A Monograph of the Lichen Genus Parmelina Hale - Smithsonian ...
A Monograph of the Lichen Genus Parmelina Hale - Smithsonian ...
A Monograph of the Lichen Genus Parmelina Hale - Smithsonian ...
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SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 33<br />
A <strong>Monograph</strong> <strong>of</strong> <strong>the</strong> <strong>Lichen</strong> <strong>Genus</strong><br />
<strong>Parmelina</strong> <strong>Hale</strong> (Parmeliaceae)<br />
Mason E. <strong>Hale</strong>, Jr.<br />
SMITHSONIAN INSTITUTION PRESS<br />
City <strong>of</strong> Washington<br />
1976
ABSTRACT<br />
<strong>Hale</strong>, Mason E., Jr. A <strong>Monograph</strong> <strong>of</strong> <strong>the</strong> <strong>Lichen</strong> <strong>Genus</strong> <strong>Parmelina</strong> <strong>Hale</strong> (Par-<br />
meliaceae). <strong>Smithsonian</strong> Contyibutions to Botany, number 33, 60 pages, 21<br />
figures, 1976.-The 47 species <strong>of</strong> <strong>Parmelina</strong> are revised on <strong>the</strong> world level. Two<br />
sections are recognized: section Parmelzna with 30 species widely distributed in<br />
temperate to tropical montane regions and section Myelochroa with 17 terpene-<br />
containing species concentrated in eastern and sou<strong>the</strong>rn Asia. The genus is<br />
most closely related to Parmotrema hlassalongo. Five new species, P. crassata<br />
<strong>Hale</strong>, P. degelii <strong>Hale</strong> P. indica <strong>Hale</strong>, P. rhytidodes <strong>Hale</strong>, and P. schindleri <strong>Hale</strong>,<br />
are described, and six new combinations proposed, P. amagiensis (Asahina)<br />
<strong>Hale</strong>, P. damaziana (Zahlbruckner) <strong>Hale</strong>, P. endoleuca (Taylor) <strong>Hale</strong>, P. irrugans<br />
(Nylander) <strong>Hale</strong>, P. jamesii (<strong>Hale</strong>) <strong>Hale</strong>, and P. pastillifera (Harmand) <strong>Hale</strong>.<br />
New combinations are also made for Hypotrachyna baguioensis (<strong>Hale</strong>) <strong>Hale</strong> and<br />
Parmotrema nylanderi (Lynge) <strong>Hale</strong>.<br />
OFFICIAL PUBLICATION DATE is handstamped in a limited number <strong>of</strong> initial copies and is recorded<br />
in <strong>the</strong> Institution’s annual report, Srnithsonian Year. SERIES COVER DESIGN: Leaf clearing from <strong>the</strong><br />
katsura tree Cercidiphyllum juponicum Siebold and Zuccarini.<br />
Library <strong>of</strong> Congress Cataloging in Publication Data<br />
<strong>Hale</strong>, Mason E.<br />
A monograph <strong>of</strong> <strong>the</strong> lichen genus <strong>Parmelina</strong> <strong>Hale</strong> (Parmeliaceae)<br />
(<strong>Smithsonian</strong> contributions to botany ; no. 33)<br />
Bibliography: p.<br />
Includes index.<br />
Supt. <strong>of</strong> Docs. no.: SI 1.29:33<br />
1. <strong>Parmelina</strong>. I. Title. 11. Series: <strong>Smithsonian</strong> Institution. <strong>Smithsonian</strong> contributions to<br />
botany : no. 33.<br />
QKl.SZ747 no. 33 [QK585.P2] 581’.08s [589’.1] 76-608065
Contents<br />
Page<br />
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1<br />
Morphology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2<br />
Chemistry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8<br />
Phytogeography and Speciation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10<br />
Classification <strong>of</strong> <strong>Parmelina</strong> . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14<br />
Key to <strong>the</strong> Species <strong>of</strong> <strong>Parmelina</strong> . . . . . . . . . . . . . . . . . . . . . . . . . 16<br />
Species Treatment . . , . . . . . . . . . . . . . . 18<br />
Doubtful and Rejected Names . . . . . . . . . . . . . . . . . . . . . . ... 53<br />
Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55<br />
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59<br />
iii
A <strong>Monograph</strong> <strong>of</strong> <strong>the</strong> <strong>Lichen</strong> <strong>Genus</strong><br />
<strong>Parmelina</strong> <strong>Hale</strong> (Parmeliaceae)<br />
Introduction<br />
The genus <strong>Parmelina</strong> <strong>Hale</strong> is a segregate <strong>of</strong> Par-<br />
melia Acharius comprising 47 species widely dis-<br />
tributed in temperate and tropical regions. It is<br />
recogni7ed by <strong>the</strong> narrow, adnate lobes, marginal<br />
cilia, and absence <strong>of</strong> usnic acid in <strong>the</strong> cortex (<strong>Hale</strong>,<br />
1974:438). Some <strong>of</strong> <strong>the</strong> species included here were<br />
formerl) classified in <strong>the</strong> ill-defined (and invalid)<br />
genus Zmbitca? ia (Schreber) Michaux (<strong>Hale</strong> and<br />
Kurokawa, 1964: 130). This world level monograph<br />
is based on <strong>the</strong> study <strong>of</strong> all available type speci-<br />
mens, collections preserved in <strong>the</strong> major herbaria,<br />
and personal field work. The format follows my<br />
earlier treatments <strong>of</strong> Bulbothrtx (<strong>Hale</strong>, 1976b),<br />
Hypotrachyna (<strong>Hale</strong>, 1975a), Pseudopawnella<br />
(<strong>Hale</strong>, 1976a), and Rellcina (<strong>Hale</strong>, 197513).<br />
I wish to thank curators <strong>of</strong> <strong>the</strong> following insti-<br />
tutions who sent specimens on loan so promptl) and<br />
<strong>of</strong>ten allowed extensions <strong>of</strong> loan periods: BM, BO,<br />
BP, COLO, DUKE, F, FH, FI, G, GLAM, H, KAN,<br />
KR, L, LD, Rl, LlICH, NSC, RlVAf, NSLV, NY,<br />
PC, PH, REX;, S, SI, TNS, TRH, TUR, UC, UPS,<br />
W, LVIS, WU, and ZT. Specimens in TNS were<br />
annotated during 1964-65 while I studied in Tokyo.<br />
The priyate herbaria <strong>of</strong> Dr. D. D. Awasthi, Dr.<br />
Gunnar Degelius, Dr S Nakanishi, Dr. Clyde Reed,<br />
Mason E <strong>Hale</strong>, Jr, Department <strong>of</strong> Botan), National Mu-<br />
seum <strong>of</strong> Aatiital Histor), <strong>Smithsonian</strong> Institution, Washang-<br />
ton, D. C. 20160.<br />
Mason E. <strong>Hale</strong>, Jr.<br />
I<br />
and Dr. R. Santesson were also geneiously placed<br />
at my disposal. The material in <strong>the</strong>se various her-<br />
baria was chemically tested and annotated between<br />
1958 and 1975, but not all <strong>of</strong> it has been re-<br />
examined for possible name changes in <strong>the</strong> light <strong>of</strong><br />
new synonymies, species concepts, or improved<br />
chemical techniques.<br />
Any monograph is bound to be more complete if<br />
<strong>the</strong> author has had an opportunity to observe and<br />
collect specimens in <strong>the</strong> field. I am especially grate-<br />
ful to <strong>the</strong> following colleagues for assistance in<br />
arranging excursions: blr, J. Anderson (Sarawak),<br />
Alrs. Sheila Collenette (Sabah), Dr. S. Kurokawa<br />
(Japan), Dr. 51. L6pez-Figueii-as (l’enezuela), Dr.<br />
W. Meijer (Sabah), Dr. hI. Nakanishi and Dr. S.<br />
Nakanishi (Japan), Dr. P. G. Patwardhan (India),<br />
Dr. Stella Thrower (Hong Kong), and Dr. Flora<br />
Uyenco (Philippines). I have also conducted field<br />
studies in Llexico and Central America, <strong>the</strong> Lesser<br />
Antilles, and Malaya.<br />
The scanning-electron photographs were pre-<br />
pared by <strong>the</strong> <strong>Smithsonian</strong> Scanning-Electron Rficro-<br />
scope Laboratory. Photographs <strong>of</strong> <strong>the</strong> specimens<br />
were taken by <strong>the</strong> <strong>Smithsonian</strong> Photographic Ser-<br />
vices.<br />
Grants in support <strong>of</strong> this ret,earch have been re-<br />
ceived from <strong>the</strong> National Science Foundation (Sys-<br />
tematic Biology and <strong>the</strong> U. S-Japan Cooperative<br />
Science Program), hforden-Smi thsonian Expedition<br />
to Dominica, National Geographic Society, and <strong>the</strong><br />
<strong>Smithsonian</strong> Research Foundat ion.
2<br />
Morphology<br />
THALISIS STRLCTURE.-Pa?-melina basically has a<br />
closely appressed to adnate foliose thallus with narrow<br />
lobes (1-4 mm wide). Most species have more<br />
or less subirregular lobation and apically rotund<br />
lobes (Figures I&, 18b), although narrow lobed<br />
species are sublinear (Figures 12d, 16c,). There is,<br />
however, a wide range <strong>of</strong> variation in lobe configuration<br />
and width, much more than exists, for example,<br />
in Hypotmchynn (<strong>Hale</strong>, 1975a).<br />
The internal anatomy is similar to that <strong>of</strong> o<strong>the</strong>r<br />
epicorticate genera (<strong>Hale</strong>, 1973a). A thin, generally<br />
palisade plectenchymatous cortex is overlain by a<br />
pored epicortex (Figures 1 and 2). The medulla<br />
consists <strong>of</strong> loosely packed hyphae, <strong>of</strong>ten encrusted<br />
with lichen substances (Figure 2f). The lower cortex<br />
is p”p1ectenchymatous (Figure 20-d).<br />
The upper surface is strongly white maculate in<br />
Pwmelina con.son (Figure 3a), P. miielleii, and P.<br />
pilostr, less conspiciiously so in P. nzelanochaetn, P.<br />
pa.stillifei.a, P. qiiercina, and P. tiliacea, and faintly<br />
or not at all in <strong>the</strong> remaining species.<br />
The lower surface is black with only three exceptions.<br />
Pai.melina cnormis has a uniformly pale<br />
brown lower surface, and P. expallida and P. versiforrnis<br />
are darker brown tending toward black at<br />
<strong>the</strong> center.<br />
CILIA.-The marginal cilia which characterize<br />
Pnrrnelina are usually distinct but may be variably<br />
dispersed around <strong>the</strong> lobe margins. In many species<br />
<strong>the</strong>y occur more or less regularly both on lobe tips<br />
and in <strong>the</strong> axils (Figures 3a, lia, 18b). In o<strong>the</strong>rs<br />
cilia may be lacking at <strong>the</strong> lobe tips and confined<br />
to <strong>the</strong> axils (Figure 3b) or may be so sparse and<br />
inconspicuous in a few species, such as P. .rimplicioy,<br />
that <strong>the</strong>y are overlooked. In <strong>the</strong>se cases one might<br />
tend to classify <strong>the</strong>m in <strong>the</strong> genus Pseutlopaimelia<br />
(<strong>Hale</strong>, 1976a), which is differentiated from <strong>Parmelina</strong>,<br />
among o<strong>the</strong>r ways, by <strong>the</strong> total absence <strong>of</strong><br />
marginal cilia. Great care must be taken to establish<br />
<strong>the</strong> presence or absence <strong>of</strong> cilia in <strong>the</strong> lobe<br />
axils. Ano<strong>the</strong>r example <strong>of</strong> this problem is Pa?-melin(i<br />
endoleum and P.rezidopnrmelin siibtiliacea (Nylander)<br />
<strong>Hale</strong>, two very similar species in Australia that<br />
differ in <strong>the</strong> production <strong>of</strong> cilia. Both have fatty<br />
acids and cannot be separated easily by a chemical<br />
test.<br />
Confusion may also arise with species <strong>of</strong> Hypot~a-<br />
SSIITHSONIAPi COSTRIBUTIONS TO BOTANY<br />
chyna that lack cilia but may have a very dense<br />
rhizinal mat below. The mat may assume a marginal<br />
position and resemble cilia. These “cilia,”<br />
however, will be dichotomously branched. Such a<br />
distinction fails in many specimens <strong>of</strong> H. rcvoluta<br />
(Floerke) <strong>Hale</strong> having ra<strong>the</strong>r sparsely branched<br />
rhizines and a few marginally positioned “cilia.” If<br />
incorrectly interpreted as having furcate rhizines<br />
and sparse marginal cilia, such a lichen might be<br />
identified as <strong>Parmelina</strong> cryptochlom, which has<br />
more powdery capitate soralia, or as pustulate P.<br />
spiimosn, which has a pale yellowish medulla. All<br />
<strong>of</strong> <strong>the</strong>se species have gvrophoric acid.<br />
One fur<strong>the</strong>r example <strong>of</strong> parallelism involving<br />
Pam e 1 in a rl is sec t a and Hypo t ru c h y n a n eod issec t a<br />
(<strong>Hale</strong>) <strong>Hale</strong> can be mentioned. The latter has<br />
clearly dichotomously branched rhizines and lacks<br />
cilia, yet both contain <strong>the</strong> same chemical, gyrophoric<br />
acid, and have virtually identical lobe configuration.<br />
It is difficult, if not impossible, to decide at<br />
this time whe<strong>the</strong>r intergeneric hybridization has<br />
occurred in <strong>the</strong>se cases and resulted in “hybrid”<br />
species.<br />
RHIzIms.-?’he rhiLines <strong>of</strong> <strong>Parmelina</strong> are <strong>of</strong> two<br />
types: simple to sparsely furcate (Figure 3c) and<br />
squarrose. Squarrose rhizines, that is, those with a<br />
main axis and short lateral branches, are confined<br />
mostly to some species in section Myelochroa. This<br />
pattern is also known in a few species <strong>of</strong> Parmelia<br />
(e.g., P. ,szilcata Taylor) and Pai.motl-ema (<strong>the</strong> P.<br />
7.cticnlOtzLm group).<br />
ISIDIA AND LosuLEs.-Isidia are normally cylindrical<br />
and erect as in o<strong>the</strong>r parmelioid genera and<br />
occur in <strong>the</strong> following 13 species: P. antillensis, P.<br />
expallidla, P. di.csecta (Figure 34, P. horrescens, P.<br />
jamesii, P. indica, P. lindmanii, P. melanochaeta,<br />
P. obsessn, P. perisidians, P. tiliacea, P. usambarensis,<br />
and P. urnllichinna. Isidia are distinctly lobulate<br />
in P. degelii ant1 P. spatlziilata (Figure 44 and<br />
uniquely peltate in P. pastillifem (Figures 3e, 4b), a<br />
close relative <strong>of</strong> P. tiliacea, Apical cilia almost always<br />
occur in P. howescens and P. melanochaeta<br />
(Figure 4c) and resemble those in PaYmotTema crin i-<br />
tiinz (Acliarius) Choisy.<br />
Lobules not originating from isidia are found in<br />
P. schindle7.i and P. xantholepis (Figure 4e,f); <strong>the</strong>y<br />
are mostly marginal.<br />
PvsTuLEs.-These are characteristic <strong>of</strong> four species,<br />
P. Iiaynchinensis, P. leucotylizn (Figure 5g), P.
NUMBER 33<br />
FIGURE 1.-Epicortical pores oti <strong>the</strong> surface <strong>of</strong> Parnze[irzn species: a,b, P. ettornzis (X 400 and<br />
1600) (Jeilicoe 150 in US); c,d, P. muelleri (X 400 and 1600) (<strong>Hale</strong> 42219); e,!, P. indica (X 400<br />
atid 1600) (<strong>Hale</strong> 43884). (Scatitiing-electron microphotographs.)<br />
3
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
FIGURE<br />
2.-Internal structure <strong>of</strong> <strong>Parmelina</strong>; a, vertical cross section <strong>of</strong> P. degelii (Degelius in<br />
US) (x 800); b, upper cortex <strong>of</strong> P. degelii (same section as in a) (x 1600); c, vertical cross sec-<br />
tion <strong>of</strong> P. galbina (<strong>Hale</strong> 23415) (x 800); d, upper cortex <strong>of</strong> P. galbina (same section as d) (x<br />
1600); e, epicortex (arrow) <strong>of</strong> P. pastillifera (Schroppel 143) (x 5000); f, crystals <strong>of</strong> lecanoric<br />
acid on hyphae <strong>of</strong> P. pastillifera (Schroppel 143) (x 4000).
NUMBER 33 5<br />
FIGURE 3.-Morphological structures <strong>of</strong> <strong>Parmelina</strong>: a, white maculae and marginal cilia <strong>of</strong> P.<br />
consors (Malme 1282 in US) (x 10); b, axillary cilia <strong>of</strong> P. lindmanii (Nee and Mori 4258 in US)<br />
(x 10); c, rhizines <strong>of</strong> P. galbina (<strong>Hale</strong> 18930) (x 70 with scanning-electron microscope); d,<br />
isidia <strong>of</strong> P. dissecta (Montes 10121 pro parte) (x 90 with scanning-electron microscope): e,<br />
peltate isidia <strong>of</strong> P. pastillifera (Schroppel 143) (x 90 with scanning electron microscope):<br />
f, cross-section <strong>of</strong> a peltate isidium <strong>of</strong> P. pastillifera (Schroppel 143) (x 400 with scanning-<br />
electron microscope).
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
FIGURE<br />
4.-Morphological structures <strong>of</strong> Parwelina: a, isidia <strong>of</strong> P. dissecta (<strong>Hale</strong> 33375) (x 10); b,<br />
peltate isidia <strong>of</strong> P. pastillifera (Schroppel 143) (x 10); c, ciliate isidia <strong>of</strong> P. horrescens (Imshaug<br />
22174) (X 10); d, procumbetit isidia <strong>of</strong> P. spathulata (Koefler, in US) (x 10); e, lobules <strong>of</strong> P.<br />
schindleri (Schindler 4569 in US) (x 10); f, 1ol)ules <strong>of</strong> P. xantholepis (<strong>Hale</strong> 25993) (x 10).
NUMBER 33 7<br />
FIGURE<br />
5.-Morphological structures <strong>of</strong> Parrnelina: a, soralia <strong>of</strong> P. cc~irulenta (Weber anti McYean<br />
L-51280 in US) x 10); b, soralia <strong>of</strong> P. eryptochlora (<strong>Hale</strong> 35357) (x 10); c, pustules <strong>of</strong> P. su0-<br />
fatiTcens (isotjpe in US) (x 10); tl, coarse coralia <strong>of</strong> P. swinsco7~'ii (Swinscow K 31/13 in US)<br />
(x 10); e, pustules <strong>of</strong> P. spiiniosa (<strong>Hale</strong> 19897) (x 10): f, pustule <strong>of</strong> P. ~piiniosa (<strong>Hale</strong> 19897)<br />
(x 100 with scanning-electron microscope); g, pustules <strong>of</strong> P. leitcotyliza (<strong>Hale</strong> 29.544) (x 10).
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
spumosa (Figure 5e,f), and P. szibfatiscens (Figure<br />
5c). Soredia are not produced on <strong>the</strong>se pustules.<br />
<strong>Parmelina</strong> denegans, on <strong>the</strong> o<strong>the</strong>r hand, is clearly<br />
pustulate initially but produces coarse soredia at<br />
maturity.<br />
SOREDIA.-SOralia are produced in seven species <strong>of</strong><br />
<strong>Parmelina</strong>: P. auwlenta, P. cryptochlora, P. dene-<br />
guns, P. metawvoluta, P. muelleri, P. pilosa, and<br />
P. swinscowii (Figures 5a,b,d, 17b, 18b). They are<br />
orbicular and lamina1 or subterminal but never<br />
marginal and linear as in Parmotrema.<br />
The remaining 17 species <strong>of</strong> <strong>Parmelina</strong> produce<br />
no vegetative propagules, although <strong>the</strong> very<br />
wrinkled and <strong>of</strong>ten fragile upper cortex <strong>of</strong> P.<br />
ento<strong>the</strong>iochroa or P. rhytidodes might be mistaken<br />
for pustules (Figure 6).<br />
APoTHEcIA.-The apo<strong>the</strong>cia are typically adnate<br />
or sessile and less than 5 mm in diameter, though<br />
Pal-nzelina irrugniis has discs up to 10 mm broad.<br />
The disc is occasionally perforate in P. co1zsors. No<br />
species has a coronate rim, and only two, P. qzier-<br />
cina and P. tiliacea, sometimes have retrorse<br />
rhizines on <strong>the</strong> lower part <strong>of</strong> <strong>the</strong> amphi<strong>the</strong>cium.<br />
No fertile collections are known as yet for P.<br />
ci.yptochlora, P. degelii, P. hayachinensis, P. iradica,<br />
P. jamesii, P. spathulatcl, or P. ~winscowi~.<br />
Spore siLe is very uniform within a given species<br />
and not greatly different between unrelated species.<br />
For example, <strong>the</strong> species in <strong>the</strong> P. dissecta-P. hor-<br />
wscens group (Figure 9) have spores 8-12 X 12-18<br />
pm. The P. aurzilenta-P. subaurulenta group (sec-<br />
tion Myelochroa) has spores 5-10 X 7-15 pm. The<br />
largest spores have been measured in P. versiformis<br />
(10-14 X 18-28 pm), but no o<strong>the</strong>r species in <strong>the</strong><br />
genus have spores greater than 19 pm long. The<br />
smallest spores (3-6 X 7-8 pm) occur in <strong>the</strong> two<br />
anamolous norstictic acid-containing species, P.<br />
antillensis and P. phlyctina.<br />
Chemistry<br />
The chemistry <strong>of</strong> <strong>Parmelina</strong> species was de-<br />
termined with microcrystal tests before 1965 (<strong>Hale</strong><br />
and Kurokawa, 1964) and later with thin-layer<br />
chromatography (Merck F-254 silica gel pre-coated<br />
plates). Two solvent systems, hexane-e<strong>the</strong>r-formic<br />
acid and benzene-dioxane-acetic acid, have been<br />
routinely employed using C. Culberson’s methods<br />
(1972). I have had <strong>the</strong> benefit <strong>of</strong> Dr. Culberson’s<br />
advice on <strong>the</strong> identification <strong>of</strong> many <strong>of</strong> <strong>the</strong> lichen<br />
substances, and she has very kindly communicated<br />
to me <strong>the</strong> results <strong>of</strong> her work on <strong>the</strong> “horrescens”<br />
unknown. It should be assumed, incidentally, that,<br />
unless o<strong>the</strong>rwise stated, <strong>the</strong> components <strong>of</strong> each<br />
species in tlie taxonomic section refer to <strong>the</strong> liolo-<br />
type or lectotype specimens with additional com-<br />
ments on tlie results from o<strong>the</strong>r specimens.<br />
Before listing <strong>the</strong> lichen substances discovered<br />
so far in Pal-melinn, I might point out a most re-<br />
markable feature in its chemical pr<strong>of</strong>ile. Orcinol<br />
FIGURE<br />
6,-MorpholoLgy <strong>of</strong> <strong>Parmelina</strong>: a, flaking cortex <strong>of</strong> P. ento<strong>the</strong>iochroa (<strong>Hale</strong> 29.534) (x 4);<br />
b, wrinkles <strong>of</strong> P. rhytidodes (Numnriri 156 in US) (x 10).
