STUDIES ON PARMELIA SUBGENUS PARMELIA 
By Mason E. Hale, Jr., and Syo Kurosawa 
Introduction 
At least 250 distinct species, most of which occur in the tropics, 
are included in Parmelia subgenus Parmelia. They are generally 
characterized by lobes that are narrow (0.5-4.0 mm. wide), sublinear 
to subirregular, and often apically truncate. The apothecia are adnate 
and usually imperforate, and there are rhizines over all or most of 
the lower surface. This broadly delimited group has usually been 
called section Hypotraehyna Vain., but we propose to recognize it as 
a subgenus, subgenus Parmelia, typified by P. saxatilis (L.) Ach. 
Subgenus Parmelia is coordinate with subgenus Amphigymnia 
(Vain.) Dodge, the broad-lobed Parmelias, and subgenus Xaiitko- 
parmelia (Vain.) Hale, comb, no v., based on section Xanthoparmelia 
Vain. (1890, p. 60), with Parn elia conspersa (Ach.) Ach. as the type 
species. The Xanthoparmelias are similar in configuration to many 
species in subgenus Parmelia, but they differ in being saxicolous, 
always containing usnic acid, and having simple rhizines. The brown 
Parmelias (sections Melanoparmelia Zahlbr., Vainioellae Gyel., and 
Olivascentes (Hue) Hillm.) are excluded from subgenus Parmelia, 
although we have not yet decided on their exact position in the genus. 
In the course of preparing a world monograph of subgenus Parmeliat 
we have examined the types of most of the described species and 
reevaluated the subgeneric classification. This preliminary study 
will present the outlines of a new sectional classification, descriptions 
of 52 new species, 2 new combinations, and 4 new names, and pre- 
liminary keys to the species in the major sections. A final mono- 
graph must await more extensive study of general herbarium material 
as well as field work, since many species are still known only from their 
type localities. 
This study has been made possible by the prompt and generous 
cooperation of the curators and directors of museums and university 
herbaria in lending type specimens and other valuable collections and 
in rendering assistance and providing facilities for study during visits. 
They include Dr. Sten Ahlner (Naturhistoriska Riksmuseet, Stock- 
121 
122 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
holm), Dr. Reino Alava (Botanical Institute, Turku), Dr. Ove 
Almborn (Botanical Museum, Lund), Dr. Y. Asahina (Research 
Institute for Natural Resources, Tokyo), Dr. Charles Baehni (Con- 
servatoire et Jardin Botaniques, Geneva), M. P. Bourrelly (Museum 
National d'Histoire Naturelle, Paris), M. Skytte Christiansen (Botan- 
ical Museum, Copenhagen), Dr. G. Cufodontis (University of Vienna, 
Vienna), Dr. W. L. Culberson (Duke University, Durham), Dr. H. 
des Abbayes (University de Rennes, Rennes), Dr. Henry Imshaug 
(Michigan State University, East Lansing), Mr. Peter James (British 
Museum, London), Dr. A. Kostermanns (Herbarium Bogoriense, 
Bogor), Dr. I. M. Lamb (Farlow Herbarium, Cambridge), Dr. R. A. 
Maas Geesteranus (Rijksherbarium, Leiden), Dr. J. Millar (Chicago 
Natural History Museum, Chicago), Dr. G. Moggi (Istituto Botanico, 
Firenze), Dr. E. Mfiller (Institut fur Spezielle Botanik, Zurich), C. E. 
Palmar (Glasgow Art Gallery and Museums, Glasgow), Dr. J. Poelfc 
(Botanische Staatssammlung, Munich), Dr. K. Rechinger (Natur- 
historisches Museum, Vienna), Dr. C. Rogerson (New York Botanical 
Garden, New York), Dr. H. Roivainen (Botanical Museum, Helsinki), 
Dr. Rolf Santesson (Institute for Systematic Botany, Uppsala), Sir 
George Taylor (Royal Botanic Gardens, Kew), Dr. J. W. Thomson 
(University of Wisconsin, Madison), Dr. K. Verseghy (Museum of 
Natural History, Budapest), Dr. R. Vincenzo (Istituto Botanico, 
Rome), and Dr. R. Woodson (Missouri Botanical Garden, St. Louis). 
Drs. G. Degelius and D. D. Awasthi (abbreviated DEGEL and 
A WAS respectively in the list of species) have kindly sent us their 
valuable private collections. Dr. Z. Chernohorsk^ arranged for loans 
of Gyelnik's collections and kindly provided facilities for study in 
Prague. Mr. M. Wirth assisted in the early work of chemical testing. 
Drs. Rolf Santesson and Ove Almborn have kindly read various parts 
of the manuscrip t and given invaluable aid in problems of sectional 
classification. 
This study has been supported in part by a grant from the National 
Science Foundation. 
Morphological Characters 
The following are the more important morphological characters 
that we have found in subgenus Parmelia. Some of them have been 
overlooked or poorly understood by previous workers and will be 
discussed in detail. 
1. Rhizines: Are produced more or less uniformly over the entire 
lower surface in the majority of species. Certain species with sub- 
rotund subirregular lobes, however, may have a narrow though distinct 
bare or papillate zone around the margins. Three major types of 
rhizines may be distinguished: simple and unbranched or rarely 
HALE & KUROKAWA—SUBGENTJS PARMELIA 123 
sparsely furcate; dichotomously and usually richly branched; and 
squarrosely branched (fig. X). Branching of the rhizines has proved 
Figure 1.—Schematic drawings of the three types of rhizines: Simple, squarrosely branched, 
and dichotomously branched (left to right). 
to be one of the most important characters on which our proposed 
sectional classification is based. 
2. Cilia: Two types of ciliate outgrowths occur on the margins of 
lobes, simple cilia and bulbate cilia. Simple cilia, comparable to those 
known in Anaptychia or Cetraria, occur in at least 34 species of sub- 
genus Pannelia (pi. 1). These same species usually have a black lower 
surface and simple rhizines (e.g., P. dissecta Nyl. and P. tiliacea 
(Hoffm.) Ach.). Bulbate cilia differ from simple cilia in having a 
conspicuously inflated base. This structure (pi. 1) is known in some 
45 species, all of them tropical. It has not been consistently recog- 
nized by lichenologists, although commemorated in names such as 
P. circumnodata Nyl. Bulbate species may have branched or un- 
branched rhizines and a pale or black lower surface. Bulbate cilia 
may merely be modifications of simple cilia, but they are distributed 
around the lobes with much greater regularity. Chemical data 
(table 1) indicate that simple-ciliate and bulbate-ciliate species are 
far more closely related to each other than to any other groups in 
the subgenus. Both types of cilia are probably related to rhizines 
and, as in the case of rhizines, are of very great importance in sub- 
generic classification. 
3. Lobules: These structures have usually been confused with 
isidia. They apparently originate as isidia but soon begin to flatten 
and grow horizontally (cf. Parmelia ensijolia, pi. 7). At maturity 
they are more or less dorsi ventral and often cilia te. Lobules axe 
rather rare and best seen in P. cultnigena Zahlbr., P. digitate Lynge, 
P. horrescens Tayl., or P. ensifolia Kurokawa. 
4. Pustules: Are coarse papillar outgrowths from the upper cortex. 
They are extremely fragile and seem to lack the thalloid structure of 
isidia. They may remain intact, as in P. dactylijera Vain., or at 
length erupt or burst open apically and form dense pustular masses, 
with little or no tendency to become sorediate (pi. 1). If coarse 
124 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
soredia form, they are not so dense as to obscure the basic pustular 
structure. Typically pustulate species include P. aurulenta Tuck., 
P. endocMora Leight., P. formosana Zahlbr., and P. spumosa Asah. 
5. Maculae: Are submacroscopic white dots uniformly scattered in 
the upper cortex. They are apparently caused by irregular clumping 
of algal colonies in the gonidial layer. They are at times difficult to 
recognize, and most lichenologists have in fact overlooked them. 
Maculae are nevertheless important, especially at the species level. 
They occur in such well-known species as P. laevigata (Sm.) Ach. and 
P. tiliacea (Hoffm.) Ach, 
6. Pseudocyphellae: Here we refer to actual pores in the upper 
cortex. There are apparently two different kinds. One is found in 
species related to P. barren (Sm.) Turn., where the pores are distinctly 
orbicular or elliptical and scattered irregularly in the upper cortex 
(pi. 1). They are correlated with the presence of simple rhizines 
and P— acids (gyrophoric acid, lecanoric acid, and fatty acids). 
A second type of pore is found in species related to P. sazatUis (L.) 
Ach., where the pores are angular or effigurate and are arranged in 
more or less definite patterns on the cortex and margins of lobes 
(pl. 1). These species usually contain a P+ acid (protocetraric or 
salacinic acid) and have a black lower surface with squarrosely 
branched rhizines. 
7. Reticulation: The upper cortex of a small group of species 
related to P. reticulata Tayl. is finely reticulately maculate and 
fissured (pl. 1). Initially the cortex is maculate to the margins of 
the lobes and at maturity actual Assuring usually occurs. Many 
species of Parmelia are irregularly or more or less reticulately cracked 
on older lobes, but young lobes lack any reticulation. Reticulate 
species always have a black lower surface and simple to squarrosely 
branched rhizines. 
8. Isidia and soredia: These two characters are well known to 
lichenologists through the work of Du Rietz (1924a) and require no 
further discussion here. They are valuable species characters in 
Parmelia but have no value at the sectional level. 
9. Apothecia: These characters offer little basis for classification 
in subgenus Parmelia. Spores are uniformly small, generally only 
6-18 n in length, and attain a length of 30 p in only a few species such 
as P. leucopis Kremplh., P. mntata Vain., and P. versijormis Kremplh. 
Except for P. cetrata Ach., P. homotoma Nyl., and P. reticulata Tayl., 
the apothecial disc is never perforate. There is little significant varia- 
tion in degree of adnation on the thallus, except in such unusual 
pedicellate species as P. peruviana Nyl. Coronate exciples and basal 
retrorse rhizines are often found on the apothecia of species with 
w 
HALE & KUROKAWA—SUBGENUS PARMELIA 125 
bulbate cilia. Parmelia carporrhizans Tayl., a ciliate species, is 
densely rhizinate at the base of the apothecia. 
10. Anatomy: The internal anatomy is for the most part uniform. 
The total thickness of the thallus varies from about 60 & to 250 ft; 
the cortex is often thinner and more fragile than in subgenus 
Ampkigymnia. Peculiar moniliform cells are found in the medulla 
of P. galbina Ach., P. obsessa Ach., and P. metarevolvta Asah., three 
closely related species (cf. Asahina, 1952, p. 98). Future workers 
may place greater reliance on internal characters, but for the time 
being this approach seems unrewarding. 
Chemical Characters 
The taxonomy of subgenus Parmelia cannot be studied fully with- 
out the use of chemical tests. This does not imply that we are 
accepting or deliberately describing so-called "chemical species." 
Anyone who works with a genus such as Parmelia, which has a wide 
Table 1.—Number of species in the various sections and subsections containing 
19 important chemical substances. 
Section 
Parmelia 
Section 
Irregu- 
lares 
Section 
Imbrlcarla 
Section 
Cyclo- 
chella 
Section 
Hypotra- 
chyna 
o 
o 23 
ft ^ 
sa 
|s» 
00 
h3 
ii 
to 
ii 
a 
11 
P 
la 
; s1-1 
: 00 
Number of species 
Gyrophoric acid 
Lecanoric acid 
Salacinic acid 
Stictic acid 
Norstictic acid 
Protocetraric acid 
Usnic acid 
Rhodophyscin 
K— pigments 
Olivetoric acid 
Crypto clorophaeic acid 
Evernic acid 
Divaricatic acid 
Perlatolic acid 
Alectoronic acid 
Barbatic acid 
Lichexanthone 
KC+ unknown ("livida") 
30 
28 
2 
25 
5 
10 
10 
7 
1 
46 
7 
3 
10 
2 
4 
3 
18 
33 
4 
4 
4 
2 
2 
1 
1 
11 
46 
2 
3 
5 
12 
9 
3 
5 
1 
8 
1 
1 
1 
83 
6 
» [5] 
8 
1 
5 
15 
12 
Q 
5 
4 
1 
5 
7 
11 
7 
5 
• Lecanoric acid in section Hypotrachyna la an accessory component with evemic acid. 
126 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
variety of color tests and chemical constituents, soon realizes that 
the lichen acids are absolutley indispensable aids in species identi- 
fication. We have tested all specimens with the usual microchemical 
methods of Asahina and Shibata (1954). Detailed discussions of 
problematic or unidentified substances encountered will be delayed 
until a final monograph is completed. In the meantime, however, 
we have prepared a preliminary table showing the number of species 
within each section or subsection containing each of 19 identifiable 
lichen substances (table 1). Certain noteworthy features of this 
table will be discussed below under the various sectional headings. 
The color of the thallus and medulla is described for many of the 
species according to Ridgway's Color Standards and Color Nomen- 
clature (1912); such colors are indicated by the notation R in paren- 
theses. 
Subgeneric Classification 
A classification for subgenus Parmelia is based on results from our 
own morphological and chemical studies and on reviews of previously 
proposed classifications, especially those of Vainio (1890, 1923), 
Gyelnik (1932), and Asahina (1952). 
1. Section Parmelia 
Subsection Parmelia 
Subsection Simplices Hale & Kurokawa 
2. Section Irregvlarea (Vain.) Vain. 
3. Section Imbricaria (Schreb.) Fr. 
Subsection I mbricaria 
Subsection Bicornvtae (Lynge) Hale & Kurokawa 
Series Bicornutae 
Series Relicinae Hale & Kurokawa 
4. Section Cyclocheila (Vain.) Rag. 
5. Section Hypotrachyna Vain. 
Typification, descriptions, and discussions of these taxa are 
presented below under appropriate sectional headings. Keys to 
species in the major sections are included whenever possible in order 
to show the relationships of the new species to those already known, 
The keys do not include a small number of rare problematic species 
that have not yet been characterized. It is anticipated of course 
that more new species and revisions of names will be made in the final 
monograph. It should be emphasized that all species names in the 
keys are based on the actual study of holotype or lectotype specimens. 
Key to Sections 
1. Cortex pseudocyphellate, with or without effigurate maculae; rhizines simple 
or squarrose 1. Section Parmelia (p. 127) 
HALE & KUROSAWA—SUBGENUS PARMELIA 127 
1. Thallus lacking pseudocyphellae; rhizines simple, dichotomously or squarrosely 
branched, 
2. Lobes more or less rotund; upper cortex uniformly reticulately maculate 
or fissured to the margin ... 2. Section Irregulares (Vain.) Vain. (p. 129) 
2. Lobes sublincar and truncate to subrotund; upper cortex continuous, 
fissured only on older lobes. 
3. Cilia or bulbate cilia present on margins of lobes. 
3. Section Imbricaria (Schreb.) Fr, (p. 130) 
3. Cilia absent. 
4. Rhizines simple 4. Section Cyclocheila (Vain.) Ras. (p. 147) 
4. Rhizines dichotomously branched. 
5. Section Hypotrachyna Vain, (p. 159) 
Subgenus Parmelia 
Type species: Parmelia saxatUis (L.) Ach. 
Thallus appressed to adnate, rarely subascending; lobes sublinear 
to subirregular, rarely broad and rotund, the margins entire and 
smooth, ciliate, or bulbate-ciliate; under surf ace black or brown, 
uniformly rhizinate to the margins, or, in species with rotund lobes, 
with a narrow bare or papillate zone, the rhizines simple and un- 
branched, dichotomously branched, or squarrosely branched. 
Apothecia more or less adnate, rarely pedicellate, the disc imperforate 
(except in section Irregulares); spores simple, suborbicular to ellip- 
soid, usually less than 20 ft long. 
Parmelia has been proposed as a nomcn genericum conservandum 
with P. saxatUis (L.) Ach. as the lectotype species (cf. International 
Code of Botanical Nomenclature, p. 220, 1961). This has been the 
unanimous choice as a lectotype for the genus (cf. Dodge, 1959), 
and we see no objections to it. Parmelia sa&atUis is therefore the 
type of subgenus and section Parmelia. 
1. Section Parmelia 
Thallus adnate; lobes sublinear, rarely subirregular and subrotund; 
upper cortex pseudocyphellate, with or without effigurate maculae; 
lower surface black or brown, the rhizines simple or squarrosely 
branched. 
Imshaug (1957) placed section Hypotrachyna Vain, in synonymy 
under section Parmelia. As will be shown below, Vainio's section 
is based on P. brasiliana Nyl., a tropical species with dichotomously 
branched rhizines and no pseudocyphellae. 
As we previously indicated, there are two quite distinct types of 
pseudocyphellae in this section, one typically elliptical and correlated 
128 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
with simple rhizines and the other effigurate-maculate and correlated 
with squarrosely branched rhizines. They are best treated as two 
subsections, Parmelia and Simplices, as follows: 
Subsection Parmelia 
Section Hypotrachyna subsection Myeloleuca Asah. (1952, p. 24). Lecto- 
type: P, eaxatilis (L.) Ach. 
Upper cortex with cffigurate maculae; lower surface black, the 
rhizines squarrosely branched or simple. 
Asahina's subsection Myeloleuca included all species of subgenus 
Parmelia lacking medullary pigments. We feel that the most suitable 
choice of a lectotype for this extremely broad subsection would be 
P. saxatUis. According to the International Code of Botanical 
Nomenclature (Art. 22), the rule of tautonomy applies only to the 
subgeneric and sectional ranks. Notwithstanding, we prefer to retain 
the spirit of the Code and use Parmelia as a subsectional epithet as 
well, even though Asahina's subsection Myeloleuca is validly published 
and synonymous. 
As we have delimited this subsection, it contains about 30 species. 
The upper cortex is often ridged with the effigurate pseudocyphellae 
forming patterns along the ridges (cf. pi. 1, fig, 4). As far as is known, 
all species contain salacinic acid, except P. insensitiva (Magn.) Anders, 
and P. psevdosulcata Gyel., which contain protocetraric acid. Most 
of the species are confined to montane or subboreal areas in the 
Northern and Southern Hemispheres, with the greatest abundance in 
eastern Asia, where at least 15 species have been reported. Most of 
the Asian species have been summarized by Asahina (1952). There 
is a small group of Australian species, including P. tenuirima Tayl. 
and P. signata Nyl., which have not yet been fully investigated. The 
common American and European species P. jraudans Nyl., P. 
omphalodes (L.) Ach., P. eaxatilis (L.) Ach,, and P. svlcata Tayl. are 
well known. 
Subsection Simplices Hale & Kurokawa* subsect* nov. 
Thallus superno pseudocypliidlatus, pseudocyphellis suborbiculari- 
bus vel ovalibus, subtus niger vel pallide castaneus, rhizinis 
simplicibus. 
Type species: Parmelia borreri (Sm.) Turn. 
This subsection contains at least 25 species, but their taxonomy has 
not yet been completely worked out. They are found commonly in 
savannas or dry temperate forests. The pseudocyphellae are similar 
to those of P. cetrarioides (Duby) Nyl., an Amphigymnia species. 
The rhizines are always simple and quite long. The chemistry is 
extremely simple, the medulla reacting P—,C+ rose or red (gyrophor- 
HALE & KUROKAWA—SUBGENXJS PARMELIA 129 
ic or lecanoric acids), or C— (fatty acids). There is very strong 
correlation between the color of the lower surface and chemical 
constitution. All species with lecanoric acid, including the well- 
known P. subrudecta Nyl. (P. dubia (Wulf.) Schaer.) and P. rudecta, 
Ach., have a pale brown lower surface; and all species with gyrophoric 
acid, such as P. borreri (Sm.) Turn. (P. pseudoborreri Asah.) and P. 
borrerina NyL, have a jet-black lower surface. Those with fatty acids 
have either a pale (e.g., P. bolliana Mull. Arg. and P. canalicidata 
Lynge) or a black lower surface (e.g., P. appalachensis Culb. and 
P. microsticta Mull. Arg.) (cf. Culberson, 1962). 
2. Section Irregulares (Vain.) Vain. (1923, p. 34) 
Section Hypotrachyna * Irregularis Vain. (1890, p. 38). 
Section Reticulatae Du Rietz (1924b, p. 331). 
Lectotype species: Parmelia reticulata Tayl. 
Section Hypotrachyna subsection Irregulares (Vain.) Gyel. (1932, p. 224). 
Section Hypotrachyna subsection Irregulares (Vain.) Hillm. (1934, p. 188). 
Superfluous combination. 
Type species: Parmelia cetrata Ach. 
Thallus loosely adnate; lobes subirregular, usually broad and rotund ; 
upper cortex reticulately maculate and at maturity reticulately fissured 
to the margins; lower surface black, the rhizines simple or at maturity 
squarrosely branched. Apothecia more or less substipitate, the disc 
perforate or imperforate, 
Vainio first proposed Irregulares in 1890 as a group name under 
section Hypotrachyna without a designated rank. The obvious lecto- 
type is Parmelia cetrata Ach. He proposed two other groups of 
equivalent rank, Cyclocheila (lectotype P. amazonica Nyl.) and 
Sublinearis (lectotype P. brasiliana Nyl.). These three groups were 
separated by subtle and overlapping differences in lobe configuration 
and presence of rhizines, papillae, or a narrow bare zone below along 
the margins. No lichenologist has successfully employed this classi- 
fication and most recent workers in fact have rejected it (cf. Asahina, 
1952; Maas Geesteranus,1947). 
Almost all lichenologists except Rasanen (1943), however, have 
overlooked the fact that Vainio radically revised his own classification 
in a study of Philippine lichens (1923). He eliminated section 
Hypotrachyna and proposed in its place two sections, section Irregu- 
lares (including the former Cyclocheila and Irregularis) and section 
SvMineares. This realignment more or less successfully circumvented 
the difficulties of his former classification by recognizing only two 
general types of lobe configuration, subirregular and apically rotund 
and sublinear and truncate. 
Du Rietz* section Reticulatae was set up parallel with section 
Hypotrachyna Vain, and section Amphigymnia Vain. Du Rietz based 
130 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
the section not only on P. cetrata and P. reticulata but also on P. 
perforate, (Jacq.) Ach., which is heavily maculate but lacks reticulate 
fissures. If P. perforate, is excluded and placed in subgenus Am- 
phigymnia, section Reticnlatae corresponds exactly with section 
Irregvlares. 
This small section is close to Amphigymnia in lobation, but the 
lower surface is usually rhizinate or papillate to the margins. Fur- 
thermore, the rhizines are squarrosely branched at maturity, whereas 
the rhizines of Amphigymnia species are always simple and often 
sparse. There are about ten species in section Irregvlares, some very 
widely distributed in temperate and subtropical areas. The com- 
monly collected species are P. bedansae Mull. Arg., P. cetrata Ach., 
P. homotoma Nyl., P. macrocarpoides Vain., P. pilosa Stein., P. 
retievlata Tayl., and P, subisidiosa (Mull. Arg.) Dodge. 
3. Section Imbricaria (Schreb.) Fr. (1825, p. 242) 
Lichen sect. Imbricaria Schreb. (1791, p. 767). 
Lectotype species: Parmelia tiliacea (Hoffm.) Ach. 
Thallus adnate; lobes sublinear to subirregular, the margins ciliate 
or bulbate-ciliate; lower surface black or brown, the rhizines simple 
or branched. Apothecia adnate, the exciple frequently coronate. 
Section Imbricaria was first proposed by Schreb er under the genus 
Lichen and characterized as follows: "Syn. Squamaria Hoffm. Frondes 
subfoliaceae, membranaceae, imbricatae, depressae, flexiles. Fr. 
scutellae." Squamaria is a genus set up by Hoffmann to accommodate 
the species now recognized as Cetraria pinastri (Scop.) Rohl. and C. 
juniperina (L.) Ach. Although Schreber cites only the Hoffmann 
name, we are not obliged to use it in typifying Imbricaria. It seems 
more appropriate to retain Imbricaria in the sense that it was so 
commonly used in the 19th century, that is, for the narrow lobed 
species of Parmelia. Acharius (1794) was the first to adopt Schreber's 
name and included under it species now classified in the genera 
Physcia, Parmeliopsis, Xanthoria, and Parmelia. Of these, Parmelia 
tiliacea is the most suitable choice as a lectotype for section Imbricaria. 
Section Imbricaria may be divided into two subsections, Imbricaria 
and Bicornutae (p. 135), on the basis of whether marginal cilia or 
marginal bulbate cilia are present. 
Subsection Imbricaria 
Subsection Myeloekroa As&h. (1952, p. 74). 
Lectotype species: Parmelia aurulcnta Tuck. 
Thallus with simple marginal cilia, bulbate cilia lacking; lower 
surface usually black, the rhizines simple or sparsely branched or 
rarely squarrosely branched. Apothecia adnate, ecoronate. 
HALE & KTJROKAWA—SUBGENUS PARMELIA 131 
In his study of the Japanese Parmelias, Asahina (1952) divided 
section Hypotrachyna Vain, into two subsections, Myeloehroa (pig- 
ments present in the medulla) and Mydohuca, (medulla white). We 
have already reduced subsection Myeloleuca to synonymy under sub- 
section Parmelia. The six pigmented species that Asahina included 
in subsection Myeloehroa are all related to P. aurulenta Tuck. They 
all also have marginal cilia, a character that we consider to be of 
basic importance in classification. As far as we have been able to 
determine, medullary pigments have no value above the species level. 
Actually there are a number of pigmented species outside of Japan, 
unknown to Asahina at the time (e.g., P. isidiocera Nyl. and P. sil- 
vatica Lynge), which have branched rhizines and lack cilia and are 
classified below in section Hypotrachyna. 
