiata, 30 

subreuolitta, 19 

subsz~lphitrata, 39 

tilracea. 48 

tiliacea \ ar. afjixa, 42 

tilicicea Tar. efflorescens, 19 

tiiiacva var. exirnin, ,51 

tiliacen ~ar. hsmthri\, 42 

tiiiacra \ar. leuciiia, 53 

tiliacea var. minor, 30 

ti/iacen \ar. rimiilosn, 53 

t iliacea su bq ii r rci folio, 30 

tilinrea var. siiipliurouz,, 30 

tiliaren I ar. siilphurosa f. asperata, 36 

usariibarensis, 50 

versiforniis, 5 1 

wailichiaiia, 51 

wettsteinii, 51 

yalungana, 42 

vnnthorarpa, 34 

santholefiis, 53 

Parmelina amagiensis, 18 

antillensis, 18 

aurulenta, 19 

baguioensis, 53 

carporrhizans, 54 

consors, 22 

crassata. 22 

cr) ptochlora, 23 

crystallorum, 54 

tlamaziana, 23 

degelii, 25 

tlenegans, 25 

dissecta, 25 

endoleuca, 27 

enormis, 27 

entotheiochroa, 28 

expallida, 28 

galbina, 30 

hayachinensis, 31 

heteroloba, 31 

homogenes, 47, 54

horrescens, 31 

immiscens, 32 

indica, 34 

jamesii, 35 

leucotyliza, 35 

lindmanii, 36 

melanochaeta, 36 

metarevoluta, 36 

muelleri, 38 

nylanderi, 54 

obsessa, 38 

pastillifera, 39 

perisidians, 39 

phlyctina, 40 

pilosa, 40 

pruinata, 42 

quercina, 42 

rhytidodes, 43 

schindleri, 44 

simplicior, 44 


spathulata, 46 

spumosa, 46 

suhaurulenta, 46 

suhfatiscens, 47 

swinscowii, 48 

tiliacea, 48 

usamharensis, 50 

versiformis, 51 

wallichiana, 51 

xantholepis, 53

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