NUMBER 33<br />
depsides (except for gyrophoric and lecanoric acids),<br />
meta-depsides, orcinol depsidones (except for rare<br />
lobaric acid), and <strong>the</strong> p-orcinol depsides related to<br />
barbatic acid are completely lacking. Two o<strong>the</strong>r<br />
cortical substances, lichexanthone and usnic acid,<br />
are also missing. The general impression is that<br />
<strong>Parmelina</strong> has a relatively “primitive” assemblage<br />
<strong>of</strong> substances and would fall on a lower evolutionary<br />
scale than Hypotrachyna, Parmotrerna, and<br />
Pseudopamelia but significantly higher than Bzilbothrix,<br />
Painzelia, and Relicina (<strong>Hale</strong>, 1967).<br />
ALIPHATIC L\cIDs.-The identification <strong>of</strong> fatty<br />
acids presents many problems and no entirely satisfactory<br />
techniques have been worked out to deal<br />
with <strong>the</strong>m. The most direct method is to spray <strong>the</strong><br />
chromatographic plates ivith water and carefully<br />
watch for opaque, white, water-repellant spots as <strong>the</strong><br />
plate dries. These are <strong>the</strong> fatty acids. Terpenes have<br />
a similar reaction but differ in turning bluish, purple,<br />
or I~roivn when sprayed with H,SO, and heated.<br />
Dr. AIyles Keogh <strong>of</strong> <strong>the</strong> Universitlatl de 10s Andes,<br />
hlerida, Venefuela, has also informed me (pers.<br />
comm.) that <strong>the</strong> fatty acids turn brownish when<br />
heated for 12 hours or more at 100°C (in contrast<br />
to <strong>the</strong> 10 minutes usually needed for development<br />
<strong>of</strong> phenolic substances). In any event, protolichesterinic<br />
acid has been identified in P. expallida<br />
(Kurokawa, 1968a), and unidentified fatty acids<br />
are <strong>the</strong> principle components in P. endolezica and<br />
P. !let ei.ochloti.<br />
hPsmEs.-These well-known substances are common<br />
in section Pamwlina (those species <strong>of</strong> <strong>Parmelina</strong><br />
lacking terpenes) and occur in most <strong>of</strong> <strong>the</strong><br />
pantemperate-montane pantropical species.<br />
Atranorin is <strong>the</strong> primary cortical substance found<br />
in all species <strong>of</strong> Paymelina.<br />
Gyrophoric acid, a tridepside, occurs in Paimelinn<br />
cryptochlora, P. dissecta, P. melanochaeta, P. spnthirhta,<br />
and P. spztrnosu. It is a probable minor component<br />
in P. tlninnziana, P. horl-escens, P. .schindlei~i,<br />
ancl P. subfritiscen.s along with <strong>the</strong> “horrescens” unknown.<br />
Gyrophoric acid is not easily chromatographed<br />
because <strong>of</strong> <strong>the</strong> tendency for streaking antl<br />
variable R, values that are highly influenced by<br />
solvent composition. In addition, one or more associated<br />
spots, still unidentified, may appear on<br />
<strong>the</strong> plates.<br />
“Horrescens” unknown is actually several substances<br />
presumed to be depsides related to lecanoric<br />
acid. The spots react with H,SO, <strong>the</strong> same as<br />
orcinol depsides and fall close to gyrophoric and<br />
lecanoric acids in <strong>the</strong> usual solvent systems. The<br />
best separation is achieved in benzene-dioxane<br />
where a single large spot, <strong>the</strong> main “horrescens”<br />
unknown, falls well above gyrophoric acid. This<br />
unknown has now been detected in P. damaziana,<br />
P. horrescens, P. schindleri, and P. subfatiscens. I<br />
have also recently discovered this substance in an<br />
undescribed species <strong>of</strong> Hypoti.achyna.<br />
The widespread medullary component lecanoric<br />
acid occurs in P. pastillifem, P. pnrinata, P. qiiercina,<br />
and P. tiliacea. It falls Close to gyrophoric<br />
acid on chromatographic plates antl must be carefully<br />
distinguished from it by running controls.<br />
?’he toluene-acetic acid solvent system should be<br />
used in addition to <strong>the</strong> o<strong>the</strong>r t\vo systems.<br />
DEPsmosiis.-?’liese substances occur in many<br />
lichen genera but are at best s~~oradically developed<br />
in Paim el ina. Pliy logene t icall y “primitive ” salazinic<br />
acid and its close acetate galbinic acid predominate<br />
in section Pai.melina.<br />
Constictic acid is an accessory substance with<br />
stictic acid ancl salazinic acid (see below).<br />
Fumarprotocetraric acid is extremely rare in <strong>the</strong><br />
genus, occurring in only one species, <strong>the</strong> Australian-Sew<br />
Zealantl endemic P. jnmesii. It is accompanied<br />
by protocetraric acid. Succinprotocetraric<br />
acid is lacking.<br />
Galbinic acid is confined to a small group <strong>of</strong><br />
very closely related species: P. grilbincr, P. haynchinensi,~,<br />
P. nic/awvolufa, arid P. ob.scsscr, accompanied<br />
by secalonic acid X and <strong>the</strong> ”s~~baui~i~le~ita” terpene<br />
series. It is easily identified as ;I deep H,SO, orange<br />
spot below norstictic acid in benzene-tlioxane.<br />
Lobaric acid is <strong>the</strong> only orcinol tlepsidone in<br />
Pni.melinci and occurs as an accessory substance<br />
with salwinic acid in P. su~inscou~ii.<br />
Korstictic acid is <strong>the</strong> main component in two<br />
closely related Caribbean species, P. antillensis and<br />
P. phlyctina. A lower spot <strong>of</strong> <strong>the</strong>, as yet, unidentified<br />
connorstictic acid may be present.<br />
Protocetraric acid occurs only in P. jnmesii with<br />
fumarprotocetraric acid (see above).<br />
Salazinic acid, a very common substance in <strong>the</strong><br />
Parmeliaceae, occurs in P. enotmis, P. .rimplicior,<br />
P. ,\ ZL’ in .sc ow i i, P. 1 I sa ?n have n.c is, P. ve~s i f 07-rn is, ant1<br />
P. zuallichinna. It is a rare accessory with leucotylin<br />
in P. el-assata antl may be accompanied by constictic<br />
9
10 S\IITHSONIAS CONTRIBUTIONS TO BOTANY<br />
acid in P. metarevoluta. All <strong>of</strong> <strong>the</strong>se species, ex-<br />
cepting P. versiformis, are distributed in <strong>the</strong> Old<br />
World following a pattern seen in Bulbothrix<br />
(<strong>Hale</strong>, 1976b).<br />
Stictic acid has been discovered in only one spe-<br />
cies, P. mnelleri, where it is accompanied by con-<br />
stictic acid and two unknown compounds falling<br />
between stictic and constictic acids in benezene-<br />
dioxane.<br />
TERPENEL-A closely related series <strong>of</strong> triterpenes<br />
with a liopane skeleton have been described for<br />
species in section Myelochroa. Their structures have<br />
been determined by Yosioka and his group in Ja-<br />
pan using nuclear magnetic resonance (NhIR)<br />
spectra, mass spectrometry, etc. The tliin-layer<br />
chromatography is not yet standardized or well un-<br />
derstood, and I have not tried to identify individual<br />
spots. Typical pr<strong>of</strong>iles for <strong>the</strong> terpene-containing<br />
species <strong>of</strong> <strong>Parmelina</strong> are illustrated in Figure 7.<br />
Leucotylic acid, a colorless compound closely<br />
related to both zeorin and leucotylin, was first elu-<br />
cidated by Yosioka (1966). It was isolated from Pa?.-<br />
melina leucotyliza, although my results indicate that<br />
leucotylin is, in fact, <strong>the</strong> main component <strong>of</strong> this<br />
species. I suspect that <strong>the</strong>ir material included a<br />
mixture <strong>of</strong> P. crassata, which does contain leuco-<br />
tylic acid. The species <strong>of</strong> Pnrmelina which contain<br />
leucotylic acid (or at least have <strong>the</strong> same terpenic<br />
pr<strong>of</strong>ile as P. airrulentn, as illustrated in Figure 7)<br />
are P. aurzilenta, P. degelii, P. il-rugans, and P.<br />
rhytidodes.<br />
Leucotylin, <strong>the</strong> progenitor <strong>of</strong> leucotylic acid, is<br />
<strong>the</strong> main terpene component in P. amagiensis, P.<br />
crassata, P. denegans, P. ento<strong>the</strong>iochroa, P. galbina,<br />
P. hayachinensis, P. indica, P. leucotyliza, P. meta-<br />
revoluta, P. obsessa, P. perisidians, P. subaurulenta,<br />
and P. xantholepis. It forms a low major spot in<br />
benzene-dioxane, but I have generally identified it<br />
as a pr<strong>of</strong>ile <strong>of</strong> several terpene spots as illustrated in<br />
Figure 7.<br />
Yosioka (Yosioka and Nakanislii, 1963; Yosioka<br />
and Nakanishi, 1966) has isolated at least five o<strong>the</strong>r<br />
compounds related to leucotylin in “Parmelia ento-<br />
<strong>the</strong>iochroa.” I presume that at least some <strong>of</strong> <strong>the</strong>se<br />
make up <strong>the</strong> numerous spots resolved in hexane-<br />
e<strong>the</strong>r (see Figure 7), but no attempt has been made<br />
to identify any <strong>of</strong> <strong>the</strong>m on <strong>the</strong> plates. There seems<br />
to be great variation in <strong>the</strong> intensity <strong>of</strong> <strong>the</strong> spots,<br />
reflecting different concentrations in <strong>the</strong> thallus. It<br />
remains to be seen whe<strong>the</strong>r this variation has any<br />
taxonomic value.<br />
Zeorin is a well known lichen substance. The<br />
stereochemistry <strong>of</strong> its hopane skeleton was only recently<br />
studied by Yosioka et al. (1968b). It occurs in<br />
all species in section 1\4pelochroa and has been reported<br />
in Acroscyplirrs, Cladonin spp., Hppotrachpa<br />
majoris (Vainio) <strong>Hale</strong>, Lecanom, Sephroma,<br />
Peliigera, and <strong>the</strong> Physciaceae. It may be identified<br />
as <strong>the</strong> highest blue spot on <strong>the</strong> chromatographic<br />
plates in Iiotli solvent systems (Figure 7).<br />
Prc~iE.\’Ts.-.\Iedullary pigments are especially<br />
characteristic <strong>of</strong> species in section hfyelochroa. The<br />
chromatography <strong>of</strong> <strong>the</strong>se is extremely difficult and<br />
~is~iallp unsuccessful. There is considerable streaking<br />
and individual components cannot be distinguished.<br />
Secalonic acid X was first ideniifietl by Yosioka<br />
et al. (1968a:2090), who established its identity with<br />
ento<strong>the</strong>in. It also occurs in ergot (Shibata et al.,<br />
1964). Yosioka found secalonic acid A in <strong>Parmelina</strong><br />
nii~rilcntm, P. enfo<strong>the</strong>iochroa, P. pel-isiclians, and<br />
P. subauixlenta, and I presume it is <strong>the</strong> major<br />
pigment in all species <strong>of</strong> section ilfpelochroa with<br />
a yellowish orange medulla, as well as in P. immiscens<br />
and P. lintlmanii, both lacking terpenes. Unidentified<br />
reddish pigments accompany secalonic<br />
acid 4 in <strong>the</strong> lower medullary layer <strong>of</strong> P. amagien-<br />
,pis and P. dcnegans.<br />
Phytogeography and Speciation<br />
Pawnelinti occurs primarily on trees in secondary<br />
forests in lemperate zones and at higher elevations<br />
in <strong>the</strong> tropics. A number <strong>of</strong> <strong>the</strong> commoner species<br />
may also occur on rocks. There are, however, only<br />
four oliligately saxicolous species, P. enownis, P.<br />
indica, P. obsessa, and P. usambaiensi.r. The Pnr-<br />
melina floras <strong>of</strong> various geopolitical units are<br />
enumerated below. It is altoge<strong>the</strong>r obvious that<br />
many countries are underrepresented because <strong>the</strong>y<br />
have not been visited by lichen collectors.<br />
XORTH AMERIC.~<br />
United States: P. antillensis, P. aurulenta, P. dissecta, P.<br />
galbi17n, P. hoi 1 escciis, P. ~ne/ar~~volii/a, P. obsessa, and P.<br />
sfiu in osa.<br />
MEXICO ALD C~NTRAL AMERIC~<br />
Mexico: P. antillensis, P. aurulenta, P. dissecta, P. hor-
NUMBER 33 11<br />
rescens, P. ivztniscens, P. Zindnlanii, P. niuelieri, and P. WEIT ISDIE s<br />
spumosa.<br />
Guatemala: P. horrescens. Cuba: P. antiliensis, P. dissecta, P. horrescens, and P.<br />
Panama: P. dissect(/, P. Izorre,ce,ls, and P. subfutiscens. pizlyctinn.<br />
FIGURE<br />
7.-Chromatographic pr<strong>of</strong>iles <strong>of</strong> Pnrnielinu species: pr<strong>of</strong>iles in <strong>the</strong> hexane-e<strong>the</strong>r solvent<br />
system (top) and benzene-dioxaiie solvent system (bottom). (1 1 P. uicndenla, 2 = P. degelii,<br />
3 = P. irrugans, 4 = P. rhytidot/u.i, 5 = P. ci)nrrgie!i,iis, G = P. ci-nssata, 7 = P. denegnns, 8 = P.<br />
ento<strong>the</strong>iochroa, 9 = P. indlica, 10 = P. iettcotyliza, 11 = P. perisidians, 12 = P. subaurulentn,<br />
13 = P. santholepis, 14 = P. plbi!rcr, 1.5 = P. tnetarevoltrtrr, lG = P. ohsessa, Z = zeorin spot,<br />
G = galbinic acid, S = salarinic acid, and C = constictic acid: leucotylic acid is presumed to<br />
be <strong>the</strong> series <strong>of</strong> spots in 1 to 4 just below rerorin, and leucotylin is <strong>the</strong> lowest series in 5-10<br />
in benzene-dioxane; atranorin falls just above <strong>the</strong> point where <strong>the</strong> photographs were trimmed.)
Jamaica: P. antillensis, P. dissecta, P. horrescens, P.<br />
phlyctina, P. spumosa, and P. subfatiscens.<br />
Hispaniola: P. antillensis, P. dissecta, P. horrescens, and<br />
P. phlyctina.<br />
Puerto Rico: P. phlyctina.<br />
Lesser Antilles (including Trinidad): P. anlillensis, P.<br />
cryptochlora, P. dissecta, and P. subfatiscens.<br />
SOUTH AMERICA<br />
Colombia: P. lindmanii, P. melanochaeta, and P. spumosa.<br />
Venezuela: P. antillensis, P. dissecta, P. horrescens, P.<br />
lindmanii, P. muelleri, and P. spumosa.<br />
Ecuador: P. pilosa.<br />
Peru: P. muelleri.<br />
Brazil: P. antillensis, P. aurulenta, P. consom, P. damaziana,<br />
P. dissecta, P. heteroloba, P. lindmanii, P. melanochaeta,<br />
P. muelleri, P. schindleri, P. spumosa, and P.<br />
wersiformis.<br />
Uruguay: P. cons or,^, P. horrescens, P. lindmanii, and P.<br />
pi losa .<br />
Paraguay: P. consors, P. lindmanii, and P. melanochaeta.<br />
Argentina: P. consors, P. lindrnanii, P. muelleri, P. pilosa,<br />
and P. uersiformis.<br />
Chile: P. horrescens, P. pilosa, P. spumosa, and P. swinscowii.<br />
EUROPE<br />
Western Europe: P. dissecta, P. horrescens, P. pastillifera,<br />
P. quercina, and P. tiliacea.<br />
Russia: P. aurulenta, P. qziercina, and P. tiliacea.<br />
Israel: P. tiliacea.<br />
AFRIC4<br />
Morocco: P. tiliacea.<br />
Tunisia. P. tiliacea.<br />
Ivory Coast: P. usambarensis and P. wallichiana.<br />
Guinea: P. usambarensis and P. wallichiana.<br />
Uganda: P. usambarensis and P. wallichiana.<br />
Kenya: P. dissecta, P. pilosa, P. swinscowii, and P. wallichiana.<br />
Angola: P. wallichiana.<br />
Rhodesia: P. wallichiava.<br />
Zambia: P. enormis.<br />
Tanzania: P. aurulenta, P. dissecta, P. usambarensis, and<br />
P. wallichiana.<br />
Union <strong>of</strong> South Africa (including Swaziland): P. dissecta,<br />
P. horrescens, P. pilosa, P. spathulata, P. spumosa, P. subfatiscens,<br />
and P. wallichiana.<br />
Madagascar-RPunion: P. aurulenta, P. spumosa, and P.<br />
wallichiana.<br />
ASIA AND PACIFIC AREAS<br />
Pakistan: P. aurulenta, P. quercina, and P. tiliacea.<br />
India (including Nepal): P. aurulenta, P. cryptochlora, P.<br />
denegans, P. dissecta, P. expallida, P. horrescens, P. indica,<br />
P. perisidians, P. rhytidodes, P. simplicior, P. spumosa, P.<br />
SAIITHSONIAN CONTRIBUTIONS TO BOTANY<br />
subaurulenta, P. tiliacea, P. wallichiana, and P. xantholepis.<br />
Sri Lanka: P. denegans, P. dissecta, P. perisidians, and P.<br />
subaurulenta.<br />
Thailand: P. expallida, P. perisidians, P. usambarensis, P.<br />
wallichiana, and P. xantholepis.<br />
Indochina: P. aurulenta.<br />
China (including Hong Kong): P. aurulenta, P. irrugans,<br />
P. quercina, P. subaurulenta, and P. wallichiana.<br />
Korea: P. ento<strong>the</strong>iochoroa.<br />
Japan: P. amagiensis, P. aurulenta, P. crassata, P. dissecta,<br />
P. ento<strong>the</strong>iochroa, P. galbiria, P. hayachiizensis, P. horrescens,<br />
P. irrugans, P. leucotyliza, P. metareuoluta, P. perisidians, P.<br />
quercina, P. rhytidodes, P. spumosa, and P. wallichiana.<br />
Taiwan: P. azcrulenta, P. dissecta, P. expallida, P. horrescens,<br />
P. spirmosa, P. subaurulenta, and P. wallichiana.<br />
Philippines: P. aurulenta, P. dissecta, P. horrescens, P.<br />
perisidians, P. wallichiana, and P. xantholepis.<br />
Malaya: P. wallichiana.<br />
Indonesia: P. aurulenta, P. dissecta, P. horrescens, P. spu-<br />
mom, P. wallichiana, and P. xantholepis.<br />
Sabah: P. denegans, P. leucotyliza, and P. wallichiana.<br />
New Guinea: P. aurulenta.<br />
Australia (including New Zealand): P. degelii, P. endoleuca,<br />
P. horrescens, P. jamesii, P. pruinata, P. qziercina, and<br />
P. spumosa.<br />
Hawaiian Islands: P. aurulenta.<br />
To summarize, t.he New World has a total flora<br />
<strong>of</strong> 20 species, 12 <strong>of</strong> <strong>the</strong>m endemic, 3 (P. pilosa, P.<br />
subfatiscens, and P. swinscowii) shared with Africa,<br />
and <strong>the</strong> remaining 5 (P. aurzilenta, P. dissecta, P.<br />
horrescens, P. quercina, and P. spumosa) essentially<br />
pantemperate or montane pantropical.<br />
Europe has a small Paimelina flora. Dahl and<br />
Krog (1973) list only P. tiliacea in Scandinavia. The<br />
checklist by James (1965) for Great Britain includes<br />
P. dissecta, P. horrescens, P. qziercina, and P.<br />
tiliacen. For <strong>the</strong> rest <strong>of</strong> Europe, Poelt (1969) adds<br />
only P. caipoirhizans (= P. qziercina) and P. pastilli-<br />
fera.<br />
Africa has only two endemic species, P. enormis<br />
and P. spnthdatn, and nine o<strong>the</strong>r species ei<strong>the</strong>r<br />
shared with <strong>the</strong> Americas, Asia, or pantemperate in<br />
distribution.<br />
Asia is especially rich in <strong>Parmelina</strong> with 15 spe-<br />
cies on <strong>the</strong> Indian subcontinent and 16 in Japan<br />
and eastern Asia. Tropical Sou<strong>the</strong>ast Asia has nine<br />
species, none <strong>of</strong> <strong>the</strong>m endemic and all occurring<br />
in <strong>the</strong> higher elevation cloud forests. The genus is<br />
not represented at all in <strong>the</strong> lowland dipterocarp<br />
rain forests from Sri Lanka to Indonesia.<br />
The Australia-New Zealand region has seven<br />
species, <strong>of</strong> which four, P. degelii, P. endoleuca, P.<br />
jamesii, and P. pruinata, are endemic. No Parme-
NUMBER 33<br />
FIGURE 8.-Number <strong>of</strong> species in section Myeloclztm in each major geographical division.<br />
linae have been collected on <strong>the</strong> Pacific Islands ex-<br />
cept for P. cl~irttlenta at higher elevations in Ha-<br />
waii. The total Old World flora (Africa, Asia, and<br />
Australia-New Zealantl) comprises 26 species.<br />
One distrihution pattern <strong>of</strong> considerable phyto-<br />
geographic interest stands out. The species <strong>of</strong> sec-<br />
tion Myeloch ma, a ra<strong>the</strong>r homogeneous group, are<br />
concentrated in India and eastern Asia (Figure 8)<br />
and only one <strong>of</strong> <strong>the</strong>m, P. aurulentn, occurs in all<br />
geographic areas (Figure 10). The species in section<br />
Pasmelina, on <strong>the</strong> o<strong>the</strong>r hand, are not strongly<br />
Concentrated in any region.<br />
TVithin section Myelochl-oa <strong>the</strong> species related to<br />
P. galbina form a particularly interesting group. As<br />
pointed out by Kurokawa (1972), P. galbina, <strong>the</strong><br />
fertile progenitor species, behaves as an Arcto-<br />
tertiary plant which migrated southward and has<br />
survived in Japan and eastern Korth America. A<br />
rare sorediate morph, P. metawvoluta, has <strong>the</strong><br />
same range, whereas <strong>the</strong> isidiate morph, P. ob.tessa,<br />
is restricted to North America, and <strong>the</strong> pustulate<br />
morph, P. hayachinensis, to Japan. W. Culberson<br />
(1972) also regards P. aimilenla as a Tertiary relict.<br />
Pai.melina has several complex groups <strong>of</strong> species<br />
that have evidently resulted from combined chem-<br />
ical and morphological evolution. The species in<br />
section iMyelochroa, for example, all <strong>of</strong> which con-<br />
tain terpenes and almost always have secalonic<br />
acid A, a yellow pigment, have evolved from two<br />
chemically dissimilar ancestors, one, <strong>the</strong> smaller<br />
group, with leucotylic acid and resembling P. ir-<br />
g.iigans, and <strong>the</strong> o<strong>the</strong>r, a larger series with leuco-<br />
tylin, resembling <strong>the</strong> present-day P. subazirzilenfa.<br />
The numerous morplis derived from <strong>the</strong>se presump-<br />
tive parents or <strong>the</strong>ir ancestors, P. nmagiensis, P.<br />
aiti.itlenta, P. crassata, P. clegelii, P. denegnns, P.<br />
ento<strong>the</strong>iochroa, P. leucotyliza, P. perisidians, P.<br />
Thytidodes, and P. xnntholcpis, have diversified<br />
mostly by production <strong>of</strong> soretlia, isidia, pustules,<br />
and wrinkles, variation in apotliecial diameter, etc.<br />
AIorph forniation (see <strong>Hale</strong>, 1975a:13) is not<br />
strongly delineated for <strong>the</strong> most part since <strong>the</strong> par-<br />
ents cannot be traced directly. \Ve are left to con-<br />
clude that section Alyelochron is an ancient group<br />
13
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
evolved over a long period. The P. galbina group,<br />
as mentioned above, forms <strong>the</strong> only exception to<br />
this rule.<br />
ILIorph formation has also been an important<br />
mode <strong>of</strong> speciation in section <strong>Parmelina</strong>. The fol-<br />
lowing parent morph-vegetative morph series are<br />
indisputable: P. consors-P. pilosa (sorediate), P.<br />
quercina-P. tiliacea (isidiate), and P. imrniscens-P.<br />
lindmanii (isidiate). <strong>Parmelina</strong> phlyctina and<br />
isidiate P. antillensis are extremely close but less<br />
clear-cut morphs. A large complex <strong>of</strong> species have<br />
been derived from now extinct parents in <strong>the</strong> P.<br />
dissecta-P. horrescens group, as illustrated in Figure<br />
9. Like <strong>the</strong> P. subaurulenta group <strong>of</strong> section Mye-<br />
lochroa, <strong>the</strong> parent morphs <strong>of</strong> <strong>the</strong>se numerous<br />
vegetative morphs are ei<strong>the</strong>r extinct or have yet to<br />
be discovered.<br />
The remaining species in <strong>the</strong> genus appear to<br />
have no discernible interrelationships or morphs.<br />
These include P. endoleuca, P. enormis, P. expal-<br />
lida, P. heteroloba (part <strong>of</strong> <strong>the</strong> P. darnaziana se-<br />
ries?), P. pruinata, P. sirnplicior, P. swinscowii, P.<br />
usambarensis, P. veisiformis, and P. wallichiana.<br />
Classification <strong>of</strong> <strong>Parmelina</strong><br />
The taxon <strong>Lichen</strong> section Imbricaria was first<br />
proposed by Schreber (1791:767). He cited no spe-<br />
cies, only Squamaria H<strong>of</strong>fmann, which is invalid as<br />
<strong>the</strong> later homonym <strong>of</strong> Squamah Ludwig (Phane-<br />
rogamae). Acharius (1794:250) adopted <strong>the</strong> name as<br />
a tribe <strong>of</strong> <strong>Lichen</strong> and included in it L. olivaceus L.<br />
and L. tiliaceus H<strong>of</strong>fmann, as well as several o<strong>the</strong>r<br />
species now recognized as belonging to Hypogymnta,<br />
Parmeliopsis, Physcia, and Xanthoria. XIichaux<br />
(1803) raised <strong>the</strong> name to generic rank and cited<br />
one species, a lichen now called Anzia colpodes<br />
(Acharius) Stizenberger. Fries (1825:242) later trans-<br />
ferred it to Parmelia as Parmelia section Imbricaria,<br />
and Koerber (1855:68) used it at <strong>the</strong> generic rank,<br />
even though he realized that <strong>the</strong> name was a later<br />
homonym <strong>of</strong> Imbricaria Decandolle (Phaneroga-<br />
mae). Koerber and contemporary lichenologists<br />
placed many species that we now consider parme-<br />
lioid in Imbricaria, while at <strong>the</strong> same time using<br />
Parmelia for many species now placed in Physcia.<br />
After <strong>the</strong> delimitation <strong>of</strong> Parrnelia in a modern<br />
sense by authors such as Sluller and Nylander,<br />
Imbricaria was relegated to synonymy under it.<br />
<strong>Hale</strong> and Kurokawa (1964: 130) grouped <strong>the</strong> narrow-<br />
lobed, marginally ciliate Parrneliae in Parmelia<br />
subgenus Parmelia section Imbricaria (Schreber) E.<br />
Fries, and it is essentially this group that I have seg-<br />
regated as a distinct genus, <strong>Parmelina</strong> (<strong>Hale</strong>, 1974:<br />
482).<br />
<strong>Parmelina</strong> is a ra<strong>the</strong>r heterogeneous assemblage <strong>of</strong><br />
47 species differing in lobe width, production <strong>of</strong><br />
cilia, pigmentation, and rhizine branching. Two<br />
major groups can be recognized, one including<br />
those species lacking terpenes and one with terpene-<br />
containing species. They may be conveniently re-<br />
garded as sections.<br />
Section <strong>Parmelina</strong><br />
Parmelia section Hypotrachyna subsection Myelolezica Asa-<br />
hina, 1952:74 [type-species: Purrnelina tiliacea (H<strong>of</strong>fmann)<br />
<strong>Hale</strong>].<br />
The species in this section are characterized by<br />
<strong>the</strong> lack <strong>of</strong> any triterpenes. Except for two species,<br />
P. immiscens and P. lindmanii, <strong>the</strong> medulla is white<br />
and unpigmented. Several internally homogeneous<br />
groups can be recognized. For example, <strong>the</strong> P.<br />
dissecta-P. horrescens group (Figure 9) includes<br />
seven species with very similar lobe configuration,<br />
abundant marginal cilia, and gyrophoric acid or<br />
<strong>the</strong> closely related “horrescens” unknown. Heavily<br />
white-niaculate P. consow, P. pilosa, and P. rnuelleri,<br />
having coarse, lea<strong>the</strong>ry thalli and thick, furcate<br />
marginal cilia, are obviously related.<br />
The predominantly European complex <strong>of</strong> spe-<br />
cies, P. qztcrcina-P. iiliacea-P. pastillifera, contains<br />
lecanoric acid and forms an easily recognized but<br />
isolated group. <strong>Parmelina</strong> antillensis and P. phlyc-<br />
tina, both unusual in containing norstictic acid,<br />
have no close relatives in <strong>the</strong> genus. Two o<strong>the</strong>r<br />
New World species, P. immiscens and P. lindmanii<br />
with a yellow medulla, form an isolated branch <strong>of</strong><br />
this section.<br />
The species with salazinic acid, P. enormis, P.<br />
simplicior, P. swinscowii, P. usambarensis, P. versi-<br />
formis, and P. wallichiana, have little in common<br />
except <strong>the</strong>ir chemistry. Finally, <strong>the</strong> remaining spe-<br />
cies, P. endoleuca, P. expallida, P. heteioloba, P.<br />
jamesii, and P. pririnata, have no obvious common<br />
ancestry or affinities with o<strong>the</strong>r species in <strong>the</strong> sec-<br />
tion.