Subsection Imbricaria is a rather heterogeneous group of 33 species 
widely distributed in temperate and subtropical regions. The well- 
known species P. dissecta Nyl., P. quercina (Willd.) Vain., and P. 
tUiacea (Hoffm.) Ach. belong here. The distribution of chemical sub- 
stances is very similar to subsection Bicornutae (cf. table 1) and the 
almost complete absence of sorediate species is similar. There seems 
to be no further natural subdivision of the subsection. 
Key to Species in Subsection Imbricaria 
1. Thallus isidiate or isidiate-Iobulate. 
2. Isidia tabulate at maturity, dorsi ventral. 
3. Medulla C+ rose (gyrophoric acid) ... 2. P. spathulata Kurokawa 
3. Medulla C—, KC+ rose (unknown) P. horrescens Tayl. 
2. Thallus isidiate, the isidia cylindrical. 
4. Medulla pigmented yellow orange, K—. 
5. Upper surface distinctly maculate P perisidians Nyl. 
5. Upper surface not maculate. 
6. Thallus saxicolous; lobes sublinear, 1.0-1.5 mm. wide, 
P. obsessa Ach. 
6. Thallus corticolous; lobes subrotund, 2-5 mm. wide. 
P. lindmannii Lynge 
4. Medulla white. 
7. Medulla K+ yellow or red, C—. 
8. Medulla K+ persistent yellow; stictic acid present. 
P. internexa Nyl. 
8. Medulla K+ yellow turning red. 
9. Norstictic acid present; tropical America . . . P. antillensis Nyl. 
9. Salacinic acid present; Old World. 
10. Lobes subrotund, the marginal cilia short. 
P. wallichiana Tayl. 
10. Lobes sublinear, the marginal cilia often 1-3 mm. long. 
P. usambarensis Stein. & Zahlbr. 
7. Medulla K—, C+ rose or red or C—. 
11. Medulla C— P. ilsomao Asah. 
11. Medulla C+ rose or red. 
132 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
12. Cortex strongly maculate; medulla C+ red (lecanoric acid). 
P. tiliacea (Hoffm.) Ach. 
12, Cortex weakly maculate or e maculate; medulla C+ rose (gyro- 
phoric acid). 
13. Lobes subrotund, 2-5 mm, wide. 
1. P. melanochaeta Kurokawa 
13. Lobes sublinear, 1-2 mm. wide P. dissecta Nyl. 
1. Thallus lacking isidia or lobules. 
14. Thallus pustulate; soredia scarcely if at all developed. 
15. Medulla white 3. P. subfatiscens Kurokawa 
15. Medulla pale yellow or orange yellow. 
16. Lobes 1-2 mm. wide; pustules fine, not sorediate; medulla C-f rose 
(gyrophoric acid) P. spumosa Asah. 
16. Lobes 2-4 mm. wide; pustules coarsely sorediate; medulla C—. 
P. aurulenta Tuck. 
14. Thallus lacking pustules. 
17. Thallus sorediate. 
18. Medulla yellow orange P, aurulenta Tuck, 
18. Medulla white. 
19. Thallus yellowish green (usnic acid) P. nylanderi Lynge 
19. Thallus mineral gray (usnic acid lacking). 
20. Soredia subterminal; moniliform cells in the medulla. 
P. metarevoluta Asah. 
20. Soredia laminal; moniliform cells lacking ... P. muelleri Vain. 
17. Thallus lacking soredia. 
21. Thallus pale brown below P. vergiformis Kremplh. 
21. Thallus jet black below. 
22. Medulla white (even under the apothecia). 
23. Medulla C+ blood red (lecanoric acid). 
24. Lobes about 1 mm. wide P. pruinata Miill. Arg. 
24. Lobes 2-5 mm. wide, 
25. Thallus strongly maculate; apothecia with retrorse rhi- 
zines  . P. carporrhizans Tayl. 
25. Thallus weakly maculate; retrorse rhizines lacking. 
P. quercina (Willd.) Vain. 
23. Medulla C—, P-f orange red. 
26. Medulla K—, P+ (protocetraric acid). 
P. michauxiana Zahlbr. 
26. Medulla K+ red. 
27. Thallus fragile, the cortex flaking off; spores 7-8 ju long; 
norstictic acid present P.phlyctinaHale 
27. Thallus coriaceous, the cortex firm; spores 20-28 long; 
salacinic acid present P. mutata Vain, 
22, Medulla yellow or yellow orange (especially under the apothecia). 
28. Pigment K+ purple P. denegans Nyl. 
28. Pigment K— or K+ more deeply yellow. 
29. Lobes sublinear, 0.5-1.5 mm. wide. 
30. Medulla P+reddish (unknown); thallus firm. 
P. gaLblna Ach. 
30. Medulla P—; thallus fragile. 
P. xantholepis Mont. & v.d. Bosch. 
29. Lobes subrotund, 3-6 mm. wide. 
HALE & KUROKAWA—SUB GENUS PARMELIA 133 
31. Upper cortex without maculae; apothecia numerous; Mexi- 
co F. immiscena Nyl. 
31. Upper cortex more or less distinctly maculate; Asia. 
32. Cortex fragile, flaking off . . . P. entotheiochroa Hue 
32, Cortex continuous, firm. 
33. Spores 7-12 ft long . 
33. Spores 13-17 p long . 
P. subaurulenta Nyl. 
. P. homogenes Nyl. 
1. Parmelia melanochaeta Kurokawa, sp. no v. PLATE 3 
Thallus laxe adnatus, olivacco-albicans, 4-7 cm. latus, lobis ir- 
regularibus vel sublinearibus, subrotundatis, 2-6 mm. latis, margine 
crenatis, cilia tis, sup erne albomaculatus, isidiatus, subtus niger, 
rhizinosus, rhizinis nigris, simplicibus. Apothecia adnata vel sub- 
stipitata, 1-3 mm, diametro; hymenium 80-90 p altum; sporae 
8-10X13-15 p. 
Thallus loosely attached to bark, turning olive buff or cream buff 
(R) in the herbarium, 4-7 cm. in diameter; lobes irregularly branched, 
irregular to sublinear, 2-6 mm. wide, 110-150 fi thick, the margins 
crenate, ciliate, the cilia mostly simple, coarse, 1-2 mm. long (cf. pi. 1); 
upper surface shiny, rather distinctly white-maculate, moderately to 
densely isidiate, the isidia thin, cylindrical, often branched and with 
black spinules or short cilia; medulla white; undersurface black, dark 
brown in a rather wide zone near the tips, moderately rhizinate, the 
rhizines black to blackish brown, simple. Apothecia adnate to sub- 
stipitate, 1-3 mm. in diameter, amphithecium isidiate, spinulate, disc 
vandyke brown (R); hymenium 80-90 high; spores 8-10X13-15 n. 
Reactions: Thallus K+ yellow; medulla K—, C-J- rose, KC+ red, 
P—, atranorine and gyrophoric acid present. 
Type in the Naturhistoriska Riksmuseet, Stockholm, collected at 
Santa Anna da Chapada, Ma to Grosso, Brazil, Jan. 18, 1894, by 
G. A. Malme (no. 2243; isotype in US). 
This is an unusual species that could be mistaken for P. dissecta 
Nyl, In fact, Lynge identified the same material as P. minarum 
Vain., a synonym of P. dissecta (cf. Hale, 1960). Although the 
chemistry of the two species is identical, P. melanochaeta is much 
larger and loosely attached and has broad rotund lobes. It seems 
to be common in southern Brazil and adjacent Paraguay. 
Additional specimens examined: Brazil: Buriti, Cerra da Chapada, 
Malme 2243, 2397B (S); Corumb&, Mato Grosso, Malme, August 
1894 (S). Paraguay: Gran Chaco, Malme, Sept. 14, 1893 (S); 
Colonia Risso, Malme 1802 Aa*, 1862 Ba*, 1911 Ba (S). 
2. Parmelia spathulata Kurokawa, sp. nov. 
Thallus adnatus, corticola, olivaceo-albicans, 2-5 cm. diametro, 
lobis sublinearibus, plus minusve imbricatis, 1-3 mm. latis, margine 
dissectis, superne sparse ciliatis, nitidus, emaculatus, isidiato-lobulatus, 
134 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
lobulis dorsiventralibus, apice dilatatis, subtus niger, rhizinosus, 
rhizinis nigris, sparse ramosis. Apothecia ignota. 
Thallus adnate, growing over mosses on bark, pale glaucous green 
(R), 2-5 cm. in diameter; lobes irregularly branched, subimbricate, 
1-3 mm. wide, 90-130 p thick, the margins sparsely ciliate, dissected; 
upper surface shiny, without maculae, covered with numerous small 
isidioid lobules, the lobules dorsiventral, subascending, often branched 
and dilatated towards the apices; medulla white; undersurface black, 
rhizinate, the rhizines black, shiny, simple to sparsely branched. 
Apothecia not seen. 
Reactions: Thallus K+ yellow; medulla K—, C+ rose, EG-}- red, 
P—, atranorine and gyrophoric acid present. 
Type in the Botanical Museum, Lund University, collected north- 
west of Tea-room, Skeleton Gorge, Wynberg, Cape Province, Union 
of South Africa, July 18, 1953, by Ove Almborn (no. 305; isotype in 
US). 
This species is characterized by having numerous spathulate lobules 
and simple to sparsely branched rhizines. It is closely related to 
P. dissecta Nyl., which has similar chemistry but cylindrical isidia 
rather than lobules. Parmelia spathulata is apparently restricted to 
South Africa. 
Additional specimens examined: Union of South Africa: Polela 
Forest, Polela, Natal, Almborn 9516 (LD, US); Kirstenbosch, Cape 
Province, Arnell 437 (LD). 
3, Parmelia gubfatiscens Kurokawa, sp. nov. 
Thallus adnatus, albido-cinerascens, 4-7 cm, latus, lobis subline- 
aribus, 0.5-1.5 mm. latis, margine ciliatis, sup erne planus, emaculatus, 
pustulatus, subtus niger, dense rhizinosus, rhizinis nigris, simplicibus. 
Apothecia adnata, 1.5-4.0 mm. diametro; hymenium 60-75 altum; 
sporae 8-9X12-14 /*. 
Thallus adnate on bark, pale olive gray to light mineral gray (R), 
4-7 cm. in diameter; lobes dichotomously or irregularly branched, 
sublinear-elongate, 0.5-1.5 mm. wide, 80-100 ja thick, the margins 
crenate, ciliate, the cilia black, simple, 0.5-1.0 mm. long; upper 
surface plane and smooth, shiny, not maculate, pustulate, the pustules 
not becoming sorediate, often ciliate; medulla white; undersurface 
black, densely rhizinate, the rhizines black, simple, about 1.0 mm. 
long. Apothecia adnate, 1.5-4.0 mm. in diameter, margins smooth, 
disc clove brown (R), radially split; hymenium 60-75 n high; spores 
8-9 X12-14 p. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ rose, 
P—, atranorine and an unknown KC+ substance present. 
Type in the Botanical Museum, Lund University, collected at 
Punch Bowl Inn, north of Louis Trichardt, Zoutpansberg, Transvaal, 
135 
Union of South Africa, Oct. 10, 1953, by Ove Almborn (no. 6443; 
isotype in US). 
This species is very close to P. dissecta Nyl. but it is distinctly 
pustulate rather than isidiate and the medulla is C—, whereas P. 
dissecta is C+ rose (gyrophoric acid). The pustules are similar to 
those of P. spumosa Asah., a smaller species with a pale yellow 
medulla and gyrophoric acid. Parmelia subfatiscens has the same 
West Indies-Africa distribution characteristic of P. ezsplendens Hale, 
P. suffixa Stirt., and P. ventricosa Hale & Kurokawa. 
Additional specimens examined: Jamaica: Blue Mountains: North 
slope below Corn Puss, Imshaug 14550 (MSC); New Haven Gap, 
Imshaug 15137 (MSC, US). Union of South Africa: Natal: Bosch- 
fontein Forest, Lions River Distr., Almborn 8717 (LD); Transvaal: 
Forest Drive from Houtbosch to Tzaneen, Pietersburg, Almborn 
6797 (LD); Cape Province: Deepwalls, Knysna, Almborn 3433 (LD). 
Subsection Bicornutae (Lynge) Hale & Kurokawa, comb. nov. 
Section Bicornvia Lynge (1913, p. 17). Type species: P. semilunata Lynge. 
Subgenus Bicornuta (Lynge) Gyel. (1932, p. 219). 
Thallus with marginal bulbate cilia; lower surface black or brown, 
the rhizines simple or branched. Apothecia frequently coronate. 
Lynge first proposed section Bicornuta on the basis of two species, 
P. semUunata and P. schiffneri Zahlbr., both of which have peculiar 
bicornute spores. As a matter of fact, there is apparently only one 
other species in Parmelia with such spores, P. bieomuta Mtill. Arg. 
However, these species also have marginal bulbate cilia, one of the 
most fundamental characters that we have found in subgenus Parmelia 
Normally ciliate species (subsection Imbriearia) appear to be closely 
related in chemistry (see table 1). 
This subsection of 46 primarily tropical species may be further 
subdivided into two series, based on the presence or absence of usnic 
acid, as follows: 
Series Bicornutae 
Thallus mineral gray; usnic acid lacking. 
Series Relicinae Hale & Kurokawa, ser, nov. 
Thallus stramineus vel viridiflavicans, acidum usnicum continens. 
Type species: Parmelia relicina Fr. 
When the bulbate-ciliate species are divided on the presence or 
absence of usnic acid, there is significant geographical correlation. 
In the series Bicornulae seven of the nine species with salacinic acid 
occur only in Africa. Of the nine C+ species, six are restricted to 
tropical America. Among the remaining species, five occur only in 
South America, two in Africa, one in America-Africa, and one 
722-891—64 2 
136 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
in Asia. In other words, the species in this series are most common 
in Africa and South America. 
In the series Relicinae there are no species at all in Africa. Thirteen 
occur only in southeastern Asia and Australia, two in tropical America 
and the Pacific, two in tropical America only, and one in America- 
Asia, This series then shows very strong representation in south- 
eastern Asia. 
Key to Species in Subsection Bicornutae Series Bicornutae 
1. Thallus isidiate or lobulate-isidiate. 
2. Lower surface pale brown. 
3. Medulla K+ red (salacinic acid). 
4. Thallus large, the lobes 2-5 mm. wide; upper surface not distinctly 
maculate P. tsidiza Nyl. 
4. Thallus smaller, the lobes 1.0-2.5 mm. wide; upper surface distinctly 
maculate P. subglandulifera Hue 
3, Medulla K—, C+ rose (gyrophoric acid) P. acortella Nyl, 
2. Lower surface jet black. 
5. Isidia becoming dorsiventral, lobulate; medulla C+ rose (gyrophoric 
acid), 
6. Apothecia ecoronate P, suffixa Stirt. 
6. Apothecia coronate P. fungicola Lynge 
5. Isidia normal, cylindrical. 
7. Medulla K+ red. 
8. Norstictic acid present 6. P. ventricosa Hale & Kurokawa 
8. Salacinic acid present. 
9. Plants saxicolous 4. P. decurtata Kurokawa 
9. Plants corticolous P. tabacina Mont. & v. d. Bosch. 
7. Medulla K-. 
10. Medulla C—, KC-f red (unknown present). 
2. P. apophysata Hale & Kurokawa 
10. Medulla C + rose or red. 
11. Medulla 0+ deep red (lecanoric acid) ... P. laevigatula Nyl. 
11. Medulla C+ rose (gyrophoric acid). 
12. Thallus tightly adnate, the lobes 0.5-1.0 mm. wide; Asia. 
P. subdissecta Nyl. 
12. Thallus loosely adnate, the lobes 1.0-3.0 mm. wide; tropical 
America P. papyrina F£e 
1. Thallus without isidia. 
13. Lower surface pale brown. 
14. Apothecia coronate; norstictic acid present . P. subcoronata Mull. Arg. 
14. Apothecia ecoronate; salacinic acid present. 
15. Upper surface distinctly maculate P, hypocraea Vain. 
15. Upper surface dull, not maculate, 
16. Thallus small, the lobes 0.7-2.0 mm. wide; South America. 
P continua Lynge 
16. Thallus medium sized, the lobes 2.0-5.0 mm. wide; Asia. 
P. setschwanensia Zahlbr. 
13. Lower surface Jet black. 
17. Medulla K+ red (salacinic acid). 
HALE & KUROKAWA—SUBGENUS PARMELIA 137 
18. Upper surface coarsely pustulate, pustules breaking open, esorediate. 
5. P. puBtulata Hale 
IS. Upper surface smooth, pustules lacking. 
19. Spores 7-9 » long; Africa ... P. sensibilis Stein, & Zahlbr. 
19. Spores 14-19 p long; eastern Asia ... P. meizospora (Nyl.) Nyl. 
17. Medulla K—. 
20. Medulla C—. 
21. Thallus adnate, the lobes 1-2 mm. wide. 
22. Thallus coriaceous; bulbate cilia conspicuous; India. 
3. P. bulbochaeta Hale 
22. Thallus fragile; bulbate cilia small; South America. 
P, viridescens Lynge 
21. Thallus appressed, the lobes less than 1 mm. wide. 
23. Lobes 0.3-1.0 mm. wide; spores elliptical. 
1. P affixa Hale & Kurokawa 
23. Lobes 0.1-0.2 mm. wide; spores bicornute. 
24. Spores 4-5 (i long P. schiflneri Zahlbr. 
24. Spores 15-20 /i long P. semilunata Lynge 
20. Medulla C+ rose or red. 
25. Medulla C+ deep red (leeanoric acid). 
26. Spores bicornute P. bicomuta Mull. Arg. 
26. Spores elliptical  P. confoederata Culb. 
25. Medulla C+ rose (gyrophoric acid). 
27, Apothecia coronate; upper cortex without maculae P. coronata F6e 
27. Apothecia ecoronate; upper cortex distinctly maculate. 
P, atrichella Nyl. 
1. Parmelia affixa Hale & Kurokawa, stat. et nom. nov. 
Parmelia coronata F6e var. denudata Vain, in Welw. Cat. Afr. Pl. 2, no. 2:401. 
1901. Type collection: Morro de Lopollo, Huilla, Angola, Welwitsch 
33 (TUR, holotype; BMt isotype). 
Thallus very closely adnate on bark, turning olive buff to tea green 
in the herbarium, about 3 cm. in diameter; lobes dichotomously 
branched, sublinear-clongate, 0.3-1.0 mm. wide, 70-90 p thick, the 
margins more or less crenatc, with dense bulbate cilia; upper surface 
plane, shiny, emaculate, cracked on older lobes, isidia and soredia 
lacking; medulla white; undersurface black, densely rhizinate, rhi- 
zines black, shiny, simple. Apothecia numerous, adnate, 0.5-1-5 
mm. in diameter, exciple smooth to crenate, coronate, the base 
retrorsely rhizinate; hymenium 40-50 ju. high; spores 4-5X8-10 p, 
episporium 1 ;u thick. 
Reactions: ThallusK-f yellow; medullaK—, C—, KC+ rose, P—, 
atranorine and a KC+ rose unknown substance (not further identified 
because of paucity of material). 
The name Parmelia denudata has already been used by Hampe 
(Linnaea 17:121. 1843) for a species now recognized as Everniopsis 
trvlla (Ach.) Nyl. Parmelia affixa is a small easily overlooked species 
that is probably more common than suspected in southwest Africa. 
It is one of the few coronate species in Africa. 
138 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Additional specimen examined: Africa: 10 km, north of S& da 
Bandeira, Huila, Angola, Degelius, Mar. 2, 1960 (DEGEL, US). 
2. Parmelia apophysata Hale & Kurokawa, sp. nov. 
Thallus arete adnatus, cortieola, albido-olivascens, 3-6 cm. dia- 
metro, lobis sublinearibus, 0.7-1.5 mm. latis 90-130 jjl crassis, margine 
bulbato-ciliatis, superne nitidus, isidiatus, isidiis cylindricis, subtus 
niger, rhizinosus, rhizinis dichotome ramosis. Apothecia ignota. 
Thallus closely adnate on bark, turning light olive buff in the her- 
barium, 3-6 cm. in diameter; lobes sublinear-elongate, more or 
less dichotomously branched, 0.7-1.5 mm. wide, 90-130 n thick, 
the margins densely bulbatc-ciliate; upper surface plane, shiny, 
sometimes faintly maculate, often irregularly cracked on older lobes, 
sparsely to moderately isidiate, isidia simple, cylindrical, less than 
0.3 mm. high; undersurface black, with a broad pale brown zone 
near the tips of lobes, short rhizinate, rhizinesj pale to black, shiny, 
densely branched. Apothecia unknown. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ reddish, 
P—, atranorine and an unknown KC+ substance (near alectoronic 
acid?) present. 
Type in the U.S. National Herbarium, collected on mangrove tree 
in pine barrens, vicinity of Piedra Blanca, La Vega, Dominican 
Republic, Oct. 14, 1947, by II. A, Allard (no. 16073). 
Parmelia apophysata is characterized by having a medium-sized 
thallus, simple cylindrical isidia, bulbate cilia and branched rhizines, 
and faintly maculate lobes. The unknown KC+ substance is notice- 
ably fluorescent in ultraviolet light. This species resembles P. sub- 
dissecta Nyl., which has more adnate smaller lobes and a C+ pale-red 
reaction caused by gyrophoric acid, Parmelia apophysata is known 
only from the type collection but it is certain to be found in other 
collections from tropical America. 
3* Parmelia bulbochaeta Hale, sp. nov. Plate 1 
Thallus adnatus vel laxe adnatus, albido-cinerascens, coriaceus, 
4-8 cm. diametro, lobis sublinearibus, 1.5-2.5 mm. latis, margine 
bulbato-ciliatis, superne plan us, nitidus, emaculatus, isidiis sorediisque 
destitutus, subtus nigricans, dense rhizinosus, rhizinis nigricantibus, 
dichotome ramosis. Apothecia adnata, coronata, 2-3 mm. diametro; 
hymenium 40-50 n altum; sporae 4X5 p. 
Thallus adnate to loosely adnate, dark mineral gray, coriaceous, 
4-8 cm. in diameter; lobes sublinear, 1.5-2.5 mm. wide, 160-210 m 
thick, the margins entire, conspicuously bulbate-ciliate, bulbules with 
simple or branched apical cilia; upper surface plane, shiny, without 
maculae, usually heavily pyenidiate, soredia and isidia lacking; 
medulla white, dense; undersurface black, becoming brown at the 
HALE <fc KUROSAWA—SUBGENUS PARMELIA 139 
tips, densely rhizinate, rhizines black, brownish black near the 
margins, dichotomously branched. Apothecia adnate, 2-3 mm. in 
diameter, coronate; hymenium 40-50 n high; spores 4X5 jt. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P—, 
only atranorine present. 
Type in the herbarium of D. D. Awasthi, collected at Perumal 
coffee plantation, Shembaganur, Madurai district, India, elev. 5300- 
5700 ft., Dec. 23, 1959, by D. D. Awasthi (no. 4347; isotype in US). 
Parmelia bulbockaeta resembles the South American species P. 
coronata F6e and P. viridescens Lynge in the presence of coronate 
apothecia, black undersurface, and branched rhizines, and in the 
absence of maculae. It differs in having a more robust coriaceous 
thallus and more conspicuous bulbate cilia and no chemicals other 
than atranorine. It is known only from two collections made in 
South India. 
Additional specimen examined: College compound, Shembaganur, 
Madurai, India, Awasthi 4275 (AWAS). 
4. Parmelia decurtata Kurokawa, sp. BO v. 
Thallus adnatus, saxicola, oh vaceo-cin erase ens, 3-7 cm. latus, lobis 
subirregularibus, subimbricatis, 1-3 mm. latis, margine crenatis, 
bulbato-ciliatis, superne planus, nitidus, emaculatus, isidiatus, subtus 
nigricans, rhizinosus, rhizinis nigris, simplicibus. Apothecia ignota. 
Thallus closely adnate on rock, pale olive buff to pale olive gray 
(R), 3-7 cm. in diameter; lobes irregularly branched, subirregular to 
sublinear, more or less imbricate, 1-3 mm. wide, 220-300 ju thick, the 
margins crenate, moderately bulbate-ciliate; uppersurface plane, 
shiny, often cracked on older lobes, isidiate, the isidia cylindrical, 
often more or less granular, rarely branched; medulla white; under- 
surface dark brown to black, moderately to densely rhizinate, the 
rhizines black, simple, about 1 mm. long. Apothecia not seen. 
Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid present. 
Type in the Botanical Museum, Lund University, collected 10 
miles southeast of Lydenburg, Transvaal, Union of South Africa, 
Oct. 20, 1953, by Ove Almborn (no. 7388; isotype in US). 
Parmelia decurtata bears a very close resemblance to P. tabacina 
Mont. 6 v.d. Bosch., a widespread corticolous species. They both 
have isidia, bulbate cilia, and salacinic acid. Parmelia decurtata 
differs in being strictly saxicolous, lacking any trace of maculae, and 
having smaller, often papillate, black-tipped isidia. 
Additional specimens examined: Union of South Africa: Natal: 
Cathedral Peak Area, Bergville Distr., elev. 6000 ft., Almborn 9328 
140 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
(LD); Cape Province: Bains Kloof, Wellington Distr., Almborn 5053 
(LD). 
5, Parmelia pustulata Hale, sp. nov. 
Thallus adnatus, corticola, membranaceus, 6-10 cm. diametro, 
olivaceo-albicans, lobis subirregularibus, 4-6 mm. latis, apice subro- 
tundatis, margine bulbato-ciliatis, superne planus vel rugosus, nitidus, 
isidiato-pustulatus, pustulis infiatis, subtus niger, modice rhizinosus, 
rhizinis simplicibus, ambitu castaneus, nudus vel papillatus. Apo- 
thecia ignota. 