NUMBER 33 15<br />
Progenitor<br />
I-<br />
Progenitor Progenitor<br />
(gyrophoric acid) (“horrescens” unkn.)<br />
P. horrescens IP. schindleri P. sirbfutiscens<br />
Progenitor ,(white Progenitor rnedullaj (isidiate) (lobulate) (pustulate)<br />
(yellow medulla)<br />
I P. spir ntosu<br />
(pustulate)<br />
r-l-l<br />
P. dissei.tu P. sputhulatu P. cryptochloru<br />
(isidiate) (lobulate-isidiate) (sorediate)<br />
FIGURE 9.-Hypo<strong>the</strong>tical derivation <strong>of</strong> species in <strong>the</strong> <strong>Parmelina</strong> dissecta-P. horrescens group.<br />
hrot shoivn are P. damaziana, a presumptile nonisidiate, nonsorediate derivative <strong>of</strong> <strong>the</strong> “hor-<br />
rescens” progenitor, and P. melanochaetn, a possible relative <strong>of</strong> P. dissecta but derived from a<br />
different progenitor.<br />
Section Myelochroa (Asahina) <strong>Hale</strong>, new status<br />
Parmelia section Hypotrachyna subsection Myelochroa Asa-<br />
hina, 1952374 [type-species: <strong>Parmelina</strong> aurdenta (Tucker-<br />
man) <strong>Hale</strong>].<br />
All <strong>of</strong> <strong>the</strong> species in this section contain zeorin<br />
and ei<strong>the</strong>r leucotylin or leucotylic acid and asso-<br />
ciated terpenes, and all, excepting P. indica, produce<br />
varying amounts <strong>of</strong> <strong>the</strong> yellow-orange pigment<br />
secalonic acid A and possibly o<strong>the</strong>r, as yet, unidenti-<br />
fied pigments. This group is highly restricted to<br />
Asia, and <strong>the</strong> 17 species have very close affinities, in<br />
sharp contrast to section <strong>Parmelina</strong>.<br />
Two apparently unrelated groups are recognira-<br />
ble in <strong>the</strong> section. One, <strong>the</strong> P. galbina group (P. gal-<br />
bina, P. hayachinensis, P. metarevoluta, and P. ob-<br />
sessa), is characterized by galbinic acid in addition<br />
to leucotylin and secalonic acid A. The o<strong>the</strong>r group<br />
includes two closely related subgroups, <strong>the</strong> P. auru-<br />
lenta subgroup (P. aul-ulenta, P. degelii, P. irru-<br />
guns, and P. rhytidodes) with leucotylic acid, and<br />
<strong>the</strong> P. subazirulenta subgroup (P. amagiensis, P.<br />
crassata, P. denegans, P. ento<strong>the</strong>iochroa, P. leuco-<br />
tyliza, P. perisidzans, P. subauiulenta, and P. xan-<br />
tholepis) with leucotylin. <strong>Parmelina</strong> indica can<br />
probably be classified in <strong>the</strong> P. szi baurulenta sub-<br />
group although it has a small saxicolous thallus<br />
and lacks secalonic acid A.<br />
The affinities <strong>of</strong> <strong>Parmelina</strong> lie primarily with<br />
Parmotrema Alassalongo, a broader lobed, loosely<br />
attached, <strong>of</strong>ten ciliate genus (<strong>Hale</strong>, 1965). Palme-<br />
Zina, however, has a far less evolved chemistry, nar-<br />
rower adnate lobes, consistently small spores, and<br />
adnate or sessile, usually imperforate apo<strong>the</strong>cia.<br />
Where soredia are produced, <strong>the</strong>y are lamina1<br />
ra<strong>the</strong>r than marginal in contrast to most Parmo-<br />
trema species. All in all, however, <strong>the</strong> apparent<br />
intergradation between broad-lobed <strong>Parmelina</strong>e and<br />
<strong>the</strong> smaller lobed species <strong>of</strong> Parmot?-emu, involving<br />
perhaps six or eight species, poses difficult problems<br />
in <strong>the</strong> correct generic identification <strong>of</strong> individual<br />
specimens and for <strong>the</strong> “lumper,” at least, raises<br />
questions on <strong>the</strong> distinctness <strong>of</strong> <strong>the</strong> two genera as<br />
circumscribed here. I do not expect that <strong>the</strong> prob-<br />
lem can be resolved to <strong>the</strong> satisfaction <strong>of</strong> all lichen-<br />
ologists, but <strong>the</strong> basic differences should become<br />
clearer as more collections are made in tropical
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
countries and our knowledge <strong>of</strong> soredial formation, species lacking isidia, soredia, and pustules; (2)<br />
significance <strong>of</strong> chemical variation, etc., broadens. those with isidia, lobulate isidia, or lobules; and<br />
The key below is divided into three sections: (1) (3) those with soredia or pustules.<br />
Key to <strong>the</strong> Species <strong>of</strong> <strong>Parmelina</strong><br />
1. THALLUS LACKING ISIDIA, SOREDIA, AND PUSTULES<br />
(Cortex flaking <strong>of</strong>f in <strong>Parmelina</strong> ento<strong>the</strong>tochroa and P. phlyctina, and heavily rugose in P.<br />
crassata and P. rlzytidodes)<br />
1. Medulla more or less completely yellow to yellow-orange (pigmented mostly below <strong>the</strong><br />
apo<strong>the</strong>cia in P. galbina).<br />
2. Upper surface hea\ily rugose (Figure 6b) 35. P. rhytidodes<br />
2. Upper surface more or less plane.<br />
3. Upper cortex vei) fragile, flaking awa) over extensive areas (Figure 6a) .<br />
4. Medulla deep yellow 13. P. ento<strong>the</strong>iochroa<br />
4. Medulla white (turning reddish if impropeily curated) 3 1, P. phylctina<br />
3. Upper cortex entire and continuous.<br />
5. Lower half <strong>of</strong> medulla darker reddish orange 1. P. amagiensis<br />
5. Loner half <strong>of</strong> medulla not darkly pigmented.<br />
6. Lobes narrow and sublinear, 1-2 mm wide 15. P. galbina<br />
6. Lobes broader, subirregular, 2-4 mm wide.<br />
7. Collected in Mexico<br />
19. P. immiscens<br />
7. Collected in Asia.<br />
8. Apo<strong>the</strong>cia 3 mm or less in diameter<br />
40. P. subaurulenta<br />
8. Apo<strong>the</strong>cia 3-10 mm in diameter.<br />
9. Thallus coarse, about 250 pm thick: leucotylin present 5. P. crassoto<br />
9. Thallus tnembranous, about 150 pm thick: IeuLotylic acid present<br />
21. P. irrugans<br />
1. Medulla entirely white.<br />
10. Lower surface pale brown.<br />
11. Collected on rocks in Africa 12. P. enormis<br />
11. Collected in treer in South America 45. P. versiformis<br />
10. Lone1 surface black.<br />
12. Medulla K+ yellow turning red.<br />
13. Lobes sublinear, 1-2 mm wide 15. P. galbina<br />
13. Lobes broader, 2-5 mm wide.<br />
14. Upper cortex fragile, flaking <strong>of</strong>f<br />
31. P. phlyctina<br />
14. Upper cortex firm, continuous.<br />
15. Collected in India 37. P. simplicior<br />
15. Collected in South America 45. P. versiformis<br />
12. Medulla K negatike.<br />
16. Medulla Cf red.<br />
17. Cortex distinctly white maculate<br />
34. P. quercina<br />
17. Cortex emaculate 33. P. pruinnta<br />
16. Medulla C-.<br />
18. Thallus coriaceous, heavily whlte-maculate; lobes 2-5 mm jvide<br />
4. P. consors<br />
18. Thallus thin, emaculate: lobes 1-3 mm i\ide.<br />
19. Medulla KCf rose; “horrescens” unknor\n present 7. P. damaziana<br />
19. Medulla KC-: fatty acids present.<br />
20. Collected in Australia<br />
11. P. endoleuca<br />
20. Collected in South America<br />
17. P. heteroloba<br />
11. THALLUS ISIDIATE, LOBULATE-ISIDIATE, OR LOBULATE<br />
I. Medulla yellow to pale orange-yellow.
NUMBER 33<br />
2. Thallu\ densely lobulate uithout formation <strong>of</strong> isidial initials (Figure 4f)<br />
47. P. xantholepis<br />
2. Thallus noimally isidiate or lobulate-isidlate.<br />
3. Isidia large, doisiientral and lobulate<br />
8. P. degelii<br />
3. Isidia normal, cylindrical and erect.<br />
4. Lobes subirregular and apicall) rotund, 3-6 mm wide 24. P. lindmanii<br />
4. Lobes sublinear, 1-2 mni Ttide.<br />
5. Plants saxicolous (rarel) coi ticolous) 111 eastetn Sorth America 28. P. obsessa<br />
5. Plant5 corticolous in Sou<strong>the</strong>ast A5ia 30. P. perisidians<br />
1. Medulla M hite.<br />
6. Thallus densel) lobulate I\ ithout isidial initlals (Fignre 4e)<br />
6. Thallus isidiate 01 lobulate-isidiate.<br />
7. lsidia in part ptocumbent and lobulate (Fignre 4c,rl).<br />
36. P. schindleri<br />
8. Medulla C+ rose 38. P. qpathulata<br />
8. Medulla C-<br />
18. P. hori escens<br />
7. lsitlia erect, cylintliical.<br />
9. Medulla Kf jellou turning red.<br />
10. Collected in tiopical Ameiica, norstictic acid piesent<br />
10. Collected in Asia and Aflica; salazinic acid present.<br />
2. P. antillensis<br />
11. Lobes subirregulal, cioitded, apicall) iotund<br />
46. P. urallichiana<br />
1 I . Lobes bublineai, looselp dii aricate 44. P. usambarensis<br />
9. Medulla I(- or faintly )elloit.<br />
12. Medulla Cf rose 01 led.<br />
13. Isidia apically flattened and peltate<br />
13. Isidia cylindiical, not peltate.<br />
29. P. pastillifera<br />
14. Medulla C+ intense red (lecanoric acid)<br />
14. Medulla C+ lose (g)iophoiic acid).<br />
43. P. tiliacea<br />
15. Lobes narrow, 1-2 mni nide; upper surface emaculate: isidia<br />
eciliate 10. P. dissecta<br />
15. Lobes broader, 2 4 mm itide; upper surface maculate; isidia<br />
ciliate 25. P. melanochaeta<br />
12 4iedulln C-.<br />
16. Medulla P+ red<br />
16. Medulla P-.<br />
22. P. jamesii<br />
17. Louei surface bronn to hlacLening in palt<br />
17. Lower suiface unifoimly black.<br />
14. P. expallida<br />
18. Isidia apically ciliate; medulla KC+ rose 18, P. hort escens<br />
18. Isidia eciliate; medulla KC- 20. P. indico<br />
111. THALLUS SOREDIATE, SOREDIATE-PUSTULATE, OR PU5TULATE<br />
1. Medulla entirely white.<br />
2. Lobes broad and apicall) rotund; coi tex con
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
10. Pustules forming in capitate masscs (Figuie 5g) 23. P. Zeucotyliza<br />
10. Pustules irregular, intermixed with wrinkles (Figure 6b) 35. P. rhytidodes<br />
9. Lobes narrow and sublinear, less than 2 mm wide.<br />
11. Medulla K+ reddish, P+ orange 16. P. hayachinensis<br />
11. Medulla K-, P-, C+ rose, pigmented bery faint yellow 39. P. spumosa<br />
8. Thallus sorediate or pustulate-sorediate at maturit).<br />
12. Medulla deeper reddish orange In <strong>the</strong> lower half 9. P. denegans<br />
12. Medulla uniforiiil) pigmented to nhite and pigmented only under <strong>the</strong> soralia.<br />
13. Lobes 2-4 mm wide; medulla P- 3. P. aurulenta<br />
13. Lobes less than 2 mm wide; medulla P+ orange 26. P. metarevoluta<br />
Species Treatment<br />
The 47 species <strong>of</strong> <strong>Parmelina</strong> are arranged below<br />
in alphabetical order. Location <strong>of</strong> specimens listed<br />
under “Specimens Examined” are indicated by <strong>the</strong><br />
standard herbarium acronyms except for those col-<br />
lected by <strong>Hale</strong>, all <strong>of</strong> which are preserved in <strong>the</strong><br />
<strong>Smithsonian</strong> Institution (US).<br />
1. <strong>Parmelina</strong> amagiensis, new combination<br />
FIGURE Ila<br />
Parnzelia arriagiensis Asahina, 1951a:228 [type collection: Mt.<br />
Amagi, Prov. Izu, Japan, Asahina 95 (TNS, lectotype)].<br />
DEScRIPTIoN.-Thallus loosely adnate on bark or<br />
mosses over bark, buff mineral gray, 4-8 cm broad;<br />
lobes more or less subirregular, imbricate, 2-3 mm<br />
wide, <strong>the</strong> marginal cilia numerous, simple, up to<br />
1 mm long; upper surface plane, white-maculate,<br />
becoming heavily pycnidiate, isidia and soredia<br />
lacking; medulla deep salmon orange; lower sur-<br />
face black, densely rhizinate, <strong>the</strong> rhizines black,<br />
simple or sparsely squarrosely branched, 0.5-1 .O mm<br />
long. Apo<strong>the</strong>cia sessile to subpedicillate, 2-4 mm<br />
in diameter; spores 8, 6-7 X 9-1 1 pm.<br />
CHEMISTRY.-cOrteX K + yellow (atranorin); rne-<br />
dulla more intensively yellow with color tests<br />
(zeorin, leucotylin and related “subaurulenta” ter-<br />
penes, secalonic acid A, and unidentified pigments).<br />
DIsTRIBuTIoN.-Japan.<br />
REMARKS.-ThiS rare Japanese species is very<br />
close to P. denegans, a pustulate or sorediate-<br />
pustulate species in tropical Asia. Both have a<br />
more deeply pigmented medulla, especially in <strong>the</strong><br />
lower half, than o<strong>the</strong>r members <strong>of</strong> section Myelo-<br />
chroa. I have not been able to resolve <strong>the</strong>se pig<br />
ments with <strong>the</strong> usual chromatographic solvent<br />
systems. <strong>Parmelina</strong> amagiensis appears to be some-<br />
what more robust than P. denegans, judging from<br />
<strong>the</strong> few specimens available, and does not seem to<br />
be a parent morph for <strong>the</strong> latter.<br />
SPPCIMENS Ex \MIsFD.-Japan: Prov. Ettchu, Yanagisawa<br />
599a (TNS); Prov. Hyuga, Kurokawa 50058 (TNS, US).<br />
2. <strong>Parmelina</strong> antillensis<br />
FIGURE 11 b<br />
Parmelirin antillensis (Nylander) <strong>Hale</strong>, 1974:482.<br />
Parmelia antillensis Nylander, 1868:264 [type collection:<br />
Matouba, Guadeloupe, Husnot 445 (H. Nylander herbar-<br />
ium number 35119, lectotype; isolectotypes in G, P)].<br />
Parinelia blastica ’I’ainio, 1896:32 [type collection: Shawford<br />
Estate, Dominica, Elliott 899 (TUR, lectotype; isolectotype<br />
in BM)].<br />
DEscRIPTIoN.-Thallus adnate on bark, fragile<br />
and membranous, up to 10 cm across, greenish min-<br />
eral gray, <strong>of</strong>ten more or less white-pruinose at <strong>the</strong><br />
lobe tips; lobes irregularly branched, subimbricate<br />
to convoluted, <strong>the</strong> marginal cilia sparse, mostly in<br />
lobe axils, to 0.5 mm long; upper surface shiny,<br />
plane, cortex continuous or cracked with age;<br />
densely isidiate, <strong>the</strong> isidia fine, to 0.4 mm high,<br />
sparingly branched or simple; lower surface black,<br />
brown and shiny in a narrow zone along <strong>the</strong> mar-<br />
gins, densely rhizinate, <strong>the</strong> rhizines simple. Apo-<br />
<strong>the</strong>cia adnate, 3-5 mm in diameter, <strong>the</strong> amphi-<br />
<strong>the</strong>cium isidiate; spores 8, 5-6 X 7-8 pm.<br />
CHEMISTRY.-cOrteX K + yellow (atranorin); rne-<br />
dulla K+ yellow turning red, C-, KC-, P+<br />
orange (norstictic and connorstictic acids).<br />
DIsTRIsuTIoN.-Caribbean region.<br />
REMARKs.-Parmelia antillensis and its close Ca-<br />
ribbean relative P. phlyctina are <strong>the</strong> only two spe-<br />
cies in <strong>the</strong> genus containing norstictic acid. This<br />
trait, along with <strong>the</strong> ra<strong>the</strong>r broad, apically rotund<br />
lobes, might tempt one to place <strong>the</strong>m in Parmo-<br />
trema Massalongo, where, in fact, I had originally<br />
suggested <strong>the</strong>y might belong (<strong>Hale</strong>, 1959). The
NUMBER 33<br />
lower surface, however, has at most a narrow bare<br />
or papillate zone, <strong>the</strong> lobes are generally adnate at<br />
<strong>the</strong> tips, not ascending, and <strong>the</strong> cilia are mostly in<br />
<strong>the</strong> axils <strong>of</strong> <strong>the</strong> lobes. Both P. antillensis and P.<br />
phlyctinn share traits intermediate between Parme-<br />
Zina and Parmotrema and have evolved in a very<br />
restricted region, <strong>the</strong> rain forests <strong>of</strong> <strong>the</strong> Caribbean<br />
basin (excepting <strong>the</strong> unusual record <strong>of</strong> P. until-<br />
1en.yi.r. from <strong>the</strong> sou<strong>the</strong>astern United States discussed<br />
in <strong>Hale</strong>, 1971b:46). O<strong>the</strong>r parmelioid groups have<br />
evolved here, eg., Pseudoparmelia martinicnnn<br />
(Nylander) <strong>Hale</strong> and P. raunkiaeri (Vainio) <strong>Hale</strong><br />
and perhaps also Relicinn eximbricnta (Gyelnik)<br />
<strong>Hale</strong> (1975b, 1976a), but <strong>the</strong>se are all lowland spe-<br />
cies in dry, disturbed habitats. Although <strong>Parmelina</strong><br />
ontillensis is <strong>of</strong>ten collected on citrus trees at 600-<br />
700 m elevation, its primary habitat is <strong>the</strong> canopy<br />
branches <strong>of</strong> trees in virgin rain forest.<br />
SrECI\rEss Ex4\rrXED.-L?nited States: Tennessee, Yoshi-<br />
viiira et al. 660872 (US). Mexico: Chiapas, <strong>Hale</strong> 20238.’<br />
Cuba: Oriente, Iinsliairg 250.51 (MSC). Jamaica: Cztlberson<br />
13648, 13777, 13787 (DUKE), Znishaug 14088, 14627, 14943,<br />
1.5321 (MSC), 14593, 15295 (MSC, US), Orcult 3551b, 5626b<br />
(US), Plitt (US). Haiti: Oueit, Inishaug 22642 (MSC). Puerto<br />
Rico: Stinisoti 1440 (DUKE). Guadeloupe: Culberson 14511<br />
(DUKE), Oitestal 1916 (US). Martinique: Citlberson 14682<br />
(DUKE), Uegeliuc (Degelius herbarium). Dominica: Irnshaug<br />
32726 (MSC), <strong>Hale</strong> in <strong>Lichen</strong>es Selecti Exsiccati 942 (H, LD,<br />
US) (for additional records see <strong>Hale</strong> 1971a:lO). St. Lucia:<br />
Inishaug 29712 (MSC). St. Vincent: Znishaug 30504 (MSC).<br />
Grenada: <strong>Hale</strong> 38298, Inishaug 16092, 16335 (MSC). Trini-<br />
dad: <strong>Hale</strong> 38291, Inisliaug 32042 (MSC). Barbados: Elliott<br />
(TUR). \’enezuela: MC-rida, <strong>Hale</strong> 42751, 42941. Brazil: Minas<br />
Gerais, T’ninio in <strong>Lichen</strong>es Brasilienses Exsiccati 675 (TUR)<br />
(di7rributed as Parnielia a~tiazonica).<br />
3. <strong>Parmelina</strong> aurulenta<br />
<strong>Parmelina</strong> aurulenta (‘Tuckerman) <strong>Hale</strong>, 1974:482.<br />
Parmelia auruletzta Tuckerman, 18583424 [type collection:<br />
Harpers Ferry, Virginia, Tuckernia?i (FH-Tuck, lectotype)].<br />
Parmelia tiliacea var. eporescens Miiller Argoviensis,<br />
1887:316 [type collection: Siberia, Russia, Lahm 5 and 6<br />
(G, lectotype: isolectotype in W)].<br />
Pnrnzelia albido-strarninea Hue, 1899:161 [type collection:<br />
Sanctum Dionysium, Reunion, Rodriguez (PC, lectotype)].<br />
Parnielia subrevoluta Harmand, 1928:326 [type collection:<br />
Cha Pon, Indochina, Demnnge (PC, lectotype)].<br />
Pornielia hitnunensis Zahlbruckner, 1930: 187 [type collection:<br />
’ Entries without herharium designations <strong>of</strong> specimens<br />
collected by <strong>Hale</strong> are in US.<br />
Tschangscha, Hunan, China, Hnndel-Mazzetti 11454 (WU,<br />
lectotyie)].<br />
Parnielia sihestris Degelius, 1940:47 [type collection: Togue<br />
Pond near Mt. Katahdin, Maine, Degelius (Degelius her-<br />
barium [not seen]: isolectotype in US)].<br />
Pnrmelia aurulenta var. silrwstris (Degelius) Degelius, 1941 :58.<br />
DEscRIPrIoN.-Thall~is adnate on hark or rock,<br />
pale greenish mineral gray, 4-10 cni broad; lobes<br />
sublinear to subirregular, apically subrotund, 2-4<br />
mni wide, <strong>the</strong> marginal cilia irregularly dispersed<br />
but mostly in <strong>the</strong> lobe axils, to 0.8 mni long; upper<br />
surface shiny, plane to rugulose, soon sorediate to<br />
i)iistLilate-sorecliate, <strong>the</strong> soralia up to 1 mm in diameter,<br />
coalescing into large subcapitate clumps, <strong>the</strong><br />
soretlia coarse (Figure 5a); medulla white and becoming<br />
orange sulfur yellow only beneath soralia<br />
antl near exposed cracks in <strong>the</strong> upper cortex or<br />
entirely sulfur yellow; lower surface black and<br />
moderately to densely rhizinate, <strong>the</strong> rhizines black,<br />
simple or becoming sparsely f urcate or squarrosely<br />
branched. Apo<strong>the</strong>cia rare, adnate to substipitate,<br />
<strong>the</strong> amphi<strong>the</strong>cium sorediate-pListulate, 2-5 mm in<br />
diameter; spores poorly developecl, 7 x 12 pm.<br />
CmxmmY.-Cortex K + yellow (atranorin); medulla<br />
negative with color reagents or if pigmented<br />
areas tested becoming more intensely yellow (zeorin,<br />
leucotylic acid and related terpenes, antl secalonic<br />
acid A).<br />
DIsTRIsvi-Io~.-Panteni~)er~~e antl montane pantropical<br />
(Figure 10).<br />
REMARKs.-<strong>Parmelina</strong> auritlenta is one <strong>of</strong> <strong>the</strong> most<br />
widespread species in <strong>the</strong> genus and, at least in <strong>the</strong><br />
United States, is one <strong>of</strong> <strong>the</strong> most commonly collected<br />
foliose lichens. It is also unusual in being <strong>the</strong><br />
only species <strong>of</strong> <strong>the</strong> subaurulenta complex that occurs<br />
in <strong>the</strong> New World, perhaps suggesting merely<br />
that efficient dispersal by soredia is <strong>the</strong> prime reason<br />
for <strong>the</strong> pantemperate distribution.<br />
Soredial formation is extremely variable. The<br />
lectotype, for example, is densely pustulate with<br />
well developed soredia that obscure most <strong>of</strong> <strong>the</strong><br />
pustular initials (Figure 5a). At o<strong>the</strong>r times, <strong>the</strong><br />
pustules become only sparsely and coarsely sorediate,<br />
this being especially true <strong>of</strong> saxicolous forms.<br />
In no case, however, do <strong>the</strong> pustules remain intact<br />
as in P. leucotyliza, a Japanese species. In Japan <strong>the</strong><br />
two species may also be separated by chemistry<br />
since P. leztcotyliza produces <strong>the</strong> “subaurulenta”<br />
terpenes. There is a very minor amount <strong>of</strong> variability<br />
in chromatograms <strong>of</strong> P. aul-zdenta from different<br />
19
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
FIGURE 10.-Distribution <strong>of</strong> Parmefina aurzrlenta hased on all available herharium specimens.<br />
geographic areas but it does not seem significant or<br />
consistent. I did not, incidentally, recheck <strong>the</strong><br />
terpene pr<strong>of</strong>iles <strong>of</strong> Pawnelia tiliacea var. efiol-esrens,<br />
P. albido-strarninea, P. subrevoluta, or P. hunanen-<br />
sis, but assumed that <strong>the</strong>y have <strong>the</strong> “aurulenta” ter-<br />
penes since <strong>the</strong>y are sorediate and fall outside <strong>the</strong><br />
range <strong>of</strong> P. leucotyliza. Should any <strong>of</strong> <strong>the</strong>m be<br />
found to contain “subaurulenta” terpenes, <strong>the</strong> no-<br />
menclature <strong>of</strong> P. aurulenta would not change.<br />
The parent morph <strong>of</strong> Pal-melina awulenta might<br />
well be P. irl-ugans, which is identical in chemistry.<br />
It would be desirable, however, to examine more<br />
specimens <strong>of</strong> P. irrugans in relation to P. aurulenta<br />
in Japan before making a final decision.<br />
The ecology <strong>of</strong> P. aurulenta has been intensively<br />
studied in Wisconsin, an area <strong>of</strong> typical deciduous<br />
and conifer forests well within its range. For ex-<br />
ample, I found in 1955 that it occurred on 30y0 <strong>of</strong><br />
a 2800 tree sample at <strong>the</strong> 1.3 m level on <strong>the</strong> trunk<br />
and on 21y0 at base level in mesic deciduous forests.<br />
It was most common in closed oak-hickory stands<br />
and much rarer in both open, savanna-like oak<br />
stands and in heavily shaded maple (Acer saccha-<br />
l-urn) climax forests. <strong>Parmelina</strong> aul-ulenta occurs<br />
frequently with Parmelia ruderta Acharius and<br />
Pseudopal-rnelia capel-ata (L.) <strong>Hale</strong>.<br />
In nor<strong>the</strong>rn Wisconsin, an area <strong>of</strong> hardwood-<br />
conifer forests more comparable in climate to<br />
nor<strong>the</strong>rn Europe, P. aurulenta had an overall fre-<br />
quency <strong>of</strong> only Il.670 at <strong>the</strong> 1.3 m level and 1.3%<br />
at <strong>the</strong> base. It avoided Pinus spp. and grew almost<br />
exclusively on Acel- and Querczis (W. Culberson,<br />
1955). This avoidance <strong>of</strong> Pinus was demonstrated<br />
even more graphically by W. Culberson (1958) in<br />
an ecological survey exclusively <strong>of</strong> Pinus in North<br />
Carolina. Although o<strong>the</strong>rwise common in this state,<br />
not a single collection was made on conifers.<br />
SPECI~IESS<br />
Ex.\xIINED.-United States and Canada: See <strong>Hale</strong><br />
(1958384) for a map <strong>of</strong> distribution based on specimens in<br />
various American herbaria. Mexico: Michoacin, Wirth 329<br />
(US). Brazil: Minas Gerais, Eiten 6932 (US). Europe: Russia,<br />
Vasiljeva 9 (US). Tanzania: Tanga Province, Santesson 23358,<br />
23359 (Santesson herliarium, US). Union <strong>of</strong> South Africa:<br />
Satal, Alniboria 8071, 8079 (LD); Transvaal, Alrnborn 7225<br />
(LD). Madagascar: des .4 bboyes (TUR), Lernaitre (H). Paki-
NUMBER 33<br />
FIGURE 11.-Species <strong>of</strong> <strong>Parmelina</strong>: a, P. nmagiensis (Kurokawa 550058 in US); b, P. antillensis<br />
(<strong>Hale</strong> 38291); c, P. aurulentn (<strong>Hale</strong> 18941); d, P. consors (Reit- and Klein 15973 in US); e, P.<br />
crassutn (Kur<strong>of</strong>mwu 550446 in US); 1, P. cryfitochlora (<strong>Hale</strong> 37782). (Scale in mm.)<br />
21
22 SMITHSOSIAN CONTRIBUTIONS TO BOTANY<br />
Stan: Ahmad 211 (L), Iqbnl 831, 839, 843 (US). Nepal: Noikett<br />
9368 (BM). India: Almora District, Arvasthi 3970 (Aivasthi<br />
herbarium, US); Kangra District, Hoeg 1492, 1497 (Awasthi<br />
herbarium); Punjab: Schelpe 3260 (BM); Tamil Natlu: <strong>Hale</strong><br />
43664, 43812, 43816; Uttar Pradesh, Awasthi 3374 (Awasthi<br />
herbarium); !Vest Bengal, Togashi (TNS). Sri Lanka: Degel-<br />
ius As407 (Degelius herbarium), Thwaites 32 (BM). China:<br />
Kiangsi Province, Tai (BM). Taiwan: Kurokawa 1359, 1390,<br />
2694 (TNS). Korea: Den (TNS). Hong Kong: without collec-<br />
tor, 1897 (BM, US); <strong>Hale</strong> 46393. Japan: Prov. Aki, Yan-<br />
agisatca 746-1 (TNS); Prov. Awa, Inobe 23 (TNS); Prov.<br />
Higo, Mayebnra 169 (TNS) Togashi in <strong>Lichen</strong>es Japoniae<br />
Exsiccati 177 (US); Prov. Hizen, Kurokawa 6261.5 (TNS):<br />
Prov. Kii, Kurokawa 59129 (TNS, US), Sasnki 481 (TNS);<br />
Prov. Settsu, <strong>Hale</strong> 29417, 29428; Prov. Shimotsuke, Kiirokaroa<br />
56509 (TNS). Philippines: Benguet, Degeliiis As-888 (Degelius<br />
herbarium). Indonesia: Java, Groenhart 5872 (L, US). New<br />
Guinea: IMorobe District, Weber and McVean L-51280<br />
(COLO, US). Hawaiian Islands: Maui, Faurie 514 (LD), 866<br />
(BM, PC), Hnle 31543; Kauai, <strong>Hale</strong> 31747, 31755.<br />
4. <strong>Parmelina</strong> consors<br />
FIGUKE lld<br />
Pnrmelinci consors (N) lander) <strong>Hale</strong>, 1974:482.<br />
Pnrnielia co?isors Sylander, 1885:613 [type collection: Minas<br />
Gerais, Brazil, Weddell (H, Nylander herbarium number<br />
35277, lectotype)].<br />
Parmelia balansae Miiller Argoviensis, 1888a: 1 [type collection:<br />
AsunciOn, Paraguay, Balansn 8 (G, lectotype)].<br />
Parmelin sampaiana Hue, 1899: 170 [type collection: S%o<br />
Paulo, Brazil, Sampaio (PC, lectotype)].<br />
Parmelia continentalis Lynge, 1914:lll [type collection:<br />
Corumba, Mato Grosso, Brazil, hfalme 48 (S, lectotype)].<br />
DEsCRIPTIoN.-Thallus adnate on bark, coria-<br />
ceous greenish mineral gray and turning deep olive-<br />
buff in <strong>the</strong> herbarium, 5-10 cm in diameter; lobes<br />
irregularly branched, sublinear-elongate to subir-<br />
regular, apically rotund, <strong>of</strong>ten imbricate, 1-4 mm<br />
wide, <strong>the</strong> margins crenate, moderately to densely<br />
ciliate, <strong>the</strong> cilia stout, blackish brown to black,<br />
becoming furcate, 0.2-1 .O mm long; upper surface<br />
shiny, becoming heavily maculate, sometimes<br />
pruinose towards <strong>the</strong> tips, isidia and soredia<br />
lacking; medulla white; lower surface black, mod-<br />
erately rhizinate, <strong>the</strong> rhizines black, thick and thin<br />
intermixed, simple or squarrose. Apo<strong>the</strong>cia substi-<br />
pitate, 2-8 mm in diameter, amphi<strong>the</strong>cium smooth,<br />
<strong>the</strong> disc cinnamon brown, rarely perforate; spores<br />
8,S-lZ X 14-19 pm.<br />
CHEMISTRY.- Cortex K+ yellow (atranorin);<br />
medulla negative with color tests (no substances<br />
demonstrated).<br />
DIsTRIBuTroN.-South America.<br />
REhfARKs.-This robust, coriaceous lichen is char-<br />
acterized by its heavily white-maculate cortex (Fig-<br />
ure 3a), thick, furcate cilia (Figure 3a), and lack <strong>of</strong><br />
medullary chemistry. It is also <strong>the</strong> only species as-<br />
signed to <strong>Parmelina</strong> with perforate apo<strong>the</strong>cia, a<br />
trait usually associated with Parmotrema hfassa-<br />
longo. The dense rhizine mat below to <strong>the</strong> margin<br />
and <strong>the</strong> generally close adnation, however, do not<br />
conform to Parmotrema. Paymelina consors is evi-<br />
dently a common lichen in dry, scrubby forests and<br />
on trees along roads. The presumptive sorediate<br />
morph, P. pilosa, has a broader distribution in<br />
South America and also occurs in Africa.<br />
SPECIMENS Ex4MIsEo.-Brazil: WawrU 669 (M); Mato Groqso,<br />
Monte\ 10146 (DUKE, LD, US), Minaq Gerais, hfalme 201B<br />
(S), I’ainio in <strong>Lichen</strong>es Brasilienses Exsiccati 398 (BM, M,<br />
TUR); Rio de Janeiro, Burchell 2303 (BM), Gardner 37<br />
(BM), Glnziou 1821, 1836 (M), 1840 (M, S), Hemmendorff<br />
5386 (S, UPS), Milne (BM); Rio Grande do Sul, Malme 1282<br />
(LD, S, UC, UPS, US); Santa Catarina, Reitz and Klein<br />
12901, 15973, 16032a (US): SZo Paulo, Eiteti 5731 (US), Gehlt<br />
(US), 5924 (MICH), Puiggari (G). Paraguay: Itapua, Malme<br />
1462 (S, UPS, US). Uiuguay: Latellaja, Lamb (H). Argentina:<br />
Buenos Aires, Kuhnemann 22 (S), Santesson 76, 77a (S);<br />
Misiones, Montes 36 (US), 3353 (WIS).<br />
5. <strong>Parmelina</strong> crassata, new species<br />
FIGURE lle<br />
DEscRIPTIoN.-Thallus laxe adnatus, corticola,<br />
crassatus coriaceusque, cinereo-albidus vel obscurascens,<br />
usque ad 15 cm latus, lobis sublinearibus<br />
vel subirregularibus, contiguis vel congestis, 3-6<br />
mm latis, margine ciliatis, ciliis irregulariter dispersis,<br />
nigris, 0.3-0.8 mm longis, superne planus vel<br />
aetate rugulosus, nitidus, continuus, isidiis sorediisque<br />
destitutus, cortex superior ca. 12 pm crassus,<br />
epicorticatus, stratum gonidale 16-18 pm crassum,<br />
medulla omnino sulphureo-salmonea, 160-2 10 pm<br />
crassa, cortex inferior fuscus, 14-16 pm crassus,<br />
subtus niger, dense rhizinosus, rhizinis nigris, nitidis,<br />
1-2 mm longis, simplicibus vel pro parte<br />
furcatis vel squarroso-ramosis. Apo<strong>the</strong>cia numerosa,<br />
subpedicillata, ainphi<strong>the</strong>cio rugoso, usque ad 10<br />
mm diamentro, sporis octonis, 6-8 X 14-15 pm.<br />
CHEnmTR\r.-Cortex K + yellow (atranorin);<br />
medulla more intensively yellow with color tests<br />
(zeorin, leucotylin and associated terpenes, and<br />
secalonic acid A).
NUMBER 33<br />
HoLOTYPE.--;\ft. Akagi, Japan, S. Kzirokazun<br />
550466, 23 August 1955 (US; isotype in TNS).<br />
DIsTRrBLJTr0N.-Japan.<br />
REivARKs.-This is ano<strong>the</strong>r member <strong>of</strong> <strong>the</strong> P.<br />
sitbaic~iilenfa group so abundantly developed in<br />
Japan. It has <strong>the</strong> same chemistry as P. subazirulenta<br />
but is well separated by <strong>the</strong> larger, lea<strong>the</strong>ry thallus,<br />
averaging 218 pm in thickness (sample <strong>of</strong> 10 speci-<br />
mens), as opposed to 150 pm for P. subaurzilenta.<br />
It also has significantly larger, substipitate apo<strong>the</strong>-<br />
cia. Parmelia in-ugans, which has large apo<strong>the</strong>cia,<br />
can be distinguished by chemistry (<strong>the</strong> “aurulenta”<br />
terpenes) and thinner thallus.<br />
This species previously had been identified<br />
(along with P. irrugans) as “Parmelia homogenes”<br />
in Japan, but, as explained below, P. homogenes<br />
was misinterpreted and is in reality <strong>the</strong> same as<br />
P. .siibnicrulentn, Puvmelia cmssatn is a common<br />
species in Japan, but since I have not had an<br />
opportunity to re-examine tlie chemistry <strong>of</strong> <strong>the</strong> rich<br />
collections in TNS, I have cited below only those<br />
specimens also represented in US.<br />
SIV CIMFX Ex \~is~o.--Japaii: Prov. Kozuke, Kurokarc~a<br />
.58583 (TNS, US) ; Prov. Sagami, Kurokarca 58039 (TNS, US) :<br />
I’rov. Suruga, ‘4saliina 539, 2G27 (US).<br />
6. <strong>Parmelina</strong> cryptochlora<br />
FIGURE llf<br />
PorrridiTia crjptochloro (Vainio) <strong>Hale</strong>, 19743482.<br />
Pnrrnelin crjptoclilorn 1-ainio, 1896:34 [type collection: Lau-<br />
dat, Dominica. Elliott 912 (BM, lectot!pe)].<br />
DEscRIPTIos.-Thallus closely adnate on bark,<br />
whitish mineral gray, 1-3 cm broad; lobes sublinear,<br />
separate to crowded, 1-2 mm wide, <strong>the</strong> marginal<br />
cilia sparsely developed, 0.1-0.3 mm long;<br />
upper surface plane to convex, sorediate toward <strong>the</strong><br />
tips, tlie soralia capitate, up to 1 mm in diameter<br />
(Figure 5b), <strong>the</strong> sorediate lobe tips becoming<br />
revolute: lower surface black, sparsely rhizinate, tlie<br />
rliizines black, simple. Apotliecia not seen.<br />
CHEhIISTRY.-COrteX K + yellow (atranorin):<br />
medulla K-, C+ rose, KC+ red, P- (gyroplioric<br />
acid).<br />
DISTRIBUTIOS.-~YeSt Indies and India (?).<br />
REMARKS.--AS I explained in my study <strong>of</strong> <strong>the</strong><br />
Pawneline <strong>of</strong> Dominica (<strong>Hale</strong>, 1971a: 12), Vainio’s<br />
type collection is so poor that I had not been able<br />
to interpret it until I collected additional speci-<br />
mens from <strong>the</strong> type-locality in Dominica. It is<br />
ra<strong>the</strong>r common at one locality (base <strong>of</strong> Morne<br />
Anglais) on trees in a pasture at 600-800 m elevation,<br />
One collection that I made in South India<br />
also appears to lie this species. It is so small and<br />
inconspicuous that it has undoubtedly been overlooked<br />
elsewhere or, iC collected, has been set aside<br />
as unidentifiable. It has <strong>the</strong> same chemistry as<br />
P. dissecta, a common species with which it is<br />
clearly allied as <strong>the</strong> sorediate morph <strong>of</strong> <strong>the</strong> extinct<br />
or still undiscovered parent morpli. It also has <strong>the</strong><br />
same chemistry as sorediate Hypotl-achynn l-evoliita<br />
(Floerke) <strong>Hale</strong>, which woulcl normally be amply<br />
distinguished by a larger tliallus, more ascending<br />
lobes, dichotomously branched rhizines, and<br />
broader soretliate pustules. A poorly developed<br />
specimen <strong>of</strong> H. iavolicta, however, and a large specimen<br />
<strong>of</strong> P. c~pfochlol-a would have to be separated<br />
with extreme care.<br />
QI’ECIMFNS ExA\rINl.o.-~ominic;i: <strong>Hale</strong> 3535i, 37782. India:<br />
Tamil Nadu, <strong>Hale</strong> 40203.<br />
7. <strong>Parmelina</strong> damaziana, new combination<br />
FIGURE 120<br />
Partrielin danzaziatia Zahlbruckner, 1905:.541 [type collection:<br />
Mt. Ituculumi, Brazil, Daniazio 137j (W, lectotype; isolectotype<br />
in G)].<br />
Partnelin brachpconidia Zahlbruckner, 1908:465 [type collection:<br />
\‘ellozo, Serra (lo Our0 Preto, Braiil, Daniazio 1741<br />
(LV, lectot!pe)].<br />
Pnririelin brnchyrovidin var. cliloroccirpn Zahlbruckner,<br />
1908:466 [type collection: Cachoeira do Campo, Brazil,<br />
Dnmnzio I740 (W, lectotype)].<br />
Pnrtnelici crystallorum Lynge, 1914: 128 [t\pe collection: Corcovado,<br />
Rio de ,janeiro, Brazil, Mnlme 59* (S, lectotype)].<br />
D~sc~~~l~o~.-Thallub<br />
closely adnate on twigs,<br />
whitish ashy mineral gray, 3-6 cm broad; lobes<br />
short, sublinear-elongate to subirregular, 1-3 mm<br />
wide, tlie marginal cilia evenly dispersed, about 0.5<br />
mm long; upper surface plane, shiny; lower surface<br />
Black, moderately rhizinate, <strong>the</strong> rhizines simple,<br />
black. Apo<strong>the</strong>cia common, sessile to substipitate, to<br />
12 mm in diameter, <strong>the</strong> disc flat, <strong>of</strong>ten radially split;<br />
spores 8,s-12 X 12-18 pm.<br />
CHEmsTRY.-Cortex K + yellow (atranorin);<br />
medulla K-, C-, KCf rose, P- (“horrescens”<br />
unknown).<br />
DIsrRIBuTIo,v.-So~itli !imerica.<br />
RE~r,~RIts.-AlthoUgll clearly distinct because <strong>of</strong><br />
23
24 SSIITHSONIAN CONTRIBUTIONS TO BOTANY<br />
FIGURE<br />
12.4pecies <strong>of</strong> <strong>Parmelina</strong>: a, P. damaziana (Dainazio 1375 in W); b, P. degelii (Degelius<br />
A-I82 in Degelius herbarium); c, P. de?iegans (<strong>Hale</strong> 43855); d, P. dissecta (<strong>Hale</strong> 14999); e, P.<br />
endoleuca (Weber in <strong>Lichen</strong>es Exsiccati 244 in US): f, P. enorinis (Jellicoe in US). (Scale in mm.)
NUMBER 33<br />
<strong>the</strong> chemical constitutents, P. damaziana is a rare<br />
and not well comprehended species. The lectotype<br />
was collected on tree branches whereas Lynge’s<br />
species was apparently saxicolous. Burchell 1105-06<br />
is tentatively placed here since <strong>the</strong> chemistry, while<br />
not clear, is closest to <strong>the</strong> “horrescens” type. Parme-<br />
lina damaziana has ra<strong>the</strong>r large spores, just as <strong>the</strong><br />
o<strong>the</strong>r members <strong>of</strong> <strong>the</strong> P. horrescens group. It could<br />
be regarded <strong>the</strong>oretically as <strong>the</strong> parent morph for<br />
P. howescens, P. schindleria, and P. subfatiscens, but<br />
all <strong>of</strong> <strong>the</strong>se vegetative morphs have smaller, more<br />
fragile thalli.<br />
SPECIXIENS ExAalrNED.-BraLil: Burchell 1105-06 (BM, US).<br />
8. <strong>Parmelina</strong> degelii, new species<br />
FIGURE 12h<br />
DEscRIPrroN.-Thallus adnatus, corticola, fragilis,<br />
pallide viridi-albicans, 6-8 cm latus, lobis plus<br />
minusve subirreqularibus, 2-3 mm latis, margine<br />
aetate lobulato-dissectis, ciliatis, ciliis usque ad 0.8<br />
mm longis, superne planus vel aetate rugulosus,<br />
nitidus, sparse vel modice isidiatus, isidiis primum<br />
cylindricis, mox procumbentibus, dorsiventralibus,<br />
expansis, ad 0.6 mm longis, cortex superior 12 pm<br />
crassus, epicorticatus, stratum gonidiale 12-14 pm<br />
crassum, medulla pallide sulfurea, 60-90 pm<br />
crassa, cortex inferior paraplectenchymatus, brunneus,<br />
12 p,m crassus, subtus niger, dense rhizinosus,<br />
rhizinis nigris, simplicibus, 0.5-09 mm longis.<br />
Apo<strong>the</strong>cia ignota.<br />
CHEMIsTRY.-Cortex K + yellow (atranorin);<br />
medulla more intensively yellow with color tests<br />
(zeorin, leucotylic acid, apparently secalonic acid<br />
A, and traces <strong>of</strong> o<strong>the</strong>r pigments and terpenes).<br />
HoLoTYPE.-Rain forest, Cairns Cove, Queensland,<br />
Australia, elevation 600 m, G. Degelius A-I82<br />
(Degelius herbarium; isotype in US).<br />
DI~TRIRUTI~N.-A~S traha.<br />
REMARKS.-This is <strong>the</strong> only species <strong>of</strong> section<br />
Myelochroa that has been discovered in Australia.<br />
It is remarkable that such a conspicuous lichen<br />
should be collected so recently in Queensland. The<br />
terpene chemistry is close to P. aurulenta except<br />
for <strong>the</strong> absence <strong>of</strong> one spot (Figure 7). Regardless<br />
<strong>of</strong> chemistry, however, it is distinct from all o<strong>the</strong>r<br />
species in <strong>the</strong> P. subaurulenta complex because <strong>of</strong><br />
<strong>the</strong> production <strong>of</strong> large lobulate isidia. <strong>Parmelina</strong><br />
degelii is named in honor <strong>of</strong> Dr. Gunnar Degelius,<br />
who has enriched our knowledge <strong>of</strong> lichen distribu-<br />
tions through his careful collecting efforts in <strong>the</strong><br />
tropics.<br />
9. <strong>Parmelina</strong> denegans<br />
FILLRF 12C<br />
Parmelzna denegans (Nylander) <strong>Hale</strong>, 1974,482.<br />
Parnelta denegans Njlander, 1900 6 [tjpe collection: Ram-<br />
podde, Ceylon, Almquzst (H, Njlander herbarium number<br />
35129, lectotype; isolectotype in S)].<br />
DE~CRIPTI~X .-Thallus adnate to closely adnate<br />
on bark, light greenish mineral gray, 5-9 cm broad;<br />
lobes irregularly branched, sublinear, imbricate, 1-3<br />
mm wide, <strong>the</strong> marginal cilia irregularly dispersed,<br />
0.4-0.7 mm long; upper surface shiny, weakly maculate,<br />
plane but soon developing small pustulate<br />
areas, <strong>the</strong> pustules eventually breaking open<br />
apically and forming coarse soredia; medulla pale<br />
orange-salmon, in part K+ purple; lower surface<br />
black, densely rhizinate, <strong>the</strong> rhizines black, simple<br />
to sparsely branched. Apo<strong>the</strong>cia sessile, 2-3 mm in<br />
diameter; spores 8, 5-7 X 7-9 p.m.<br />
CHEhmTRY.-Cortex K + yellow (atranorin);<br />
medulla more intensively yellow-orange with color<br />
tests (zeorin, leucotylin and associated terpenes,<br />
secalonic acid A, and possibly anthraquinones).<br />
DIsTRIsuTIoN.-India, Sri Lanka, and Sabah.<br />
REMARKS.-Except for <strong>the</strong> darker reddish orange<br />
pigment in <strong>the</strong> lower part <strong>of</strong> <strong>the</strong> medulla, this species<br />
is <strong>the</strong> pustulate morph <strong>of</strong> P. subaurulenta.<br />
They have similar lobe configuration and habitats,<br />
occurring at about 2000 m elevation in open forests.<br />
The lectotype specimen is fertile and has few<br />
pustules. The specimens that I collected are heavily<br />
pustulate-sorediate by contrast.<br />
SPECIMENS ExAM1xFD.-India: Tamil Nadu, Foreau 4128<br />
(Awa3thi herbarium, US), <strong>Hale</strong> 40239, 43636, 43762, 43855.<br />
Sri Lanka: Thruaites (UPS). Sabah: <strong>Hale</strong> 29020.<br />
10. <strong>Parmelina</strong> dissecta<br />
FIGCJRE 12d. 13<br />
<strong>Parmelina</strong> dissecta (Nylander) <strong>Hale</strong>, 1974:482.<br />
Parmelia dissecta h’ylander, 18823451 [type collection: Fon-<br />
tainebleau, France, A’ylander (H, Nylander herbarium<br />
number 35131, lectotype; isolectotype in PC)].<br />
Parinelia laeuigata var. gracilis f. furfuracea Miiller Argovien-<br />
25
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
sis, 1888c:529 [type collection: Faxina, Brazil, Puiggari 47<br />
(G, lectotype)].<br />
Parmelia minarum Vainio, 1890:48.<br />
Parmelia amazonica var. hwnotii Hue, 1899:158.<br />
Parmelia puiggari Gyelnik, 1931:288 [based on Parmelia<br />
lamigata var. gracilis f. furfuracea Muller Argoviensis].<br />
Parmelia camtschadalis var. epiphylla Cengia Sambo,<br />
1938:379.<br />
Parmelia hubrichtii Berry, 1941:102.<br />
[For full citation <strong>of</strong> synonymy, see <strong>Hale</strong>, 1971a:6.]<br />
DEScRIPTIoN.-Tha1Ius adnate on bark or rock,<br />
yellowish glaucous to pale greenish mineral gray,<br />
3-7 cm broad: lobes sublinear-elongate, contiguous,<br />
1-3 mm wide, <strong>the</strong> marginal cilia irregularly dis-<br />
persed, mostly simple, to 0.7 mm long: upper sur-<br />
face shiny, emaculate, plane to convex, moderately<br />
to densely isidiate, <strong>the</strong> isidia cylindrical, erect,<br />
<strong>of</strong>ten branched, less than 0.5 mm high; medulla<br />
white; lower surface black, moderately rhizinate,<br />
<strong>the</strong> rhizines black, shiny, simple or sparsely<br />
branched. Apo<strong>the</strong>cia adnate, <strong>the</strong> rim crenate, <strong>the</strong><br />
amphi<strong>the</strong>cium isidiate: 1-4 mm in diameter: spores<br />
8,8-10 X 12-17 pm.<br />
CHEMIsmY.-Cortex K+ yellow (atranorin);<br />
medulla K-, C+ rose, KC+ red, P- gyrophoric<br />
acid with or without associated unknown sub-<br />
stances).<br />
DIsTRIBuTI0N.-Pantemperate and montane pan-<br />
tropical (Figure 13).<br />
REMAws.-This is one <strong>of</strong> <strong>the</strong> most widespread<br />
and commonly collected species in <strong>the</strong> genus,<br />
especially in temperate zones, occurring on a wide<br />
variety <strong>of</strong> substrates in open, secondary forests. In<br />
tropical regions it is strictly montane, usually being<br />
most abundant in cloud forests up to 2300 m eleva-<br />
tion. Morphological variation is wide, even though<br />
<strong>the</strong> basic characters, isidia, ciliate lobes, and gyro-<br />
phoric acid, are constant. The lobes, for example,<br />
are narrow, sublinear, and separate in <strong>the</strong> lectotype<br />
specimen. At <strong>the</strong> o<strong>the</strong>r extreme are plants such as<br />
those collected in Dominica (<strong>Hale</strong>, 1971a) with<br />
broader, contiguous to subimbricate lobes.<br />
FIGURE 13.-Distribution <strong>of</strong> <strong>Parmelina</strong> dissecta based on all available herbarium specimens.