Thallus adnate on bark, membranaceous, 6-10 cm. in diameter, 
olivaceous to light mineral gray; lobes more or less subirregular, 4-6 
mm. wide, the apices subrotund, the margins broadly crenate, in- 
conspicuously bulbate-ciliate; upper surface plane to rugose, shiny, 
becoming moderately isidlate-pustulate, pustules Inflated, irregular, 
breaking open apically, soredia absent; medulla white; lower surface 
black and moderately rhizinate at the center, the rhizines black, 
simple, the marginal zone bare or papillate, brown. Apothecia not 
seen. 
Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid present. 
Type in the U.S. National Herbarium, collected in a bamboo forest, 
Urundi, Africa, elev. 2200 m., by Deuse (communicated by Dr. J. L. 
Ramaut, Li&ge, Belgium). 
This species is unique in subsection Bicornutae in having large 
distinct pustules. Isidiate P. tabacina Mont. & v.d. Bosch, has similar 
lobe configuration and chemistry and is probably related to it. Par- 
melia pustulata is known only from the type collection. 
6. Parmelia ventricosa Hale & Kurokawa, stat. et nom. no v. 
Parmelia isidiza var. domingensis Vain. Ann. A cad. Sci. Fenn. 6, no. 7:17. 
1915. Type collection: La Cumbra, Santo Domingo (Dominican Re- 
public), Raunkiaer 492 (TUB,, liolotype; C, isotype). 
Thallus adnate on bark, pale glaucous green, 3-5 cm. in diameter; 
lobes irregularly branched, 1-5 mm. wide, 65-80 p. thick; margins 
crenate, with bulbate cilia; upper surface plane and smooth, shiny, 
weakly maculate, more or less wrinkled on older lobes, isidiate, the 
isidia mostly simple, fine and short; medulla white; undersurface 
uniformly black, short-rhizinate, the rhizines black, simple. Apo- 
thecia adnate, coronate, more than 2.5 mm. in diameter, exciple 
crenate, amphithecium smooth, weakly maculate, disc russet, imper- 
forate; hymenium 50-60 \i high; spores 7-8X13-17 ju, the episporium 
1.0-1.5 j\i thick. 
Reactions: Thallus K+ yellow; medulla K-j- yellow turning red, 
C—, KC—, P+ pale orange, atranorine and norstictic acid present. 
HALE & XUROKAWA—SUBGENTJS PARMELIA 141 
The name Parmelia domingemis is preoccupied by P. domingensis 
Ach., which is now recognized as a species of Anaptychia. Parmelia 
ventricosa and P. isidi2a Nyl. share isidia and bulb ate cilia, but P. 
ventricosa has coronate apothecia and norstictic acid. Parmelia 
isidizat a common African lichen, has ecoronate apothecia and salacinic 
acid. Parmelia ventricosa is a rather rare species in the Caribbean 
region with a disjunct locality in South Africa. 
Additional specimens examined: Mexico: Chiapas: Near ranch, El 
Suspiro, Hale 20166 (US). Union of South Africa: Transvaal: Drive 
from Houtbosch to Tzaneen, Pietersburg, Almborn 6813 (LD). 
Key to Species in Subsection Bicornutae Series Relicinae 
1. Thallus isidiate or isidiate-lobulate. 
2. Isidia dorsiventral, tabulate. 
3. Medulla P + red (unknown) 7. P. schizospatha Kurokawa 
3. Medulla P—, C+ rose (gyrophoric acid) . 4. P. luteoviridis Kurokawa 
2. Isidia normal, cylindrical. 
4. Lower surface pale to dark brown. 
5. Upper surface maculate; rhizines simple. 
9. P. tumescens Hale & Kurokawa 
5. Upper surface emaculate; rhizines branched . P. circumnodata Nyl. 
4. Lower surface jet black. 
6. Lobes more or less subirregular, to 3 mm. wide; medulla P-f, K— 
(unknown) 5. P. planiuscula Kurokawa 
6. Lobes sublinear, 1-2 mm. wide; medulla P+, K-f- red (norstictic acid). 
P. abstruga Vain. 
1. Isidia and lobules lacking. 
7. Lower surface pale to dark brown. 
8. Base of apothecia retrorsely rhizinate; plants saxicolous; stictic acid 
present P. limbata Laur. 
8. Base of apothecia without retrorse rhizines; plants corticolous; stictic acid 
lacking, 
9. Rhizines black; protocetraric acid present. 
6. P. ramosissima Kurokawa 
9. Rhizines uniformly pale brown. 
10. Medulla K+ red (salacinic acid) . . . 1. P. acrobotrys Kurokawa 
10. Medulla K— (protocetraric acid) . . . 8. P. sublanea Kurokawa 
7. Lower surface jet black. 
11. Medulla P—, K—. 
12. Lobes appressed, 0.2-0.8 mm. wide ... P. relicinula Miill. Arg. 
12. Lobes adnate, 0.7-2,0 mm. wide. 
13. Rhizines mostly branched; protolichesteric acid present. 
2. P. connivens Kurokawa 
13. Rhizines mostly simple; fatty acids absent .... P. relicina Fr. 
11. Medulla P+, K+ red Or K—. 
14. Medulla K+ red (norstictic acid) .... P. subabstrusa Gyel. 
14. Medulla K—. 
15, Apothecia ecoronate; P+ unknown present . P. sublimbata Nyl. 
15. Apothecia coronate; fumarprotocetraric acid present. 
142 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
16. Asci 16-32-epored P. relicinella Nyl. 
16. Asci 8-spored. 
17. Marginal bulbules ciliate, 0.3-1.0 mm, long; Pacific area. 
P. samoensis Zahlbr. 
17. Marginal bulbules not ciliate, 0.1-0.2 mm. long; tropical 
America . . . 3. P. eximbricata (Gyel.) Hale & Kurokawa 
1. Parmclia acrobotrys Kurokawa, sp. nov. Plate 2 
Thallus adnatus, viridi-flavicans, 6-12 cm. diametro, lobis subline- 
aribus, 1.0-2.5 mm. latis, margine bulbato-ciliatis, superne plerumque 
convexus, albomaculatus, isidiis sorediisque destitutus, subtus pallide 
castaneus, modice rhizinosus, rhizinis dense ramosis, pallidis. Apo- 
thecia substipitata, 1,0-3.5 mm. diametro; hymenium 50-60 /i altum; 
sporae 4-5 X 7-8 p. 
Thallus adnate on bark, olive buff (R), 6-12 cm. in diameter; lobes 
dichotomously branched, sublinear-elongate, 1.0-2.5 mm. wide, 240- 
320 n thick, the margins smooth, with bulb ate cilia; upper surface 
more or less convex, shiny, maculate, isidia and soredia lacking; me- 
dulla white; undersurface pinkish buff (R), moderately rhizinate, 
rhizines densely branched towards the tips, pale. Apothecia sub- 
stipitate, without retrorse rhizines, 1.0-3.5 mm. in diameter, exciple 
crenate, amphithecium smooth, disc burnt amber (R); hymenium 50- 
60 fi high; spores 4-5X7-8 fi. 
Reactions: Thallus K+ yellowish; medulla K+ yellow turning red, 
C—, KC—, P-f pale orange red, atranorine, usnic acid, and salacinic 
acid. 
Type in Herbarium Bogoriense, collected at Tjibodas, Java, May 
1949, by Neervoort (no. 60 pr. p.; isotype in US). 
Parmelia acrobotrys is closely related to P. sublanea Kurokawa. 
Both species have a pale undersurface, pale brown rhizines, ecoronate 
apothecia, and similar spores. Parmelia acrobotrys has coarser rhi- 
zines and contains salacinic acid, whereas P. sublanea contains proto- 
cetraric acid. The saxicolous species P, limbata Lain*, is superficially 
similar but contains stictic acid. Parmelia acrobotrys is known only 
from the type collection. 
2. Parmelia coimivens Kurokawa, sp. nov. Plate 2 
Thallus adnatus, viridiflavicans, 3-5 cm. diametro, lobis subline- 
aribus, 1.0-2.5 mm. Iatis, margine bulbato-ciliatis, superne planus vel 
convexus, sorediis et isidiis destitutus, subtus niger, dense rhizinosus, 
rhizinis nigris, ramosis. Apothecia adnata, 1.0-2.5 mm. diametro, 
retrorso-rhizinata; hymenium 45-60 p altum; sporae 2-3 X4-6 p. 
Thallus adnate, deep sea-foam green (R), turning cinnamon buff 
(R) in the herbarium, 3-5 cm. in diameter; lobes dichotomously 
branched, sublinear-elongate, 1.0-2.5 mm. wide, 120-160 ju thick, the 
HALE & KUROKAWA—SUBGENTJS PARHELIA 143 
older lobes becoming lobulate, the margins smooth, narrowly black- 
rimmed near the tips, with bulbate cilia; upper surface plane to 
slightly convex, smooth but becoming rugulose and cracked on older 
lobes, soredia and isidia lacking; undersurface almost uniformly black, 
densely rhizinate, the rhizines black, irregularly branched, sometimes 
projecting beyond the margins. Apothecia adnate, 1.0-2.5 mm. in 
diameter, retrorsely rhizinate, exciple crenate, disc seal brown (R); 
hymenium 45-60 n high; spores 2-3 X4-6 ju. 
Reactions: Thallus K+ yellowish; medulla K—, C—, KC—, P—, 
atranorine, usnic acid, and protolichesteric acid present. 
Type in the Kew Herbarium, collected in the Milne Bay district, 
Papua, New Guinea, Sept. 11, 1954, by R. D. Hoogland (no. 4726; 
isotypes in L, US). 
At first glance this species resembles P. acrobotrys Kurokawa and 
P. ramosissima Kurokawa but it is easily distinguished by the black 
undersurface and the negative reaction with P. No other species in 
series Relicinae produces a fatty acid. It is a typical Indonesian 
species. 
Additional specimens examined: Molucca Islands: Tanimbar Island, 
P. Jamdena, Buwalda 4246 (BO). Guam: McGregor s.n. (H). 
3* Parmelia eximbricata (Gyel.) Hale & Kurokawa, comb. nov. 
Parmelia samoensis Zahlbr. var. eximbricata Gyel. Ann. Myool. 36:288. 
1938. Type collection: Monte Rug, Cuba, Hioram 10506 (BP, holotype). 
Thallus closely adnate on bark, massicot yellow (R), 3-5 cm. in 
diameter; lobes more or less dichotomously branched, sublinear, 
0.7-1.5 mm. wide, 160-200thick; margins crenate, with small 
bulbate cilia, 0.1-0.2 mm. long; upper surface plane, smooth and 
shiny emaculate, isidia and soredia lacking; medulla white; under- 
surface black, densely rhizinate, the rhizines black, simple. Apothecia 
common, adnate, 0.5-1.5 mm, in diameter, exciple coronate, the base 
of the amphithecium often retrorsely rhizinate; hymenium 45-55 jtt 
high; spores 4-5 X 7-9 p, the episporium 1 jli thick. 
Reactions: Thallus K—; medulla K—, C—, KC—, P+ brick red, 
fumarprotocetraric acid and usnic acid present. 
Parmelia eximbricata resembles the Pacific species P. samoensis 
Zahlbr. in having coronate apothecia and fumarprotocetraric acid. 
The marginal bulbate cilia of P. samoensis however are longer and 
more conspicuous, and tipped with distinct cilia. This species is 
widely distributed in the Caribbean as far north as southern Florida, 
usually occurring at low elevations. 
Additional specimens examined: Florida: Big Pine Key, Florida 
Keys, Sivanson <& Dowling C-795 (US). Jamaica: Wareika House, 
summit of Long Mountain, St. Andrew Imshaug 13492 (MSC); north 
144 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
slope of Montpelier, St. Catherine, Imshaug 14261 (MSG, US). 
Grand Cayman: Near Georgetown, Imshaug 24385 (MSC); center of 
Island, Imshaug 24518 (MSC, US). Dominican Republic: Vicinity 
of Ciudad Trujillo, Santo Domingo, AUard 16186, 16192 (US); near 
Rio Chav6n, east of La Romana, La Altagracia, Hassler, July 31, 
1935 (WISC). Cuba: Without locality, Wright, Lieh. Cub. ser. II, 
10 (US). 
4. Purinelia luteoviridis Kurokawa, sp. no v. PLATE 2 
Thallus adnatus, corticola, viridiflavicans, ca. 7 cm. diametro, 
lobis sublinearibus, 1.5-3.0 mm. latis, margine dense lobulatis, 
bulbato-ciliatis, superne nitidus, lobulatus, lobulis vulgo ramosis, 
subtus niger, rhizinosus, rhizinis nigris, sparse ramosis. Apothecia 
ignota. 
Thallus adnate on bark, deep sea-foam green to light dull green 
yellow (R), about 7 cm. in diameter; lobes dichotomously branched, 
sublinear-elongate, 1,5-3.0 mm. wide, 160-200 p thick, the margins 
lobulate, with shiny bulbate cilia; upper surface plane and smooth, 
shiny, faintly maculate, lobulate, the lobules dorsiventral, not as- 
cending, often dichotomously branched, rhizinate below; undersurface 
black, rhizinate, the rhizines black, shiny, sparsely branched. Apoth- 
ecia not seen. 
Reactions: Thallus K+ yellowish; medulla K—, C+ rose, KC+ 
red, P~, atranorine, usnic acid, and gyrophoric acid. 
Type in the Herbarium Bogoriense, collected on peak of B. Papan, 
Beu territory, E. Kutai, Borneo, July 8, 1952, by W. Meijer (no. 
B1948; isotype in US). 
This species is characterized by the conspicuous yellow richly lobu- 
late thallus and by the production of gyrophoric acid. It is the only 
species in series Relicinae with a C+ substance. It resembles P. schizo- 
spatka Kurokawa in having lobules and a black lower surface but 
has wider lobes and greater development of lobules. Parmelia schizo- 
spatha is C—, P+ red. Parmelia luteoviridis is still known only from 
the type locality. 
Additional specimen examined: Borneo: Same data as the holotype, 
Meijer B1548a (B0). 
5. Parmelia plnniuscula Kurokawa, sp. nov. 
Thallus adnatus, corticola, viridiflavicans, usque ad 9 cm. latus, 
lobis sublinearibus, 1-3 mm. latis, margine subcrenatis, bulbato- 
ciliatis, superne planus, nitidus, sparse isidiatus, subtus niger, modice 
rhizinosus, rhizinis nigris, simplicibus. Apothecia ignota. 
Thallus adnate to loosely attached over mosses on bark, rather 
rigid, turning deep olive buff (R) in the herbarium, about 9 cm. in 
diameter; lobes irregularly branched, sublinear-elongate, 1-3 mm. 
HALE & KUROKAWA—SUBGENUS PARMELIA 145 
wide, 150-190 ju thick, the margins more or less crenate, with bulbate 
cilia; upper surface plane and smooth, shiny, faintly maculate, sparsely 
isidiate, isidia simple, cylindrical, less than 0.7 mm. high; medulla 
white; undersurface black, moderately rhizinate, the rhizines black, 
shiny, simple. Apothecia not seen. 
Reactions: Thallus K~b yellowish; medulla K—, C—, KC—, P+ 
orange red, atranorine, usnic acid, and an unknown P+ substance. 
Type in the Herbarium Bogoriense, collected at Kandang Badak, 
Java, May 27,1949, by Neervoort (no. 427; isotype in US). 
Externally this species resembles P. tumescem Hale & Kurokawa. 
It differs in having a black undersurface and aP-f unknown substance 
instead of stictic acid. Parmelia circumnodata Nyl., a much smaller 
lichen, has short sparse isidia, a pale undersurface with branched 
rhizines, and protocetraric acid. Parmelia abslrusa Vain., the fourth 
isidiate species in series Relicinae, has a thick rigid thallus and pro- 
duces norstictic acid. Parmelia planiuscula is still known only from 
the type locality. 
6* Parmelia ramosissima Kurokawa, sp. no v. Plate 2 
Thallus adnatus, corticola, obscure viridiflavicans, ca. 3 cm, dia- 
metro, Iobis sublinearibus, 0.7-2.0 mm. latis, margine integris, bulbato- 
ciliatis, superne nitidus, albomaculatus, isidiis sorediisque destitutus, 
subtus pallide castaneus, rhizinosus, rhizinis nigricans, dense ramosis. 
Apothecia adnata, ecoronata, retrorso-rhizinata; hymenium 40-50 p 
altum; sporae 2 X 3 p. 
Thallus adnate on bark, turning dark olive buff (R) in the herbar- 
ium, more than 3 cm. in diameter; lobes dicliotomously branched, 
sublinear-elongate, 0.7-2.0 mm. wide, 200-240 y. thick, the margins 
smooth, with bulbate cilia; upper surface plane and smooth, rather 
shiny, maculate, isidia and soredia lacking; medulla white; under- 
surface pale brown to brown, moderately rhizinate, the rhizines black- 
ening, finely and densely branched. Apothecia adnate, ecoronate, 
1.0-2.5 mm. in diameter, exciple crenate, often basally retrorsely 
rhizinate, disc warm blackish brown (R); hymenium 40-50 n high; 
spores about 2X3 p. 
Reactions: Thallus K+ yellowish; medulla K—, C—, KC+ red- 
dish, P+ orange red, usnic acid and protocetraric acid present. 
Type in Herbarium Bogoriense, collected on Tanimbar Island, 
Moluccas, Apr. 2, 1938, by P. Buwalda (no. 4583; isotype in US). 
This species resembles P. acrobotrys Kurokawa in habit and in the 
pale undersurface. However, P. ramosissima is distinctly maculate 
and contains protocetraric rather than salacinic acid. It has the 
smallest spores of any species in section Relicinae. It is presently 
known only from the type locality. 
146 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
7. Parmelia schizospatha Kurokawa, sp. no v. Plate 2 
Thallus adnatus, corticola, obscure yiridiflavicans, ca. 5 cm. di- 
ametro, lobis sublinearibus, 1-2 mm. latis, margine subcrenatis, 
bulbato-ciliatis, lobulatis, lobulis ramosis, superne plan us, nitidulus, 
lobulatus, subtus niger, modice rhizinosus, rhizinis nigris, simplicibus. 
Apothecia ignota. 
Thallus adnate on bark, turning deep olive buff (R) in the herbar- 
ium, about 5 cm. in diameter; lobes irregularly branched, sublinear- 
elongate, 1-2 mm. wide, 100-120 ju thick, the margins more or less 
crenate, with black bulbate cilia, lobulate, the lobules dor si ventral, 
often dichotomously branched, not ascending; upper surface plane 
and smooth, faintly maculate, somewhat shiny, lobulate, the lobules 
the same as the marginal ones; undersurface black, moderately rhi- 
zinate, the rhizines black, shiny, simple. Apothecia not seen. 
Reactions: Thallus K+ yellowish; medulla K—, C—, KC-, P+ 
orange red, usnic acid and an unknown P+ substance present. 
Type in Herbarium Bogoriense, collected at Gegerbentang, Java, 
June 27, 1949, by Neervoort (no. 1062; isotype in US). 
This new species is identical with P. sublimbata Nyl. in chemistry 
but it is distinctly lobulate. The lobules are quite fragile and fre- 
quently branched. At present the species is known only from the 
type locality. 
8. Parmelia sublaaea Kurokawa, sp. no v. 
Thallus adnatus, rigidulus, obscure viridiflavicans, ca. 10 cm. 
diametro, lobis sublinearibus, subimbricatis, 1-2 ram. latis, margine 
integris, bulbato-ciliatis, ciliis apice ramosis, superne planus vel 
convexus, nitidus, emaculatus, isidiis sorediisque destitutus, subtus 
pallide castaneus, dense rhizinosus, rhizinis pallide castaneis, dense 
ramosis. Apothecia substipitata, 1.0-4.5 mm. diametro; hymenium 
40-50 fi altum; sporae 4-5X7-9 m- 
Thallus rather rigid, olive buff to deep olive buff (R), corticolous, 
about 10 cm. in diameter; lobes dichotomously branched, sublinear- 
elongate, subimbricate, 1-2 mm. wide, 220-300 n thick, the margins 
entire, often brown rimmed, with bulbate cilia; upper surface plane 
to more or less convex, smooth and shiny, without maculae, isidia and 
soredia lacking; medulla white; undersurface pale brown, densely 
rhizinate, the rhizines pale brown, densely branched, often forming a 
woolly mat. Apothecia substipitate, 1.0-4.5 mm. in diameter, exciple 
crenate, more or less undulate, amphithecium smooth, disc seal brown 
(R); hymenium 40-50n high; spores 4-5X7-9p 
Reactions: Thallus K+ yellowish; medulla K—, C—, KC+ 
reddish, P+ orange red, usnic acid and protocetraric acid present. 
Type in Herbarium Bogoriense, collected on route between Ake 
HALE & KUROKAWA—SUBGENUS PARMELIA 147 
Biaur and Tolewang, Halmahera Island, Oct. 12, 1951, by P. Groen- 
hardt (no. 8409; isotype in US). 
As previously discussed, P. sublanea is very similar to P. acrobotrys 
except in chemistry. Parmelia ramosissima has similar chemistry 
but the rhizines are darker. The bulb ate cilia of P. sublanea are 
unusual in being almost spherical and quite large. The species is 
known only from the type locality. 
9. Parmelia tumescens Hale & Kurokawa, stat. et nom. nov. 
Parmelia limbata Laur. f. isidiosa Mull. Arg. Flora 70:59. 1887. Type 
collection: Richmond River, Australia, Hodgkinson s.n. (G, holotype). 
Thallus adnate on bark, turning deep olive buff in the herbarium 
about 4 cm. in diameter; lobes dichotomously branched, sublinear, 
1.0-1.5 mm. wide, 190-230 ju, thick, 'the margins subcrenate, with 
bulb ate cilia; upper surface more or less convex, irregularly cracked 
on older lobes, white-maculate, isidiate, the isidia cylindrical, simple or 
branched, up to 0.5 mm. high; medulla white; undersurface brown, 
moderately rhizinate, the rhizines simple, rather thick, dark brown to 
black. Apothecia not seen. 
Reactions: Thallus K—; medulla K-f yellow, C—, KC—, P+ pale 
orange red, usnic acid and stictic acid present. 
The specific epithet isidiosa is preempted by Parmelia isidiosa 
(Mull. Arg.) Hale, a species in subgenus Xanihoparmelia. Except for 
the presence of isidia, P. tumescens is very similar to P. limbata Laur., 
another Australian species that occurs commonly on rocks rather 
than bark. 
Additional specimens examined: Australia: Eastern Creek, New 
South Wales, Wilson (H); Brisbane, Queensland, Bailey (BM). 
4. Section Cyclocheila (Vain.) R&s. (1943, p. 10) 
Section Hypotrachyna *Cyclocheila Vain. (1890, p. 47). 
Pseudoparmelia Lynge (1913, p. 15). Type species: P. cypheUata (Lynge) 
Sant. 
Section Subfiavescentes (Vain.) Gyel. subsection Edliatae Gyel. (1932, p. 
225). Lectotype: P, caperata (L.) Ach. 
Section Hypotrachyna subsection Cyclocheila (Vain.) Gyel. (1932, p. 224). 
Section Hypotrachyna subsection Cyclocheila (Vain.) Hillm. (1934, p. 187). 
Superfluous combination. 
Section Pseudoparmelia (Lynge) Ras. (1943, p. 10). 
Lectotype species: Parmelia amazonica Nyl. 
Thallus adnate to loosely attached; lobes sublinear and truncate 
to subirregular and subrotund; lower surface black or brown, the 
rhizines simple, often coarse. 
As we previously mentioned (p. 129), Vainio first recognized Oyclo- 
cheila as a group under section Hypotrachyna, characterized by 
148 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
subrotund lobes and including three species, P. amazonica Nyl., P, 
minarum Vain, {—P. dissecta Nyl.), and P. muelleri Vain. The latter 
two species are ciliate and should be classified in our section Imbricaria 
subsection Imbricaria, Vainio (1923) later combined this group and 
Irregulares into a single section Irregulares (Vain.) Vain. We have 
amended section Irregulares to include only species with a reticulate 
upper cortex, marginal cilia, and usually squarrosely branched rhizines. 
The type species of section Cyelocheiia, Parmelia amazonica, has 
simple unbranched rhizines and lacks any marginal cilia. There are 
45 additional eciliate species with simple rhizines that can be classified 
here. These include typical sublinear-lobed species (e. g., P. sphaero- 
spora Nyl., P. texana Tuck.) and a number of subirregular, subrotund- 
lobed species, such as P. amazonica Nyl., P. caperafa (L.) Ach., P. 
caroliniana Nyl., and P. soredians Nyl. Some of these species have 
a narrow bare or papillate zone below at the margins and could be 
confused with species in subgenus Amphigymnia. Section Cyelocheiia 
is best represented in Africa where 18 of the species are endemic; they 
are often saxicolous. Divaricatic acid is especially common along 
with a number of other K—, P— acids also found in section Hypo- 
trackyna (see below, also cf. table 1). 
Lynge first proposed the genus Psevdoparmelia on the supposed 
discovery of "pseudocyphellae" on the lower surface of Parmelia 
cyphellata, a mistake followed by R as an en (1943). Santesson (1944) 
has conclusively shown that these structures are merely breaks or 
gashes in the lower cortex where rhizines have been torn away. 
Gyelnik (1933) used this genus name to accommodate the pseudo- 
cyphellate Amphigymnia species related to Parmelia cetrar ioides (Duby) 
Nyl., but this application of the name seems to be incorrect. 