NUMBER 33<br />
<strong>Parmelina</strong> dissecta has been correctly identified<br />
by European lichenologists both on <strong>the</strong> Continent<br />
and in tropical regions, Degelius (1941:60) was <strong>the</strong><br />
first to identify <strong>the</strong> species correctly in North<br />
America. He noted two populations differing in C<br />
test (C+ rose or C-) and presence or absence <strong>of</strong><br />
cilia on <strong>the</strong> isidia. <strong>Hale</strong> and Kurokawa (1962)<br />
determined that <strong>the</strong> C- ciliate population is P.<br />
horrescens. No parent morph has been discovered<br />
for P. dissecta, but it forms a convenient nucleus<br />
<strong>of</strong> <strong>the</strong> “dissecta” group, including isidiate but sig-<br />
nificantly larger P. melanochaeta, sorediate P. cryp-<br />
tochlora, and isidiate-lobulate P. subfatiscens (Fig-<br />
ure 9).<br />
SPECIMENI EXAMINED (selected).-United States: Pennsylvania,<br />
<strong>Hale</strong> 17140; Marjland, <strong>Hale</strong> 14367; West Virginia,<br />
<strong>Hale</strong> 11470: Ohio, <strong>Hale</strong> 13910, 15735; Kentucky, <strong>Hale</strong> 13770;<br />
I’irginia. <strong>Hale</strong> 38419, Liittrell 3799 (US); Tennessee, <strong>Hale</strong><br />
31124, 37038; North Carolina, Culberson 7099 (DUKE), <strong>Hale</strong><br />
30645; South Carolina, Culberson 7459 (DUKE), <strong>Hale</strong> 16601;<br />
Alabama, <strong>Hale</strong> 33937, Skorepa 4547 (US); Georgia, Culberson<br />
7249 (DUKE), <strong>Hale</strong> 7509; Florida, Nelson (US) (see<br />
Moore, 1968:220 for additional records in Florida); Mississippi,<br />
<strong>Hale</strong> 7977, Hubricht 1532 (US): Texas, <strong>Hale</strong> 5242.<br />
Mexico: Chiapas, <strong>Hale</strong> 20304; \‘era Cruz, <strong>Hale</strong> 21231. Panama:<br />
Chiriqui, <strong>Hale</strong> 38879. Cuba: Oriente, Imshaug 24720<br />
(MSC). Jamaica: Inishaug 14172 (MSC). Guadeloupe: Dim<br />
1030 (TUR). Dominica: <strong>Hale</strong> 35569 (see <strong>Hale</strong>, 1971a:13, for<br />
additional records). Martinique: Degeliits (Degelius herbarium).<br />
St. T’incent: Guilding 46 (BM). Trinidad: Imshaug<br />
32103 (MSC). Colombia: Antioquia, Nee and Mori 4258 (US).<br />
l’enezuela: Distrito Federal, Dennis 1568 (BM); Mkrida, <strong>Hale</strong><br />
42543; TAchira, <strong>Hale</strong> 42540, 45112. Brazil: Minas Gerais,<br />
Eiten 6867 (US); Parana, Montes 10121 (hIVM). Europe:<br />
Portugal, Persson (UPS), Sampaio in <strong>Lichen</strong>es de Portugal<br />
252 (LD, US), Tauares 97 (H, LD, WIS), 632 (US); Spain,<br />
Degelius (S), Santesson 19428 (UPS), Schauer in <strong>Lichen</strong>es<br />
Selecti Exsiccati 265 (LD, US); France, des Abbayes in<br />
<strong>Lichen</strong>es Armoricani Spectabiles Exsiccati 88 (LD, W), Hasselrot<br />
(S); Italy, Gresino (S), Sbarbaro (US). Kenya: Rift<br />
Valley Province, Maas Geesteraniis 6067 (L). Tanzania: Tanga<br />
Prov., Santesson 23137 (UPS). Union <strong>of</strong> South Africa, Cape<br />
Province, Almborn 301, 670, 3201 (LD); Natal, Almborn<br />
9192, 9525 (LD), Hoeg (TRH). Swaziland: Almborn 7904<br />
(LD). India: Tamil Nadu, Degelius As-238 (Degelius herbarium),<br />
<strong>Hale</strong> 40222, 43715, 43852; West Bengal, Degelius As-<br />
21 1 (Degelius herbarium). Sri Lanka: Degelius As-466 (Degelius<br />
herbarium, US). Malay: Pahang, <strong>Hale</strong> 30137, 30483.<br />
Taiwan: Kurokawa 2400 (TNS). Japan: Prov. Aki, <strong>Hale</strong><br />
29368: Prov. Higo, Mayebara 1.58 (TNS); Prov. Kazusa, Kurokawa<br />
64096 (TNS); Prov. Ohmi, <strong>Hale</strong> 29484; Prov. Owari,<br />
Asahina 146 (TNS); Prov. Settsu, <strong>Hale</strong> 29411. Philippines:<br />
Mountain Province, Degelius As-% (Degelius herbarium),<br />
<strong>Hale</strong> 26040. Indonesia: Java, Groenhart 6022 (L), 8483<br />
(BOR); Sumatra, Groenhart 967 (L). Hawaii: <strong>Hale</strong> 31340,<br />
32933.<br />
11. <strong>Parmelina</strong> endoleuca, new combination<br />
FIGURE 12e<br />
Parmelia endole.uca TaFlor, 1847: 167 [type collection: Swan<br />
Rixer, Australia, Drurnmond (FH-Tajl, lectotjpe)].<br />
DEscRIP’rIos.-Thallus closely adnate on bark,<br />
whitish mineral gray, 2-4 cm broad; lobes sublinear<br />
to subirregular, short and crowded, 1-2 mm wide,<br />
<strong>the</strong> marginal cilia sparse, less than 0.3 mm long;<br />
upper surface shiny, plane to minutely rugulose,<br />
becoming more strongly rugose and lobulate toward<br />
<strong>the</strong> center, heavily pycnidiate, soredia and isidia<br />
lacking; medulla white: lower surface dark brown<br />
at <strong>the</strong> margins and black toward <strong>the</strong> center, mod-<br />
erately rhizinate, <strong>the</strong> rhizines simple, brown but<br />
blackening at maturity. Apo<strong>the</strong>cia very numerous,<br />
sessile, 1-2.5 mm in diameter, <strong>the</strong> disc dark brown,<br />
plane; spores 8, 6-7 X 11-12 pm.<br />
CHExmrRY.--Cortex K + yellow (atranorin);<br />
medulla negative with color tests (an unidentified<br />
fatty acid).<br />
DIsTRIsuTIoN.-Austra~ia.<br />
REalARKs.-In my earlier work on Parmelia I<br />
placed this species in synomymy under <strong>Parmelina</strong><br />
galbina, a Japanese-North American species. Since<br />
<strong>the</strong> Australian locality seemed at variance with <strong>the</strong><br />
Arcto-Tertiary distribution pattern <strong>of</strong> P. galbina<br />
and since my first chemical tests had been done<br />
with crystal techniques, I have checked <strong>the</strong> type <strong>of</strong><br />
Parmelia endoleiica again and found that it con-<br />
tains nei<strong>the</strong>r galbinic acid nor any <strong>of</strong> <strong>the</strong> associated<br />
terpenes that characterize P. galbina. I have con-<br />
cluded, <strong>the</strong>refore, that Parmelia endoleuca is a good<br />
species in spite <strong>of</strong> <strong>the</strong> very close external resem-<br />
blance to P. galbina. The fatty acid falls higher on<br />
<strong>the</strong> chromatographic plates than ei<strong>the</strong>r caperatic<br />
acid or protolichesterinic acid. The species is<br />
endemic to <strong>the</strong> dry scrub forests <strong>of</strong> eastern<br />
Australia.<br />
SPECIVENS ExA\rIN~.o.-Australia: Australian Capital Terri-<br />
tory, Weber in <strong>Lichen</strong>es Exsiccati 244 (COLO, US).<br />
12. <strong>Parmelina</strong> enormis<br />
FIGURE 12f<br />
<strong>Parmelina</strong> enormis (<strong>Hale</strong>) <strong>Hale</strong>, 1974:482.<br />
Parmelia enormis <strong>Hale</strong>, 1972c:344 [type collection: Nyaka<br />
Plateau, Zambia, Jellicoe, September 1968 (BM, holotype;<br />
isotype in US)].<br />
27
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
DESCRIPTIoN.-Thahs expanded, loosely attached<br />
on rock, coriaceous, whitish to ivory mineral gray,<br />
10-30 cm broad; lobes sublinear, dichotomously<br />
branched, imbricate and crowded toward <strong>the</strong> cen-<br />
ter, 5-8 mm wide, <strong>the</strong> margins entire, ciliate mostly<br />
in <strong>the</strong> lobe axils, <strong>the</strong> cilia to 2.0 mm long; upper<br />
surface plane to convex, continuous, lacking soredia<br />
and isidia; medulla white; lower surface pale<br />
brown, densely rhizinate, <strong>the</strong> rhizines pale brown,<br />
simple. Apo<strong>the</strong>cia numerous, subpedicillate, 3-8<br />
mm in diameter; spores 8, 6-7 x 8-11<br />
CHEMIsTRY.-Cortex K + yellow (atranorin);<br />
medulla I
NUMBER 33<br />
FIGURI: 14.--Species <strong>of</strong> Parmeliiza: a, P. ento<strong>the</strong>iochroa (<strong>Hale</strong> 29456a); b, P. expollida (Kurokair-a<br />
2930 in TKS); c, P. gnlbina (Hole 2341.5); tl, P. hoyachitzensis (Kurokawa 67081 in TNS): e,<br />
P. hetemloba (Schiflner in \\'); f, P. liorrescens (Hnle 36999).<br />
29
90 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
CHEMIs-rRY.-Cortex K + yellow (atranorin);<br />
medulla negative with color tests (protolichesterinic<br />
acid and an unidentified fatty acid).<br />
DIsmInuTIoN.-India, Thailand, and Taiwan.<br />
As pointed out by Kurokawa (1968a:191), an<br />
unusual feature <strong>of</strong> this species is <strong>the</strong> brown lower<br />
surface, indicating some relationship with P. versi-<br />
forrnis, a South American species. It is also one <strong>of</strong><br />
<strong>the</strong> few Parrnelinae to produce a fatty acid.<br />
SrEcIarENs EXAMINF‘.D.-See Kurokawa (1968a:193) for records<br />
from India, Taiwan, and Japan.<br />
15. <strong>Parmelina</strong> galbina<br />
FIGURES 14c, 15<br />
<strong>Parmelina</strong> galbina (Acharius) <strong>Hale</strong>, 1974:482.<br />
Parmelia galbina Acharius, 1814: 195 [type collection: North<br />
America (?Pennsylvania), Muhlenberg (H-Ach, lectotype;<br />
isolectotype in PH)].<br />
Parmelia tiliacea var. minor Muller Argoviensis, 1877:X<br />
[type collection: near Dallas, Texas, Boll (G, lectotype;<br />
isolectotypes in FH, M, W)].<br />
Parmelia tiliacea var. sulphurosa Tuckerman, 1882:57 [type<br />
collection: Illinois, Hall (FH-Tuck, lectotype)].<br />
Parmelia deminuta Hue, 1899:156 [type collection: Texas,<br />
Boll (PC, lectotype)].<br />
Parmelia subquercifolia Hue, 1899: 157 [type collection: Ohio,<br />
Sullivant (PC, lectotype)].<br />
Parmelia subquercifolia var. rugosa Hue, 1899: 175 [type col-<br />
lection: Oyama, Japan, Faurie (not seen)].<br />
Parmelia tiliacea subquercifolia (Hue) Merrill and Burnham<br />
in Burnham, 1922:75 [rank not designated].<br />
Parmelia quercina var. sulphurosa (Tuckerman) Zahlbruck-<br />
ner, 1929:192.<br />
Parmelia sulphurosa (Tuckerman) Fink, 1935:328.<br />
Parmelia laeuigata ssp. extremi-orientalis f. rugosa (Hue)<br />
Asahina, 1951c:291.<br />
Parmelia subquercifolia f. subradiata Asahina, 1952:98 [type<br />
collection: base <strong>of</strong> Mt. Fuji, Japan, Yamamoto (TNS, lecto-<br />
type) (nomen nudum in Asahina, 1951c:291)].<br />
Parmelia galbina var. rugosa (Hue) Asahina, 1963:225.<br />
Parmelia galbina var. subradiata (Asahina) Asahina, 1963:225.<br />
DEscRIPTIoN.-Thallus closely adnate on bark,<br />
yellowish-glaucous to greenish mineral gray, 3-10<br />
cm in diameter; lobes short, sublinear, contiguous,<br />
1-2 mm wide, marginal cilia mostly in axils, some-<br />
times only sparsely developed, less than 0.5 mm<br />
long; upper surface plane to rugulose, continuous,<br />
pycnidia usually numerous; medulla white or pale<br />
yellow to orange, especially beneath <strong>the</strong> apo<strong>the</strong>cia;<br />
lower surface densely rhizinate. <strong>the</strong> rhizines black,<br />
simple. Apo<strong>the</strong>cia very common, adnate, 2-5 mm in<br />
diameter, <strong>the</strong> disc cinnamon brown; spores 8, 7-9 X<br />
10-13 pm.<br />
CHEMIsrRY.-Cortex K + yellow (atranorin);<br />
medulla K+ yellow, C- or C+ yellow, KC-,<br />
P + pale orange (galbinic acid, zeorin, secalonic<br />
acid A, trace <strong>of</strong> salazinic acid, and leucotylin and<br />
associated terpenes).<br />
DIsTRIsvTIo;v.-Japan and eastern North America.<br />
REMARKs.-During a visit to <strong>the</strong> Academy <strong>of</strong><br />
Natural Sciences in Philadelphia in 1958 I found a<br />
specimen in <strong>the</strong> hfuhlenberg herbarium without a<br />
name but with a penciled number “15.2.” I later<br />
found <strong>the</strong> same plant in <strong>the</strong> Acharian collection<br />
(H) also numbered 15.2 and described by Acharius<br />
as Parrnelia galbina. This discovery clarified <strong>the</strong><br />
status <strong>of</strong> one <strong>of</strong> <strong>the</strong> very common corticolous lichens<br />
in eastern North America. W. Culberson (1961: 173)<br />
subsequently studied <strong>the</strong> problem and lectotypified<br />
P. subquercifolia Hue as synonymous with P. gal-<br />
bina. The second syntype <strong>of</strong> Hue’s species was<br />
determined by Culberson to be Parmelia livida<br />
(= Hypotrachyna livida (Taylor) <strong>Hale</strong>), ano<strong>the</strong>r<br />
very common corticolous lichen in eastern North<br />
America distinguished by more or less distinctly<br />
dichotomously branched rhizines and <strong>the</strong> lividic<br />
acid complex (<strong>Hale</strong>, 1975a:45). As a rule, H. livida<br />
has a whiter cast than P. galbina, is more robust,<br />
and lacks pigments in <strong>the</strong> medulla or under <strong>the</strong><br />
apo<strong>the</strong>cia. Still, care must be taken when identi-<br />
fying <strong>the</strong>se two species. One o<strong>the</strong>r distinguishing<br />
character is <strong>the</strong> unique moniliform cells in <strong>the</strong><br />
medulla <strong>of</strong> P. galbina and its morphs, P. hayachin-<br />
ensis, P. metarevoluta, and P. obsessa, first described<br />
by Asahina (1951~). The cells, however, are not<br />
easy to find and <strong>the</strong>ir exact significance is unknown<br />
at this time.<br />
SPECIMENS EXAMINED.-United States: See W. Culberson<br />
(1961:lil) for a discussion <strong>of</strong> <strong>the</strong> distribution and map <strong>of</strong><br />
localities in North America based on specimens in DUKE<br />
and US (Figure 15). Japan: Prov. Aki, Kurokawa 64431<br />
(TXS); Prov. Bungo, Kurokawa 63191 (TNS); Prov. Hizen,<br />
Kurokawa 62588 (TNS); Prov. Hoki, Yasuda (TNS); Prov.<br />
Hyogo, Tagawa L15 (TNS); Prov. Kozuke, Degelius As-1072<br />
(Degelius herbarium); Prov. Mikawa, Takaki 353 (TNS):<br />
Prov. Musashi, Kurokawa 64279 (TNS); Prov. Mutsu, Kuro-<br />
kawa 550385 (TNS); Prov. Shimotsuke, Kurokawa 64057<br />
(TNS); Prov. Shinano, Kurokawa 51758 (TNS, US), 59179<br />
(TNS); Prov. Totomi, h’akanishi 27 (Kobe University); Prov.<br />
Ugo, Suzuki 272 (TNS).
NUMBER 33 31<br />
FIGURE 15.-Distribution <strong>of</strong> Parrnelina galbina in Korth<br />
America (taken froin W. Culberson, 1961).<br />
16. <strong>Parmelina</strong> hayachinensis<br />
FIGURE 14d<br />
Parrnelina hayaclzinensis (Kurokait-a) <strong>Hale</strong>, 1971:482.<br />
Prrrnlelia ha)achinensis Kurokawa, 1968b:350 [type collection:<br />
hit. Hayachine, Proy. Rikuchu, Japan, Kurokawa 67081<br />
(TSS, holotype)].<br />
DEscRIPnoN.-Thallus adnate on bark, whitish to<br />
greenish mineral gray, 5-8 cm broad; lobes sub-<br />
linear, 1.5-3 mm wide, marginal cilia simple, about<br />
0.2 mm long; upper surface shiny, faintly maculate,<br />
becoming densely pustulate, <strong>the</strong> pustules becoming<br />
aggregated, erupting but not forming soredia;<br />
medulla white; lower surface black, densely rhizin-<br />
ate, <strong>the</strong> rhizines black, simple. Apo<strong>the</strong>cia not seen.<br />
CHE~IsTRY.-Cortex K + yellow (atranorin);<br />
medulla K+ yellow turning red, C-, KC-, P+<br />
orange (galbinic acid, leucotylin, and zeorin).<br />
DxsmIsuTIoN.-Japan.<br />
REMARKS.-AS pointed out by Kurokawa (1968b:<br />
350), this species is a member <strong>of</strong> <strong>the</strong> P. galbina<br />
complex, representing <strong>the</strong> pustulate morph. It is<br />
known only from <strong>the</strong> type collection growing on<br />
Cryptomeria in Japan.<br />
17. <strong>Parmelina</strong> heteroloba<br />
FIGURE 14e<br />
Parrnelina heteroloba (Zahlbruckner) <strong>Hale</strong>, 1974:482.<br />
Parrnelia heteroloba Zahlhruckner, 1909: 171 [tlpe collection:<br />
Mt. Itatiaya, Rio de Janeiro, Brazil, Schiflner (W, lecto-<br />
type)l.<br />
DEscRIPTIoN.-Thallus adnate on bark, light buff<br />
mineral gray in <strong>the</strong> herbarium, 5-8 cm broad; lobes<br />
more or less sublinear, short and imbricate, be-<br />
coming lobulate toward <strong>the</strong> center, 2-3 mm wide,<br />
<strong>the</strong> marginal cilia mostly in <strong>the</strong> lobe axils, to 0.4<br />
mm long; upper surface shiny, plane to rugulose,<br />
becoming somewhat lobulate; lower surface black<br />
except for a narrow brown zone at <strong>the</strong> tips, densely<br />
rhizinate, <strong>the</strong> rhizines black, simple or sparsely<br />
furcate. Apo<strong>the</strong>cia numerous, substipitate, <strong>the</strong> disc<br />
splitting, 2-9 mm in diameter; spores 8, 8 X 12 pm.<br />
CHEMIsTRY.-Cortex K + yellow (atranorin); me-<br />
dulla negative with color tests (an unidentified fatty<br />
acid and a faint unknown spot, perhaps related to<br />
one <strong>of</strong> <strong>the</strong> “horrescens” unknown).<br />
DIsmIsuTIoN.-Brazil.<br />
REMARKS.-siXICe this species is represented only<br />
by <strong>the</strong> single type collection, it is difficult to gen-<br />
eralize on its affinities to o<strong>the</strong>r <strong>Parmelina</strong>e. It would<br />
probably be mistaken for P. damaziana, which has<br />
a different chemistry (“horrescens” unknown) and<br />
smaller spores. It is only one <strong>of</strong> three species in<br />
Paimelina with fatty acids, <strong>the</strong> o<strong>the</strong>rs being P.<br />
endolezica and P. expallida. Parmelilza heteroloba<br />
has not been recollected at Alt. Itatiaya, a well<br />
known site that has been visited by several experi-<br />
enced lichen collectors, and it may be an extinct<br />
species.<br />
18. <strong>Parmelina</strong> horrescens<br />
FIGURE 14f<br />
Paririelina horresce?is (.laylor) <strong>Hale</strong>, 1974:482.<br />
Pnr,,relin korrescens Taylor in Mackay, 1836:144 [t!Iie collec-<br />
tion: Dunkerron Mountains, Kerry, Ireland, Taylor (FH-<br />
Tayl, lectotype)].<br />
Dk:scRrp?.xos.-Thallus closely adnate to adnate on<br />
bark, rocks, or mosses over rocks, whitish to green-<br />
ish mineral gray, 2-5 cm broad; lobes more or less<br />
dichotomously branched, sublinear, <strong>of</strong>ten crowded<br />
and imbricate, 0.5-2.0 mm wide, <strong>the</strong> margins cre-
nate, <strong>of</strong>ten becoming lobulate, ciliate, <strong>the</strong> cilia more<br />
or less evenly dispersed, black, simple, 0.3-0.8 mm<br />
long; upper surface shiny, emaculate, densely isidi-<br />
ate, <strong>the</strong> isidia cylindrical, <strong>of</strong>ten branched and<br />
apically spinulate or short-ciliate, in part becoming<br />
procumbent; medulla white; lower surface black,<br />
moderately rhizinate, <strong>the</strong> rhizines black, simple.<br />
Apo<strong>the</strong>cia rare, sessile, 2-4 mm in diameter, <strong>the</strong><br />
amphi<strong>the</strong>cium isidiate, <strong>the</strong> disc splitting at ma-<br />
turity; spores 8, 10-12 X 16-18 pm.<br />
CHExmTRY.-Cortex K + yellow (atranorin); me-<br />
dulla K -, C - or C + faint rose, KC + rose or red,<br />
P - (trace <strong>of</strong> gyrophoric acid, “horrescens” un-<br />
known falling above gyrophoric acid on chroma-<br />
tographic plates, and one or two o<strong>the</strong>r unidentified<br />
s PO t S) .<br />
DIsTRIsuTIozr.-Pantemperate and montane pan-<br />
tropical.<br />
REMARK\.-,& discussed under P. dissecta, this<br />
species only recently was differentiated correctly<br />
from P. dissecta (<strong>Hale</strong> and Kurokawa, 1962:2), an<br />
isidiate lichen with gyrophoric acid. <strong>Parmelina</strong><br />
horrescens is characterized by dense fine isidia with<br />
greater or lesser development <strong>of</strong> short apical cilia<br />
(Figure 4c). The isidia sometimes become procum-<br />
bent and lobulate. On <strong>the</strong> whole, <strong>the</strong> lobes are<br />
somewhat narrower and more appressed than in<br />
P. dissecta. Parinelina horrescens has essentially <strong>the</strong><br />
same geographic range as P. dissecta and occurs<br />
most abundantly in <strong>the</strong> temperate deciduous<br />
forests <strong>of</strong> <strong>the</strong> eastern United States and Japan. It is<br />
montane in <strong>the</strong> tropics, occurring as high as 3000 m<br />
in <strong>the</strong> paramos <strong>of</strong> Venezuela.<br />
The chemistry <strong>of</strong> P. horrcscens is now being<br />
studied by several lichen chemists. A number <strong>of</strong><br />
unidentified spots appear in both hexane and ben-<br />
zene solvent systems, one <strong>of</strong> which is probably a<br />
trace <strong>of</strong> gyrophoric acid. The o<strong>the</strong>r spots fall above<br />
and below gyrophoric acid and seem to represent<br />
closely related depsides.<br />
The parent morph <strong>of</strong> P. horrescens is probably<br />
extinct. Chemically identical P. damaziana, a non-<br />
isidiate Brazilian species, is larger and more robust<br />
although obviously from <strong>the</strong> same stock as <strong>the</strong><br />
parent <strong>of</strong> P. horrescens. Among <strong>the</strong> parallel morphs,<br />
both pustulate-sorediate P. subfatisceiis and lobulate<br />
P. schindzeri are very close in lobe configuration,<br />
adnation, and thallus texture.<br />
SPECIMENS EX&MINED.-United States: Illinois, Skorepa 4627<br />
SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
(US); Kentucky, <strong>Hale</strong> 13743a; West Virginia, <strong>Hale</strong> 10612,<br />
11772, 11878; Virginia, <strong>Hale</strong> 18388, 33153, Reed 9091 (Reed<br />
herbarium), Roller 400 (US); h‘orth Carolina, Culberson<br />
5114, 5706, 5744, 7134 (DUKE), <strong>Hale</strong> 18042, 15058, Imshaug<br />
22174, 22178, 22353 (MSC, US); Tennessee, <strong>Hale</strong> 31106, 36921,<br />
36959, 36970, Moore 284 (US), Phillips 358, 377 (US), Skorepa<br />
5527 (US); South Carolina, <strong>Hale</strong> 7723; Georgia, <strong>Hale</strong> 7405,<br />
7537, 16761, 16776, 30882, 30885; Alabama, <strong>Hale</strong> 7186, 7216,<br />
31164, 33779, 33918, 34121, 34174, McCullough 2195 (US);<br />
Florida, <strong>Hale</strong> 21685 (for additional records see Moore,<br />
1968:220). Mexico: Chiapas, <strong>Hale</strong> 20204, 20223, 20400a, 20414,<br />
20549, 21086. Guatemala: Baja Vera Paz, <strong>Hale</strong> 45828. Pan-<br />
ama: Darien, >Vori and Gentry 4309 (US); Panami, <strong>Hale</strong><br />
38451. Cuba: Oriente, Imshaug 24737, 24810, 24932 (MSC).<br />
Jamaica: Zmshnug 14216 (MSC). Dominican Republic: Cordil-<br />
lera Central, Imshaug 23516 (MSC), Wetmore 3739 (MSC); La<br />
T’ega, Allnrrl 17695a (US). Haiti: Ouest, Fahius 2-4 (US),<br />
Ini.rhaug 22768, 22837, 22860 (MSC, US), Wetmore 3222<br />
(MSC); Sud, Imshaug 23233 (MSC, US). Venezuela: Distrito<br />
Federal, Deiinis 2394 (BM), Santesson 6679 (S); Wrida, <strong>Hale</strong><br />
42038, 42067, 42952, 45121, 45201. Urugna): Trienta y Tres,<br />
Osorio 5931 (MT‘M). France: Harmand (DUKE). Spain: Ponte-<br />
vedra, Schnurr (XI); Tenerife, Zmshazcg 34476, 35677 (MSC).<br />
Union <strong>of</strong> South Africa: Cape Province, Alniborn 1442 (LD).<br />
India: Tamil Nadn, <strong>Hale</strong> 43784. Philippines: Mountain Pror-<br />
ince, <strong>Hale</strong> 26531. Indonesia: Java, Groenhart 2903 (L, US),<br />
6279 (L), Kurokau,a 2071 (TNS). Taiwan: Kurokawa 693<br />
(TSS). Japan: Proy. Buzen, Kiirokawa 62468, 63166 (TNS);<br />
Prov. Kii, Kurokawa 64127 (TNS); Prov. Ohmi, <strong>Hale</strong> 29465.<br />
Australia: Sew South \Vales, Cizeel L1708 (NSW), Craigie<br />
(NSW). Sew Zealand: Wade 85 (BM, US).<br />
19. <strong>Parmelina</strong> immiscens<br />
FIGURE 16a<br />
Pnrmelina irnmiscens (Nylander) <strong>Hale</strong>, 1974:482.<br />
Parmelia immiscens Nylander, 1885:606 [type collection:<br />
Orizaba, Mexico, Galeotti 6879 (PC, 1ectot)pe; isolectotype<br />
in H, Xilander herbarium number 35674)l.<br />
Pnrmelia michoacanensis Body de Lesdain, 1914:7 [type<br />
collection: Jeslis del Monte, Morelia, Michoacan, Mexico,<br />
Arsene 4456 (US, lectotype; isolectotypes in COLO, DUKE,<br />
G, LE, and UPS)].<br />
DEscRIPTIoN.-Thallus adnate on bark, pale tur-<br />
tle green, 5-10 cm in diameter; lobes subirregular<br />
and apically rotund, 2-6 mm wide, <strong>the</strong> marginal<br />
cilia mostly in <strong>the</strong> axils; upper surface plane, con-<br />
tinuous, <strong>of</strong>ten pruinose near <strong>the</strong> tips; medulla<br />
sulphur yellow; lower surface densely rhizinate,<br />
short rhizinate and pale brown along <strong>the</strong> margins,<br />
<strong>the</strong> rhizines simple or squarrosely branched. Apoth-<br />
ecia numerous, adnate, <strong>the</strong> disc pale, 2-7 mm in<br />
diameter; spores 8, 4-6 X 7-12 pm.<br />
CmmsmY.-Cortex K + yellow (atranorin);
NUMBER 33<br />
FIGURE 16.4pecies <strong>of</strong> Parrnelina: a, P. irnrniscens (Andne 4456 in US); b, P. indica (<strong>Hale</strong><br />
43884); c, P. irrugans (Kurokauu 55342); ti, P. jarnesii (Du Rieti 3117:3 in US); e, P. leucotyliza<br />
(<strong>Hale</strong> 29402); f, P. lindrnnnii (Maline 450 in S). (Scale in mm.)<br />
33
34 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
medulla K+, C+, KC+ more intensely yellow,<br />
P - (unidentified yellow pigments).<br />
DIsTRIsuT1oN.-Mexico.<br />
REMARKs.-<strong>Parmelina</strong> immiscens is <strong>the</strong> only spe-<br />
cies in <strong>the</strong> genus endemic to Mexico. It occurs<br />
ra<strong>the</strong>r rarely on trees in open oak-pine forests in<br />
<strong>the</strong> arid highlands (2000-2400 m elevation). Al-<br />
though <strong>the</strong> medulla is distinctly yellow, no terpenes<br />
are produced. The pigment seems to be secalonic<br />
acid A but chromatographic tests are inconclusive.<br />
This species and its presumptive isidiate morph<br />
P. Zindmanii, <strong>the</strong>refore, are unrelated to P. sub-<br />
aurulenta and similar Asian species with a yellow<br />
medulla.<br />
SPECIMENS EXAMINED.-MeXicO: Durango, Cramer 1998<br />
(KAN, US); Jalisco, Pringle 10706 (US); Oaxaca, <strong>Hale</strong> 20825.<br />
20. <strong>Parmelina</strong> indica, new species<br />
FIGURE 166<br />
DEscRIPTIoN.-Thallus arcte adnatus, saxicola,<br />
obscure albo-cinereus, 2-3 cm latus, lobis sub-<br />
linearibus, contiguis, 0.8-1.2 mm latis, ciliis mar-<br />
gine irregulariter dispersis, simplicibus, 0.1-0.3 mm<br />
longis, superne planus, continuus vel rimosus,<br />
nitidus, modice vel dense isidiatus, isidiis cylin-<br />
dricis vel leviter inflatis, praecipue simplicibus,<br />
usque ad 0.3 mm altis; cortex superior 12 pm<br />
crassus, epicorticatus, stratum gonidiale 15-18 pm<br />
crassum, medulla alba, 100-120 pm crassa, cortex<br />
inferior paraplectenchymatus, 12-14 pm crassus;<br />
subtus niger, dense rhizinosus, rhizinis nigris, sim-<br />
plicibus, 0.4 mm longis. Apo<strong>the</strong>cia ignota.<br />
CHEMIsmY.-Cortex K + yellow (atranorin);<br />
medulla negative with color tests zeorin, leucotylin<br />
and traces <strong>of</strong> associated “subaurulenta” terpenes).<br />
HOL0TYPE.-on rocks in river below Silver Cas-<br />
cade, Kodaikanal, Tamil Nadu, India, elevation<br />
about 1800 m, M. E. <strong>Hale</strong> 43884, 24 January 1975<br />
(US: isotype in Poona).<br />
DIsmIsuTIoN.-India.<br />
REMARKS.-This small saxicolous lichen was col-<br />
lected on a large rock outcropping in an undis-<br />
turbed stream bed. Externally it is close to <strong>the</strong><br />
American P. obsessa but lacks both yellow pig-<br />
ments and galbinic acid. It is <strong>the</strong> only terpene-<br />
containing <strong>Parmelina</strong> without pigments.<br />
21. Parmelia irrugans, new combination<br />
FIGURE 16c<br />
Parmelia irrugans Nylander, 1890:26 [type collection: Simon-<br />
oseki, Japan, A lmquist (H, Nylander herbarium number<br />
35551, lectotype; isolectotype in S)].<br />
Parmelia insinuata Hue, 1899: 158 [type collection: Ta-Pin-<br />
Tze, Yunnan, China, Delavay 3008 (PC, lectotype); not P.<br />
insinuata ”).lander, 1856: 3243.<br />
Parmelia xanthocarpa Hue, 1899:178 [type collection: Ta-<br />
Long-Tan near Ta-Pin-Tze, Yunnan, China, Delavay 23<br />
(PC, lectotype)].<br />
Parmelia insinuatula Zahlbruckner, 1929: 169 [based on P.<br />
insinuata Hue].<br />
DEsCRIPTIoN.-Thallus adnate to loosely adnate,<br />
corticolous, firm, 4-10 cm broad; lobes sublinear<br />
to subirregular with more or less rotund apices,<br />
contiguous, 2-5 mm wide, <strong>the</strong> marginal cilia dis-<br />
tinct, mostly in <strong>the</strong> axils, to 0.8 mm long; medulla<br />
pale orange or yellow; lower surface black, densely<br />
rhizinate, <strong>the</strong> rhizines simple or becoming squar-<br />
rosely branched. Apo<strong>the</strong>cia very common, adnate<br />
to sessile, 3-10 mm in diameter; spores 8, 8-10 X<br />
12-15 pm, <strong>of</strong>ten poorly developed.<br />
CHEMIsmY.--Cortex K + yellow (atranorin);<br />
medulla more intensively yellow with color tests<br />
(zeorin, leucotylic acid and associated terpenes, and<br />
secalonic acid A).<br />
DIsmIsuTIoN.-China and Japan.<br />
REMARKS.-This species has not been recognized<br />
by contemporary lichenologists or included in lists<br />
<strong>of</strong> Asian <strong>Parmelina</strong>e. It is characterized by <strong>the</strong><br />
presence <strong>of</strong> <strong>the</strong> “aurulenta” terpene series, <strong>the</strong> thin,<br />
expanded thallus, and large apo<strong>the</strong>cia (3-10 mm in<br />
diameter), invariably larger than those <strong>of</strong> P. sub-<br />
aurulenta (3 mm or less). The spores are slightly<br />
but not significantly larger than those in P. SUB-<br />
aurulenta. As a rule, <strong>the</strong> size <strong>of</strong> apo<strong>the</strong>cia will<br />
separate <strong>the</strong> two species. Two <strong>of</strong> Nylander’s syn-<br />
types <strong>of</strong> P. subaurulenta (see “Specimens Exam-<br />
ined” below) are actually P. irrugans.<br />
<strong>Parmelina</strong> irrugans has a more restricted range<br />
than P. subaurulenta. It does not occur in India<br />
where P. subaurulenta is so common. It is, how-<br />
ever, more frequent in Japan than one might<br />
expect from <strong>the</strong> specimens examined because I did<br />
not borrow <strong>the</strong> specimens at TNS for redetermina-<br />
tion. Many annotated by me in 1964 in Tokyo as<br />
P. homogenes are probably this species.<br />
SPECIMENS EXAMINED.-China: Montiguy (H. Nylander her-
NUMBER 93 35<br />
barium number 35668, a syntype <strong>of</strong> Parmelia subaurulenta<br />
Kllander); Hunan, Handel-Mazzetti 2469 (US, \V); Yunnan,<br />
Delaruy (H, K)lander herbarium number 35666). Japan:<br />
illaries (H, Sylander herbarium number 35667, probable syn-<br />
type <strong>of</strong> Pormelia subaurulenta Nylander); Prov. Kii, Kuro-<br />
kawn 59064 (TNS, US); Prov. Kikuchu, Kzirokazua 59322<br />
(TNS, US); Prov. Kozuhe, Kurokazua 58581 (TNS, US); Prov.<br />
Mutsu, Kurokawa 550341, 55342, 56162, 58582 (TNS, US);<br />
Prov. Ohmi, <strong>Hale</strong> 29456, 29470; Prov. Rikuchu, Kurokawa<br />
59322 (TNS, US); Prov. Shimane. Omura 23 (US); Prov.<br />
Suruga, Culberson 10738 (DUKE, US).<br />
22. <strong>Parmelina</strong> jamesii, new combination<br />
FIGURE 16d<br />
Parmelia jamesii <strong>Hale</strong>, 1972b:179 [type collection: Welling-<br />
ton, New Zealand, James SZ2118 (US, holotype; isotypes in<br />
BM, TNS)].<br />
DEscRIPTros.-Thallus adnate on bark, ra<strong>the</strong>r<br />
fragile, whitish mineral gray, 5-10 cm broad; lobes<br />
sublinear, contiguous, 1.5-3.0 mm wide, <strong>the</strong> mar-<br />
gins sparsely ciliate, <strong>the</strong> cilia irregularly dispersed,<br />
long; upper surface plane but becoming rugulose<br />
with age, shiny or becoming opaque and white-<br />
pruinose at <strong>the</strong> tips, sometimes faintly white-<br />
reticulate, moderately isidiate, <strong>the</strong> isidia cylindrical,<br />
thin, erect, up to 0.3 mm high; lower surface black,<br />
moderately rhizinate, <strong>the</strong> rhizines simple or sparsely<br />
squarrosely branched. Apo<strong>the</strong>cia unknown.<br />
CHEmsTRY.-Cortex K+ yellow (atranorin);<br />
medulla K-, C-, KC-, P+ red (fumarproto-<br />
cetraric acid and a trace <strong>of</strong> protocetraric acid).<br />
DIsTRIBuTI0N.-Eastern Australia and New<br />
Zealand,<br />
REMARKs.-<strong>Parmelina</strong> jamesii is <strong>the</strong> only species<br />
producing ,B-orcinol depsidone fumarprotocetraric<br />
acid. It is also one <strong>of</strong> <strong>the</strong> few species endemic to<br />
<strong>the</strong> Australian region. It might be mistaken super-<br />
ficially for a Hypotrachyna except that <strong>the</strong> lobe<br />
margins are distinctly ciliate and <strong>the</strong> rhizines<br />
simple.<br />
23. <strong>Parmelina</strong> leucotyliza<br />
FIGURE 16e<br />
<strong>Parmelina</strong> leucotylizn (Sylander) <strong>Hale</strong>, 1974:482.<br />
Parmelia leucotyliza Nylander, 1890:27 [type collection: Rock-<br />
osan, Japan, Almquist (H, Nylander herbarium number<br />
35196, lectotype: isolectotype in S)].<br />
Parmelia fmudans ssp. subfraudans Zahlbruckner, 1927:352<br />
[type collection: Inokashira, near Tokyo, Japan, Asahina<br />
23a (W, lectotype)].<br />
Parmelia leucotyliza f. rugulosa Asahina, 1952:94 [tvpe col-<br />
lection: Mt. Higane, Prov. Suruga, Japan, Yamashita 17<br />
(TNS, lectotype)].<br />
Parmelia leucotyliza f. sziblaevis Asahina, 1952:95 [type collec-<br />
tion: Kadoike, Pro\.. Suruga, Japan, Asahina (TSS, lecto-<br />
type)].<br />
DEscRIPTIo;u.-Thallus adnate to loosely attached<br />
on bark or rocks, light greenish mineral gray, 8-12<br />
cm broad; lobes sublinear to subirregular, con-<br />
tiguous, 2-4 mm wide, <strong>the</strong> marginal cilia mostly<br />
in lobe axils, about 0.5 mm long; upper surface<br />
shiny or becoming dull white pruinose at <strong>the</strong> tips,<br />
plane but soon becoming pustulate, <strong>the</strong> pustules<br />
breaking open and coalescing into large clumps<br />
without formation <strong>of</strong> soredia; medulla very pale<br />
salmon colored to white; lower surface black ex-<br />
cept for a dark brown zone at <strong>the</strong> tips, densely<br />
rhizinate, <strong>the</strong> rhizines black and shiny, simple or<br />
sparsely squarrosely branched. Apo<strong>the</strong>cia rare, sub-<br />
stitpitate, <strong>the</strong> amphi<strong>the</strong>cium pustulate, 1-3 mm in<br />
diameter; spores 8, 6 X 11-12 pm.<br />
CHEmsTRY.-Cortex K + yellow (atranorin);<br />
medulla more intensively yellow with color tests<br />
(zeorin, leucotylin and related “subaurulenta” ter-<br />
penes, secalonic acid A, and traces <strong>of</strong> unidentified<br />
substances).<br />
DIsTRIBuTIoN.-Japan and hlalaysia (Sabah).<br />
REMARKS.-Originally when studying <strong>the</strong> Jap<br />
anese Paymeliae, I synonymized P. leucotyliza under<br />
P. aurulenta. These two species, however, are differ-<br />
ent in several important respects. First, <strong>the</strong>y have<br />
different terpene chemistry. Second, <strong>the</strong> pustules <strong>of</strong><br />
Paymelina leucotyliza do not become sorediate,<br />
whereas those in P. aurulenta are <strong>of</strong>ten densely<br />
sorediate. Lastly, P. leucotyliza is restricted to Japan<br />
except for a tentatively identified specimen from<br />
Sabah. It seems to be much more common than<br />
P. aurulenta in Japan, occurring on rocks and trees<br />
in open forests, trees along roads, and rocks in rice<br />
fields.<br />
SPECIXIE~S<br />
ExAaIisED.-Japan: Prov. Aki, <strong>Hale</strong> 29535, 29445,<br />
29544; Prov. Hyuga, <strong>Hale</strong> 29677; Prov. Iwaki, Kurokarua<br />
58079 (TNS, US); Prov. Izu, Asahina in <strong>Lichen</strong>es Japoniae<br />
Exsiccati 139 (US), Kurokawa 58004, 58600 (TNS, US); Prov.<br />
Kazusa, Kurokawa 59002 (TSS, L’S); Prov. Kii, Kurokawa<br />
59087, 59131, 59132 (TNS, US); Prov. Musashi, Kurokawa<br />
51040 (TNS, US); Prov. Settsu, <strong>Hale</strong> 29402, 29433; Shizuoka<br />
Prefecture, Omura 157 (US). Malaysia: Sabah, <strong>Hale</strong> 29102.
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
24. Pamtelina lindmanii<br />
FIGURE 16f<br />
<strong>Parmelina</strong> lindmanii (Lynge) <strong>Hale</strong>, 1974:483.<br />
Parrnelia lindrnanii Lynge, 1914:74 [type collection: Porto<br />
Alegre, Rio Grande do Sul, Brazil, Maime 450 (S, lecto-<br />
type)l.<br />
Parmelia tiliacea var. sulphurosa f. asperata Miiller Argoviensis,<br />
1883:46 [type collection: Argentina, Balansa 5 (G,<br />
lectotype)].<br />
DESCRIPTION.- Thallus adnate on bark, pale buff<br />
to yellowish mineral gray, 5-8 cm broad; lobes subirregular,<br />
apically rotund, 3-5 mm wide, <strong>the</strong> marginal<br />
cilia mostly in lobe axils, less than 0.5 mm<br />
long; upper surface plane to rugose with age, shiny,<br />
moderately to densely isidiate, <strong>the</strong> isidia cylindrical,<br />
unbranched, to 0.3 mm high; medulla uniformly<br />
pale sulfur-yellow tinged with orange; lower surface<br />
black except for a narrow marginal brown<br />
zone, moderately rhizinate, <strong>the</strong> rhizines black, simple.<br />
Apo<strong>the</strong>cia rare, sessile, 2-5 mm in diameter;<br />
spores 8,6-10 X 9-14 pm.<br />
CHExmTRY.-Cortex K + yellow (atranorin);<br />
medulla more intensely yellow with K and C, P-<br />
(probably secalonic acid A and unidentified<br />
pigments).<br />
DISTRIBUTION.-RleXiCO and South America.<br />
REhlARKS.-This corticolous lichen is character-<br />
ized by <strong>the</strong> pale yellow-orange medulla and <strong>the</strong><br />
isidia. It is probably <strong>the</strong> isidiate morph <strong>of</strong> <strong>the</strong><br />
Mexican endemic P. immiscens. When I first began<br />
study <strong>of</strong> Pal-nzelia, I confused Pal-melina lindmanii<br />
and Pal-melia endosulphur-ea ( = Parmotl-ema endo-<br />
sulphur-eum (Hillmann) <strong>Hale</strong>) (<strong>Hale</strong>, 1960:20). Both<br />
species have a yellow-orange medulla and isidia, but<br />
P. lindmanii has more adnate lobes, cilia in <strong>the</strong><br />
axils, and no gyrophoric acid. Both species lack<br />
terpenes.<br />
SPECIMENS Ex.4JrINEo.-Mexico: Queretaro, Nakanishi 257<br />
(US); Tamaulipas, Nakanishi 106 (US): \'era Cruz, <strong>Hale</strong><br />
19389, 19685. Colombia: Antioquia, Nee and Mori 4238 (US).<br />
Venezuela: Merida, Rodrigues 168 (US). Brazil: Rio Grande<br />
do Sul, Lima 3 (US). Paragua:; Caacupe, Balansa (G, H).<br />
Uruguay: Durazno, Osorio 2772 (DUKE, UPS): Lavalleja,<br />
Osorio 3822 (MVM, US): Paysandu, Lamb (BM, H); Soriano,<br />
Rosengurtt (H); Trinidad, Osorio 3603 (LD). Argentina:<br />
James (BM): Buenos Aires, Santesson 58, 100, 446 (S).<br />
25. <strong>Parmelina</strong> melanochaeta<br />
FIGURE 17a<br />
<strong>Parmelina</strong> nielanochaeta (Kurokawa) <strong>Hale</strong>, 1974:483.<br />
Parmelia inelanochaeta Kurokaira in <strong>Hale</strong> and Kurokawa,<br />
1964:133 [type collection: Santa Anna de Chapada, Mato<br />
Grosso, Brazil, Malme 2243 (S, lectot) pe: isolectotypes in<br />
UC, US)].<br />
DEscRIrTIos.-Thallus adnate on bark, turning<br />
olive-buff to cream-buff in <strong>the</strong> herbarium, 4-7 cm<br />
in diameter; lobes irregularly branched, sometimes<br />
sublinear-elongate, 2-6 mm wide, <strong>the</strong> margins more<br />
or less crenate, ciliate, <strong>the</strong> cilia black, mostly simple,<br />
1-2 mm long; upper surface shiny, maculate, mod-<br />
erately to densely isidiate, isidia thin, cylindrical,<br />
usually branched, <strong>of</strong>ten with black spinules or short<br />
cilia; medulla white; lower surface black, dark to<br />
pale brown in a ra<strong>the</strong>r wide zone near <strong>the</strong> tips,<br />
moderately rhizinate, rhizines black to blackish<br />
brown, simple. Apo<strong>the</strong>cia adnate to substipitate,<br />
1-3 mm in diameter, <strong>the</strong> amphi<strong>the</strong>cium isidiate,<br />
spinulate, <strong>the</strong> disc Vandyke-brown; spores 8, 8-10 x<br />
13-15 pm.<br />
CHEiwsTRY.-Cortex K + yellow (atranorin); me-<br />
dulla K-, C+, KC+ rose, P- (gyrophoric acid).<br />
DIsTRIBuTIoN.-South America.<br />
REMARKS.-This species is very close to P. dissecfa<br />
in general morphology and has <strong>the</strong> same chemistry.<br />
The lobes, however, are much broader, <strong>the</strong> isidia<br />
ciliate, <strong>the</strong> marginal cilia long and distinct, and<br />
<strong>the</strong> upper cortex strongly white-maculate. It occurs<br />
in a ra<strong>the</strong>r restricted area from Brazil into Para-<br />
guay with one specimen recorded from Colombia.<br />
SPECIMF\S ExA\IINED.-Colombia: Cundinamarca, Flenniken<br />
1745 (US). Brazil: Mato Grosso, Malme 2397B (S). Paraguay:<br />
Asuncih, Schinini 17333 (M\'M): Gran Chaco, hlalme (S).<br />
Additional records from Brazil and Paraguai are listed in<br />
<strong>Hale</strong> and Kurokana, 1964:133.<br />
26. <strong>Parmelina</strong> metarevoluta<br />
FIGURE 17b<br />
Par??7e[i?7a rnetaraloluta (Asahina) <strong>Hale</strong>, 1974:483.<br />
Pnrmelia tnetarevolirtn Asahina, 1960:97 [type collection:<br />
AzusaJama, Prov. Shinano, Japan, h'uno and Kurokawa<br />
59243 (TSS, lectot) pe; isolectotype in US)].<br />
DEscRIPTIo;u.-Thallus adnate on bark or rock,<br />
pale olive-buff, 2-6 cm in diameter; lobes dicho-<br />
tomously branched, sublinear-elongate, more or<br />
less ascending at <strong>the</strong> tips, 1-4 mm wide, lobules<br />
sometimes present on older lobes, <strong>the</strong> margins more<br />
or less crenate, narrowly black rimmed, ciliate, <strong>the</strong><br />
cilia black, shiny, simple, 0.2-0.5 mm long; upper
NUMBER 33<br />
FIGURE 17.4pecies <strong>of</strong> Parinelina: a, P. melanochaeta (21ialnie 2243 in S); b, P. inetarevoluta<br />
(Kurokawa 59180 in US); c, P. niiielleri (<strong>Hale</strong> 42277): d, P. obsessa (<strong>Hale</strong> 19186); e, P. pnsr~lltfera<br />
(specimen in US); f, P. perisidrans (Togashi in US). (Scale in mm.)<br />
37
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
surface smooth and shiny, not maculate, more or<br />
less rugose on older lobes, sorediate, soralia subterminal,<br />
capitate, pale orange below; lower surface<br />
black, densely rhizinate; <strong>the</strong> rhizines black, shiny,<br />
simple, about 0.5 mm long. Apo<strong>the</strong>cia very rare,<br />
adnate, 1-5 in diameter, <strong>the</strong> rim more or less<br />
crenate, sorediate, <strong>the</strong> disc dark brown; spores 8,<br />
7 X 10-12pm.<br />
CmMIsmY.-Cortex K + yellow (atranorin); medulla<br />
K + reddish, C -, KC -, P + orange (galbinic<br />
acid, rarely constictic acid, trace <strong>of</strong> salazinic acid,<br />
zeorin, leucotylin and associated terpenes, and<br />
secalonic acid A).<br />
DIsTR1suTIoN.-Eastern United States, Japan, and<br />
China.<br />
REMARKL-AS pointed out by Kurokawa (1968b:<br />
351) this is <strong>the</strong> sorediate morph <strong>of</strong> P. galbina. It<br />
was first collected and correctly identified in <strong>the</strong><br />
United States by Dr. Satoshi Nakanishi in 1971.<br />
Two additional collections have since been identified,<br />
one <strong>of</strong> <strong>the</strong>m (Plitt 236) filed under Physcia<br />
orbicularis, which is very superficially similar in<br />
lobe configuration and capitate soralia. It has <strong>the</strong><br />
same Arcto-Tertiary distribution pattern as P. galbina<br />
but is extremely rare in North America.<br />
SPECIMENS ExAMmEu.-United States: Ohio, Wetmore 18051<br />
(MI", US); Maryland, Plitt 236 (US); Tennessee, Nakanishi<br />
238 (US). Japan: Prov. Bungo, Kurokawa 62329 (TR'S); Prov.<br />
Ise, Asahina (TNS); Prov. Omi, Asahina (TNS); Prov.<br />
Shinano, Asahina (TNS). Kurokawa 58326 (TNS); 59180<br />
(TNS, US) 59243, 59244 (TNS); Prov. Suruga, Asahina (TR'S);<br />
Prov. Totomi, Asahina 160 (TNS); Prov. Yamashiro, Asahina<br />
(TNS). China: Manchuria, Asahina 29 (TNS).<br />
27. <strong>Parmelina</strong> muelleri<br />
FIGURE 17c<br />
<strong>Parmelina</strong> muelleri (Vainio) <strong>Hale</strong>, 1974:483.<br />
Parmelia muelleri Vainio, 1890:49 [type collection: Sitio,<br />
Minas Gerais, Brazil, Vainio in <strong>Lichen</strong>es Brasilienses<br />
Exsiccati 948 (TUR, Vainio herbarium number 2677, lecto-<br />
type; isolectotypes in BM, FH, M, UPS)].<br />
DEScRIPTIoN.-Tha11us closely adnate on bark,<br />
lichen green, turning honey yellow in <strong>the</strong> her-<br />
barium, 5 cm or more in diameter; lobes irregularly<br />
branched, rounded at <strong>the</strong> tips, 2-3 mm wide, <strong>the</strong><br />
margins evenly ciliate, <strong>the</strong> cilia simple or rarely<br />
branched, black, shiny, up to 1.5 mm long; upper<br />
surface strongly white-maculate, irregularly cracked<br />
on older lobes, sorediate, soredia laminal, more or<br />
less granular, <strong>the</strong> soralia round, separate; lower<br />
surface uniformly black, moderately rhizinate, <strong>the</strong><br />
rhizines simple, black. Apo<strong>the</strong>cia rare, sessile, <strong>the</strong><br />
amphi<strong>the</strong>cium sorediate, disk brick red, 2-5 mm in<br />
diameter; spores 8,7-I0 X 12-15 pm.<br />
CHEMIsTRY.-Cortex K + yellow (atranorin); me-<br />
dulla K+ persistent yellow, C-, KC-, P+ pale<br />
orange (stictic acid with or without constictic acid).<br />
DISTRIRUTI0N.-Mexico and South America.<br />
REMARKS.--When 1 first collected this species on<br />
shade trees in a c<strong>of</strong>fee plantation in Venezuela, I<br />
identified it tentatively as P. pilosa because <strong>of</strong> <strong>the</strong><br />
laminal soralia and distinct white maculae. Chem-<br />
ical tests, however, showed that it contained stictic<br />
acid and would have to be identified with P.<br />
muelleri. After comparing <strong>the</strong> two species in <strong>the</strong><br />
herbarium, I concluded that P. pilosa is larger and<br />
more robust on <strong>the</strong> average, but <strong>the</strong> two are very<br />
closely related. Vainio (1890) had in fact compared<br />
P. muelleii with Parmelia balansa f. sorediata (=<br />
<strong>Parmelina</strong> pilosa) which he distinguished with a<br />
negative K test. They have different geographic<br />
ranges, P. muelleri being more common in nor<strong>the</strong>rn<br />
South America and possibly occurring at a higher<br />
elevation than P. pilosa. There is no esorediate,<br />
stictic acid-containing parent morph comparable to<br />
<strong>the</strong> P. consors-P. pilosa species pair.<br />
SPECIMENS ExAMINED.-Mexico: Michoackn, Wirth 329 (US);<br />
Vera Cruz, <strong>Hale</strong> 21124. Venezuela: MCrida, <strong>Hale</strong> 42219,<br />
42230, 42256, 42277, Oberroinkler and Poelt 7529 (M, US);<br />
Thchira, <strong>Hale</strong> 45716. Peru: Cuzco, lltis 3214 (WIS). Argentina:<br />
TucumPn, Culberson 14909 (DUKE, US).<br />
28. <strong>Parmelina</strong> obsessa<br />
FIGURE 17c<br />
<strong>Parmelina</strong> obsessa (Acharius) <strong>Hale</strong>, 1974:483.<br />
Parmelia obsessa Acharius, 1814:195. [type collection: North<br />
America (?Pennsylvania), Muhlenberg (H-Ach, lectotype;<br />
isolectotype in PH)].<br />
Parmelia finkii Zahlbruckner in Hedrick, 1934: 162 [type<br />
collection: Williamsville, Wayne County, Missouri, Russell<br />
(W, lectotype; isolectotype in MICH)].<br />
DEScRIPTIoN.-Tha11us closely adnate on rock<br />
(rarely on bark), pale smoke gray to light mineral<br />
gray, 3-6 cm broad; lobes irregularly branched,<br />
sublinear, contiguous, 1-2 mm wide, <strong>the</strong> margins<br />
dissected, sparsely ciliate, especially in axils, <strong>the</strong><br />
cilia black, simple, to 0.3 mm long; upper surface<br />
densely isidiate, <strong>the</strong> isidia cylindrical, sometimes
NUMBER 88 39<br />
branched; medulla white to pale dull green-yellow;<br />
lower surface black, sparsely to moderately rhi-<br />
zinate, dark brown and short rhizinate near <strong>the</strong> tips,<br />
<strong>the</strong> rhizines simple. Apo<strong>the</strong>cia adnate, 1-3 mm in<br />
diameter, <strong>the</strong> disc brown, <strong>the</strong> amphi<strong>the</strong>cium isidi-<br />
ate; spores 8,4-5 X 6-10 pm.<br />
CHEmsTRY.-Cortex K + yellow (atranorin); me-<br />
dulla more intensively yellow with color tests<br />
(zeorin, galbinic acid, trace <strong>of</strong> salazinic acid, leuco-<br />
tylin and associated terpenes, and secalonic acid A).<br />
DIsTRIsuTI0N.-Eastern United States.<br />
REMARKs.-Parme~ina obsessa is ano<strong>the</strong>r Acharian<br />
species that had been lost from <strong>the</strong> literature since<br />
1814. Not only were extremely few collections<br />
available, but also many were incorrectly identified.<br />
One in US was listed as “Parmelia prolixa,” and as<br />
recently as 1958 P. obses,ra was identified as Parme-<br />
lia finkii (<strong>Hale</strong>, 1958:82). Recent collecting by<br />
American lichenologists, however, has shown that<br />
it is extremely common on sandstone outcrops in<br />
closed oak forests and can tolerate considerable<br />
shade. As a closely appressed saxicolous lichen it is<br />
not easy to collect. Kurokawa (1968b) correctly<br />
recognizes it as <strong>the</strong> isidiate morph <strong>of</strong> P. galbina. No<br />
comparable isidiate population has evolved in<br />
Japan.<br />
SPECIMENS EXAMINED (Selected).-New Hampshire, Willey<br />
(US): Pennsylvania, <strong>Hale</strong> 16288, 17486; Maryland, <strong>Hale</strong><br />
14422; West Virginia, <strong>Hale</strong> 10311, 10660, 12408, 15003; Kentucky,<br />
<strong>Hale</strong> 13866; Indiana, <strong>Hale</strong> 14182; Illinois, <strong>Hale</strong> 13935,<br />
14022; Michigan, <strong>Hale</strong> 33889; Wisconsin, <strong>Hale</strong> 23529: Minnesota,<br />
<strong>Hale</strong> 23602; Virginia, <strong>Hale</strong> 12018, 18803, Luttrell<br />
3864 (US), Tucker 6588 (US); North Carolina, Culberson<br />
6491, 10418 (DUKE, US), <strong>Hale</strong> 16358; Tennessee, <strong>Hale</strong> 37083;<br />
South Carolina, <strong>Hale</strong> 7698; Georgia, <strong>Hale</strong> 7404, 30605; Alabama,<br />
<strong>Hale</strong> 7098; Louisiana, <strong>Hale</strong> 33862; Arkansas, <strong>Hale</strong><br />
2972, 4030; Kansas, <strong>Hale</strong> 4749; Oklahoma, <strong>Hale</strong> 4886, 5054,<br />
Keck 1493 (US); Texas, <strong>Hale</strong> 5452.<br />
29. <strong>Parmelina</strong> pastillifera, new combination<br />
FIGURE 17e<br />
Parmelia scortea var. pastillifera Harmand, 19103558 [type<br />
collection: Bussang, Vosges, France, Claude1 and Harmand<br />
in <strong>Lichen</strong>es Gallici Exsiccati 491 (H, lectotype; isolectotypes<br />
in BM, BPI)].<br />
Parmelia pastillifera (Harmand) Schubert and Klement,<br />
1966:58.<br />
DEscRIPTIoN.-Thallus largely as in P. tiliacea<br />
(p. 48) except isidia dark tipped, peltate, and<br />
apically flattened (Figures 3e,f, 4b).<br />
CHEMISTRY.-cOrteX K + yellow (atranorin); me-<br />
dulla K-, C+, KC+ red, P- (lecanoric acid).<br />
DIsTRrsuTIoN.-Great Britain and Europe.<br />
REMARKS.-while this species has long been<br />
recognized as a distinct variety (Poelt, 1961:194), it<br />
was only recently raised to species level. Dobson<br />
and Hawksworth (1976) have compared it in detail<br />
with <strong>the</strong> normally isidiate P. tiliacea, finding that<br />
P. pastillifera has, on <strong>the</strong> average, somewhat nar-<br />
rower lobes (2-6 mm) than P. tiliacea (3-8 mm) and<br />
denser rhizines.2 <strong>Parmelina</strong> pastillifera also has a<br />
somewhat smaller, slightly more bluish thallus. The<br />
ecological differences are pronounced, for P. pastil-<br />
Zifera occurred in England west <strong>of</strong> <strong>the</strong> 813 mm<br />
rainfall isopleth and <strong>the</strong> 2OC January mean iso-<br />
<strong>the</strong>rm. This agrees with Schauer’s (1965:80) defini-<br />
tion <strong>of</strong> P. pastillifera as an oceanic species in<br />
central Europe.<br />
SPECIMFNS EXAMINED.-EUrOpe: France, Lambinon in<br />
<strong>Lichen</strong>es Selecti Exsiccati 363 (LD, US); Germany, Schroppel<br />
in <strong>Lichen</strong>es Alpium 143 (US); Austria, Strasser in Crypto-<br />
gamae Exsiccatae Vindobonensi 3062b (US); Yugoslavia,<br />
Vtzda in <strong>Lichen</strong>es Selecti Exsiccati 762 (US).<br />
30. <strong>Parmelina</strong> perisidians<br />
FIGURE 17f<br />
Parmelia perisidians (Nylander) <strong>Hale</strong>, 1974:483.<br />
Parmelia perisidians (h’jlander, 1900:6 [type collection: Ram-<br />
podde, Ceylon, Almquist (S, 1ectot)pe; isolectotype in H,<br />
Nylander herbarium number 35673)l.<br />
Parmelia subsulphuuata Asahina, 1951a:228 [t)pe collection;<br />
Higashi-Shirakawa, Prov. Mino, Japan, Yasue (TNS, lecto-<br />
type)l.<br />
DEscRIPTIoN.-Thallus adnate on bark, greenish<br />
mineral gray, 3-6 cm in diameter; lobes sublinear,<br />
0.5-2 mm wide, <strong>the</strong> marginal cilia mostly in <strong>the</strong><br />
axils; upper surface plane, continuous, densely<br />
isidiate, <strong>the</strong> isidia simple or branched; medulla<br />
sea-foam yellow; lower surface densely rhizinate,<br />
<strong>the</strong> rhizines black, simple or squarrosely branched.<br />
Apo<strong>the</strong>cia adnate, 2-5 mm in diameter, <strong>the</strong> am-<br />
phi<strong>the</strong>cium isidiate; spores 8,5-7 X 7-1 1 pm.<br />
CHEMISTRY.-cOrteX K + yellow (a tranorin); me-<br />
dulla K+, C+ KC+ more intensely yellow, P-,<br />
(zeorin, leucotylin and related terpenes, and<br />
secalonic acid A).<br />
*I wish to thank Dr. Hawksworth for allowing me to see<br />
his manuscript before publication.