In his original publication, Rasanen (1943) did not actually indicate 
himself as the author of section Cyelocheiia but listed Vainio alone, 
apparently on the mistaken assumption that Vainio had recognized 
this taxon at the sectional level previously. Rasanen clearly cites 
Cyelocheiia as a section, however, and we believe he should be cited as 
the author of the new combination. 
Key to Species in Section Cyelocheiia 
1. Thallus isidiate or pustulate. 
2. Thallus distinctly pustulate; aoredia absent or only sparsely developed. 
3. Thallus corticolous; lobes rather broad, 2-5 mm. wide. 
4. Medulla P— (divaricatic acid)  4. P. eruptens Kurokawa 
4, Medulla P+ red (protocetraric acid) P. raunkiaeri Vain. 
3. Thallus saxicolous; lobes narrow, to 1 mm, wide. 
5. Medulla P+ red (unknown) 6. P impcrfecta Kurokawa 
5. Medulla P—. 
HALE & KUROKAWA—SUBGENUS PARMELIA 149 
6. Caperatic acid present ...... 8. P* pustulescens Kurokawa 
6. Divaricatic acid present  P. owariensis Asah. 
2. Thallus with cylindrical isidia. 
7. Lower surface pale brown. 
8. Medulla pale yellow . . P. cyphellata (Lynge) Bant. 
8. Medulla white. 
9. Lobes broad, 2-5 mm. wide; subrotund . . 9. P. salacinifera Hale 
9. Lobes narrow, 0.5-2.0 mm. wide, sublinear. 
10. Thallus mineral gray, usnic acid lacking . 2. P. arcana Kurokawa 
10. Thallus yellowish green (usnic acid). 
11. Lobes 1.5-2.0 mm. wide; rhizines black. 
P. abstrusoides des Abb. 
11. Lobes 0.5-1.5 mm. wide; rhizines mostly pale. 
12. Medulla P—, KC+ orange (barbatic acid). 
P. rahengensis Vain. 
12. Medulla P+red (protooetraric acid) . . P. malaccensis Nyl. 
7. Lower surface jet black. 
13. Thallus yellowish green (usnic acid) .... P. ecaperata Mull. Arg. 
13. Thallus mineral gray (usnic acid lacking). 
14. Thallus saxicolous. 
15. Medulla C-f- deep red (lecanoric acid) . 1. P. annexe Kurokawa 
15. Medulla C—, P+ pale orange (stictic acid). 
7. P. ischnoides Kurokawa 
14. Thallus corticolous. 
16. Medulla P+ orange red. 
17. Medulla K-f red (salacinic acid).... P. cinerascene Lynge 
17. Medulla K — or brownish (protooetraric acid). 
18. Isidia rather irregular, often becoming granular. 
P. martinicana Nyl. 
18. Isidia cylindrical, normal P. amazonica Nyl. 
16. Medulla P—. 
19. Upper cortex finely cracked; perlatolic acid present. 
P. caroliniana Nyl. 
19. Upper cortex continuous or irregularly cracked with age. 
20. Lobes subrotund, 2-5 mm. wide; KC-f red unknown present. 
P. meiosperma (Hue) Dodge 
20. Lobes sublinear, 1.5-2.0 mm. wide; divaricatic acid present. 
P. concresccns Vain. 
1. Thallus lacking pustules or isidia. 
21. Thallus sorediate. 
22. Thallus yellowish green (usnic acid). 
23. Medulla K+red (salacinic acid); soredia farinose . P. soredians Nyl. 
23. Medulla K— (protocetraric acid); soredia coarse, originating from 
pustules  P. caper a ta (L.) Ach. 
22. Thallus mineral gray (usnic acid lacking). 
24. Upper cortex reticulately ridged and wrinkled; stictic acid present. 
25. Lobes subrotund, 2-5 mm. wide ... P. crozalsiana B. de Lesd. 
25. Lobes sublinear, 1.0-2.5 mm. wide . . P. carneopruinata Zahlbr. 
24. Upper cortex plane, not ridged. 
26. Soredia strongly capitate, submarginal, 
P. cryptochlorophaea Hale 
26. Soredia laminal, not strongly capitate. 
150 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
27. Yellowish pigment present under soralia; KC+ unknown 
present P. aptata Kremplh. 
27. Pigments absent; divaricatic acid present . . . P. texana Tuck. 
21. Thallus lacking soredia. 
28. Lower surface pale to dark brown. 
29. Medulla pigmented, the pigment K+ purple black. 
30. Pigment present throughout the medulla. 
11. P. violacea Kurokawa 
30. Pigment present only in lower half of the medulla. 
P. hypomilta F6e 
29. Medulla white or pale yellow, the pigment K—. 
31. Medulla pale yellow. 
32. Thallus saxicolous; lobes 1 mm, wide or less. 
P. chapadensis Lynge 
32. Thallus corticolous; lobes 2-5 mm. wide . P. sphaerospora Nyl. 
31. Medulla white, without any pigments. 
33. Thallus corticolous; medulla P+ orange red (protocetraric acid). 
34. Usnic acid present .... P. intertexta Mont. & v.d. Bosch. 
34. Usnic acid absent. 
35. Spores 5-7X8-11 p; Africa ... P. somaliensis Mull. Arg, 
35. Spores 14-18X26-28p; South America. 
P. leucopis Kremplh, 
33. Thallus saxicolous; medulla P —. 
36, Medulla 04- deep red (lecanorie acid) . . P. molybdiza Nyl. 
36. Medulla C—  10. P. tortula Kurokawa 
28. Lower surface jet black. 
37, Medulla pigmented, the pigment K+ purple black. 
38, Thallus yellowish green (usnic acid) . . P. rutidota Hook. & Tayl. 
38. Thallus mineral gray (usnic acid lacking), 
3D. Thallus saxicolous, the lobes subareolate, appressed. 
P. lecanoracca Mull. Arg, 
39. Thallus corticolous, the lobes loosely adnate. 
5. P. lieterochroa Hale & Kurokawa 
37. Medulla white; pigments absent. 
40. Thallus saxicolous. 
41. Lobes appressed to the substratum. 
42. Medulla C+ red (lecanorie acid) P. molybdiza Nyl. 
42. Medulla C—. 
43, Medulla K+ yellow (stictic acid). 
P* xanthomelaena Mull. Arg. 
43. Medulla K— (divaricatic acid) P. rupicola Lynge 
41. Lobes loosely adnate. 
44. Medulla P+ red (protocetraric acid) . „ . 3. P. caribaea Hale 
44. Medulla P— (divaricatic acid) .... P. rodriguesiana Hue 
40. Thallus corticolous. 
45. Thallus yellowish green (usnic acid) . P. rutidota Hook. & Tayl. 
45. Thallus mineral gray (usnic acid lacking). 
46. Medulla K+ yellow (stictic acid); upper cortex retieulately 
ridged   . P. scrobicularis Kremplh. 
46. Medulla K—; upper cortex plane P. subtiliacea NyL 
151 
1. Pannelia annexa Kurokawa, sp. nov. Plate 4 
Thallus arete adnatus, saxicola, albido-cinerascens, 3-7 cm. latus, 
lobis sublinearibus, centrum versus subimbricatis, 0.7-3.5 mm. latis, 
margin© crenatis, superne nitidus, emaculatus, isidiatus, subtus niger 
sparse rhizinosus, rhizinis nigris, simplicibus. Apotbecia adnata, 
1.5-5.0 mm. diametro; hymenium 70-80 n altum; sporae 5X7-8 m- 
Thallus tightly adnate on rock, pale olive gray to light mineral gray 
(R), 3-7 cm. in diameter; lobes irregularly branched, sublinear- 
elongate, more or less imbricate towards the center, 0.7-3.5 mm. 
wide, 180-240 fi thick, the margins crenate; upper surface plane, 
shiny, not maculate, irregularly cracked with age, isidiate, the isidia 
cylindrical, mostly simple, darkening at the tips, about 0.2 mm. high; 
medulla white; undersurface black, sparsely rhizinate, the rhizines 
black, simple. Apothecia adnate, 1.5-5.0 mm. in diameter, exciple 
undulate, amphithecium isidiate, disc snuff brown (R); hymenium 
70-80 p high; spores 5X7-8 fi. 
Reactions: Thallus K+ yellow; medulla K—, C+ deep red, KC+ 
red, P—, atranorine and lecanoric acid present. 
Type in the Botanical Museum, Lund University, collected at 
Mossel River, 3 miles east of Hermanns, Caledon, Cape Province, 
Union of South Africa, Sept. 29, 1953, by Ove Almborn (no. 5683; 
isotype in US). 
This is a typical South African saxicolous lichen clearly related to 
P. molybdiza Nyl., the only other species of the section Cyclockeila 
with lecanoric acid. Parmdia molybdiza differs in lacking isidia and 
in having a pale brown, rarely black, undersurface. Both species 
grow tightly adnate on exposed acidic rocks over a wide area in 
southern Africa. 
Additional specimens examined: Angola: Malanje: Between Ben- 
guela and Coporolo, Degelim, Feb. 2, 1960 (DEGEL), Duque de 
Braganca, Degelius, Mar. 5, 1960 (DEGEL, US). Union of South 
Africa: Natal: 8 miles south of Lomkile, Nongoma Distr., Hoegt 
August 1929 (TRH); Cape Province: Cape Town, Hdeg, May 31,1929 
(TRH). Mossel River, Caledon Distr., Almborn 5701 (LD), Camps 
Bay, Cape Distr., Almborn 4402 (LD). 
2. Pannelia arcana Kurokawa, sp. nov. Plate 4 
Thallus arete adnatus, saxicola, albido-cinerascens, 3-7 cm. Iatus, 
lobis sublinearibus, 0.3-1.0 mm. latis, margine subcrenatis, superne 
nitidus, emaculatus, isidiatus, medulla pro parte pallide flavescens, 
subtus castaneus, sparse rhizinosus, rhizinis castaneis, simplicibus. 
Apothecia ignota. 
Thallus tightly adnate on rock, pale to dark olive gray (R), 3-7 cm. 
in diameter; lobes dichotomously or irregularly branched, sublinear- 
722-891—64 3 
152 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
elongate, areolate toward the center, 0.3-1.0 mm. wide, 100-150m 
thick, the margins more or less crenate; upper surface plane to convex 
rather shiny, not maculate, tangentially cracked on older lobes 
isidiate, the isidia cylindrical, mostly simple, about 0.2 mm, high; 
medulla white, partly pale yellow in the upper half; undersurface 
brown to pale brown, sparsely rhizinate, the rhizines brown, simple. 
Apothecia not seen. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P—, 
pigment if present K—, atranorine, an unidentified fatty acid, and an 
unidentified pale yellow pigment present. 
Type in the herbarium of G. Degelius, collected in desert northeast 
of Baba, Mocamedes, Angola, Feb. 5,1960, by G. Degelius (isotype in 
US). 
Parmelia arcana is a tightly adnate saxicolous species with the 
same habit and lobe configuration as P. annexa Ivurokawa and P. 
molybdiza Nyl. These latter two species contain lccanoric acid. 
The pale undersurface of P. molybdiza is reminiscent of P. arcana, 
and the isidia of P. annexa are similar to those of P. arcana except 
that they are generally more papillate. 
Additional specimen examined: Union of South Africa: Konings- 
kroon, Elliott Distr., Cape Prov., Hoeg, Dec. 1, 1929 (TRH). 
3. Parmelia caribaea Hale, sp. nov. Plate 3 
Thallus adnatus, eborinus vel cinereo-albicans, saxicola, 6-10 cm. 
diametro, lobis parum lobatis, irregulariter sublinearibus, subimbri- 
catis, margine integris, superne planus, demum rugosus, sorediis 
isidiisque destitutus, subtus niger, modice rhizinosus, rhizinis crassis, 
nigris, simplicibus, ambitu fusco-castaneus, anguste nudus vel papil- 
latus, Apothecia subpedicellata, urceolata, usque ad 5 mm. diametro; 
hymenium 50-60 n altum; sporae 4-5X10-13 p. 
Thallus adnate, saxicolous, ivory to whitish gray, 6-10 cm. wide, 
rather coriaceous; lobes irregularly sublinear, infrequently branched, 
moro or less imbricate, 3-6 mm. wide, 200-250 /x, the margins entire 
to undulate, more or less black-rimmed, soredia and isidia lacking; 
upper surface plane, rugulose with age, cracked on older lobes; medulla 
white; undersurface black, moderately rhizinate, the rhizines thick, 
simple, the marginal zone brown, narrowly naked or papillate. Apo- 
thecia subpedicellate, at first globose, urceolate at maturity, up to 
5 mm. in diameter, amphithecium rugose, the disc imperforate; 
hymenium 50-60 n high; spores 4-5X10-13 p. 
Reactions: Thallus K+ yellow; medulla K—, KC+ rose, C—, 
P+ orange red, atranorine and protocetraric acid present. 
Type in the Missouri Botanical Garden, collected at Flanc Nord, 
Morne de Villet, St. Barthelemy, West Indies, Oct. 23, 1952, by 
C. Le Gallo (no. 494; isotype in US). 
HALE & KUROKAWA—SUBGENUS PARMELIA 153 
This remarkable species has been collected only on the island of 
St. Barthelemy. It is a saxicolous lichen without any obvious 
affinities. The mode of branching and the undersurface resemble 
P. nairobiensis Stein. & Zahlbr. and related South African species 
that contain divaricatic acid. Parmelia caribaea also has the black- 
rimmed margins typical of these species but the upper surface lacks 
any distinct white-netted reticulation. The apothecia are peculiar 
in that the disc may remain closed until quite late in the development 
of apothecia, so that young apothecia appear like spherical buds. 
Additional specimens examined: St. Barthelemy: Saline, Le Gallo 
491 (MO, US); Morne de Villet, Le Gallo 473 (MO, US); without 
specific locality, Le Gallo 2604 (MSC). 
4, Parmelia eruptens Kurokawa, sp. nov. Plate 3 
Thallus adnatus, corticola, olivaceo-albicans, 5-8 cm. diametro, 
lobis subirregularibus, 2-8 mm. latis, margine crenatis, superne 
emaculatus, isidiatus, isidiis crassis, inflatis, apice demum pustules- 
centibus, subtus niger, sparse rhizinosus, ambitu castaneus, sub- 
nudus, rhizinis nigricans, simplicibus. Apothecia adnata, 1-3 mm. 
diametro; hymenium 90-100 fi altum; sporae 5-7X10-12 n. 
Thallus adnate on bark, marguerite yellow to olive buff (R), 5-8 
cm. in diameter; lobes irregularly branched, subrotund, 2-8 mm. 
wide, 190-240 jw thick, the margins crenate, dark brown-rimmed near 
the tips; upper surface smooth and shiny, without maculae, moderately 
to densely isidiate, the isidia irregularly inflated, narrower at the base, 
at length breaking open at the tips; medulla white; undersurface 
black, brown and naked in a narrow zone near the tips, sparsely 
rhizinate, the rhizines pale near the margins, blackening at the center, 
simple. Apothecia adnate, 1-3 mm. in diameter, exciple more or less 
crenate, amphithecium isidiate, disc vandyke brown (R); hymenium 
90-100m high; spores 5-7X10-12 ju. 
Reactions: Thallus K-j- yellow; medulla K—, C—, KC—or KG + 
fleeting purple violet, P—, atranorine and divaricatic acid present. 
Type in the Botanical Museum, Lund University, collected 5 miles 
east of Mokobulaan, Lydenburg, Transvaal, Union of South Africa, 
Oct. 21, 1953, by Ove Almborn (no. 7498; isotype in US). 
Parmelia eruptens is characterized by rather large subirregular 
lobes and unusually large pustules which break open apically without 
soredial formation. It has much broader and more loosely attached 
lobes than other corticolous species containing divaricatic acid, such as 
P. nairobiensis Stein. & Zahlbr. and P. texana Tuck., though it is 
obviously related to these species. Parmelia eruptens is apparently 
endemic to South Africa. 
Additional specimens examined: Union of South Africa: Transvaal: 
Mount Anderson, Sabie-Lydenburg, Mam Qeesteranus 6453 (L, US), 
154 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
6455 (L); Mozambique: Near Rock Pueshang, west of Martins Falls, 
Manica and Sofala distrs., Mitchell 332 (US). 
5. Parmelia hetcrocliroa Hale & Kurokawa, nom, nov. 
Parmelia hypoxantha Stirt. Trans. New Zealand Inst. 32:76. 1899. Type 
collection: Warwick, Queensland, Australia, Gwytker s.n. (BM, holotype). 
Non P. hypoxantha Mtill. Arg. Flora 64:85. 1881 (= Ckondropsis semi- 
viridts Nyl.). 
Parmelia tiliacea (Hoffm.) Ach. var. stenophylla Miill. Arg. Flora 66:46. 
1883. Type collection: Toowoomba, Queensland, Australia, Hartmann 
s.n. (G, holotype). 
P. tiliacea var. rugulata Mull. Arg. Nuov. Giorn. Bot. Ital. 23:388. 1891. 
Type collection: Brisbane, Australia, Bailey s.n. (G, lectotype). 
P. tiliacea var. convexula Miill. Arg. Bull. Herb. Boiss. 4:90. 1896. Type 
collection: Brisbane, Australia, Shirley 1665 (G, holotype) [collector on 
label given as Bailey], 
Type collection: Warwick, Queensland, Australia, Qwyther s.n. 
(BM). 
Thallus adnate, mineral gray, turning olive buff in the herbarium, 
4-12 cm. in diameter; lobes more or less dichotomously branched, 
sublinear-elongate, 1-4 mm. wide, 160-210 thick, the margins more 
or less crenate, often black-rimmed near the tips; upper surface plane 
to convex, smooth, shiny, irregularly rugose and cracked on older 
lobes, isidia and soredia lacking; medulla white above, capucine 
buff to zinc orange in the lower half; undersurface black, moderately 
to densely rhizinate, the rhizines black, simple, irregularly to squar- 
rosely branched, 0.5-1.7 mm. long. Apothecia adnate, 1-5 mm. in 
diameter, exciple smooth to crenate or undulate, disc imperforate; 
hymenium 70-80 y, high; spores 7-8X10-12 n, episporium 1 n thick. 
Reactions: Thallus K-f- yellow; medulla K—, C—, KC-{- rose, 
P-f- orange red, pigment K+ purple, atranorine, protocetraric acid, 
and an unidentified anthraquinone pigment present. 
This remarkable species is known only from Queensland, Australia, 
where it appears to be common. Superficially, it resembles P. 
livida Tayl. or even P. brasUiana Nyl. but the lower medulla is con- 
spicuously pigmented. The rhizines are mostly simple, though 
occasionally squarrosely branched. It is the only species outside of 
section Parmelia and section Irregulares that has squarrose rhizines, 
but it is clearly unrelated to these two sections. On a gross morpho- 
logical basis we would prefer to put P. heteroehroa in section Hypo- 
trackyna but the peculiar rhizines indicate a closer affinity to section 
Cyclocheila. 
Additional specimens examined: Australia: Toowoomba, Queens- 
land, Hartmann 45 (G) (syntype of P. tiliacea var. rugulata); Queens- 
land, Bailey 334 (BM). 
* 
HALE & KUROKAWA-—STJBGENTJS PARMELIA 155 
6. Parmelia imperfect# Karokawa, sp. nov. 
Thallus arete adnatus, saxicola, albido-cinerascens, 2-4 cm. dia- 
metro, lobis sublinearibus, 0.5-2.0 mm. latis, margin© crenatis, 
superne nitidus, isidiato-pustulatus, subtus niger, sparse rhizinosus, 
rhizinis nigris, simplicibus. Apothecia ignota. 
Thallus closely or loosely adnate on rock, pale gull gray to light 
mineral gray (R), 2-4 cm. in diameter; lobes irregularly branched, 
sublinear-elongate, 0.5-2.0 mm. wide, 120-170 ju thick, the margins 
crenate; upper surface plane to more or less concave, shiny, irregularly 
cracked on older lobes, isidiate-pustulate, the pustules short, erupting; 
medulla white; undersurface black, sparsely rhizinate, the rhizines 
simple, about 0.2 mm. long. Apothecia not seen. 
Reactions: Thallus K+ yellow; medulla K— or K+ yellowish, 
G—, KC—, P+ orange red, atranorine and an unknown P+ sub- 
stance (forming small balls in G. A. o-T.) present. 
Type in the Botanical Museum, Lund University, collected at the 
Cathedral Peak Area, Bergville, Natal, Union of South Africa, 
Nov. 2, 1953, by Ove Almborn (no. 8815; isotype in US). 
Parmelia imperjeeta is another saxicolous lichen endemic to South 
Africa. It is apparently related to P. annexa Kurokawa, P. arcana 
Kurokawa, and P. molybdiza Nyl. The pustules are rather sparse, 
thick and short and burst open apically without formation of soredia. 
Almost identical pustules are found in the saxicolous Japanese species 
P. owariensis Asah., which differs principally in containing divaricatic 
acid. 
Additional specimens examined: Union of South Africa: Transvaal: 
Punch Bowl Inn, north of Louis Trichardt, Almborn 6491 (LD); 
Louis Trichardt, near Punch Bowl, Zoutpansberg, Almborn 6108 
(LD, US). 
7. Parmelia isclinoides Kurokawa, Bp. nov. Plate 4 
Thallus arete adnatus, saxicola, cinereo-albicans, 4-10 cm, latus, 
lobis sublinearibus, 0.5-2.0 mm. latis, superne nitidus, emaculatus, 
isidiatus, subtus niger, sparse rhizinosus, rhizinis nigris, simplicibus. 
Apothecia adnata, 1-2 mm. diametro; hymenium 70-80 p altum; 
sporae 5-6X7-8 /*. 
Thallus closely adnate on rock, pearl gray to mineral gray (R), 
4-10 cm. in diameter; lobes irregularly branched, sublinear-elongate, 
more or less imbricate with age, 0.5-2.0 mm. wide, 200-280 n thick, 
the margins entire to subcrenate; upper surface plane to convex, 
shiny, not maculate, isidiate, the isidia cylindrical, short and simple, 
darkening apically; medulla white; undersurface black, sparsely 
rhizinate, the rhizines black, simple. Apothecia adnate, 1-2 mm. 
156 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
in diameter, exciple more or less crenate, amphithecium isidiate, disc 
light seal brown (R); hymenium 70-80 n high; spores 5-6X7-8 fi. 
Reactions: Thallus K+ yellow; medulla K+ yellow, C—, KC—, 
P-f- pale orange, atranorine and s tic tic acid present. 
Type in the Botanical Museum, Lund University, collected on 
rocks near Window Stream, Kirstenbosch, Wynberg, Cape Province, 
Union of South Africa, Aug. 3, 1953, by Ove Almborn (no. 1698; iso- 
type in US). 
Parmelia ischnoides adds still another member to that unique group 
of saxicolous species so common in the Cape region (see discussions 
under P. annexa and P. imperfecta). It is characterized by adnate, 
linear and more or less separate lobes as illustrated in plate 4 to broad 
and more crowded lobes up to 2 mm. wide. Isidia are often quite 
dense but always rather small or even at times papillate. The 
chemical component, stictic acid, is quite rare in the section CycLockeila, 
Additional specimens examined: Union of South Africa: Cape 
Province: Kirstenbosch, Wynberg Distr., Almborn 124, 1443 (LD); 
Window Gorge, Wynberg Distr., Almborn 788 (LD); near Kloof Neck, 
Camps Bay Road, Cape Distr., Almborn 864 (LD). 
8. Parmelia pustulescens Kurokawa, sp. no v. Plate 4 
Thallus arete adnatus, saxicola, albido-cinerascens, 3-5 cm. latus, 
lobis sublinearibus, 0.3-1.2 mm. latis, margine crenatis, superne 
nitidus, emaeulatus, isidiatus, isidiis irregulariter inflatis, demum 
apice pustulescentibus, subtus niger, sparse rhizinosus, rhizinis nigris, 
simplicibus. Apothecia adnata, 0.7-2.0 mm. diametro; hymenium 
45-55 n altum; sporae 4-5X7-9 p. 
Thallus tightly adnate on rock, pearl gray to light mineral gray (R), 
3-5 cm. in diameter; lobes dichotomously to irregularly branched, 
sublinear-elongate, 0.3-1.2 mm. wide, 100-130 n thick, the margins 
crenate, narrowly black-rimmed near the tips; upper surface plane 
and shiny, without maculae, irregularly cracked on older lobes, isid- 
iate, the isidia cylindrical, irregularly inflated, at length bursting 
open apically; medulla white; undersurface black, sparsely rhizinate, 
the rhizines simple, black, 0.1-0.2 mm. long, Apothecia adnate, 
0.7-2.0 mm. in diameter, exciple isidiate-pustulate, disc clove brown 
(R); hymenium 45-55 y. high; spores 4-5X7-9 ju. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P—, 
atranorine and caperatic acid present. 
Type in the herbarium of G. Degelius, collected near Fazenda 
Canjangue, Vila Flor, Humbo, Angola, Feb. 18, 1960, by G. Degelius 
(isotype in US). 
Parmelia pustulescens is an adnate saxicolous lichen virtually in- 
distinguishable from P. imperfecta Kurokawa and P. owariensis Asah. 
HALE & KUROKAWA—SUB GENUS PAEMELIA 157 
in the development of pustules. (See discussion under P. imperjecta.) 
Parmelia pusttdescens differs chiefly in containing caperatic acid. It 
is known only from the type specimen. 