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
DIsTRIBuTI0N.-Sou<strong>the</strong>ast Asia from India to<br />
Japan.<br />
REMARKS.-This densely isidiate species is closely<br />
related to P. amagiensis and may represent its isidi-<br />
ate morph, although differences in pigments, so far<br />
unresolved, may exclude this possibility. Both<br />
species have <strong>the</strong> typical “subaurulenta” terpene<br />
series. <strong>Parmelina</strong> perisidians has a broad distribu-<br />
tion in <strong>the</strong> higher elevation forests <strong>of</strong> tropical Asia<br />
and <strong>the</strong> temperate forests <strong>of</strong> Japan.<br />
SPECIMENS ExAMIKED.-India: Karnataka, Patwardhan<br />
74.3283 (Poona, US): Tamil Nadu, <strong>Hale</strong> 40101, 40.102, Togashi<br />
(TNS, US). Sri Lanka: see Kurokawa and Mineta (1973:74)<br />
for records in Ceylon. Thailand: Kurokawa 1818 (TNS, US).<br />
Philippines: Rlountain Province, <strong>Hale</strong> 26166a. Japan: Prov.<br />
Kozuke, Asahina (TNS); Proc. Ifusashi, Asahina (TNS);<br />
Kurokawa 550009 (TKS, LS), 64284, 64285 (TNS); Prov.<br />
Settsu, Togashi (TNS, US).<br />
3 1. <strong>Parmelina</strong> phlyctina<br />
FIGURE 18a<br />
<strong>Parmelina</strong> phlyctina (<strong>Hale</strong>), <strong>Hale</strong>, 1974.483.<br />
Parmelia phlyctina <strong>Hale</strong>, 1959:129 [tjpe collection, Blue<br />
Mountains, Jamaica, lnzshaicg 14908 (MSC, holotjpe; iso-<br />
qpe in US)].<br />
DEscRIPTIoN.-Thallus adnate to loosely adnate<br />
on bark, membranous and fragile, light mineral<br />
gray but turning pink in <strong>the</strong> herbarium if improp-<br />
erly dried, 5-15 cm broad; lobes subirregular,<br />
apically rotund, 5-10 mm wide, <strong>the</strong> margins entire<br />
to lobulate, sparsely ciliate in <strong>the</strong> lobe axils, <strong>the</strong><br />
cilia up to 0.5 mm long; upper surface plane,<br />
smooth to rugose in older parts, upper cortex<br />
fragile and flaking <strong>of</strong>f in small pieces, isidia and<br />
soredia lacking; lower surface black except for a<br />
narrow brown zone at <strong>the</strong> tips, sparcely rhiAnate,<br />
<strong>the</strong> rhizines simple. Apo<strong>the</strong>cia rare, adnate, 3-8 mm<br />
in diameter; spores 8, 3-5 X 7-8 pm.<br />
CHEMIsmY.-Cortex K + yellow (atranorin); me-<br />
dulla K+ yellow turning red, C-, KC-, P+<br />
orange (norstictic and connorstictic acids).<br />
DISTRIBUTION.-hkXiCO and <strong>the</strong> West Indies.<br />
REMARKS.-sinCe my discription <strong>of</strong> this species in<br />
1959, additional collections have been made in<br />
Mexico, Cuba, and o<strong>the</strong>r islands in <strong>the</strong> West Indies.<br />
It occurs in wet upland rain forests or secondary<br />
mist forests. The main distinguishing features are<br />
<strong>the</strong> fragile thallus with a flaking cortex, presence<br />
<strong>of</strong> norstictic acid, and small spores. It is closely re-<br />
lated to isidiate P. antillensis but is not <strong>the</strong> direct<br />
parent morph since P. antillensis has a continuous<br />
cortex. It is not, as implied in my original descrip-<br />
tion, a member <strong>of</strong> Amphigymnia (= Parmotrema)<br />
because <strong>the</strong> thallus is adnate overall, cilia are pro-<br />
duced in <strong>the</strong> axils, and <strong>the</strong> bare zone at <strong>the</strong><br />
margins below is very narrow. Both P. phlyctina<br />
and P. antzllensis, however, constitute an anomolous<br />
element in <strong>Parmelina</strong>, not only because <strong>of</strong> <strong>the</strong><br />
somewhat amphigymnioid lobation but also be-<br />
cause <strong>of</strong> <strong>the</strong> production <strong>of</strong> norstictic acid.<br />
SPECIW ?s Exs~i~\k~.-hfexico: Chiapas, <strong>Hale</strong> 20202 (S,<br />
US), Cuba: Oriente, Imshnug 27174 (MSC, US), 25146, 25148<br />
(MSC), ,\lorton 9394 (US). Jamaica: Imshaug 14614, 14714,<br />
15211, 15362, 15375, 15458, 15502, 15524, 15536 (MSC), 13179,<br />
14727, 15146, 15453, 15513 (MSC, US). Haiti: Sud, Wetmore<br />
3373 (MSC). Dominican Republic: Cordillera Central, Wetmore<br />
3728 (LD, MSC), 3738 (MSC, US). Puerto Rico: Imshaug<br />
29562 (MSC).<br />
32. <strong>Parmelina</strong> pilosa<br />
FIGURE 18b<br />
<strong>Parmelina</strong> pilosa (Stizenlxrger) <strong>Hale</strong>, 1974:483.<br />
Parmeha pilosa Stizenberger, 180: 165 [type collection:<br />
Rhenoster Ricer, Taaibosch Kranz Mountains, Orange<br />
Free State, Union <strong>of</strong> South Africa, Rehmann (ZT, lectotype;<br />
isolectotype in H)].<br />
Parrnelin halansae var. sorediata Muller Argoviensis, 1888a:Z<br />
[tjpe collection: Montevideo, Uruguaj, Arechavaleta (G,<br />
lectotype)].<br />
Parnielia subbalansae Gyelnik, 1931:288 (based on Parmelia<br />
balansae ~ar. sorediata Miiller Argoviensis.<br />
DEscRIPTIoN.-Thallus adnate to loosely attached<br />
on bark, coriaceous, light buff mineral gray, 5-12<br />
cm broad; lobes subirregular, apically subrotund,<br />
<strong>the</strong> marginal cilia coarse, becoming furcate, 0.5-1 .O<br />
mm long; upper surface shiny and heavily white-<br />
maculate, plane to minutely pitted, sorediate, <strong>the</strong><br />
soralia orbicular, about 0.5 mm in diameter, sepa-<br />
rate or coalescing into extensive lamina1 sorediate<br />
areas; medulla white; lower surface black, densely<br />
rhizinate, coarse and fine rhizines intermixed,<br />
simple to sparcely furcate. Apo<strong>the</strong>cia rare, sub-<br />
stipitate; spores not seen.<br />
CHE;MIsTRY.-Cortex K + yellow (atranorin); me-<br />
dulla negative with color reagents (no substances<br />
demonstrated).<br />
DISTRIRLTIoN.-~outh America and sou<strong>the</strong>rn<br />
Africa.
NUMBER 33<br />
FIGURE 18.-Species <strong>of</strong> <strong>Parmelina</strong>: a, P. phlyctina (Imshaug 13179 in US); b, P. pilosa (McOwan<br />
in US); c, P. pruinata (Kurokawa in <strong>Lichen</strong>es Selecti Exsiccati 81 in US); (1, P. quercina (“carporrhizans”<br />
population) (Schroppel and Poelt in <strong>Lichen</strong>es Alpiurn 8 in US); e, P. quercina<br />
(Tavares 1082 in US); f, P. rhytidodes (Kurokawa 58601 in US). (Scale in mm.)<br />
41
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
REMARKs.-<strong>Parmelina</strong> pilosa is <strong>the</strong> sorediate<br />
morph <strong>of</strong> P. consors. It occurs on a variety <strong>of</strong> trees<br />
(Celtis, Erythz’na, Scutia), even on fence posts, and<br />
in open or secondary habitats up to 2000 m eleva-<br />
tion. It must be distinguished carefully from P.<br />
muelleri, which rarely occurs in this range and con-<br />
tains stictic acid.<br />
SPECIMENS EXAMINED.-Ecuador: Pichincha, Wiggins 38945<br />
(COLO). Uruguay: Canelones, Felippone 434 (G); Durazno,<br />
Osorio 2979 (DUKE); Flores, Osorio 3519 (MVM, US);<br />
Florida, Osorio 1299 (F); Minas, Osorio 2049, 2056 (MVM,<br />
US); Soriano, Rosengurth (H). Argentina: Buenos Aires,<br />
Grassi 719 (US), Santesson 75 (S), Schnyder (G), Venturi<br />
2853 (SI); Cordoba, Giardelli 1025 (SI), Petersen 23 (S); Entre<br />
Rios, Santesson 126 (S, US); Salta, Fries 21 (S), James (BM),<br />
Lorentz (M). Chile: Valparaiso, Santesson 3084 (S). Kenya:<br />
Nyanza Province, Maas Geesteranus 4957 (L, US). Rhodesia:<br />
Hoeg (TRH). Union <strong>of</strong> South Africa: Basutoland, K<strong>of</strong>Eer<br />
(LD); Cape Province, Hoeg (LD, TRH), K<strong>of</strong>ler (LD), Mc-<br />
Owan (BM, US); Natal, Hoeg (TRH).<br />
33. <strong>Parmelina</strong> pruinata<br />
FIGURE 18c<br />
<strong>Parmelina</strong> pruinata (Muller Argoviensis) <strong>Hale</strong>, 1974:483.<br />
Parmelia pruinata Muller Argoviensis, 1883:46 [type collec-<br />
tion: Yorke Peninsula, Australia, Tepper (G, lectotype)].<br />
Parmelia tiliacea var. afixa Stirton, 1899:485 [type collection:<br />
Queensland, Australia, Bailey (?) 376 (BM, lectotype)].<br />
DEsCRIPTIoN.-ThaIIus closely adnate on twigs,<br />
ashy white, 1-5 cm broad; lobes sublinear-elongate,<br />
variable, 1-3 mm wide, marginal cilia sparse, mostly<br />
in lobe axils, up to 0.5 mm long; upper surface<br />
plane to convex, rugulose, more or less pruinose;<br />
medulla white; lower surface black, sparsely rhi-<br />
zinate, <strong>the</strong> rhizines black, simple. Apo<strong>the</strong>cia con-<br />
spicuous, adnate, 2 4 mm in diameter, <strong>the</strong> disc<br />
<strong>of</strong>ten pruinose; spores 8,7-8 X 10-12 pm.<br />
CHEMIsTRY.-Cortex K+ yellow (atranorin); me-<br />
dulla K -, C +, KC + red, P - (lecanoric acid).<br />
DISTRIBUTION.-Australia and New Zealand.<br />
REMARKS.-This species is ra<strong>the</strong>r variable in lobe<br />
configuration and width but typically grows closely<br />
adnate on small twigs and branches <strong>of</strong> trees in arid<br />
scrub land. It has no apparent relation with P.<br />
quercina, <strong>the</strong> only o<strong>the</strong>r nonsorediate, lecanoric<br />
acid-containing species in <strong>the</strong> genus, characterized<br />
by much coarser lobes and white maculae in <strong>the</strong><br />
cortex. Muller mentions “subtus alba” in his origi-<br />
nal description, but he was actually describing a<br />
Physcia species growing intermingled with <strong>the</strong> type.<br />
The marginal cilia are sparse and sometimes diffi-<br />
cult to determine. It could, in fact, be misidentified<br />
as a Pseudoparmelia but no corticolous nonisidiate<br />
species <strong>of</strong> that genus with lecanoric acid are pres-<br />
ently known.<br />
SPECIMENS ExAMINED.-Australia: Australian Capital Territory,<br />
Kurokawa in <strong>Lichen</strong>es Rariores Selecti Exsiccati 81<br />
(US), Leuthner (US); New South Wales, Boorman 1387,<br />
L1389 (NSW), Doing (L, US), McVean 6358 (COLO), Weber<br />
and McVean L-47913, L-49886 (COLO, US). Wilson 96<br />
(NSW); Queensland, Gwy<strong>the</strong>rl (BM); South Australia, Rogers<br />
886 (US); Tasmania, Bratt 1380b (US), James 2117 2125<br />
(BM, US), Weymouth 65 (BM, US), 432 (BM); Victoria, Filson<br />
6639, 7004 (US), James 223 (BM, US). New Zealand:<br />
Mason 90 (BM).<br />
34. <strong>Parmelina</strong> quercina<br />
FIGURE 18d,e<br />
<strong>Parmelina</strong> quercina (Willdenow) <strong>Hale</strong>, 1974:483.<br />
<strong>Lichen</strong> quercinw Willdenow, 1787:353 [type collection: Thier-<br />
garten and near Tegel, Germany (not seen; illustrated in<br />
pl. 7: fig. l3)].<br />
Imbricaria quercina (Willdenow) de Lamarck and de Can-<br />
dolle, 1805:389.<br />
Parmelia carporrhizans Taylor, 1847:163 [type collection:<br />
Canary Islands, Lemann (FH-Tayl, lectotype)].<br />
Parmelia sinuosa var. hypothrix Nylander, 1856:301 [type<br />
collection: Pyrenees, France, Nylander (H, lectotype)].<br />
Parmelia atricha Nylander, 1873:271 [type collection: La<br />
Preste, Pyrenees, France, Nylander 16 (H, lectotype (not<br />
seen); isolectotypes in FH, BM)].<br />
Parmelia tiliacea var. hypothrix (Nylander) Muller Argovien-<br />
sis, 1888b:196.<br />
Parmelia revoluta var. grantilata Harmand, 1897:221 [type<br />
collection: Docelles, Vosges, France, Harmand 284 (DUKE,<br />
lecto type)].<br />
Parmelia quercina (Willdenow) Vainio, 1899:279.<br />
Parmelia budapestinensis Gyelnik, 1932b3212 [type collection:<br />
Budapest, Hungary, Timkd (BP, holotype)].<br />
Parmelia carporrhizans f. malicola Gyelnik, 1932b:212 [type<br />
collection: St. George, Hungary (Austria?), Zahlbruckner<br />
(BP. holotype)].<br />
Parmelia yalungana Zahlbruckner, 1934b:206 [type collec-<br />
tion: between <strong>the</strong> Litang and Yalung rivers between Muli<br />
Gomba, Baurong, and Wa-Erh-Dje, Szechuan, China, Rock<br />
16720 (W, lectotype)].<br />
DEscRIPTIoN.-Tha1Ius closely adnate on bark,<br />
rarely on rock, ra<strong>the</strong>r coriaceous, light olive-gray,<br />
sometimes turning olive-buff in <strong>the</strong> herbarium,<br />
5-10 cm broad; lobes dichotomously or irregularly<br />
branched, sublinear, separate or becoming im-<br />
bricate, 1.5-4.5 mm wide, <strong>the</strong> margins smooth to<br />
more or less crenate, <strong>of</strong>ten narrowly black rimmed,
NUMBER 33 43<br />
moderately to densely ciliate, <strong>the</strong> cilia black, simple,<br />
0.2-1 mm long; upper surface smooth and shiny,<br />
faintly to strongly white-maculate, irregularly<br />
rugose and cracked on older lobes, sometimes partly<br />
pruinose, isidia and soredia lacking; lower surface<br />
black, densely rhizinate, <strong>the</strong> rhizines black, shiny,<br />
simple or more rarely squarrose, 1-3 mm long.<br />
Apo<strong>the</strong>cia numerous, substipitate, 1.5-5 mm in<br />
diameter, <strong>the</strong> amphi<strong>the</strong>cium smooth or retrorse-<br />
rhizinaee, <strong>the</strong> disc burnt umber; spores 8, 5-7 x<br />
6-10 pm.<br />
CHEmsmY.--Cortex K + yellow (atranorin); me-<br />
dulla K-, C+ KC+ red, P- (lecanoric acid).<br />
DISTRIBUTION.-califOrnia, Europe, Pakistan, Ne-<br />
pal, eastern Asia, and Australia.<br />
REMARKS.-AS a common member <strong>of</strong> <strong>the</strong> Euro-<br />
pean foliose lichen flora this was one <strong>of</strong> <strong>the</strong> first<br />
species <strong>of</strong> Parmelia to be described. The records<br />
from eastern North America previously identified as<br />
Parmelia quercina are now recognized as <strong>Parmelina</strong><br />
galbina or Hypotrachyna Zivida (Taylor) <strong>Hale</strong>. The<br />
interesting distribution pattern, centered in western<br />
North America and Europe, is typical <strong>of</strong> <strong>the</strong> Medi-<br />
terranean type, although <strong>the</strong> species occurs less<br />
commonly in eastern Asia and Australia. Still this<br />
is a remarkably broader range than its isidiate<br />
morph P. filiacea.<br />
The development <strong>of</strong> rhizines on <strong>the</strong> lower side <strong>of</strong><br />
<strong>the</strong> apo<strong>the</strong>cia has been used to justify a separate<br />
population, called Parmelia carporrhizans. As W.<br />
Culberson (1961: 168) points out, however, virtually<br />
all specimens in California have <strong>the</strong>se rhizines, and<br />
many European specimens identified as P. quercina<br />
are <strong>of</strong>ten found to have <strong>the</strong>m, sometimes only very<br />
sparsely developed (Dobson and Hawksworth, 1976).<br />
This trait does not seem to warrant specific status,<br />
although Schauer (1965:71) found this population<br />
to have a strongly oceanic distribution pattern in<br />
western Europe. The problem will have to be solved<br />
through careful field studies in Europe.<br />
SPECIMFNS EXAMINED (selected).-Without Retrorse Rhizines:<br />
Europe: Denmark, Rostrup (LD); Germany, Schroppel and<br />
Poelt in <strong>Lichen</strong>es Alpiurn 7 (H, LD, US); Switzerland,<br />
Mereschkovsky (US), Seifriz (US); Austria, Eggerth in Flora<br />
Exsiccata Austro-Hztngarica 1542 (US); Czechoslovakia, Vdzda<br />
in <strong>Lichen</strong>es Bohernoslouakiae Exsiccati 201 (LD, US); Hungary,<br />
Tirnkd in <strong>Lichen</strong>o<strong>the</strong>ca 135 (H, LD); Spain, Schindler<br />
4303 (KR, US); Portugal, Tavares 1082 (US). Pakistan: Iqbal<br />
180 (US). Nepal: Stainton 4098 (BM, US). China: Manchuria,<br />
Asahina 28 (TNS); Shensi Prov., Hugh (BM). Japan: Prov.<br />
Hyuga, <strong>Hale</strong> 29650, Kurokawa 65065 (TNS); Prov. Tango,<br />
Asahina (TNS); Prov. Tosa, Kurokawa 64091 (TNS), 64094<br />
(TNS, US). Australia: New South Wales, Du Rietz 577b<br />
(UPS, US), Flockton 884 (US).<br />
Mostly with Retrorse Rhizines on <strong>the</strong> Apo<strong>the</strong>cia: United<br />
States: See W. Culberson (1961:172) for a list <strong>of</strong> specimens<br />
examined and a distribution map. Europe: England, Borrer<br />
(US), Crombie (BM, US), Holl 15 (BM), James 75 (BM):<br />
France, Crozals (US), Rondon in <strong>Lichen</strong>es Selecti Exsiccati<br />
268 (US); Spain, Bauza (LD); Portugal, Tavares in <strong>Lichen</strong>es<br />
Lusitaniae Selecti Exsiccati 217 (CS); Germany, Poelt in<br />
<strong>Lichen</strong>es Alpiurn 8; Switzerland, Barkman 4294a (UPS): Austria,<br />
Stezner in Crjptogamae Exsiccatae Vindobonensi 4134<br />
(F, NY); Italy, Alrnborn (LD), Baunigartner in Cryptogarnae<br />
Exsiccatae Vindobonensi 3165 (NY, US). Tunisia: Runeinaik<br />
(LD).<br />
35. <strong>Parmelina</strong> rhytidodes, new species<br />
FIGURE 18f<br />
DEscfuPTIo;u.-Thallus adnatus ve1 laxe adnatus,<br />
corticola et saxicola, pallide olivaceo-cinereus, 8-12<br />
cm latus, lobis subirregularibus, imbricatis con-<br />
gestisque, 2-4 mm latis, margine irregulariter<br />
ciliatis, ciliis nigris, 0.3-0.6 mm longis; superne<br />
nitidus vel apicem versus albo-pruinosus, primum<br />
planus sed mox dense et omnino rugosus (Figure<br />
6b), rugis non eruptentibus, cortice integro vel rare<br />
rumpente; cortex superior 12-14 pm crassus, epicor-<br />
ticatus, stratum gonidiale 14-16 pm crassum, me-<br />
dulla pallide salmonea, 110-150 pni crassa, cortex<br />
inferior brunneus, paraplectenchymatus, 15-17 pm<br />
crassus; subtus niger, dense rhizinosus, rhizinis nigris<br />
vel apice brunneis, simplicibus vel sparse squarroso-<br />
ramosis. Apo<strong>the</strong>cia numerosa, sessilia vel substipi-<br />
tata, disco plano, 3-8 mm diametro; sporis 8,<br />
9-11 X 12-14,um.<br />
CHEMIsmY.-Cortex K + yellow (atranorin); me-<br />
dulla more intensively yellow with color tests<br />
(zeorin, leucotylic acid and related terpenes, seca-<br />
Ionic acid A, and traces <strong>of</strong> unidentified substances).<br />
HOLoTYPE.-Amagi Pass, Prov. Izu, Japan, S.<br />
Kurokawa 58601, 1 December 1958 (US, holotype;<br />
isotype in TNS).<br />
DIsTRIBuTIoN.-Japan and Nepal.<br />
REMARKS.-After I had examined specimens iden-<br />
tified as Pa? melina ento<strong>the</strong>iochron with thin-layer<br />
chromatography, <strong>the</strong>re remained a small group with<br />
<strong>the</strong> “aurulenta” terpene pr<strong>of</strong>ile. On fur<strong>the</strong>r study<br />
I found that <strong>the</strong>se are morphologically different<br />
from typical P. ento<strong>the</strong>iochroa in that <strong>the</strong> ridges<br />
which develop very densely do not burst open or
44 SSIITHSONIAX CONTRIBUTIONS TO BOTANY<br />
flake <strong>of</strong>f. While P. rhytidodes can be recognized<br />
with practice from <strong>the</strong> external morphology alone,<br />
a chemical test is desirable for confirmation. The<br />
species is probably not as common in Japan as P.<br />
ento<strong>the</strong>iochroa, but I have not had <strong>the</strong> opportunity<br />
to re-examine <strong>the</strong> numerous specimens in TNS with<br />
chromatography. The collections made by Poelt in<br />
Nepal are tentatively placed here. They are very<br />
densely rugose and have crowded lobes, modified<br />
ecologically by <strong>the</strong> high exposed elevation where<br />
<strong>the</strong>y were collected. <strong>Parmelina</strong> rhytidodes is appar-<br />
ently an <strong>of</strong>fshoot with sorediate P. aurulenta from<br />
P. irrugans or a now extinct parent morph similar<br />
to it.<br />
SPECIMENS ExAMINED.-Japan: Prov. Awa, Kurokawa 56554<br />
(TSS, US); Prov. Hyuga, <strong>Hale</strong> 29602; Hiroshima Prefecture:<br />
Nakanishi and Oshio 5731 (US); Kanto, Nurnariri 156 (US).<br />
Nepal: Poelt L-747, L-754 (M, US).<br />
36. <strong>Parmelina</strong> schindleri, new species<br />
FIGURE 19a<br />
DEscRIPrIoN.-ThallUs adnatus vel appressus,<br />
fragilis, corticola, pallide albo-cinereus, 2-4 cm<br />
latus, lobis sublinearibus, brevibus, contiguis, ca.<br />
1-1.5 mm latis, margine ciliatis, ciliis 0.2-0.4 nim<br />
longis, simplicibus, margine et pro parte superficie<br />
lobulatis, lobulis congestis, suberectis, ramosis, 0.1-<br />
0.2 mm latis et usque ad 1 mm longis, margine<br />
breve ciliatis; superne nitidus, planus; cortex supe-<br />
rior 14-15 pm crassus, epicorticatus, stratum gonidi-<br />
ale 12 pm crassum, medulla alba, 50-65 pm crassa,<br />
cortex inferior paraplectenchymatus, 12 pm crassus;<br />
subtus niger, modice rhizinosus, rhizinis nigris, sim-<br />
plicibus vel sparse furcatis. Apo<strong>the</strong>cia rara, sessilia,<br />
margine crenato, 3-4 mm diametro, sporis 8:nae,<br />
10 X 16-18pm.<br />
CHEMIsrRY.-Cortex K + yellow (atranorin); me-<br />
dulla K-, C-, KC+ rose, P- (traces <strong>of</strong> gyro-<br />
phoric acid and <strong>the</strong> “horrescens” unknown).<br />
HoLoTYPE.-Caraca, h has Gerais, Brazil, I/ainio<br />
in <strong>Lichen</strong>es Brasilienses Exsiccati 1284 (BM; isotypes<br />
in FH, RI, TUR, and UPS).<br />
DIsTRIBuTIoN.-Brazil.<br />
RExmKs.-The chemical constituents place this<br />
species in <strong>the</strong> “horrescens” group, where it is <strong>the</strong><br />
lobulate morph <strong>of</strong> a parent species represented by<br />
P. damaziana or a now extinct progenitor similar<br />
to it. The dense lobules are easily recognized (Fig<br />
Lire 4e). They do not originate from isidia and for<br />
this reason P. schindleri cannot be considered as<br />
a modified form <strong>of</strong> P. horrescens, which may occa-<br />
sionally have cylindrical and flattened isidia inter-<br />
mixed. The species is named in honor <strong>of</strong> Dr. H.<br />
Schindler, <strong>the</strong> first lichenologist to collect <strong>the</strong><br />
species since Vainio in 1885, although this region<br />
has been visited by many lichen collectors. Vainio<br />
had misidentified his exsiccate number 1284 as<br />
Parmelia coronata var. isidiosa Rluller Argoviensis<br />
(= Bztlbothrix fungicola (Lynge) <strong>Hale</strong>).<br />
SPFCIMENS ExAMIxED.-Brazil: Rio de Janeiro, Schindler<br />
4569,4577 (KR, US).<br />
37. <strong>Parmelina</strong> simplicior<br />
FIGURE 19b<br />
<strong>Parmelina</strong> simplicior (<strong>Hale</strong>) <strong>Hale</strong>, 1974:483.<br />
Parrnelia sirnplicior <strong>Hale</strong>, 1972a:99 [type collection: Panch-<br />
gani, \Vestern Ghats, India, D. D. Awasthi 4056 (US, holo-<br />
type; isotype in Awasthi herbarium)].<br />
DEscRIPTIoN.-Thallus adnate on bark, buff min-<br />
eral gray, coriaceous, 8-10 cm broad; lobes elongate,<br />
more or less subirregular, contiguous and becoming<br />
imbricate, 3-5 mm wide, <strong>the</strong> axils sparsely ciliate,<br />
<strong>the</strong> cilia 0.5 mm long; upper surface plane, contin-<br />
LIOLIS, emaculate, isidia and soredia lacking; medulla<br />
white; lower surface black, sparsely to moderately<br />
rhizinate, <strong>the</strong> rhizines simple, black. Apo<strong>the</strong>cia<br />
common, sessile, 3-6 mm in diameter; spores 8,<br />
4 X 6pm.<br />
CHEMIsmY.--Cortex K + yellow (atranorin); me-<br />
dulla K+ yellow turning red, C-, KC-, P+<br />
orange (salazinic acid).<br />
DISTRIBUTIoN.-~outh India.<br />
REMARKs.-<strong>Parmelina</strong> simphior has a ra<strong>the</strong>r<br />
lea<strong>the</strong>ry thallus and very sparsely developed axil-<br />
lary cilia. As with many o<strong>the</strong>r Indian lichens it<br />
does not exhibit clear-cut traits that enable one to<br />
place it immediately in a particular genus. I had<br />
mentioned a possible relationship to isidiate P.<br />
wallichiana, which has much larger spores and a<br />
more membranous thallus, but <strong>the</strong>se two species are<br />
not at all related. Pal-melina simplicior grows on<br />
roadside trees in <strong>the</strong> Western Ghats region <strong>of</strong><br />
India where an intense monsoon season from June<br />
to September alternates with a long period <strong>of</strong> almost<br />
total drought. <strong>Parmelina</strong> wallichiana, on <strong>the</strong> o<strong>the</strong>r
NUMBER 33<br />
FIGURE 19.-Species <strong>of</strong> Purmelina: a, P. schindleri (I’ainio 1284 in BM); 6, P. .rirnplicior<br />
(Arcnstlzi 4056 in US); c, P. spathulato (Alnzborn 931B in US); d, P. spurnma (Halt, 42934a);<br />
e, P. subaw-ulenta (<strong>Hale</strong> 43705); f, P. subfatitcens (<strong>Hale</strong> 35091). (Scale in mm.)<br />
45
46<br />
hand, occurs at higher elevations with less climatic<br />
stress.<br />
SPECIMENS ExAMIxED.-lndia: Maharashtra, <strong>Hale</strong> 40004,<br />
40007,40046, 40090,43972.<br />
38. <strong>Parmelina</strong> spathulata<br />
FIGURE 19c<br />
<strong>Parmelina</strong> spathulata (Kurokawa) <strong>Hale</strong>, 19743483.<br />
Parmelia spathulata Kurokawa in <strong>Hale</strong> and Kurokawa,<br />
1964:133 [type collection: Skeleton Gorge, Wynberg, Union<br />
<strong>of</strong> South Africa, Alrnbortz 305 (LD, holotype; isotype in<br />
Wl.<br />
DEsCRIPTIoN.-Thallus adnate on bark, pale<br />
whitish glaucous-green, fragile, 2-5 cm broad; lobes<br />
sublinear, crowded, 1-3 mm wide, <strong>the</strong> marginal<br />
cilia evenly dispersed, about 0.5 mm long; upper<br />
surface plane, continuous, moderately isidiate, <strong>the</strong><br />
isidia initially cylindrical and erect but soon be-<br />
coming procumbent and flattened; lower surface<br />
black, moderately rhizinate, <strong>the</strong> rhizines black,<br />
simple to rarely furcate. Apo<strong>the</strong>cia not seen.<br />
CmhiIsmY.-Cortex K + yellow (atranorin); me-<br />
dulla K-, C+, KC+ rose to red, P- (gyrophoric<br />
acid).<br />
DIsTRIBuTIoN.-South Africa.<br />
REMARKs.-This lobulate-isidiate species (Figure<br />
4d) is obviously a member <strong>of</strong> <strong>the</strong> P. dissecta group.<br />
It occurs only in sou<strong>the</strong>rn Africa and is much less<br />
common than P. dissrcfn, which has smaller, normal<br />
is idia .<br />
SPrcihfEss Ex.iMlsED.-~nion <strong>of</strong> South Africa: Cape Pror-<br />
ince, K<strong>of</strong>ler (LD, US). Additional records from South Africa<br />
are given in <strong>Hale</strong> and Kurokaiva (1964:134).<br />
39. <strong>Parmelina</strong> spumosa<br />
FIGURE 19d<br />
Parmrlina spumosa (Asahina) <strong>Hale</strong>, 1974:483.<br />
Parinelia spz~mosa Asahina, 1951b:259 [type collection:<br />
Higashi-Murayama, Kita-Tatna-gun, Prov. Musashi, Japan,<br />
Asahina (TNS, lectotype)].<br />
DEsCRIPTIoN.-Thallus closely adnate on bark,<br />
fragile, pale olive gray, 2-6 cm in diameter; lobes<br />
sublinear, 0.5-2 mm wide, <strong>the</strong> marginal cilia dis-<br />
tinct and evenly dispersed, about 0.5 mm long;<br />
upper surface plane, continuous, emaculate, densely<br />
pustulate-isidiate, <strong>the</strong> pustules bursting but not<br />
becoming sorediate; medulla faintly yellow; lower<br />
SMITHSONIAN COSTRIBUTIONS TO BOTANY<br />
surface moderately rhizinate, black, <strong>the</strong> rhizines<br />
simple or furcate. Apo<strong>the</strong>cia rare, adnate, 1-3 mm<br />
in diameter, <strong>the</strong> amphi<strong>the</strong>cium pustulate; spores 8,<br />
7-8 X 12-14,um.<br />
CHmiIsTRY.-Cortex K + yellow (atranorin); medulla<br />
K-, C+ rose, KC+ red, P-, (gyrophoric<br />
acid, an unidentified pigment, and frequently a<br />
white fluorescent spot).<br />
DIsTRInuTroN.-Pantropical at higher elevations.<br />
REMARKS.-T~~S widespread but ra<strong>the</strong>r rare<br />
species is characterized by <strong>the</strong> dense, erupting but<br />
nonsoretliate pustules (Figure 5e,f) and <strong>the</strong> pale<br />
yellowish medulla. The main constituent, gyrophoric<br />
acid, places it in <strong>the</strong> P. dissecta group, but<br />
it probably represents a different line <strong>of</strong> evolution<br />
from a now extinct parent morph (Figure 9). The<br />
chemistry is somewhat aberrant from P. dissecta and<br />
related species judging by <strong>the</strong> presence <strong>of</strong> a white<br />
UV-fluorescent spot in more than half <strong>of</strong> <strong>the</strong> specimens<br />
(including <strong>the</strong> lectotype) from both <strong>the</strong> New<br />
TVorltl and <strong>the</strong> Old World. The spot is not reactive<br />
with H2S0, and its identity is not known.<br />
<strong>Parmelina</strong> spurnosa has ecological requirements<br />
similar to those <strong>of</strong> P. dissecta and P. horrescens.<br />
Much less common at temperate latitudes than <strong>the</strong><br />
latter two, it grows throughout <strong>the</strong> tropics on<br />
exposed trees (hardwoods and conifers) and more<br />
rarely on rocks in forests up to 2500 m elevation.<br />
SPEcIhfENs Ex.AarIxm.-Mexico: Chiapas, <strong>Hale</strong> 19897, 20110a,<br />
20150. Cuba: Hioram 6682 (US). Jamaica: Imshaug 14279,<br />
15646 (MSC, US). Colombia: Antioquia, Nee and Mori<br />
4258a (US). Venezuela: Merida, <strong>Hale</strong> 42957, 42934a, 43037a.<br />
Brazil: Sao Paulo, Eiten and Mimura 5738 (US). Chile:<br />
Chiloe, Santesson 2264 (S, US): Valdivia, Santesson 7066 (S).<br />
St. Helena: Loveridge (AM). Union <strong>of</strong> South Africa: Cape<br />
Province, Almborn 749, 2663 (LD), 3966 (LD, US), Degelius<br />
SA-403 (Degelius herbarium, US); Natal, Almborn 10069<br />
(LD). Madagascar: Lemaitre (H). Indonesia: Java, Groenhart<br />
2218 (L), Neemoort 3354 (BOR), Oka 4089 (L). India: Tamil<br />
Nadu, <strong>Hale</strong> 43829. Japan: Prov. Aki, <strong>Hale</strong> 29529, 29540; Prov.<br />
Hizen, Kurokawa 63159 (TNS): Prov. Izu, Asahina (TNS):<br />
Prov. Mino, Asahiim (TNS); Prov. Ohmi, <strong>Hale</strong> 29472, 29482;<br />
Prov. Sagami, Asahina 40 (TNS), Kurokawa 58038 (TNS, US);<br />
Prov. Shim<strong>of</strong>usa, Asahina (TNS); Prov. Survo, Asahina 125<br />
(TNS). Taiwan: Kurokawa 494 (TNS). Australia: Victoria,<br />
Johnson (BM). New Zealand: Knight (UPS), Martin 493, 566<br />
(AM).<br />
40. <strong>Parmelina</strong> subaurulenta<br />
FIGURE 19e<br />
<strong>Parmelina</strong> siibaurulenta (Nylander) <strong>Hale</strong>, I974:483.