9- Parmelia salacinifera Hale, sp. nov. Plate 9 
Thallus adnatus, corticola, olivaceo-albicans, 6-12 cm. diametro, 
lobis subirregularibus, subrotundafcis, 3-6 mm. latis, 170—190/x crassis, 
superne planus, nitidus, modice isidiatus, subtus pallide castaneus, 
modice rhizinosus, rhizinis concoloribus, simplicibus, ambitu anguste 
nudus vel papillatus. Apothecia adnata, 2-4 mm. diametro; hyme- 
nium 30-38 fi altum; sporae non evolutae. 
Thallus adnate, corticolous, olivaceous to mineral gray, 6-12 cm, 
in diameter; lobes subirregular, 3-6 mm. wide, 170-190ft thick, 
apically subrotund, the margins entire to subcrenate; upper surface 
plane, rugose and fissured with age, shiny, moderately isidiate, the 
isidia cylindrical, usually simple, 0.04-0.08 X0.16-0.30 mm.; under- 
surface pale brown, moderately rhizinate, the rhizines pale brown, 
simple, the marginal zone bare or papillate. Apothecia adnate, 2-4 
mm. in diameter, amphithecium isidiate, disc imperforate; hymenium 
30-38m high; spores not developed. 
Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and salacinic acid present. 
Type in the U.S. National Herbarium, collected at Sanford, Semi- 
nole County, Florida, March 1925, by S. Rapp (isotype in FL AS). 
Parmelia salacinifera is a typical member of section Cyclocheila 
with irregular subrotund lobes. The thallus is usually quite closely 
adnate on bark. At first glance it seems indistinguishable from 
P. amazonica Nyl., which occurs from Florida to Brazil; P. amazonica 
has a jet black undersurface and protocetraric acid. Parmelia 
salacinifera undoubtedly occurs generally in the Caribbean region. 
A typical habitat is burned palm trunks in pastures. 
Additional specimens examined: Georgia: 5 mi. SW. of Waycross, 
Ware Co., Hale 16843 (US). Florida: Tomoka State Forest, Volusia 
Co., Hale 17063 (US); near Pensacola, Escambia Co., Hale 7994 (US); 
Dog Lake Recreation Area, Hale, 16714 (US); 6 mi. SW. of Tallahassee, 
Leon Co., Hale 16984 (US); Orange Park, Clay Co., Hale 17735 (US); 
vicinity of Fort Myers, Lee Co., Standley 13084 (US); Sanford, 
Seminole Co., Rapp 472, May 1919, February 1924 (FLAS). Mexico: 
Km. 956 on Hwy. 190, west of Ocozocoautla, Chiapas, Hale 20607 
(US). Brazil: Mato Grosso: Bocca da Serra, Malme s.n. (UPS). 
10. Parmelia tortu la Kurokawa, sp. nov, Plate 4 
Thallus adpressus, saxicola, albido-cinerascens, 5-10 cm. latus, 
lobis subirregularibus vel sublincaribus, apice subrotundatus, 1.5-4.0 
mm. latis, margine crenatis, superne nitidus, emaculatus, isidiis 
158 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
sorediisque destitutus, subtus pallide castaneus, sparse rhizinosus, 
rhizinis castaneis, simplicibus. Apothecia adnata vel substipitata, 
1-5 mm. diametro; hymenium 40-50/x altum; sporae 5-6 X 7-9 p. 
Thallus closely adnate on rock, yellowish glaucous to light olive 
gray (R), 5-10 cm, in diameter; lobes irregularly branched, sub- 
irregular to sublinear-elongate, more or less subrotund apically, often 
twisted and contorted, 1.5-4,0 mm. wide, 180-220 /a thick, the margins 
entire to crenate, black-rimmed at the tips; upper surface shiny, 
often faintly pruinose, without maculae, rugulose and distinctly 
cracked on older lobes, isidia and soredia lacking; medulla white; 
undersurface pale brown, sparsely rhizinate, the rhizines simple, 
pale brown, about 0.3 mm. long. Apothecia adnate to substipitate, 
1-5 ram, in diameter, exciple and am phi thecium smooth, disc bone 
brown (R); hymenium 40-50 fx high; spores 5-6X7-9/i. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC—, or 
KC+ faint rose, P—, atranorine and unknown colorless substances 
present. 
Type in the Botanical Museum, Lund University, collected on 
rocks between O'okiep and Springbok, Namaqualand, Cape Province, 
Union of South Africa, Sep. 15, 1953, by Ove Almborn (no. 4805; 
isotype in US). 
Parmelia tortula is a saxicolous species common in the Cape region 
and obviously related to P. molybdiza Nyl. Generally it is larger 
and the colonies more extensive than in P. molybdiza. The lobes are 
often more or less twisted and, at least near the tips, partially free of 
the substratum; P. molybdiza is less robust and contains lecanoric 
acid. The chemistry of P. tortula is not entirely clear at this time. 
The KC-J- test is variable, often weak or absent. Microchemical 
tests have not revealed the cause of this reaction. 
Additional specimens examined: Union of South Africa: Orange 
Free State: Thabanchis, Hoeg, Aug. 12, 1929 (TRH); Cape Province: 
4 miles south-southwest of Oudtshoorn, Oudtshoorn Distr., Almborn 
4199 (LD); Saldanha, Malmesbury Distr., Almborn 4988, 4996 (LD); 
Sea Point, Cape Town, Wynberg Distr., Hoeg, May 31, 1929 (TRH); 
Dordricht, Hoeg, Nov. 21, 1929 (TRI1). 
11. Parmelia violacea Kurokawa, sp. no v. 
Thallus adpressus vel adnatus, saxicola, olivaceo vel viridi-cineras- 
cens, 3-6 cm. diametro, lobis sublinearibus, 1-3 mm. latis, margine 
crenatis, superne nitidus, emaculatus, isidiis sorediisque desti tutus, 
medulla purpureus, subtus pallide castaneus, mod ice rhizinosus, 
rhizinis castaneis, simplicibus. Apothecia adnata, 1.0-1.5 mm. 
diametro; sporis non visis. 
HALE & KUROXAWA—SUBGENUS PARMELIA 159 
Thallus closely adnate on rock, smoke gray to dark olive gray 
(R), 3-6 cm. in diameter; lobes irregularly branched, sublinear- 
elongate, 1-3 mm. wide, 160-200 n thick, the margins crenate; upper 
surface shiny, without maculae, distinctly rugulose on older lobes, 
isidia and soredia lacking; medulla vinaceous purple to indian purple 
(R); undersurface pale brown, moderately rhizinate, the rhizines 
pale brown, simple, 0,5-1.0 mm. long. Apothecia adnate, 1.0-1.5 
mm. in diameter, exciple and amphithecium smooth, disc cameo 
brown to Vandyke brown (R); mature spores not seen. 
Reactions: Thallus K+ yellow; medulla K+ deep purple, P—, 
atranorine and an unidentified hydroxanthraquinone present. 
Type in the Botanical Museum, Lund University, collected at 
Blinkwater Ravine, slopes of Table Mountain, Cape Province, Union 
of South Africa, Aug. 5, 1953, by Ove Almborn (no. 1771; isotype 
in US). 
Parmelia violacea has a brilliant deep rusty-red medulla, frequently 
exposed as the upper cortex breaks away. In this respect it is very 
similar to P, endomiltoides Nyl., a Xanthoparmelia species with usnic 
acid and two unidentified pigments, one of them (of lower Rf) identical 
with that in P. violacea. Both of these species are found on Table 
Mountain in Cape Province. 
5. Section Hypotrachyna Vain. (1890, p. 38) 
Section Hypotrachyna* Sublinearis Vain. (1890, p. 50). 
Section Sublineares (Vain.) Vain. (1923, p. 34). 
Section Hypotrachyna subsection Sublineares (Vain.) Gyel. (1932, p. 223). 
Section Hypotrachyna subsection Sublineares (Vain.) Hiilm. (1934, p. 187). 
Superfluous combination. 
Type species: Parmelia brasiliana Nyl. 
Thallus adnate to loosely attached and subdivaricate; lobes sub- 
linear to linear-elongate; lower surface jet black, the rhizines more or 
less richly dichotomously branched. 
The type species of this section should be selected from the Brazilian 
species cited in Vainio's original study. We believe that he considered 
his group Sublinearis to represent the typical concept of section Hypo- 
trachyna and P. brasiliana Nyl. to be a typical species in this group. 
Two other groups described at the same time, Qyclocheila (= section 
CyclockeUa (Vain.) Ras.) and Irregularis (= section Irregulares (Vain.) 
Vain.), were described as having subirregular rotund lobes with a 
narrow bare zone below. In his study of Philippine lichens, Vainio 
(1923) put the subirregular species into a new section (Irregulares) 
and the sublinear species into section Sublineares (Vain.) Vain. He 
included Parmelia americana (Mey. & Flot.) Mont, and P. sorocheila 
160 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Vain, in section Sublineares, but these are everniiform species that 
should be transferred to the genus Pseudevernia Zopf. 
As we have delimited section Hypotraehyjia, it does not include 
P. saxatilis (L.) Ach,, the type of section Parmelia, which has pseudo- 
cyphellae and squarrosely branched rhizines. For this reason, sec- 
tion Parmelia, which includes P. saxatilis, and section Hypotrackyna 
cannot be considered as synonymous. 
Section Hypotrackyna is the largest yet surprisingly the most homo- 
geneous group of species in subgenus Parmelia, All species have a 
black lower surface with more or less distinctly dichotomously branched 
rhizines, sometimes so dense as to form a marginal mat projecting 
beyond the margins. Cilia are absent, although projecting branched 
rhizines might be mistaken for cilia. The greatest concentration of 
species is in tropical America, where 48 of the 83 species are endemic. 
Fourteen species are endemic to Asia and only six to Africa. 
Morphological and chemical diversity reach a peak in this section. 
There is an abundance of sorediate species (14). There are a number 
of acids not found in any of the other sections of subgenus Parmelia 
except section Cyclocheila (cf. table 1), These include alectoronic 
acid, barbatic acid, divaricatic acid, evernic acid, lichexanthone, olive- 
toric acid, and perlatolic acid. All of these substances are nega- 
tive with KOH, paraphenylenediamine, calcium hypochlorite (except 
for olivetoric acid), and most are KC+ red and more or less fluores- 
cent in ultraviolet light. In addition, if depsides, they have a basic 
2-methoxy substitution on the acid ring (cf, Asahina and Shibata, 
1954, p. 54), whereas the other depsides usually have a 2-hydroxy 
substitution. The full significance of this chemical variation has not 
yet been assessed, but the overall correlation between the chemical 
constituents and our sectional classification appears to be sound. 
Key to Species in Section Hypotrachyna Vain. 
1. Thallus isidiate  I. 
1. Thallus lacking isidia. 
2. Thallus sorediate or pustulate .  II. (p. 161). 
2. Thallus lacking isidia, soredia, or pustules III. (p. 163). 
I. THALLUS ISIDIATE 
1. Medulla yellow, the pigment K—. 
2. Medulla C —; isidia dense P. isidiocera Nyl. 
2. Medulla C-j- red (unknown); isidia, variable, sparse, simple to coralloid. 
23. P. prolongata Kurokawa 
1. Medulla white, or if pigmented, the pigment K+ purple. 
3. Upper cortex distinctly maculate. 
4, Medulla K—, C—, KC—, P— (fatty acids) ... P. eostariceneis Nyl. 
HALE & KUROKAWA—StTBGENUS PARMELIA 161 
4. Medulla P+ yellow to red or KC+ red or orange. 
5. Medulla K+ yellow, P-f yellow (unknown). 
9- P. dentella Hale & Kurokawa 
5. Medulla K—, KC+ red or orange. 
6, Medulla C—, KC+ orange (barbatic acid). 
7. Isidia cylindrical P. imbricatula Zahlbr. 
7. Isidia lobulate, dorsiventral P. digitata Lynge 
6. Medulla C—, KC+ rose (evernic acid). 
8. Isidia cylindrical P. bogotensis Vain. 
8. Isidia lobulate, dorsiventral ....... P. culmigena Zahlbr. 
3. Upper cortex not distinctly maculate. 
9. Medulla P + yellow to orange red. 
10. Medulla K— (protocetraric acid). 
11. Medulla ochraceous yellow in the lower half . P. consimilie Vain. 
11. Medulla entirely white P. koyaensie Asah. 
10. Medulla K+ yellow or red. 
12. Thallus yellowish green (usnic acid); norstictic acid present. 
P. microblasta Vain. 
12. Thallus grayish green (usnic acid lacking). 
13. Hhizines finely branched; isidia mostly simple; norstictic acid 
present 24, P. rhabdiformis Kurokawa 
13. Rhizines sparsely to moderately branched; isidia often branched; 
stictic acid present 5. P. crenata Kurokawa 
9. Medulla P—. 
14. Medulla C+ red or yellow or KC-f- red or orange. 
15. Medulla C+ deep red, KC+ red (unknown). 
23. P. prolongata Kurokawa 
15. Medulla C— or C+ yellow, KC+ orange (barbatic acid). 
16. Isidia cylindrical .P. imbricatula Zahlbr. 
16. Isidia lobulate, dorsiventral P. digitata Lynge 
14. Medulla C—, KC—, or KC+ red. 
17. Medulla KC+ red or rosy. 
18. Isidia inflated or pustular; medulla usually ochraceous yellow 
under the isidia P. dactylifcra Vain. 
18. Isidia simple or lobulate; pigments absent. 
19, Isidia lobulate, dorsiventral; alectoronic acid present. 
10. P. ensi folia Kurokawa 
19. Isidia cylindrical; KC+ unknown present. 
P. nodakensis Asah. 
17. Medulla C—, KC —; fatty acids present. 
20. Upper cortex weakly maculate; rhizines densely branched; 
America P. costaricensis Nyl. 
20. Upper cortex without maculae; rhizines moderately branched; Asia. 
18. P. infirm a Kurokawa 
n. THALLUS SOKEDIATE OR PUSTULATE 
1. Pustules distinct, without soredia or only very sparsely sorcdiate. 
2. Thallus yellowish green (usnic acid) P. meyeri Zahlbr. 
2. Thallus mineral gray (usnic acid lacking). 
3. Medulla pale yellow orange, pigment K— . . . P. endochlora Leight. 
3. Medulla white or in part pigmented, pigment K + purple. 
4. Pigments present, K+ purple. 
162 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
5. Thallus 2-4 cm, broad, the lobes up to 1 mm. wide . P. malmei Lynge 
5. Thallus 3-10 cm. broad, the lobes 1.0-2.5 mm, wide. 
P formosana Zahlbr. 
4. Pigments absent. 
6. Medulla C+ rose (gyrophoric acid) P. revoluta Floerke 
6. Medulla C—, KC+ orange (barbatic acid). 
7. Pustules distinct, becoming sorediate; Asia ... P. exsecta Tayl. 
7. Pustules irregular, esorediate; cortex fragile; North America. 
29. P. virginica Hale 
1. Pustules indistinct or absent; sorcdia usually abundant, farinose. 
8. Upper cortex distinctly maculate. 
9. Medulla pale yellow orange throughout .... P. endochlora Leight. 
9. Medulla white. 
10. Medulla KC-f orange (barbatic acid) . . P. laevigata (8m.) Ach. 
10. Medulla KC+ red or rose. 
11. Evernic acid present P. rockii Zahlbr. 
11. Alcctoronic acid present 13. P. exsplendcns Hale 
8. Upper cortex without maculae or at most in part faintly maculate. 
12. Thallus yellowish green (ugnic acid). 
13. Medulla K+ yellow turning red (salacinic acid). 
P. sinuosa (Sin.) Ach. 
13. Medulla K—, P+ orange red (protocetraric acid). 
14. Soralia mainly terminal; medulla ochraceous (rhodophyscin) under 
the soralia P. velloziae Vain. 
14. Soralia mainly sub terminal; pigments lacking. 
15. P flavovirens Kurokawa 
12. Thallus mineral gray (usnic acid lacking). 
15. Medulla P+ yellow to orange red. 
16. Medulla K+ yellow turning to red fsalacinic acid). 
17. Rhizines moderately branched; zeorine present. 
P. majoris Vain. 
17. Rhizines finely branched; zeorine absent. 
2. P. brevirhiza Kurokawa 
16. Medulla K—. 
18. Lichexanthone present; P+ unknown present. 
P. subaffinia Zahlbr. 
18. Lichexanthone absent; protocetraric acid present. 
19. Orange-red pigment (rhodophyscin) under the soralia. 
6. P. croceopustulala Kurokawa 
19. Pigments absent under the soralia. 
20. Lobes 1.5-2,0 mm. wide; soralia mostly subterminal. 
P. paeudosinuosa As ah. 
20. Lobes 0.5-1.0 mm, wide; soralia mostly Iaminal. 
1. P. anaptychioides Kurokawa 
15. Medulla P-. 
21. Orange-yellow pigment (rhodophyscin) under the soralia. 
19. P. leiophylla Kurokawa 
21. Pigments absent. 
22. Medulla C+ rose, red, or orange red. 
23. Medulla C+ red or orange red (olivetoric acid). 
21. P. lividescens Kurokawa 
23. Medulla C-{- rose (gyrophoric acid). 
HALE & KUROKAWA—STJBGENUS PARHELIA 163 
24. Lobes linear-elongate, subdivaricate; Mexico. 
28. P. thysanota Kurokawa 
24. Lobes loosely adnate, sublinear; Africa . . P. revoluta Flk. 
22. Medulla C—, KC+ red or orange red. 
25. Medulla KC+ orange (barbatic acid) ... P. rockii Zahlbr. 
25. Medulla KC-f rose or red. 
26. KC-f unknown present; Africa. 
17. P. immaculata Kurokawa 
26. Alectoronio acid present; tropical America. 
27. Lobes linear-elongate, subdivaricate. 
8* P. densirhizinata Kurokawa 
27. Lobes sublinear, loosely adnate to subimbricate. 
13. P. exsplendens Hale 
III. THALLUS LACKING ISIDIA, SOREDIA, AND PUSTULES 
1. Thallus yellowish green (usnic acid). 
2. Thallus saxicolous  P. flavida Zahlbr. 
2. Thallus corticolous. 
3. Medulla K—, P+ orange red P. flavida Zahlbr. 
3. Medulla K+ red, P-f orange red. 
4. Salacinic acid present 4. P. citrella Kurokawa 
4. Norstictic acid present. 
5. Thallus closely adnate; lobes sublinear; rhizines densely branched. 
P. reduceii s Nyl. 
5. Thallus loosely adnate to subdivaricate; lobes linear-elongate; 
rhizines moderately branched, 
6. Lobes 1-2 mm. wide, sparsely rhizinate, subimbricate. 
P. enderythraea Zahlbr. 
6. Lobes 2-4 mm. wide, densely rhizinate, subdivaricate. 
P. caraccensis Tayl. 
1. Thallus mineral gray (usnic acid lacking). 
7. Medulla pale yellow or in part ochraceous. 
8. Pigment K—. 
9. Apothecia stalked; rhizines richly branched . . P. pcruviana Nyl. 
9. Apothecia adnate; rhizines sparsely to moderately branched. 
25. P. rigidula Kurokawa 
8. Pigment K4- purple to purple black. 
10. Thallus orange fluorescent under TJV light (lichexanthone present), 
11. Upper cortex maculate; thallus corticolous . . P. silvatica Lynge 
11. Upper cortex not maculate; thallus saxicolous. 
12. Medulla C+ red (olive toric acid). . . P, osteoleuca Nyl. 
12. Medulla C—, P+ red (unknown). 
11. P. erythrodes (Zahlbr.) Hale & Kurokawa 
10. Thallus not fluorescent (lichexanthone absent). 
13. Medulla P+ red (protocetraric acid) .... P. bahiana Nyl. 
13. Medulla P— (fatty acids) P. Iythgoeana Dodge 
7. Medulla white, no pigments present. 
14. Medulla P+ yellow to orange red. 
15. Medulla K+ yellow turning red. 
16. Norstictic acid present  3. P. caneacens Kurokawa 
16. Salacinic acid present. 
164 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
17. Thallus corticolous; rhizines densely branched. 
P. sublaevigata (Nyl.) Nyl. 
17. Thallus saxicolous; rhizines moderately branched 
P. subsaxatilis B. de Lesd. 
15. Medulla K— or K+ faint brownish. 
18. Thallus saxicolous. 
19. Cortex orange fluorescent in UV light (lichexanthone present). 
P. brasiliana Nyl. 
19. Cortex not fluorescent (lichexanthone absent). 
P. osseoalbida Lynge 
IS. Thallus corticolous. 
20. Medulla KC+ orange (barbatic acid present). 
26. P. scytodes Kurokawa 
20. Medulla KC+ rose (protocetraric acid present). 
21. Spores 15-25 p long. 
22. Spores 20-25 n long; fatty acid present. P. keitauengis Asah. 
22. Spores 15-20 p long; fatty acids lacking. 
P. manilensis Vain. 
21. Spores 8-12 p long. 
23. Apothecia large, the disc split radially; Africa. 
14. P. fissicarpa Kurokawa 
23. Apothecia small, the disc entire; tropical America. 
24. Rhizines moderately branched lobes subrotund often, whitish 
pruinose  P. insinuans Nyl. 
24 Rhizines fine, densley branched; lobes sublinear, epruinose. 
P. bahiana Nyl. 
14. Medulla P—. 
25. Medulla C—, KC+ orange (barbatic acid). 
26. Thallus subdivaricate, the lobes linear-elongate; rhizines mostly 
moniliform 22. P. monilifera Kurokawa 
26. Thallus adnate, the lobes sublinear; rhizines normal, not moniliform. 
27. Upper cortex distinctly maculate P. boliviana Nyl. 
27. Upper cortex not distinctly maculate. 
28. Amphithecium strongly rugose; Java. 
P. bostrychodes Zahlbr. 
28. Amphithecium smooth; tropical America. 
29. Lobes sublinear to linear-elongate; rhizines densely branched. 
P. physcioides Nyl. 
29. Lobes shorter, subimbricate; rhizines sparsely to moderately 
branched P. zahlbruckncri Lynge 
25. Medulla C+ rose or red or C—, KC+ red. 
30. Medulla C+ rose (gyrophoric acid). 
31. Lobes linear-elongate, subdivaricatc. 
12. P. expor recta Kurokawa 
31. Lobes sublinear, adnate. 
32. Lobes plane to more or less canaliculate, pale brown in a rather 
wide zone at the margins below; spores 14-21 (i long. 
P. pluriformis Nyl. 
32. Lobes plane, pale brown only in a very narrow zone below; 
spores to 12 ft long. 
33. Thallus large, 4-8 cm. broad, the lobes 2-4 mm. wide. 
27. P. gcytophylla Kurokawa 
HALE & KUROSAWA—SUBGENUS PARMELIA 165 
33. Thallus smaller, 2-5 cm. broad, the lobes 1-2 mm. wide. 
P. revolutella Nyl. 
30. Medulla C+ red or C— (gyrophoric acid lacking). 
34. Medulla C+ red to orange red (olivetoric acid). 
P. intercalanda Vain. 
34. Medulla C—. 
35. Medulla KC+ red (alectoronic acid). 
36. Thallus adnate, the lobes sublinear. . . . 7. P. degelii Hale 
36. Thallus loosely attached, subdivaricate. 
37. Lobes narrow, 1-2 mm. wide. 
20. P. lineariloba Kurokawa 
37. Lobes wider, 2-4 mm. wide. 
38. Rhizines densely branched, about 1 mm. long; tropical 
America 16. P. gigas Kurokawa 
38. Rhizines moderately branched, 2 mm. or more long; 
Hawaii P. cervicornis Tuck. 
35. Medulla KC+ rose (alectoronic acid absent). 
39. Upper cortex distinctly maculate P, pulvinata F6e 
39. Upper cortex not distinctly maculate. 
40. Upper cortex flaking away, fragile; evernic acid present- 
P. taylorensis Mitch. 
40. Upper cortex continuous, not flaking; KC+ unknown sub- 
stances present. 
41. Spores uniseriate, 6-8/* long; North America. 
P. livid a Tayl. 
41. Spores biseriate, 14-18n long; South America. 
42, Thallus corticolous; apothecial disc flesh-colored. 
P. damaziatta Zahlbr. 
42. Thallus saxicolous; disc dark brown. 
P. gracilescens Vain. 
1. Parmelia anaptychioides Kurokawa, sp. no v. Plate 5 
Thallus adnatus, cinereo-albicans, 2-5 cm. diametro, lobis sub- 
linearibus, separates, 0.5-2.0 mm. latis, superne plus minusve conyexus, 
sorediatus, subtus niger, rhizinosus, rhizinis nigris, dichotome ramosis. 
Apothecia ignota. 
Thallus adnate on bark, pale olive gray to gray (R), 2-5 cm. in 
diameter; lobes dichotomously branched, sublinear-elongate, separate 
at the circumference, 0.5-2.0 mm. wide, 160-220 ft thick, the margins 
smooth, narrowly black-rimmed; upper surface more or less convex, 
shiny on younger lobes, dull and irregularly cracked on older lobes, 
sorediate, the soredia mainly laminal, 1-2 mm. in diameter, often 
fusing; medulla white; undersurface black, dark brown in a narrow 
zone at the tips, the rhizines black, branched. Apothecia not seen. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ faint red, 
P+ orange red, atranorine and protocetraric acid present. 
Type in Michigan State University, collected on low ridge connect- 
ing La Rucilla and Pico Trujillo, Maciso Central, Cordillera Central, 
166 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
elev. 8000 ft,, Dominican Republic, August 8, 1958, by C. Wetmore 
(no. 3617; isotype in US). 
This species is related to P, pseudosinuosa Asah. but the soralia are 
mainly laminal. The lobes are sublinear to linear-elongate and 
narrower than in P. pseudosinuosa. It is also close to P. subaffinis 
Zahlbr., especially in size and position of soralia, but it is differentiated 
by the absence of lichexanthone. Parmelia anaptychioides is known 
only from the West Indies, where it occurs on pine trees at higher 
elevations. 
Additional specimens examined: Jamaica: Blue Mountain Peak, 
7000 ft., Imshaug 15533 (MSC). Haiti: South of For£t des Pins, 
5500 ft., Dept. 1'Quest, Imshaug 22779 (MSC). 
2. Parmelia brevirhiza Kurokawa, sp. nov. Plate 3 
Thallus adnafcus, ca. 5 cm. diametro, olivaceo-cinerascens, lobis 
subirregularibus, 1-4 mm. latis, subrotundatis, margine crenatis, 
sup erne opacus vel nitidulus, submaculatus, partim leviter albo- 
pruinosus, sorediatus, sorediis capitatis, subtus niger, dense brevi- 
rhizinosus, rhizinis nigris, dichotome ramosis. Apothecia rara, 
adnata, usque ad 5 mm. diametro; hymenium 50-55 p altum; sporae 
male evolutae, 3-4X6-8 ju. 
Thallus adnate on bark, cream buff (R), about 5 cm. in diameter; 
lobes subirregular to sublinear, more or less subrotund and sub- 
imbricate, 1-4 mm. wide, 140-170 ti thick, the margins subcrenate; 
upper surface plane to rugulose, shiny and more or less white-maculate 
to dull and lightly white-pruinose, sorediate, the soredia mostly 
subterminal, strongly capitate, diffuse, medulla white; undersurface 
black, densely short rhizinate, dark brown and papillate in a very 
narrow zone at the tips, the rhizines black, finely branched, about 0.1 
mm. long, projecting as a narrow mat beyond the margins. Apothecia 
very rare, adnate, to 5 mm. in diameter, amphithecium sorediate, 
disc plane; hymenium 50-55 m high; spores poorly developed, 3-4X 
6-8 ft. 
Reactions: Thallus K-b yellow; medulla K+ yellow turning red, 
C—, KC— P+ pale orange red, atranorine and salacinic acid present. 
Type in the Naturhistoriska Riksmuseet, Stockholm, collected in 
Nothofagus rain forest, Isla Riesco, Mina Elena, Terr. Magallanes, 
Chile, Apr. 29, 1940, by R. Santesson (no. 2066; isotype in US). 
Parmelia brevirhiza has a distinct whitish cast, caused by faint 
pruinosity, and large subterminal and projecting soralia 2-4 mm, in 
diameter. The sorediate lobes are strongly revolute or contorted. 
This species resembles P. majoris Vain., which is known from Asia 
and Madagascar, in the presence of soredia and salacinic acid. Par- 
melia brevirhiza has more densely branched rhizines, which form a 
mat on the undersurface, Parmelia majoris contains zcorinc in 
167 
addition to salacinic acid. Parmelia brevirhiza is very common in 
southern Argentina and Chile and there is one quite unexpected record 
from Java. 
Additional specimens examined: Argentina: Lago Boca, Santesson 
1085 (S); Rio Piper, Ushuaia, Tierra del Fuego, Santesson 417 (S); 
Lago Verde, near Futalaufquen, Prov. Chubut, Lamb 5864 (CAN, 
US). Chile: Punta Guapacho, Peninsula Lacui, Isla Chilo6, Prov. 
Chilo6, Santesson 4135 (S); Cerros Divisaderos, Coyhaique, Terr. 
Ays6n, Santesson 4456 (S); Mina Elena, Isla Kiesco, Santesson 7849 
(S); Estancia Maria, Seno Skyring, Terr. Magallanes, Santesson 1974, 
1997 (S); Puerto Yartou, Canal Whiteside, Santesson 5791, 6759 (S); 
Rio Condor, south of Puerto Yartou, Canal Whiteside, Santesson 
5905 (S); Puerto Navarino, Isla Navarino, Santesson 7404 (S, US); 
Porvenir, Tierra del Fuego, Santesson 5372 (S). Java: Near Madang- 
Aer Batumbuk, Groenhart 9268 (BO). 
3. Parmelia canescens Kurokawa, sp. nov. 
Thallus adnatus, corticola, olivaceo-albicans, 3-7 cm. latus, lobis 
irregularibus, subrotundatis, 1-5 mm. latis, margine subcrenatis, su- 
perne opacus, parte albo-pruinosus, isidiis sorediisque destitutus, 
subtus niger, rhizinosus, rhizinis nigris, sparse vel modice ramosis, 
Apothecia adnata, 1.5-4.5 mm. diametro; hymenium 40-50 altum; 
sporae 5-6X8-10 m. 
Thallus adnate on twigs, pale olive buff (R), 3-7 cm. in diameter; 
lobes irregularly branched, subrotund apically, 1-5 mm. wide, the 
margins more or less crenate, rarely lobulate; upper surface plane, 
dull, sometimes faintly white-pruinose, not maculate, irregularly 
rugose or cracked on older lobes, isidia and soredia lacking; medulla 
white; undersurface uniformly black, sparsely to moderately branched; 
rhizines black, moderately branched, about 0.5 mm. long, Apothecia 
adnate, amphithecium rugulose, pruinose, disc Vandyke brown (R); 
hymenium 40-50 n high; spores 5-6X8-10 fi. 
Reactions: Thallus K+ yellow; medulla K+ yellow turning red C—, 
EC—, P+ pale orange, atranorine and norstictic acid present. 
Type in the Naturhistoriska Riksmuseet, Stockholm, collected at 
Hacienda de Cay queues, Colchagua, Chile, Aug. 22,1896, by P. Dus6n 
(no. 92; isotype in US). 
Parmelia canescens externally resembles P. sublaevigata (Nyl.) Nyl. 
in having a similar adnate thallus and irregularly branched lobes. 
Parmelia canescens contains norstictic acid, whereas P. sublaevigata 
contains salacinic acid. Moreover, the rhizines of P. canescens are 
sparsely to moderately branched while those of P. sublaevigata are 
more densely and finely branched. It is hoped that further collec- 
tions of these rare species will farther clarify their relationships. 
722-891—64 1 
168 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
4. Parmelia citrella Kurokawa, sp. nov. 
Thallus adnatus, corticola, viridiflavicans, 3-8 cm. latus, Iobis sub- 
Iinearibus, subimbricatis, 1.0-2.5 mm. Iatis, superne nitidulus, isidiis 
sorediisque desti tutus, subtus niger, rhizinosus, rhizinis nigris, furcatis. 
Apothocia adnata, 2-10 mm. diametro; liymenium 25-35 fx album; 
sporae 7-9X12-14 n. 
Thallus adnate on bark, sea-foam yellow (R), 3-8 cm. in diameter; 
lobes dichotomously branched, sublincar to linear-elongate, more or 
less imbricate, 1.0-2.5 mm. wide, 90-140p thick, the margins smooth, 
often narrowly black-rimmed; uppersurface plane to convex, rather 
shiny, isidia and soredia lacking, pyenidia forming blackish papillae 
on older lobes; medulla white; undersurface black, rhizinate, the 
rhizines black, rather long, moderately branched, often projecting 
beyond the margins. Apothecia adnate, 2-10 mm. in diameter, 
exciple crenate to undulate, disc Hay's brown to light seal brown 
(R), radially split, amphithecium smooth to rugose; liymenium 25- 
35/i high; spores 7-9X12-14/x. 
Reactions: Thallus K-f- yellowish; medulla K-f yellow turning 
red, C—, KC—, F+ pale orange red, usnic acid and salacinic acid 
present. 
Type in the Herbarium Bogoriense, collected at G. Gedeh, Java, 
elev. 2900 m., March 7, 1938, by II. van Woerden (isotype in US). 
Parmelia citrella is very close to P. reducens Nyl., but the thallus 
is more loosely attached and the rhizines are consistently longer. 
Parmelia reducens also differs chemically in producing norstictic 
acid, and on the basis of a few measurements the spores seem to be 
slightly smaller (9-1 lju long). Parmelia citrella appears primarily 
to be an Indonesian species, while P. reducens is known only from 
South America. 
Additional specimens examined: Panama: Craters edge, Volc&n 
Chiriquf, Pro v. Chiriqul, Scholander, Dec. 5-12, 1948 (MO, US). 
Colombia: Sierra do Santa Maria, elev. 8000 ft., Humbry-Troy 
313 (K). Java: G. Gedeh, van Woerden 2023 (BO); G. Pangcrango, 
Iieijnvaan 150 (BO); no locality, Ilorsficld (BM). 
5. Parmelia crenata Kurokawa, sp. nov. 
Thallus laxe adnatus, muscicola, albido-eineraseens, 3-6 cm. dia- 
metro, lobis subirregularibus, subimbricatis, apice subrotundatis 
vel subtruncatis, 1.5-5.0 mm. Iatis, margine crenatis, superne 
laevigatus, emaculatus, isidiatus, subtus niger, ambitu castaneus, 
sparse rhizinosus, rhizinis nigris, modice furcatis. Apothecia ignota. 
Thallus loosely attached, growing over mosses on rocks, yellowish 
glaucous to light mineral gray (R), 3-0 cm. in diameter; lobes ir- 
regularly branched, more or less imbricate, rounded at the tips, 
HALE & KUROSAWA—SUBGENTJS PARMELIA 169 
1,5-5.0 mm. wide, 90-110 p thick, the margins crenate; upper surface 
smooth, without maculae, moderately isidiate, the isidia short, less 
than 0.5 mm. high, often branched; medulla white; undersurface 
black, pale brown in a narrow zone at the tips, sparsely rhizinate, 
the rhizines black, sparsely to moderately branched. Apothecia not 
seen. 
Reactions: Thallus K+ yellow; medulla K+ yellow, C—, KC—, 
P-h pale orange, atranorine and stictic acid present. 
Type in the National Science Museum, Tokyo, collected on Mount 
Kuishi, Tosa-gun, Tosa-ken, Shikoku, Japan, Dec. 14, 1960, by S. 
Kurokawa (no. 60154; isotype in US). 
Parmelia crenata might at first be mistaken for P, koyaensis Asah., 
but the lobes are consistently smaller and more or less sublinear. 
It is the only species in section Hypotrachyna that contains stictic 
acid. Parmelia koyaensis contains protocetraric acid. Thus far 
P. crenata is known only from the type locality in Japan. 
6. Parmelia croceoptietulata Kurokawa, sp. nov. Pirate 9 
Thallus adnatus, albido-cinerascens, ca. 5 cm. diametro, lobis 
sublinearibus, 1.5-3.5 mm. latis, margine integris, superne nitidus, 
emaculatus, aetate rugulosus, pustulato-sorediatus, soraliis partim 
subcoloratis, subtus niger, rhizinosus, rhizinis nigris, ramosis. 
Apothecia adnata, 1-6 mm. diametro; hymenium 50-60 p altum; 
sporae non visae. 
Thallus adnate on bark, yellowish glaucous (R), 4-9 cm. in diameter; 
lobes dichotomously branched, sublinear, 1,5-3.5 mm. wide, 150- 
220 n thick, the margins entire to more or less dissected; upper surface 
shiny, rugulose with age, without maculae, sorediate, the soralia 
capitate and sub terminal, 1-5 mm. in diameter, originating from 
pustules; medulla below the soralia yellow ochre (R), otherwise 
white; undersurface black, rhizinate, the rhizines black, shiny, 
branched. Apothecia rare, adnate, 1-6 mm. in diameter, exciple 
sorediate, disc chestnut brown (R), radially split; hymenium 50-60 m 
high; mature spores not seen. 
Reactions: Thallus K-b yellow; medulla K—, C—, KC— or KC+ 
reddish, P+ orange red or in part P—, soralia P+ orange red, pig- 
ment under the soralia K-f purple, atranorine, protocetraric acid, 
and rliodophyscin present. 
Type in Michigan State University, collected on fir around summit 
of Grandfather Mountain, A very Co., North Carolina, elev. 5964 ft., 
June 23, 1958, by IT. Imshaug (no. 22275; isotype in US). 
Parmelia croceopustulata is related to species in the P. Iwida group, 
especially P. formosana Zahlbr. and P. leiophylla Kurokawa, both fo 
which also produce rliodophyscin. Parmelia formosana differs in 
having nonsorediate pustules and lichexanthone. Parmelia leiophylla 
170 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
has similar sorediate pustules but produces olivetoric acid. This 
species seems to be restricted to higher elevations in Hispaniola and 
Jamaica, with several interesting disjunct localities in the Appalachian 
Mountains. 
Additional specimens examined: United States: Virginia: White 
Top Mountain, Washington Co., elev. 5200 ft., Hale 18662 (US); 
North Carolina: Roan Mountain, Avery Co., elev. 6200 ft., Hale 18071 
(US); Mt. Mitchell, Yancey Co., elev. 6684 ft., Itmhaug 22383, 
22390 (MSC, US); Newfound Gap, Swain Co., Imshaug 22435 (MSC, 
US); Mt. Pisgah, Hay wood Co., Green in 1959 (US). Dominican 
Republic: Trail to Alto de la Bandera, Cordillera Central, La Vega, 
Jmshaug 23470 (MSC, US). Haiti: Pic La Selle, Morne La Selle, 
elev. 8844 ft., Wetmore 3120 pr. p. (MSC); above Le Refuge, Montagne 
Noire, Kenscoff, Imshaug 22538 (MSC). Jamaica: Mossmans Peak, 
Imshaug 14726 (MSC); High Peak, Blue Mountains, elev. 6800 ft., 
Imshaug 15254 (MSC). 
7. Parmelia degclii Hale, sp. no v. Plate 5 
Thallus arete adnatus, corticola, cinereo-albicans, 4-8 cm. diametro, 
lobis sublinearibus, 1-2 mm, latis, contiguis, superne rugulosus, 
isidiis sorediisque destitutus, subtus modice rhizinosus, rhizinis sparse 
vel modice dichotome ramosis. Apothecia adnata, 3-5 mm. diametro; 
hymenium 50-60 y. altum; sporae 4-5X8-11 #i. 
Thallus closely adnate on bark, 4-8 cm. in diameter, mineral gray; 
lobes sublinear, 1-2 mm. wide, 140-170 n thick, contiguous, the 
margins entire; upper surface dull to more or less shiny, rugulose, 
transversely rugose with age, isidia and soredia lacking; medulla 
white; undersurface black, moderately rhizinate, the rhizines sparsely 
to moderately branched, dichotomous. Apothecia adnate, 3-5 mm. 
in diameter, the exciple crenate, disc plane; hymenium 50-60 n high; 
spores 4-5X8-11 /t. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, P—, 
atranorine, alectoronic acid, and a-collatolic acid present. 
Type in the herbarium of G. Degelius, collected between Luso and 
Cachipoque, Moxico, Angola, about 1300 m, elevation, Feb. 16, 1960, 
by G. Degelius (isotype in US). 
Parmelia degelii is very close to P. livida Tayl. in general habit and 
color, but P. livida, a temperate American lichen with several localities 
in South Africa, has smaller spores (4-5X6-8 jx), somewhat denser 
and more richly branched rhizines, and a different chemistry (KC+ 
red unknown). Parmelia degelii is unusual in being the second known 
species of section Hypotrachyna with alectoronic acid in Africa. The 
other species is P. exsplendens Hale, which also occurs in the Caribbean 
area. The five remaining species in section Hypotrachyna with alec- 
toronic acid occur exclusively in tropical America (P. densirhizinata 
171 
Kurokawa, P. ensifolia Kurokawa, P. gigas Kurokawa, and P. 
linearUoba Kurokawa) or in Hawaii (P. cervicomis Tuck.). 
Additional specimens examined: Angola: Same locality and data 
as the holotype (DEGEL); Rio Kuiriri, near Kassuango, Bie, Goss- 
weiler 3256d (BM). 
8. Parmelia densirhizinata Kurokawa, Bp. nov. Plate S 
Thallus laxe adnatus, corticola, albido-cinerascens, 6-13 cm. latus, 
lobia lineari-elongatis, subdivaricatis, 2-6 mm. latis, margine integris, 
superne planus, emaculatus, pustulato-sorediatus, soraliis subtenni- 
nalibus, subtus nigcr, rhizinosus, rhizinis densis, dichotome ramosis. 
Apothecia ignota. 
Thallus loosely adnate on bark, pale glaucous green (R), 6-13 
mm. in diameter; lobes dichotomously branched, linear-elongate, 
subdivaricate, 2-6 mm. wide, 110-150/* thick, the margins smooth, 
narrowly black-rimmed; upper surface plane, continuous, without 
maculae, sorediate, the soralia mainly subterminal, 1-3 mm. in diam- 
eter, originating from pustules; medulla white; undersurface black, 
densely rhizinate, the rhizines black, shiny, dichotomously branched, 
1-2 mm. long, projecting in a mat beyond the margins. Apothecia 
not seen. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, P—, 
atranorine and alectoronic acid present. 
Type in the U.S. National Herbarium, collected on the east side of 
Volc&n Chiriqui, Pro v. Chiriqui, Panama, elev. 1500-3000 ft., Dec. 
5-12, 1948, by P. F. Scholander. 
Parmelia densirhizinata is a typical subdivaricate species related 
to P. gigas Kurokawa, a tropical American species, and P. cervicomis 
Tuck., a Hawaiian species. They all produce alectoronic acid, but 
P. densirhizinata is easily distinguished by the subterminal sorediate 
pustules. It is widespread in tropical America, occurring in moun- 
tains between 2000 and 3750 m. elevation. 
Additional specimens examined: Guatemala: Pacajregion of 
Desconsuelo, Totonicap&n, Standley 84522 (MO). Panama: Craters 
edge, Volc&n Chiriqui, Prov. Chiriqul, Scholander, Dec. 11, 1948 (US). 
Haiti: Ridge east of Pic La Selle, Morne La Selle, elev. 8300 ft., 
Imshaug 23026 (MSC). Dominican Republic: Trail to Alto de la 
Bandera Cordillera Central, La Vega, elev. 7200 ft., Imshaug 23486 
(MSC). Colombia: P&ramo de Guasca, Dept. Cundinamarca, Killip 
34118 (US); Chorredn San Paulina, Nevada del Cocuy, Dept. Boyac&, 
elev. 3750 m., Cuatrecasas 1348a (US). Ecuador: Mount Tunguragua, 
Spruce 175 (K). Peru: Pampalca, Dept. Ayacucho, elev. 3200 m., 
KUlip <& Smith 22237 (US). Bolivia: Unduavi Valley, Jvlw413 (US). 
Chile: Isla de Chiloe, Joseph 2245 (US); Cerro Tralcan, Lago Rinihue, 
172 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Prov. Valdivia, Santesson 3451 (S); Canal Whiteside, Puerto Yartou, 
Tierra del Fuego, Santesson 6799 (S). 
9. Parmelia dcntella Hale & Kurokawa, sp. nov. Plate 5 
Thallus laxe adnatus, saxicola, viridi-cinerascens, 3-8 cm. diametro, 
lobis sublinearibus vel subirregularibus, subimbricatis, 2-6 mm. 
latis, margine dentato-crenatis, sup erne planus, albomaculatus, dense 
isidiatus, subtus nigricans, sparse vel modice rhizinosus, rhizinis 
nigris, ramosis. Apothecia ignota. 
Thallus loosely adnate on shaded rocks, yellowish glaucous to 
glaucous green (R), 3-8 cm. in diameter; lobes irregularly branched, 
sublinear to subirregular, more or less imbricate, 2-6 mm. wide, 
150-230 n thick, the margins in part dentate-crenate; upper surface 
plane, distinctly white-maculate, densely isidiate, the isidia often 
branched, cylindrical, less than 0.7 mm. high; medulla white; under- 
surface black, pale brown in a rather broad zone at the tips, sparsely 
to moderately rhizinate, the rhizines black, shiny, moderately 
branched, about 1 mm. long. Apothecia not seen. 
Reactions: Thallus K+ yellow; medulla K+ yellowish, C—, 
KG—, P+ pale orange red, atranorine and an unknown P+ substance 
present. 
Type in the U.S. National Herbarium, collected on granite at 
Cheaha State Park, Clay County, Alabama, Mar. 16, 1962, by H. 
McCullough (no. 570; isotypes in TNS and the herbarium of Howard 
College, Birmingham, Alabama). 
Parmelia dcntella bears a striking resemblance to the endemic 
Japanese species P. crenata Kurokawa. It differs from P. crenata in 
having the upper cortex distinctly white-maculate and in producing 
a P+ unknown substance instead of stictic acid. It also resembles 
P. costaricensis Nyl., especially in the maculae, isidia, and size 
of the thallus, but the rhizines of P. costaricensis arc much more 
densely branched and its main component is caperatic acid (P—), 
Parmelia dentella is still known only from the type locality in Alabama, 
a rather dry oak forest rich in many kinds of lichens. It will probably 
be found again in the foothills of the Appalachian Mountains. 
10. Parmelia ensifolia Kurokawa, sp. nov. Plate 7 
Thallus adnatus, albido-cinerascens, 5—11 cm. latus, lobis subline- 
aribus, 1-4 mm. latis, margine lobulatis, superne nitidus, emaculatus, 
isidiato-lobulatus, subtus niger, rhizinosus, rhizinis nigris, dichotome 
ramosis. Apothecia adnata, 3—4 mm. diametro; hyraenium 50-65 p 
altum; sporae 8-10X14-18//. 
Thallus loosely adnate, corticolous, pearl gray (R), 5-11 cm. in 
diameter; lobes dichotomously branched, sublincar-elongate, 1—4 mm. 
wide, 160-200 n thick, the margins with numerous lobules, the lobules 
HALE & KUROKAWA—SUBGENUS PARMELIA 173 
ascending, dorsiventral, short-rhizinate below, often branched, up 
to 1.5 mm. long; upper surface shiny, without maculae, short-isidiate- 
lobulate to lobulate on the margins; medulla white; undersurface 
black, densely rhizinate, the rhizines black, shiny, dichotomously 
branched. Apothecia adnate, 3-4 mm. in diameter, disc tawny 
olive (R), amphithecium rugose; hymenium 50-65 m high; spores 
8-10X14-18 v. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ rose or 
red, P—, atranorine and alectoronic acid present. 
Type in Michigan State University, collected in pine forest below 
summit of Alto de la Bandera, Cordillera Central, La Vega, Dominican 
Republic, elev. 8300 ft., Aug. 5, 1958, by H. Imshaug (no. 23430; 
isotype in US). 
Parmelia ensifolia has sublinear lobes and laminal and marginal 
lobules similar to those of P. culmigena Zahlbr. and P. digitata Lynge. 
The latter two species, however, are distinctly maculate and differ 
chemically, P. culmigena having evernic acid and P. digitata having 
barbatic acid. Parmelia ensifolia is distinguished by the presence 
of alectoronic acid and lack of maculae. It seems to be rather common 
in tropical America. 
Additional specimens examined: Mexico: No data on locality, 
1890 (S). Panama: Volc&n Chiriqui, Pro v. Chiriqui, Scholander, 
Dec. 5-12, 1948 (MO, US). Jamaica: Summit of Blue Mountain 
Peak, Maxon 9865 (US). Haiti: Pic La Selle, Morne La Selle, Wetmore 
3086 (MSC, US). Dominican Republic: Below summit of Alto de 
la Bandera, Cordillera Central, La Vega, elev. 8300 ft., Wetmore 3511 
(MSC, US). Venezuela: Pan de Azticar, Merida, Jahn 241 (US); 
Laguna de Canoa, Sierra de Santo Domingo, Merida, Dennis 1924, 
1950 (K). 
11. Parmelia erythrodcs (Zahlbr.) Hale & Kurokawa, comb, nov. 
Parmelia brasiliana Nyl. var. erytkrodes Zahlbr. Denkschr. Akad. Wiss. 
Wien Math. Naturw. 83: 169. 1927. Type collection: Mt, Itatiaya, 
Minas Gerais, Brazil, Schiffner, September 1901 (WU, holotype). 
Thallus adnate on rock, ivory yellow, deep mouse gray towards 
the center, about 5 cm. in diameter; lobes dichotomously branched, 
sublinear-elongate, 1-2 mm. wide, 140-190 m thick, the margins 
entire to subcrenate; upper surface plane, shiny, emaculate, tan- 
gen tially cracked on older lobes, isidia and soredia lacking; medulla 
pale pinkish buff in the lower half, white above; undersurface uni- 
formly black, moderately rhizinate, the rhizines black, moderately 
branched, 0.5-1.0 mm. long. Apothecia adnate, 2-6 mm. in diameter, 
exciple undulate, disc imperforate, blackish brown; hymenium 50- 
60 m high; spores 5-6X9-10 At, episporium 1 /x thick. 
174 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ rose, 
P—, pigment K+ reddish purple, atranorine, a KC+ unknown 
(apparently the same as the one in P. limda Tayl.), rhodophyscin, 
and lichexanthone. 
Parmelia erythrodes is very close to P. brasiliana Nyl., differing 
chiefly in chemistry. Parmelia brasUiana lacks rhodophyscin and 
the KC+ substance but contains lichexanthone and in addition 
protocetraric acid. Both species are typically saxicolous and occur 
rather rarely in South America. 
Additional specimen examined: Peru: Without locality, Lobb s.n. 
(BM). 
12. Parmelia expo ire eta Kurokawa, sp. no v. Plate 6 
Thallus laxe adnatus, albido-cinerascens, 7-10 cm. latus, lobis 
lineari-elongatis, subdivaricatis, 2-6 mm. latis, superne planus, 
nitidus, emaculatus, sorediis isidiisque destitutus, subtus niger, 
rhizinosus, rhizinis nigris, dense ramosis. Apothecia ignota. 
Thallus loosely adnate on bark, pale glaucous green (R), 7-10 cm. 
in diameter; lobes dichotomously branched, linear-elongate, subdi- 
varicate, 2-6 mm. wide, 150-220 ju thick, the margins smooth, rarely 
lobulate; upper surface plane and smooth, shiny, often pruinose near 
the tips, without maculae, isidia or soredia; medulla white; under- 
surface black, rhizinate, the rhizines black, shiny, densely branched, 
about 1 mm. long. Apothecia not seen. 
Reactions: Thallus K+ yellow; medulla K—, C+ rose, KC+ 
red, P—, atranorine and gyrophoric acid present. 
Type in the United States National Herbarium, collected in mature 
pine-oak forest, on trail leading to Cerro San Felipe, Oaxaca, Mexico, 
elev. about 3000 m., Apr. 1, 1960, by M. E. Hale (no. 20722). 
Parmelia exporrecta seems to be the nonsorediate ally of P. thysanota 
Kurokawa (see below). It has a large subdivaricate thallus and linear- 
elongate lobes, much as in P. gigas Kurokawa, which differs in con- 
taining alectoronic acid. 
Additional specimen examined: Mexico: VeraCruz: Orizaba, Lieb- 
mann 116 (UPS). 
13. Parmelia exaplendens Hale, sp. nov, 
Thallus adnatus, viridi- vel cinereo-albicans, 4-8 cm. diametro, 
lobis sublinearibus, 1.5-2.5 mm. latis, superne nitidus, valde albomac- 
ulatus, sorediatus, soralis subterminalibus, subtus niger, dense 
rhizinosus, rhizinis dichotome ramosis. Apothecia ignota. 
Thallus adnate, greenish to whitish gray, 4-8 cm. across, corti- 
colous; lobes sublincar, dichotomously branched, 1.5-2.5 mm. wide, 
160-200 fi thick, the margins entire to subcrenate; upper surface 
plane, shiny, strongly white-maculate, sorediate, soralia capitate, 
175 
subterminal; medulla white; undersurface black, densely rhizinate, 
the rhizines black, dichotomously branched. Apothecia unknown. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, 
P—, atranorine and alectoronic acid present. 
Type in Michigan State University, collected on ridge northwest of 
Murdock's Gap, Blue Mountains, Jamaica, elev. 3600 ft., Mar. 19, 
1953, by H. Imshaug (no. 15306; isotype in US). 
This species resembles P. metarevoluta Asah. from Asia in having 
a medium-sized thallus and subterminal capitate soralia. However, 
it is distinctly white-maculate and the rhizines are densely branched. 
Parmelia exspleridens is fairly common in the Caribbean with one 
disjunct record from South Africa. 
Additional specimens examined: Mexico: Virgin upland jungle, 
El Suspiro, Chiapas, Hale 20229 (US). Guatemala: Near Tactic, 
Alta Verapaz, Standiey s.n. (MO, no. 185561). Jamaica: Gap south- 
east of Catherine's Peak, Imshaug 13894 (MSC); gap on northeast 
spur of High Peak, Imshaug 15287 (MSC); Woodcutters Gap, 
Imshaug 13423 (MSC, US); Summit, Blue Mountains, Imshaug 
13855 (MSC); west slope of John Crow Mountains, above Corn Puss 
Gap, Imshaug 14577 (MSC). Union of South Africa: 5 mi. east of 
Mokobulaan, Lydenberg, Transvaal, Almbom 7496 (LD). 
14. Parmelia fissicarpa Kurokawa, sp. nov. Plate 7 
Thallus adnatus vel laxe adnatus, albido-cinerascens, ca. 14 cm. 
diametro, lobis sublinearibus vel subirregularibus, 2-4 mm. latis, 
margine subcrenatis superne nitidulus, emaculatus, isidiis sorodiisque 
destitutus, subtus niger, dense rhizinosus, rhizinis nigris, ramosis. 
Apothecia substipitata, ca. 10 mm. diametro; hymenium 60-70 n 
altum; sporae 6-7X8-10#. 
Thallus adnate to loosely attached on rock, pale olive buff (R), 
about 14 cm. in diameter; lobes irregularly branched, sublinear to 
subirregular, 2-4 mm. wide, 210-250 /i thick, the margins subcrenate; 
upper surface rather shiny, without maculae, the older lobes rugulose, 
isidia and soredia lacking; medulla white; undersurface black, 
densely rhizinate, the rhizines black, shiny, moderately branched, 
often more than 1.5 mm. long. Apothecia substipitate, 10 mm. or 
more in diameter, exciple undulate, amphithecium rugose, disc 
carob brown (R), radially split; hymenium 60-70 m high; spores 
6-7X8-10 p. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ faint 
red, P+ orange red, atranorine and protocetraric acid present. 
Type in the Botanical Museum, Lund University, collected 2 miles 
south of forester's office, Cathedral Peak Area, Bergville, Natal, Union 
of South Africa, Nov. 7, 1953, by Ove Almbom (no. 9374; isotype in 
US). 
176 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
This peculiar species has an exceptionally large thallus and 
apothecia. It resembles P. insinuans NyL, a tropical American 
species, and P. keitauensis As ah. and P. manilensis Vain., both Asian 
species, in having rather irregularly branched lobes and protocetraric 
acid. Although P. jissicarpa and P. insinuans have similar spore size, 
the thallus of P. Jissicarpa is much larger and the rhizines are longer. 
In addition, P. insinuans often produces the pigment rhodophyscin 
along with protocetraric acid. Parmelia jissicarpa is clearly dis- 
tinguished from P. keitauensis by the absence of fatty acids and by 
smaller spores. Parmelia manilensis differs in having adnate and 
quite small apothecia (1-3 mm. in diameter) and large spores. 
15. Parmelia flavovirens Kurokawa, sp. nov. Plate 8 
Thallus adnatus, corticola, viridiflavicans, 3-6 cm. diametro, lobis 
sublinearibus, 1.0-2.5 mm. latis, margine subcrenatis, superne nitidus, 
aetate rugulosus, centrum versus lobulatus, sorediatus, soraliis sub- 
terminalibus, subtus niger, dense rhizinosus, rhizinis nigris, ramosis. 
Apothecia adnata, 2-4 mm. diametro; hymenium 60-75 n altum; 
sporae 7-10X11-15 p. 
Thallus adnate on bark, sea-foam green (R), 3-6 cm. in diameter; 
lobes more or less irregularly branched, sublinear, 1.0-2.5 mm. wide, 
130-150 n thick, the margins more or less crenate, often narrowly 
black-rimmed; upper surface shiny near the tips, more or less rugulose 
and irregularly cracked on older lobes, sometimes lobulate toward the 
center, sorediate, soralia mainly subterminal; medulla white; under- 
surface black, densely rhizinate, the rhizines black, branched. Apo- 
thecia adnate, 2-4 mm. in diameter, exciple crenate, amphithecium 
rugose, disc light seal brown (R); hymenium 60-75 p high; spores 
7-10X11-15 n. 
Reactions: Thallus K-j- yellowish; medulla K—, C—, KC+ faint 
red, P+ deep orange red, usnic acid, and protocetraric acid present. 
Type in the Naturhistoriska Riksmuseet, Stockholm, collected 
near Hotel Rio Rubens, 50 km. southeast of Natales, Magallanes, 
Chile, Jan. 14, 1941, by R. Santesson (no. 5681; isotype in US). 
Parmelia Jlavovirens is very close in external appearance to P. 
sinuosa (Sm.) Ach. and P. velloziae Vain. The last named species 
has similar chemistry but contains in addition the orange-red pigment 
rhodophyscin under the soralia. Parmelia sinuosa produces salacinic 
acid. Parmelia jlavovirens is apparently endemic to southernmost 
Chile where it occurs intermingled with P. sinuosa. 
Additional specimens examined: Chile: Puerto Navarino, Isla 
Navarino, Tierra del Fuego, Santesson 1261 (S); Ays6n: Coyhafgue, 
Santesson 8406 (S). 
HALE & KUROSAWA—SUB GENUS PARMELIA 177 
16. Parmelia gigas Kurokawa, sp, nov. Plate 8 
Thallus laxe adnatus, corticola, albido-cinerascens, 15—40 cm. 
diamet.ro, lobis lineari-elongatis, subdivaricatis, 3-10 mm. latis, 
margine integris, superne planus, emaculatus, isidiis sorediisque desti- 
tutus, subtus niger, breve rhizinosus, rhizinis nigris, dense ramosis. 
Apothecia usque ad 15 cm. diametro, substipitata; hymenium 50-65 p 
altum; sporae 8-10X16-18 /*• 
Thallus loosely attached on bark, mosses, or humus, turning from 
ivory yellow to cream buff (R) in the herbarium, 15-40 cm. in di- 
ameter; lobes dichotomously branched, linear-elongate, subdivaricate 
and separate, 3-10 mm. wide, 220-330 y, thick, the margins smooth, 
rarely short lobulate; upper surface plane and smooth, shiny, without 
maculae, isidia and soredia lacking; medulla white; undersurface 
black, short rhizinate, partly naked, the rhizines black, shiny, densely 
branched, about 1 mm. long, forming a mat projecting narrowly 
beyond the margins. Apothecia rare, to 15 mm. in diameter, sub- 
stipitate, amphithecium strongly rugose, faintly maculate; hymenium 
50-4)5 n high; spores 8-10X16-18 n, the episporium 1.5 n thick. 
Reactions: Thallus K-f- yellow; medulla K—, C—, KC-f- red, 
P—, atranorine and alectoronic acid present. 
Type in the United States National Herbarium, collected at Laguna 
Verjou, above Bogota, Colombia, 1922, by Ariste Joseph (no. B70; 
isotypes in DUKE, REN, S, TNS). 
This species has a large divaricate thallus, long linear-elongate lobes, 
and densely branched, short rhizines. Externally it is very similar 
to and has been misidentified as P. caraccensis Tayl., which differs 
significantly in producing usnic and norstictic acids. It is also 
related to the Hawaiian species Pt cervicornis Tuck, in that it produces 
the same chemical substances. The thallus of P. gigas, however, is much 
larger and the lobes ordinarily wider than in P. cervicornis. The 
rhizines of P. gigas, furthermore, are short, usually about 1 mm. long, 
and form a dense mat on the lower side, whereas the rhizines of P. 
cervicornis are often more than 2 mm. long and relatively sparse. 
Parmelia gigas is widely distributed in tropical America, being 
especially common at higher elevations (up to 3900 m.) in cloud 
forests. 
Additional specimens examined: Mexico: Oaxaca: Trail leading to 
Cerro san Felipe, elev. 2920-3000 m., Hale 20707, 20723, 20783, 21100 
(US). Panama: Craters edge, Volc&n Chirlquf, Prov. ChiriquI, Schol- 
ander, Dec. 12, 1948 (MO). Venezuela: Pan de Azticar, Cordillera de 
M&rida, elev. 4000 m., Jakn 240 (US). Colombia: Carrizales, north 
of Las Brisas, Cordillera Occidental, Dept. Valle, Cuatrecasas 20545 
(US); eastern slope of P&ramo de las Coloradas, Dept. Santander, 
elev. 3300-3900 m., Killip & Smith 18394, 18481 (US); Los Gaques, 
178 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
western slope of Pfiramo de Guasca, Dept. Cundinamarca, elev. 3250 
m., KUlip 34064B (US); without locality: Lindig 2573 (UPS). Ecua- 
dor: P&ramo de Minza Chica, Tunguragua, elev. 3800 m., Penland 
404 (MO, US); Paramo de Volc&n Cayambe, Dept. Pichincha, elev. 
13,000 ft., Prescott (MSC, US). 
17. Parmclia immaculata Kurokawa, sp. nov. 
Thallus adnatus, corticola, albido-cinerascens, 4-9 cm. latus, lobis 
sublinearibus, contiguis, 1-3 mm. latis, superne nitidulus, aetate 
rugosulus, sorediatus, soraliis laminalibus, subtus sparse rhizinosus, 
rhizinis nigris, brevibus, dichotome ramosis. Apothecia rara, sub- 
stipitata, 1.5-4.5 mm. diametro; hymenium 50-60/i altum; sporae 
5-7X9-10 jtt. 
Thallus adnate on bark, yellowish glaucous (R), 4-9 cm. across; 
lobes dichotomously branched, sublinear-elongate, contiguous, 1-3 mm. 
wide, 110-200 ju thick, the margins smooth or sparsely lobulate with 
age; upper surface rather shiny, plane, becoming rugulose and irregu- 
larly cracked on older lobes, sorediate, the soralia orbicular, laminal, 
often fusing; medulla white; undersurface sparsely rhizinato, minutely 
rugulose, black, the rhizines black, short, dichotomously branched. 
Apothecia more or less stipitate, 1.5^4.5 mm. in diameter, exciple 
crenate, amphithecium sorediate, disc vandyke brown (R), radially 
split; hymenium 50-60 p high; spores 5-7X9-10 p. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ rose, P—, 
pigment if present K+ purple, atranorine, KC+ unknown, and 
rhodophyscin present. 
Type in the Botanical Museum, Lund University, collected at 
Punch Bowl Inn, north of Louis Trichardt, Zoutpansberg, Transvaal, 
Union of South Africa, Oct. 11, 1953, by Ove Almborn (no. 6551; 
isotype in US), 
Parmelia immaculata belongs to that group of species near P. dactyl- 
ijera Vain, and P. Uvida Tayl., all of which contain the same unknown 
KC-f- substance. Parmelia daciylifera is pustulate-isidiate, whereas 
P. Uvida lacks pustules and soredia. Parmelia immaculata is appar- 
ently a very common corticolous lichen in South Africa, occurring 
frequently with P. leiophylla Kurokawa. A single specimen from 
Java also seems to belong to this species. 
Additional specimens examined: Africa: South of Felixburg, South- 
ern Rhodesia, Hoeg, April 2, 1930 (TRH); Punch Bowl Inn, north of 
Louis Trichardt, Zoutpansberg, Transvaal, Almborn 6440, 6550 (LD), 
6247 (LD, US); Polela Forest, Polela, Natal, Almborn 9511 (LD, US); 
Stormsrivier, Humansdorp, Almborn 4132 (LD); Grootvadersbosch, 
Swellendam, Cape Province, Almborn 2\§±} 2195,2196 (LD); Grahams- 
town, Albany, Cape Province, Hoeg, December 1929 (TRH). Java: 
Tjibodas, Neervoort 3140 (BO). 
HALE & KUROKAWA—SUBGENUS PARMELIA 179 
18. Parmelia infirma Kurokawa, sp. nov. Plate 7 
Thallus laxe adnatus, corticola, olivaceo-albicans, 3-8 cm. diametro, 
lobis sublinearibus, subimbricatis, 1.0-3.5 mm. latis, margine sub- 
crenatis, superne nitidus, emaculatus, isidiatus, subtus niger, breviter 
rhizinosus, rhizinis nigris, ramosis. Apothecia adnata, 1.0-2.5 mm. 
diametro; hymenium 60-85 ft altum; sporae 9-13X15-19p . 
Thallus loosely adnate on bark, yellowish glaucous (R), 3-8 cm. 
across; lobes irregularly branched, sublinear, sometimes imbricate, 
1.0-3.5 mm. wide, 130-170 n thick, the margins more or less crenate, 
sometimes blackrimmed; upper surface shiny, without maculae, 
isidiate, the isidia cylindrical, moderate, simple to coralloid branched; 
medulla white; undersurface black, short-rhizinate, the rhizines 
black, shiny, branched. Apothecia adnate, 1,0-2.5 mm. in diameter, 
exciple entire, amphithecium usually isidiate, disc seal brown (R); 
hymenium 60-85 fx high; spores 9-13X15-19 ju. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P—, 
atranorine, caperatic and protolichesteric acids present. 
Type in the National Science Museum, Tokyo, collected at Dar- 
jeeling, Kurseong, India, Apr, 12, 1960, by H. Hara et al. (isotype in 
US). 
This new species resembles P. koyaensis Asah. in external appearance, 
spore size, and presence of fatty acids. The isidia of P. infirma are 
shorter and sometimes become more or less granular. Pamnelia 
koyaensis is P+ red, containing protocetraric acid, which has not 
been demonstrated in P. infirma,. The related isidiate species P. 
nodakensis Asah. is KC+ rose and P. crenata Kurokawa contains 
stictic acid (K+ yellow). Parmelia infirma also resembles the tropical 
American species P. costaricensis NyL, which is distinctly maculate. 
Parmelia infirma is known only from the type locality. 
19. Parmelia leiophylla Kurokawa, sp. nov. Plate 9 
Thallus adnatus, corticola, cinereo-albicans, 3-7 cm. diametro, 
lobis sublinearibus, contiguis, 1-3 mm. latis, superne planus, nitidulus, 
aetate rugosus, pustulatus, medulla pro parte ochracea, subtus niger, 
rhizinosus, rhizinis nigris, dichotome ramosis. Apothecia rara, 
adnata, usque ad 1 cm. diametro; hymenium 50-55 tt altum; sporae 
5-3X10-13 ju. 
Thallus adnate, corticolous, yellowish glaucous (R), 3-7 cm. across; 
lobes dichotomously branched, sublinear, contiguous, 1-3 mm. wide, 
130-180 p thick, the margins smooth; upper surface plane, rather shiny, 
rugulose on older lobes, pustulate, not distinctly sorediate, without 
maculae; medulla white, turning yellow ochre (R) under the pustules; 
undersurface black, rhizinate, the rhizines black, shiny, dichotomously 
branched, rarely projecting beyond the margins. Apothecia very 
180 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
rare, adnate, to 1 cm. in diameter, amphithecium rugose, disc split 
with age, rarely abnormally perforate; hymenium 50-55 n high; 
spores 5-6X10-13 /i. 
Reactions: Thallus K+ yellow; medulla K~, C+ deep red, KC-f- 
red, P—, pigment K+ purple, atranorine, olive toric acid, and rhodo- 
physcin present. 
Type in the Botanical Museum, Lund University, collected on 
Forest Drive from Houtbosch to Tzaneen, Pietersburg, Transvaal, 
Union of South Africa, Oct. 12, 1953, by Ove Almborn (no. 6796; 
isotype in US). 
This species resembles P. livida Tayl. in general habitus but is 
separated by the sorediate pustules and chemistry. Parmelia im- 
maculata Kurokawa, which has similar distribution in South Africa, 
is distinctly sorediate and C—, KC+ rose. Parmelia formosana 
Zahlbr. is strictly pustulate and contains lichexanthone. The Ameri- 
can species P. croceojmstulala, Kurokawa has similar coarsely sorediate 
pustules and rhodophyscin, but the main component is protocetraric 
acid (C—, P+ red). 
Additional specimens examined: Union of South Africa: Transvaal: 
Punch Bowl Inn, north of Louis Trichardt, Zoutpansberg, Almborn 
6435 (LD); Natal: South of Nkandhla, Nkandhla, Almborn 8074 (LD), 
Impetyne Forest near Weza, Alfred, Almborn 9969 (LD), Indumeni 
Forest, Cathedral Peak Area, Bergville, Almborn 9191 (LD); Cape 
Province: Deep wall Forest, Knysna, ArneU 1632a, 1645 (LD), Table 
Mountain, Arnell 861 (LD), Maas Geesteranus 12167 (L), along 
Skeleton Stream, east slopes of Table Mountain, Wynberg, Almborn 
11153 (LD), Fern Kloof, Albany, Almborn 10735 (LD). 
20. Parmelia lineariloba Kurokawa, sp. nov. Plate 6 
Thallus adnatus, corticola, cinereo-albicans, 3-5 cm. diametro, lobis 
sublinearibus, elongatis, subdivaricatis, separatis, 0.5-2.0 mm. latis, 
superne planus, nitidus, emaculatus, isidiis sorediisque destitutus, 
subtus niger, rhizinosus, rhizinislongis, nigris, sparse ramosis. Apothe- 
cia ignota. 
Thallus adnate on bark, pale gull grey (R), 3-5 cm. in diameter; 
lobes dichotomously branched, linear-elongate, subdivaricate, sepa- 
rate, 0.5-2.0 mm. wide, 100-130 m thick, the margins smooth, narrowly 
black-rimmed; upper surface plane and smooth, shiny, without 
maculae, isidia and soredia lacking; medulla white; undersurface 
black, rhizinate, the rhizines black, long and sparsely branched, 
about 1 mm. long. Apothecia not seen. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ red, P—, 
atranorine and alectoronic acid present. 
HALE & KUKOKAWA—SUBGENUS PARMELIA 181 
Type in Michigan State University, collected in pine forest at 
summit of Alto de la Bandera, Cordillera Central, La Vega, Dominican 
Republic, elev. 2800 m., Aug. 5, 1958, by C. Wetmore (no. 3475; 
isotype in US). 
Parmelia lineariloba has sublinear lobes that are quite narrow, long, 
and usually subdivaricate. The rhizines are rather long and only 
moderately branched. The chemically related subdivaricate species 
P. gigas Kurokawa and P. cervicornis Tuck, are much larger in size. 
Parmelia lineariloba is at present known only from the Dominican 
Republic, where it is found at high elevations in pine forests. We 
might expect it to be common in similar habitats elsewhere in the 
West Indies. 
Additional specimens examined: Dominican Republic: Ridge con- 
necting La Rucilla and Pico Trujillo, Macisco Central, elev. 8000 ft., 
Wetmore 3599, 3628 (MSC, US). 
21* Parmelia lividescens Kurokawa, sp. no v. 
Thallus adnatus, corticola, cinereo-albicans, 5-10 cm. diametro, 
lobis sublinearibus, 1.5-3.0 mm. latis, margine subcrenatis, superne 
nitidulus, sorediatus, soraliis subterminalibus, subtus sparse rhizi- 
nosus, niger, rhizinis nigris, dichotome ramosis. Apothecia adnata, 
2-3 mm. diametro; hymenium 40-50 p altum; sporae 5-7X9-11 p. 
Thallus adnate on bark, yellowish glaucous (R), 5-10 cm. across; 
lobes dichotomously branched, sublinear, separate at the circumfer- 
ence, 1,5-3.0 mm. wide, 140-180 ju thick, the margins subcrenate; 
upper surface shiny on younger lobes, dull and irregularly cracked on 
older lobes, sorediate, soralia orbicular, laminal and sub terminal; 
medulla white; undersurface sparsely rhizinate, black, the rhizines 
black, dichotomously branched. Apothecia adnate, 2-3 mm. in 
diameter, disc saccardo's brown (R), radially split; hymenium 40-50 /x 
high; spores 5-7X9-11 p. 
Reactions: Thallus K+ yellow; medulla K—, C+ red, KC+ red, 
P—, atranorine and olivetoric acid present. 
Type in the Botanical Museum, Lund University, collected on bark, 
Polela Forest, Polela, Natal, Union of South Africa, Nov. 10, 1953, by 
Ove Almborn (no. 9490; isotype in US). 
This species resembles P. revolvia Flk. in the presence of sub terminal 
capitate soralia and in the absence of maculae. Parmelia revolutaf 
however, produces gyrophoric acid and has more sparsely branched 
rhizines. Parmelia leiophylla Kurokawa has the same chemical con- 
tent as P. lividescens but differs in having lamina! sorediate pustules. 
Additional specimen examined: Union of South Africa: 6 mi. 
north of Knysna, Knysna, Cape Province, Almborn 2665 (LD). 
182 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
22. Parmelia monilifera Kurokawa, sp. nov. Platb 9 
Thallus adnatus, corticola, cinereo-albicans, ca. 12 cm. diametro, 
lobis sublineari-elongatis, subdivaricatis, 2-6 mm, latis, superne 
planus, nitidus, emaculatus, isidiis sorediisque destitutus, subtus 
niger, dense rhizinosus, rhizinis nigris, ramosis, saepe moniliformibus. 
Apothecia ignota. 
Thallus loosely adnate on bark, grayish white, about 12 cm. in 
diameter; lobes dichotomously branched, sublinear-elongate, sub- 
divaricate, 2-6 mm. wide, 200-250 n thick, the margins smooth; 
upper surface smooth, plane to concave, shiny, without maculae, 
isidia and soredia lacking; medulla white; undersurface black, densely 
rhizinate, the rhizines black, shiny, branched, often moniliform or 
opuntioid, projecting as a mat beyond the margins. Apothecia not 
seen. 
Reactions: Thallus K+ yellow; medulla K—, C— or C+ pale 
orange, KC+ deep orange red, P—, atranorine, barbatic acid, and 
an unknown KC+ substance present. 
Type in the Botanische Staatssammlung, Munich, collected at 
La Aguada, Merida, Venezuela, Apr. 2, 1958, by K. Magdefrau 
(no. 651; isotype in US). 
This species resembles P. gigas Kurokawa, P. cervicornis Tuck., 
and P. exporrecta Kurokawa in having a large divaricate thallus and 
sublinear-elongate lobes. It is clearly separated by the coarse, 
branched, moniliform or opuntioid rhizines, a type that we have 
not seen in any other species of Parmelia. It is especially distinct 
chemically, since related species have alectoronic acid, or less com- 
monly gyrophoric acid. The species is known only from the type 
locality in Venezuela. 
23. Parmelia prolongata Kurokawa, ep* nov. Plate 6 
Thallus laxe adnatus, corticola, olivaceo-albicans, 5-10 cm. dia- 
metro, lobis sublinearibus, 1.5-4.0 mm. latis, margine partim isidiato- 
dissectis, superne planus, nitidus, exigue albomaculatus, sparse 
isidiatus, medulla alba vel pallide flavescens, subtus niger, dense 
rhizinosus, rhizinis nigris, dense ramosis. Apothecia ignota. 
Thallus loosely adnate on bark, sea-foam yellow (R) turning olive 
buff in the herbarium, 5-10 cm. across; lobes dichotomously branched, 
sublinear, elongate, sometimes subimbricate, 1.5-4.0 mm. wide, 100- 
200 fi thick, the margins smooth or in part with isidial branchlets 
or isidia, often elongate, ascending or suberect, dorsiventral or 
becoming cylindrical and coralloid branched; upper surface plane 
and smooth, weakly maculate, rather shiny, often sparsely isidiate, 
the isidia cylindrical, variable, sometimes branched; medulla white 
to cartridge buff (R); undersurface black, densely rhizinate, the 
HALE & KUROKAWA—SUBGENUS FARMELIA 183 
rhizines black, shiny, densely branched, often forming a mat. Apo- 
thecia not seen. 
Reactions: Thallus K+ yellow; medulla K—, C+ deep red, KC-(- 
red, P—, atranorine, a C+ unknown substance, and frequently a 
pale-yellowish pigment present. 
Type in Michigan State University, collected on ridge leading to 
Pic Macaya, Morne Macaya, Dept. du Sud, Haiti, elev. 6400-7000 ft., 
July 27, 1958, by C. Wetmore (no. 3233; isotype in US). 
Parmelia prolongata is characterized by peculiar laminal and mar- 
ginal isidia which are cylindrical at first but which at length become 
dorsiventral, suberect, and more or less branched, and as much as 
5 mm. long. However, the production of isidia and lobules is ex- 
tremely variable and they may be reduced to small marginal isidia. 
All specimens have in common a deep C+ reaction caused by an 
unknown substance apparently related to olivetoric or anziaic acid. 
The pale-yellow pigment may or may not be clearly present and its 
identity is unknown. Parmelia prolongata is found at higher eleva- 
tions, often on conifer bark, in the Caribbean and Andean regions. 
Additional specimens examined: Mexico: Chiapas: 18 km. south- 
east of San Crist6bal, Hale 20286 (US). Panama: Prov. ChiriquI: 
East side of Volc&n Chiriquf, Sckolander, Dec. 5-12, 1948 (US), 
Haiti: Ridge leading to Pic Macaya, Mome Macaya, Dept. du Sud, 
elev. 6400-7000 ft., Wetmore 3233 (MSC, US), 3286 (MSC). Colom- 
bia: Western slopes of P&ramo de Cruz Verde, Dept. Cundinamarca, 
3150 m., Cuairecasas 450Z (US). 
24. Parmelia rhabdiformiB Kuiokawa, ep. nov. 
Thallus adnatus, corticola, olivaceo-albicans in herbario, ca. 8 cm. 
diametro, lobis sublinearibus, 2-6 mm. latis, subimbricatis, margine 
plus minusve dissectis, superne planus, nitidus, emaculatus, isidiatus, 
subtus niger, dense rhizinosus, rhizinis nigris, dense ramosis. Apo- 
thecia adnata, 1-5 mm. diametro; hymenium 60-80 m altum; sporae 
6-8X15-21 n. 
Thallus adnate on bark, turning cream buff (R) in the herbarium, 
about 8 cm. in diameter; lobes irregularly branched, sublinear, 2-6 
mm. wide, subimbricate, 150-200 n thick, the margins more or less 
dissected; upper surface plane, shiny, without maculae, isidiate, the 
isidia mostly simple, club-shaped, about 0.5 mm. high; medulla white; 
undersurface black, densely rhizinate, the rhizines black, densely and 
finely branched, forming a woolly mat. Apothecia adnate, 1-5 mm. 
in diameter, exciple crenate, amphithecium isidiate, disc sayal brown 
(R), sometimes radially split; hymenium 60-80 p high; spores 6-8X15- 
21 JLt. 
722-891—C4 5 
184 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Reactions: Thallus K+ yellow; medulla K+ yellow turning red, 
C—, KC—, P+ pale orange red, atranorine and norstictic acid 
present. 
Type in the U.S. National Herbarium, collected in the vicinity of 
El Boquete, Pro v. Chiriquf, Panama, Mar. 2, 1911, by W, R. Maxon 
(no. 5128a). 
Except for the production of norstictic acid, this species resembles 
P. infirma Kurokawa and P. crenata Kurokawa, both Asian species. 
The isidia of P. rhabdiformis are mostly simple and clavate and rather 
elongate, to 0.5 mm. high. It is known only from the type locality in 
Panama. 
25. Parmclia rigidula Kurokawa, sp. nov. 
Thallus adnatus, corticola, olivaceo-cinerascens, 7-12 cm. diametro, 
lobis sublinearibus vel subirregularibus, subimbricatis, usque ad 7 
mm. latis, margine subcrenatis, superne nitidulus, exigue albomacu- 
latus vel emaculatus, sorediis isidiisque destitutus, medulla pallide 
flavicans, subtus niger, rhizinosus, rhizinis nigris, modice ramosis. 
Apothecia adnata, 2-7 mm. diametro; hymenium 70-85 n altum; 
sporae 10-12X15-18 n. 
Thallus adnate on bark, pale olive buff (R), 7-12 cm. in diameter; 
lobes irregularly branched, sublinear to subirregular, more or less 
imbricate, up to 7 mm. wide, 210-250 p thick, the margins crenate, 
often narrowly black-rimmed; upper surface plane to concave, more 
or less shiny, weakly maculate or without maculae; medulla sea-foam 
yellow to pale-pinkish buff (R), K—; under surface black, moderately 
rhizinate, the rhizines black, matted, sparsely to moderately branched. 
Apothecia adnate, 2-7 mm. diameter, exciple smooth to undulate, 
disc dark livid-brown (R), sometimes radially split; hymenium 70-85 
high; spores 10-12X15-18 p. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC—, P—, 
atranorine and a pale-yellow pigment present. 
Type in the National Science Museum, Tokyo, collected at Phalut, 
Darjeeling, India, June 6, 1960, by H. Hara (isotype in US). 
This species has a peculiar coriaceous thallus as in P. scytoOss 
Kurokawa and P. scytopkylla Kurokawa. However, it is distinguished 
by producing a pale-yellow-orange pigment in the medulla (apparently 
not entotliein). Superficially it resembles P. subaurulenta Nyl., but 
the rhizines are sparsely to moderately branched, not simple. Only 
two specimens have been collected, both in northern India, where it 
is endemic. 
Additional specimen examined: India: Tongloo. Himalayas, Watt 
7035 (BM). 
HALE & KUROKAWA—SUBGENUS PARMELIA 185 
26. Parmelia scytodes Kurokawa, sp. no v. Plate G 
Thallus adnatus vel laxe adnatus, corticola, cinereo-albicans, ca. 
20 cm. diametro, lobis sublinearibus, 2-5 mm. latis, superne planust 
emaculatus, isidiis sorediisque destitutus, subtus niger, dense rhizino- 
sus, rhizinis nigris, modice ramosis. Apothecia ignota. 
Thallus adnate to loosely adnate on bark, rather coriaceous, pale 
smoke gray (R), about 20 cm. across; lobes irregularly branched, 
sublinear, 2-5 mm. wide, 150-200 n thick, the margins subcrenatc, 
narrowly black-rimmed; upper surface plane and smooth, becoming 
rugose on older lobes, without maculae, isidia and soredia lacking; 
medulla white; undersurface black, moderately to densely rhizinate, 
the rhizines black, shiny, moderately branched. Apothecia not seen. 
Reactions: Thallus K4- yellow; medulla K—, C—, KC-f- orange, 
P+ orange red, atranorine, barbatic acid, and an unidentified P+ 
acid present. 
Type in the National Science Museum, Tokyo, collected at Dar- 
jeeling, Batasi, India, May 6, 1960, by Togashi (isotype in US). 
This new species is characterized by the large rather coriaceous 
thallus. Externally it resembles P. fissicarpa Kurokawa, an African 
species with a P+ reaction but no barbatic acid. Parmelia bostry- 
ckodes Zahlbr. and P. physcioides Nyl., both of which produce barbatic 
acid without a P+ substance, have smaller sublinear lobes. Parmelia 
scytodes is known only from the type locality in India. 
27. Parmelia scytophylla Kurokawa, sp. no v. Plate 7 
Thallus adnatus, corticola, cinereo-albicans, 5-14 cm. diametro, 
lobis sublinearibus vel subirregularibus, margine subcrenatis, superne 
pi anus, nitidus, exigue albomaculatus, isidiis sorediisque destitutus, 
subtus niger, modice rhizinosus, rhizinis nigris, sparse ramosis. Apo- 
thecia adnata, ca. 10 mm. diametro; hymenium 35^40 ft altum; 
sporae 5-6 X 8-9 p. 
Thallus adnate to loosely adnate on bark, yellowish glaucous to 
pale olive gray (R), 5-14 cm. in diameter; lobes irregularly branched, 
sublinear to subirregular, 2-6 mm. wide, 250-350 ji thick, the margins 
more or less crenate, narrowly black-rimmed; upper surface plane 
and smooth, shiny, faintly maculate, isidia and soredia lacking; 
medulla white; undersurface black, moderately rhizinate, blackish 
brown and naked or papillate in a narrow zone around the tips, the 
rhizines black, shiny, sparsely branched. Apothecia adnate to sub- 
stipitate, often more than 10 cm, in diameter, exciple undulate, 
amphithecium smooth, faintly maculate, disc warm blackish brown 
to light seal brown (R); hymenium 35-40 p. high; spores 5-6X8-9 /*. 
Reactions: Thallus K+ yellow; medulla K—, C-f- rose, KC+ 
red, P—, atranorine and gyrophoric acid present. 
186 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Type in the National Science Museum, Tokyo, collected at 
Migotang, Sikkim, at 3900 m. elevation, May 30, 1960, by H. Hara 
(isotype in US). 
Parmelia scytophylla is very similar to two other species known 
only in the Himalayan region, P. rigiduh, Kurokawa and P. seytodes 
Kurokawa. All have rather large coriaceous thalli. Parmelia 
seytodes contains barbatic acid and a P+ unknown, and P. rigidula 
contains a pale-yellow pigment and has larger spores, 15-18 n long. 
Additional specimens examined: Nepal: Langtang Village, Polunin 
M32 (BM). India: Nilgherries, Watt s.n. (BM); Darjeeling, Awasthi 
12 (H). 
28. Parmelia thysanota Kurokawa, sp. no v. 
Thallus laxe adnatus, corticola, cinereo-albicans, 5-13 cm. latus, 
lobis sublinearibus, subdivaricatis, separatis, 2-5 mm. 1 at is, superne 
planus, nitidulus, apice leviter albopruinosus, sorediatus, soraliis 
subterminalibus, subtus niger, dense rhizinosus, rhizinis nigris, 
dense ramosis, Apothecia ignota. 
Thallus loosely adnate on bark, pale-glaucous green (R), 5-13 cm. 
in diameter; lobes dichotomously branched, linear or sublinear- 
elongate, subdivaricate, separate at the circumference, 2-5 mm. wide, 
140-180 n thick, the margins entire, narrowly black-rimmed; upper 
surface plane, rather shiny, often pruinose near the tips, sorediate, 
soralia laminal and sub terminal, often protruding, coalescing; medulla 
white; undersurface densely rhizinate, black, the rhizines black, 
shiny, densely branched, usually projecting in a narrow dense mat 
beyond the margins. Apothecia not seen. 
Reactions: Thallus K-f yellow; medulla K—, C-f rose, KC-f- 
red, P—, atranorine and gyrophoric acid present. 
Type in the United States National Herbarium, collected in pine-oak 
forest on lava, 11 km. east of Las Vigas on Highway 140, Veracruz, 
Mexico, Mar. 11, 1960, by M. E. Hale (no. 20943). 
Parmelia tkysanota is very closely related to P. exporrecta Kuro- 
kawa in having a subdivaricate thallus and in producing gyrophoric 
acid. It differs in having sub terminal soralia; P. exporrecta lacks 
any soralia. It is probable that these two species represent a sore- 
diate-nonsorediate phase, but the material at hand is not sufficient 
for a final decision. 
Additional specimen examined: Mexico: Same locality data as 
the holotype, Hale 21118 (US). 
29. Parmelia virginica Hale, sp. nov. Plate 5 
Thallus adnatus, albido-cinerascens, nonnihil fragilis, corticola, 
5-7 cm. diametro, lobis sublinearibus, 1.5-2.5 mm. latis, margine 
centrum versus paulo lobulato-dissectis, superne nitidus, minute 
HALE & KUROKAWA—SUBGENUS PARMELIA 187 
rugulosus, actate valde irregulariter rugosus, cortice fragili, 
pustulescente, sorediis destitutus, subtus niger, modice rhizinosus, 
rhizinis modice dichotome ramosis. Apothecia ignota. 
Thallus adnato, mineral gray, rather fragile, corticolous; lobes sub- 
linear, 1.5-2.5 mm. wide, 110-130 ju thick, more or less subimbricate, 
towards the center lobulate-dissected; upper surface shiny, minutely 
rugulose, strongly rugose with age, the cortex fragile and breaking 
away, forming coarse pustules, soredia lacking; medulla white; under- 
surface black, brown near the tips, moderately rhizinate, the rhizines 
black, moderately branched. Apothecia not seen. 
Reactions: Thallus K+ yellow; medulla K—, C—, KC+ orange, 
P—, atranorine, barbatic acid, and an unknown KC+ substance 
present. 
Type in the U.S. National Herbarium, collected at Hawksbill, 
Shennandoah National Park, Page Co., Virginia, elev. 1200 m.( 
June 26, 1962, by M. Hale (no. 21592; isotypes in DUKE, LD, M, 
MSC, REN, S, TNS, WISC). 
Parmelia virginica is virtually identical with P. taylorensis Mitch. 
(P.rugosa Tayl. non Fr.) in regard to the fragile cortex and the develop- 
ment of pustules. The pustules originate along the ridges of wrinkles 
and are therefore usually elongate. Parmelia taylorensis (cf. Mitchell, 
1962; Hale, 1962) is typically a European species which differs in 
having evernic and lecanoric acids and more richly branched rhizines. 
The type collection of P. virginica was made on exposed dwarf spruce 
trunks. It is probably common in similar habitats in the Appa- 
lachians from Virginia through North Carolina. 
Additional specimen examined: North Carolina: Lich-Log ridge, 
Blue Ridge Parkway, McDowell Co., elev. 4660 ft., Becking 5908031 
(US). 
Bibliography 
Acharius, E. 1794. Forsok til en ftirbattrad I&fvarnes indelmng. K. Vet. Acad 
Nya Handl. 237-259. 
Asahina, Y. 1952. Lichens of Japan II. Genus Parmelia. Res. Inst. Nat. 
Resources. 162 pp. Tokyo. 
 and S. Shibata, 1954. Chemistry of lichen substances. Japan Soc. for 
Promotion of Science. 240 pp. Tokyo. 
Culberson, W. L. 1962. Some pseudocyphellate Par me lias. Nova Hedw. 
4:564-577. 
Du Rietz, G. E. 1924a. Die sore die n und isidien der Flechten. Svensk Bot. 
Tidskr. 18: 371-396. 
 . 1924b. Flechtensystematische Studien. IV. Bot. Not. 1924:329-342. 
Fries, E. 1825. Systcma orbis vegetables. Lund. 
188 CONTRIBUTIONS FROM THE NATIONAL HERBARIUM 
Gyelnik, V. 1932. Addi (amenta ad cognitionem parraeliarum III. Re pert. 
Sp. Nov. Fedde 30:209-226. 
 . 1933. Lichenes varii novi criticique. Acta Faun, Fl. Univ. II, 
1, no. 5,6:3-10. 
Hale, M. E., Jb, 1960. A revision of the South American species of Parmelia 
determined by Lynge. Contr. U.S. Nat. Herb. 36, pt. 1:1-41. 
  and S. Kurqkawa. 1962, Parmelia species first described from the 
British Isles. Lichenologiat 2:1-5. 
Hillmann, J, In Rabenhorst, Kryptogamenflora. 1934. Familie Parmeliaceae. 
9(5), part 3:1-309. 
Imshaug, H. A. 1957. Alpine lichens of the western United States and adjacent 
Canada I. The macrolichenB. Bryologist 60:177-272. 
Lynge, B. 1914. Die Flechten der ersten Regnellschen Expedition. Die 
Gattungen Pseudoparmelia gen. no v. und Parmelia Ach. Ark. Bot. 13, no. 
13:1-172. 
Maas Geesteranus, R. A. 1947. Revision of the lichens of the Netherlands. 
I. Parmeliaceae. Blumea 6:1-199. 
Mitchell, M. E. 1961. L'element eu-oceanique dans la flore lichcnique du 
sud-ouest de I'Irlande. Re vista Biol. 2, no. 3-4:177-256. 
Rasanen, V. 1943. Das System der Flechten. Act. Bot. Fenn. 33:1-82. 
Santesson, R. 1944. Pseudoparmelia Lynge, a lichen genus to be rejected. 
Svensk Bot. Tidskr. 36:471-474. 
Schreber, J. 1791. Linnd, Genera plantarum, ed. 8. 
Vainio, E. A. 1890. Etude eur la classification naturelle et la morphologie des 
lichens du Br6sil. Acta Soc. Faun. Fl. Fenn. 7, no. 7:1-240. 
— . 1923. Lichenes Insularum Philippinarum, IV. Ann. Acad. Sci. Fenn. 
Ser. A, 19, no. 15:1-84. 
Index 
(To species and varieties of subgenus Parmelia. Synonyms in 
italics) page numbers of principal entries in boldface.) 
abstrusa, 141, 145 
abstrusoides, 149 
acrobotrys, 141, 142, 143, 145, 147 
aflixa, 137 
amazonica, 129, 147, 148, 149, 157 
americana, 159 
anaptychioides, 162, 165 
annexa, 149, 151* 152, 155, 156 
antillensis, 131 
apophysata, 136, 138 
appalachensis, 129 
aptata, 150 
arcana, 149, 151, 155 
atrichella, 137, 
aurulenta, 124, 130, 131, 132 
bahiana, 163, 164 
balansae, 130 
bicornuta, 135, 137 
bogotensis, 161 
boliviana, 164 
bolliana, 129 
borreri, 124, 128, 129 
borrerina, 129 
bostrychodes, 164, 185 
brasiliana, 127, 129, 154, 159, 164, 174 
brasiliana var. erythrodes, 17 S 
bre virhiza, 162, 166 
bulbochaeta, 137, 138 
canaliculata, 129 
canescens, 163, 167 
caperata, 148, 149 
caraccensis, 163, 177 
caribaea, 150, 152, 153 
carneopruinata, 149 
caroliniana, 148, 149 
carporrhizans, 125, 132 
cerviconais, 165, 171, 177, 181, 182 
cetrarioides, 128, 148 
cetrata, 124, 129, 130 
chapadensis, 150 
cinerascens, 149 
circumnodata, 123, 141, 145 
citrella, 163, 168 
concrescens, 149 
confoederata, 137 
connivens, 141, 142 
consimilia, 161 
continua, 136 
coronata, 137, 139 
coronata var. denudata, 187 
costaricensis, 160, 161, 172, 179 
crenata, 161, 168, 172, 179, 184 
croceopuatulata, 162, 169, 180 
crozalsiana, 149 
cryptochlorophaea, 149 
culmigena, 123, 161, 173 
cyphellata, 147, 148, 149 
daetylifera, 123, 161, 178 
damaziana, 165 
decurtata, 136, 139 
degelii, 165, 170 
denegans, 132 
den&irhiziflata, 163, 170, 171 
dentella, 161, 172 
denudata, 1S7 
digitata, 123, 161, 173 
disseeta, 123, 131, 132, 133, 134, 135, 
148 
domingengjg, 141 
dubia, 129 
ecaperata, 149 
enderythraea, 163 
endochlora, 124, 161, 162 
endomiltoidee, 159 
enaifolia, 123, 161, 171, 172, 173 
entotheiochroa, 133 
eruptens, 148, 153 
erythrodes, 163, 173 
eximbricata, 142, 143 
exporrecta, 164, 174, 182, 186 
exsecta, 162 
exsplendene, 135, 162, 163, 170, 174 
189 
i 
190 
fissicarpa, 164, 175, 185 
flavida, 163 
flavovirens, 162, 176 
formosana, 124, 162, 169, ISO 
f randans, 128 
fungicola, 136 
galbina, 125, 132 
gigas, 165, 171, 174, 177, 181, 182 
gracilescens, 165 
heterochroa, 150,154 
homogenes, 133 
homotoma, 124, 130 
horrescens, 123, 131 
hypocraea, 136 
hypomilta, 150 
hypoxantha, 154 
ikomae, 131 
imbricatula, 161 
immaculata, 163, 178, 180 
iramiscens, 133 
imperfecta, 148, 155, 156 
infirma, 161, 179, 184 
insensitiva, 128 
insinuans, 164, 176 
intercalanda, 165 
interne xa, 131 
intertexta, 150 
ischnoides, 149, 155 
isidiocera, 131, 160 
isidiosa, 147 
isidiza, 136, 141 
isidiza var, domingensis, 140 
keitauensis, 164, 176 
koyaensis, 161, 169, 179 
laevigata, 124, 162 
laevigatula, 136 
lecanoracea, 150 
leiophylla, 162, 169, 178, 179, 181 
le ucopis, 124, 150 
limbata, 141, 142, 147 
limbata (. isidiosa, 147 
tindmannii, 131 
lineariloba, 165, 171, 180 
livida, 154, 165, 170, 174, 180 
lividescens, 162, 181 
luteoviridis, 141, 144 
lythgoeana, 163 
macrocarpoides, 130 
majoris, 162, 166 
malaceensis, 149 
malmei, 162 
maniJensis, 164, 176 
martinicana, 149 
meioBperma, 149 
meizospora, 137 
melanochaeta, 132, 133 
metarevoluta, 125, 132, 175 
meyeri, 161 
michauxiana, 132 
microblasta, 161 
mierosticta, 129 
minarum, 133, 148 
molybdiza, 150, 151, 152, 155, 158 
monilifera, 164, 182 
muelleri, 132, 148 
mutata, 124, 132 
nairobiensis, 153 
nodakcnsis, 161, 179 
nylanderi, 132 
obsessa, 125, 131 
omphalodes, 128 
osseoalbida, 164 
osteoleuca, 163 
owariensis, 149, 155, 156 
papyrina, 136 
perforata, 130 
perisidians, 131 
peruviana, 124, 163 
phlyctina, 132 
physcioidcs, 164, 185 
pilosa, 130 
planiuacula, 141, 144,145 
pluriformis, 164 
prolongata, 160, 161, 182 
pruinata, 132 
pseudoborreri, 129 
pseudosinuosa, 162, 166 
pseudosulcata, 128 
.pulvinata, 165 
pustulata, 137, 140 
pustulescens, 149, 156 
quercina, 131, 132 
rahengensis, 149 
ramosissima, 141, 143, 145, 147 
raunkiaeri, 148 
reducens, 163, 168 
rellcina, 135, 141 
relicinella, 142 
relicinula, 141 
reticulata, 124, 129, 130 
revoluta, 162, 163, 181 
revolutella, 165 
rhabdiformis, 161, 183 
rigidula, 163, 184, 186 
rockii, 162, 163 
rodriguesiana, 150 
INDEX 
rudectu, 120 
rugosa, 187 
rupicolu, 150 
rutidota, 150 
salacinifora, 149, 157 
samocnsis, 142, 143 
samoensis var. cximbrieata, J^/i 
saxatilis, 124, 127, 128, 100 
schiffneri, 135, 137 
schizospatha, 141, 144, 146 
scortolla, 136 
scrobicularis, 150 
seytodes, 164, 184, 185, 180 
scytophylla, 164, 184, 185 
semilunata, 135, 137 
sensibilis, 137 
setsehwanensiis, 136 
signata, 128 
silvatica, 131, 163 
sinuosa, 162, 176 
somalicnsis, 150 
soredians, 148, 149 
sorocheila, 159 
spathulata, 131, 133, 134 
sphaerospora, 148, 150 
spumosa, 124, 132, 135 
subabstrusa, 141 
subafTtuis, 162, 166 
subaurulenta, 133, 184 
subcoronata, 136 
KubdisKocta, 130, 138 
subfatiscens, 132, 134, 135 
subglandulifera, 130 
subisidiosa, 130 
sublaevigata, 164, 107 
sublanea, 141, 142. 146, 147 
sublimbata, 141, 146 
Jrubrudecta, 129 
subsaxatiii:?, 164 
Kiibtiliacca, 150 
ssufiixa, 135, 136 
sulcata, 128 
labacina, ] 36, 139, 140 
taylorensis?, 165, 187 
tcnuirinia, 128 
ti'xaua, 148, 150, 153 
thysanota, 163, 174, 186 
tiliacL-a, 123, 124, 130, 131, 1 
t iliatica var. con vexula, 154 
liliaera var. rugutata, 154 
liliaeea var. stciiophylla, Jo.j 
tortula, 150, 157 
tuiML'Kccns, 141, 145, 147 
usambari'iisis, 131 
volloaiao, 162, 176 
vontrieosa, 135, 136, 140, 141 
vrrsiformis, 124, 132 
violawa, 150, 158 
virginiea, 162, 186 
viridescens, 137, 139 
wallicliiana, 131 
xantholepis, 132 
xant homolaona, 150 
zalilbrucknori, 164 
"22-801—(54 fi 
CONTR, NAT, HERB VOL. 36 HALE 8c KUROKAWA PLATE 1 
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kaw.j (huf. ■:ypi*, HaU 20722. I S, I). 4. Parmclnt prohtuvata Kui'nkaua (lii.i|titypc\ li tl- 
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CONTR. NAT. HERB. VOL, 36 HALE 8c KUROKAWA PLATES 
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%igds Kiintkawa (liol[)typof Jristr K70, I S, 1). 
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