NUMBER 33 47<br />
Parrnelia subaurulenta Nylander, 1885:606 [type collection:<br />
Narkanda, 5. W’. Himalayas, India, Skoliczka (H, Nylander<br />
herbarium number 35672, lectotype)].<br />
Parmelia homogenes Nylander, 38853607 [type collection:<br />
India, Hooker and Thotnson 1942 (H, Nylander herbarium<br />
number 35664, lectotype)].<br />
Parmelia conspicita Hue, 1899: 1/15 [type collection: Kiao Che-<br />
Tung, above Kiang-l’n, Yunnan, China, Delavay (PC,<br />
lectotype)].<br />
Parrnelia homalotera Hue, 1899:159 [type collection: above<br />
Mo-So-Yn, Yunnan, China, Delavay (PC, lectotype)].<br />
Parrnelia fecrrnrla Hue, 1899:169 [type collection: Ta-Pin-Tze,<br />
Yunnan, China, Delavay 5 (PC, lectotype)].<br />
Parmelia srcbcrernea Zahlbruckner, 1934b:208 [type collection:<br />
Betiveen Muli Gomba, Baurong, and T\’a-Erh-Dje, Setch-<br />
Ivan, China, Rock 16720 pro parte (I%’,<br />
lectotype)].<br />
Parmelia subaurulenta var. myriocarpa Asahina, 19.5la:227<br />
[type collection: Mt. Koya, Prov. Kii, Japan, Inuinaru<br />
1232 (TSS, lectotype)].<br />
Parmelia homogenes f. mitior Asahina, 1952:78 [type collectioil:<br />
Yamanaka, Prov. Kai, Japan, Asahitla (TNS.<br />
lect o t ype)].<br />
Parmelia rnyriocarpa (Asahina) Chao, 1964:149.<br />
Parnielina hotnoge?res (Sylander) <strong>Hale</strong>, 1974:482<br />
DE~CRIPTION.-T~~~~L~S<br />
adnate to closely adnate on<br />
bark, light greenish mineral gray, ra<strong>the</strong>r fragile,<br />
4-1 0 cm broad; lobes sublinear, subimbricate, 2-4<br />
mm wide, <strong>the</strong> marginal cilia quite dense in <strong>the</strong><br />
axils, irregularly dispersed at <strong>the</strong> lobe tips, 0.4-0.7<br />
mm long; upper surface plane, faintly maculate,<br />
shiny, isidia and soredia lacking: medulla pale<br />
orange-yellow; lower surface black, densely rhizinate,<br />
<strong>the</strong> rhizines simple or becoming squarrosely<br />
branched. Apo<strong>the</strong>cia very numerous, adnate, 1-3<br />
mm in diameter; spores 8, 6-10 X 8-14 pm, usually<br />
abundantly developed<br />
CHExmTRY.-Cortex K + yellow (atranorin); medulla<br />
more intensively yellow with color tests<br />
(zeorin, leucotylin and associated terpenes, secalonic<br />
acid A, and traces <strong>of</strong> unidentified substances).<br />
DIsTRIiivTIoN.-Eastern Asia from India to Japan.<br />
RE:hfARKs.-<strong>Parmelina</strong> su bazirirlentn is characterized<br />
by a thin thallus (average thickness for 20 specimens<br />
154 pm), numerous apo<strong>the</strong>cia rarely exceeding<br />
3 mm in diameter, and well developed spores 6-9 x<br />
8-12 pm. Nylander (1885:60i) described Paimelin<br />
homogenes at <strong>the</strong> same time on <strong>the</strong> basis <strong>of</strong> a single<br />
specimen, citing as <strong>the</strong> main difference spore size<br />
(“similis siibazmilenta . . . sed sporis majoribus”).<br />
Nylander measured spores 8-10 X 14-16 pm but my<br />
own measurements on <strong>the</strong> type specimen gave<br />
7-8 X 12-13 pm, putting it well within <strong>the</strong> range<br />
<strong>of</strong> spore size for Pal-rnelina subaurulenta. Without<br />
a significant difference in spore size, P. homogenes<br />
must be regarded as a synonym <strong>of</strong> P. siibauixlenta.<br />
The species most <strong>of</strong>ten confused with P. sitbaum-<br />
lenfn, especially outside <strong>of</strong> India, is P. if-rugans,<br />
which has consistently larger apo<strong>the</strong>cia (up to 10<br />
mm in diameter), somewhat larger spores, and <strong>the</strong><br />
“aurulenta” terpene series. T’wo <strong>of</strong> Nylander’s<br />
syntypes for Parmelia sztbaurulenta are, in fact,<br />
P. imgans, as discussed above under that species.<br />
A11 specimens should be tested with chromatog-<br />
raphy, although diameter <strong>of</strong> <strong>the</strong> apo<strong>the</strong>cia is <strong>of</strong>ten<br />
adequate to separate <strong>the</strong> species in this group. All<br />
type specimens listed in <strong>the</strong> synomymy above were<br />
tested with thin-layer chromatography except for<br />
Pnmelia coiispiczta Hue, which is placed here on<br />
<strong>the</strong> basis <strong>of</strong> <strong>the</strong> small apo<strong>the</strong>cia.<br />
SPECIMEKS Ex.iMIs.ro.-Sepal: Poelt 156 (hf). India: Sikkim,<br />
Hnra et al. (TSS, US); Tamil Sadu, Degelius As352<br />
(Degelius herbarium), <strong>Hale</strong> 40214, 43705, 43761, 43834,<br />
Perinftet (H, Nylander herbarium number 35670, syntype <strong>of</strong><br />
Parrnelia srcbaurdeiita), Watt 119 (BM): TTest Bengal,<br />
.4wnsthi 3919, 3921 (Aivasthi herbarium. US). Sri Lanka:<br />
Almquist (H, Nylander herbarium number 35665). China:<br />
Yunnan, Handel-Mazzrtti 89, 2453, 2453 (US, JV), Rock 11747<br />
(US). Taiivan: Kntrokawn 873, 1208 (TSS). Japan: Afoseley<br />
(H, Nylander herbarium number 35669, syntype <strong>of</strong> Parmelia<br />
s7tbaurulen/n); .4lrnquist<br />
36671).<br />
(H, Nylantler herbarium number<br />
41. <strong>Parmelina</strong> subfatiscens<br />
FIGURE 19f<br />
Parinelina subfatiscens (Kurolawa) <strong>Hale</strong>, 1974:483.<br />
Parmelfa siibfafiscens Kuroka\\a in <strong>Hale</strong> and Kurokawa,<br />
1964:134 [t)pe collection: Louis Trichartlt, Zoutpansberg,<br />
Transiaal, Union <strong>of</strong> South Ahica, Al/nborn 6443 (LD,<br />
holotype; isotype in US)].<br />
DEscRIPTIos.-ThallLis closely adnate on bark,<br />
fragile, whitish mineral gray, 3-5 cm broad; lobes<br />
sublinear-elongate, separate to contiguous, 03-15<br />
mm wide, <strong>the</strong> marginal cilia distinct, simple, to 1.0<br />
mm long: upper surface plane, shiny, continuous,<br />
pustulate laminally and subterminally (Figure 5c),<br />
exposed medulla in center <strong>of</strong> pustules turning black,<br />
pustules remaining entire or producing very coarse<br />
sorediate or isidiate-sorediate masses, <strong>of</strong>ten short<br />
ciliate in African material but bery rarely so in <strong>the</strong><br />
New 1Yorld; medulla white; lower surface black,<br />
densely rhizinate, <strong>the</strong> rhizines simple, black.<br />
Apo<strong>the</strong>cia rare, adnate, 1.5-4.0 mm in diameter;<br />
spores 8,8-9 X 12-14 pm.
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
CHEMISTRY.-cOrteX K + yellow (atranorin); me-<br />
dulla K -, C -, KC + rose, P - (trace <strong>of</strong> gyrophoric<br />
acid and <strong>the</strong> “horrescens” unknown substances).<br />
DISTRIBUTIoN.-~outh Africa and <strong>the</strong> Caribbean<br />
region.<br />
REMARKS.-The pustules <strong>of</strong> P. subfatiscens (Fig-<br />
ure 5c) <strong>of</strong>ten erupt, leaving a central area <strong>of</strong> exposed<br />
medulla that blackens. O<strong>the</strong>r pustules become<br />
coarsely sorediate with <strong>the</strong> soredia sometimes as-<br />
suming <strong>the</strong> form <strong>of</strong> tiny isidia-like masses. The<br />
specimens from Africa usually produce short cilia<br />
on <strong>the</strong>se isidioid masses and on <strong>the</strong> surface <strong>of</strong> <strong>the</strong><br />
pustules. The tropical American population very<br />
rarely has any cilia. The chemistry <strong>of</strong> <strong>the</strong> neotrop-<br />
ical and African populations is, as far as I can<br />
determine, within <strong>the</strong> limits <strong>of</strong> <strong>the</strong> solvent systems<br />
available, identical and equivalent to that in P.<br />
horrescens. Dey (1975:433) suggests that <strong>the</strong> neo-<br />
tropical specimens, along with some from <strong>the</strong><br />
sou<strong>the</strong>rn Appalachian Mountains in <strong>the</strong> United<br />
States, may represent a new species, based in part<br />
on <strong>the</strong> absence <strong>of</strong> cilia on <strong>the</strong> pustules. It is also<br />
true that <strong>the</strong> ecological requirements seem to be<br />
different. The neotropical specimens were all col-<br />
lected in upland virgin rain forest at or above 500<br />
m elevation, whereas <strong>the</strong> African localities are in<br />
much drier forests. I prefer to keep a broader spe-<br />
cies concept until more complete morphological and<br />
ecological data can establish <strong>the</strong> case one way or <strong>the</strong><br />
o<strong>the</strong>r.<br />
SPECIMFUS EXA\IINID.-Panama. Code, <strong>Hale</strong> 43564. See <strong>Hale</strong><br />
(1971a:24) for additional records from Dominica.<br />
42. <strong>Parmelina</strong> swinscowii<br />
FIGURE 20a<br />
<strong>Parmelina</strong> swinscowii (<strong>Hale</strong>) <strong>Hale</strong>, 1974:483.<br />
Parmelia swinscowii <strong>Hale</strong>, 1973b:1 [type collection: Mt. Kenja,<br />
Central Province, Kenya, T. D. V. Swinscow K 31/33<br />
(BM, holotype; isotype in US)].<br />
DEscRIPTIoN.-Thallus growing on soil or over<br />
mosses on soil, loosely attached, whitish mineral<br />
gray or darkening, fragile, 3-5 cm broad; lobes<br />
sublinear-elongate, imbricate, 1.5-2.0 mm wide, <strong>the</strong><br />
margins sparsely ciliate, <strong>the</strong> cilia black, simple,<br />
about 0.5 mm long; upper surface plane to convex,<br />
shiny, becoming rugose and cracked toward <strong>the</strong><br />
lobe tips, <strong>the</strong> tips remaining entire or becoming<br />
minutely dissected, turning coarsely pustulate-<br />
sorediate; medulla white; lower surface black,<br />
densely rhizinate, <strong>the</strong> rhizines black, simple.<br />
Apo<strong>the</strong>cia unknown.<br />
CHEMIsTRY.-Cortex K + yellow (atranorin); me-<br />
dulla K-, C-, KC+ rose, P- (lobaric acid and<br />
salazinic acid).<br />
REMARKS.-sinCe describing P. swinscowii, 1 have<br />
identified it among collections made by Rolf<br />
Santesson in sou<strong>the</strong>rnmost Chile. The Chilean and<br />
African populations are identical in morphology<br />
and chemistry. In Kenya P. swinscowii grows in <strong>the</strong><br />
alpine zone at over 3500 m elevation, and in Chile<br />
it occupies similar barren habitats on rock and soil<br />
near sea level.<br />
As mentioned in <strong>the</strong> original description, P.<br />
swinscowii does not fit well into any <strong>of</strong> <strong>the</strong> parmeli-<br />
oid groups. It superficially resembles an “evernii-<br />
form” Parmelia, such as sorediate, salazinic acid-<br />
containing Everniastwm sorocheilum (Vainio) <strong>Hale</strong>,<br />
but <strong>the</strong> rhizines are relatively short and <strong>the</strong> lobes<br />
not at all canaliculate. It intergrades to a certain<br />
extent with abnormally small forms <strong>of</strong> Parmotrema<br />
cetratum (Acharius) <strong>Hale</strong> or P. reticulatum (Taylor)<br />
Choisy, except that <strong>the</strong> upper surface is not reticu-<br />
late and <strong>the</strong> lobes are sublinear. The species is<br />
probably best left in <strong>Parmelina</strong> where it is<br />
anomolous largely because <strong>of</strong> <strong>the</strong> presence <strong>of</strong> lobaric<br />
acid.<br />
SPECIVEM EXAhZISED.-chile: Magallanes, Santesson 2159,<br />
6381a, 6381b, 6478 (S, US); Tierra del Fuego, Santesson 1244<br />
(S, US). Specimens from Kenya are listed in <strong>Hale</strong>, 1973b:4.<br />
43. <strong>Parmelina</strong> tiliacea<br />
FIGURE 20b<br />
<strong>Parmelina</strong> tiliacea (H<strong>of</strong>fmann) <strong>Hale</strong>, 1974:483.<br />
<strong>Lichen</strong> tiliaceus H<strong>of</strong>fmann, 1784:96, pl. 16: fig. 2 [type collection:<br />
Europe (not seen but illustrated in H<strong>of</strong>fmann)].<br />
<strong>Lichen</strong> scorteus Acharius, 1798:119 [type collection: Sweden<br />
(H-Ach, lectotype)].<br />
Parmelin tiliacea (H<strong>of</strong>fmann) Acharius, 1803:215.<br />
Zmbricaria tiliacea (H<strong>of</strong>fmann) Koerber, 1855:70.<br />
Parmelia quercifolia var. scortea f. microphylla Massalongo,<br />
1856: 175 [type collection: Camp<strong>of</strong>ontana, Verona, Italy,<br />
Massalongo in <strong>Lichen</strong>es Ital. 329 (UPS, lectotype)].<br />
Parmelia scortea var. papillosa Gylenik, 1931:284 [type collection:<br />
Ndgrad, Hungary, Foriss 599 (BP, holotype)].<br />
Pnrmelia scortea var. ramificn Gyelnik, 1931 :285 [type collection:<br />
S6sko Mountains, Hungary, Timkd 4521 (BP,<br />
holotype)].<br />
Parmelia scortea f. visegradensis Gyelnik, 1932a:444 [type<br />
collection: Visegrld, Pest, Hungary, Timkd (BP, holotype)].
NUMBER 33<br />
49
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
DEscRIPTION.-Tha1lus adnate on bark or rock,<br />
light mineral gray to mineral gray, turning olive-<br />
buff to deep olive-buff in <strong>the</strong> herbarium, 5-15 cm<br />
or more in diameter: lobes irregularly branched,<br />
sublinear-elongate, <strong>of</strong>ten imbricate, rounded at <strong>the</strong><br />
apices, 2-6 mm wide, <strong>the</strong> margins more or less<br />
crenate and undulate, sometimes narrowly black<br />
rimmed, ciliate, <strong>the</strong> cilia black, coarse, shiny, simple,<br />
0.2-0.7 mm long; upper surface more or less shiny,<br />
white maculate, usually partly pruinose, irregularly<br />
cracked on older lobes, densely isidiate, <strong>the</strong> isidia<br />
cylindrical, short, rarely branched, usually black-<br />
ening at <strong>the</strong> tips; medulla white; lower surface<br />
black, moderately to densely rhizinate, rhizines<br />
black, simple, 1-2 mm long. Apo<strong>the</strong>cia rare, adnate,<br />
sometimes retrosely rhizinate, to 4 mm in diameter;<br />
spores 8, 5-6 X 8-11 pm.<br />
CHEMISTRY.-cOrteX K + yellow (atranorin); me-<br />
dulla K-, C+, KC+ red, P- (lecanoric acid).<br />
DIsTRIBuTIoN.-Europe to western India.<br />
REMARKS.-ThiS classical European species was<br />
<strong>the</strong> first <strong>Parmelina</strong> to be described. Except for <strong>the</strong><br />
nomenclatural confusion with Acharius’ Parmelia<br />
scortea, it has been correctly identified by all<br />
lichenologists, excluding, <strong>of</strong> course, those in Amer-<br />
ica, who used <strong>the</strong> name for <strong>Parmelina</strong> galbina.<br />
Hypotrachyna liuida (Taylor) <strong>Hale</strong>, and o<strong>the</strong>r<br />
narrow-lobed lichens.<br />
<strong>Parmelina</strong> tiliacea is <strong>the</strong> isidiate morph <strong>of</strong> P.<br />
q uercina, which, surprisingly, has a much broader<br />
geographic range, contrary to <strong>the</strong> situation for most<br />
species pairs. A close relative in Britain and Europe<br />
is P. pastillifera, which has peltate isidia and differ-<br />
ent ecological requirements (Dobson and Hawks-<br />
worth, 1976). Taken toge<strong>the</strong>r, <strong>the</strong> three species<br />
form a distinct group in <strong>Parmelina</strong> characterized<br />
by <strong>the</strong> white maculae and presence <strong>of</strong> lecanoric<br />
acid. Only one o<strong>the</strong>r species in <strong>the</strong> genus, P.<br />
pruinata, contains this acid.<br />
As one <strong>of</strong> <strong>the</strong> commonest parmelioid lichens in<br />
Europe, P. tiliacea has been mentioned and stud-<br />
ied in detail by many workers. Sernander-Du Rietz<br />
(1926), for example, found that it tends to occur<br />
in ornithocoprous or dust-laden habitats in Scan-<br />
dinavia. She later conducted a careful study <strong>of</strong><br />
apo<strong>the</strong>cial formation, showing that growth <strong>of</strong> new<br />
apo<strong>the</strong>cia could be correlated with heavy precipita-<br />
tion in warm periods <strong>of</strong> <strong>the</strong> summer (Sernander-Du<br />
Rietz, 1957).<br />
SxczcIhfEvs EXAMINED (selected) .-Europe: England, Crombie<br />
(BM), Davies (BM), Larbalestzer 292 (BM); Norway,<br />
Hoeg (US), Magnusson 9193 (US); Sweden, Almborn (LD),<br />
Asplund (US), Blomberg in <strong>Lichen</strong>es Sueciae Exsiccati 64<br />
(LD), Santesson 12711 (US), Vrnng in Kqptogamae Exsiccatae<br />
Vindobonensi 3718 (LD, US); Finland, Kari in <strong>Lichen</strong>es<br />
Fenniae Exsiccati 194, 1103 (H), Linkoln in <strong>Lichen</strong>es Fenniae<br />
Ewsiccati 392 (H, US); Poland, Glanc in <strong>Lichen</strong>o<strong>the</strong>ca Polonica<br />
171 (UPS); Russia, in <strong>Lichen</strong>o<strong>the</strong>ca Rossica 17 (WIS), Oxner<br />
(US): Bulgaria, Szatala (US): France, Crozals (US), Rondon<br />
in <strong>Lichen</strong>es Selecti Exsiccati 440 (US); Germany, Erichsen<br />
(US), Hiltmann in Kryptogamae Exsiccatae Vindobonensi<br />
3062 (US). Paul and Schroppel in <strong>Lichen</strong>es Alpium 9 (US);<br />
Czechosloi akia, Pisut in <strong>Lichen</strong>es Slovakiae Exsiccati 73<br />
(US): Spain, Culberson 11954 (DUKE, US); Portugal, Sampaio<br />
in <strong>Lichen</strong>es de Portugal 250 (US); Italy, Steiner in<br />
Ki yptogamae Exsiccatae Vindobonensi 4327 (US). Israel:<br />
Pharaii 11 (US). Tunisia: Runemark (LD). Morocco: Ernst<br />
2206 (US), hrewboula 198 (BM). Bahrain: Ahrnad L162 (L,<br />
US). Pakistan: Iqbal 180 (L), 511,540 (US). India: Kashmir,<br />
Kapoor 991 (Awasthi herbarium), Kaitl 4006 (Awasthi herbarium),<br />
Watt 117, 455 (BM).<br />
44. <strong>Parmelina</strong> usambarensis<br />
FIGURE 2Oc<br />
<strong>Parmelina</strong> usambarensis (Steiner and Zahlbruckner) <strong>Hale</strong>,<br />
1964:483.<br />
Parmelia usambarensis Steiner and Zahlbruckner in Zahl-<br />
bruckner, 1926:524 [type collection: Lutindi, Tanzania,<br />
Brunnthaler (W, lectotype)].<br />
Parmelia lneuigatoides des Abbayes, 1951:970 [type collec-<br />
tion: Fouta-Djalon, Dalaba, Guinea, des Abbayes (RES,<br />
lectotype)].<br />
DEscRIPTIox.-Thallus loosely attached on rock,<br />
whitish mineral gray, to 10 cm broad; lobes broadly<br />
sublinear, divaricate to contiguous, 2-5 mm wide,<br />
<strong>the</strong> marginal cilia irregularly dispersed but mostly<br />
in <strong>the</strong> lobe axils, up to 1 mm long; upper surface<br />
shiny, plane, continuous or cracking in older parts,<br />
sparsely to densely isidiate, <strong>the</strong> isidia cylindrical,<br />
simple to branched, to 0.4 mm high; medulla<br />
white; lower surface black and shiny except for a<br />
narrow dark brown zone at <strong>the</strong> tips, moderately to<br />
sparsely rhizinate, <strong>the</strong> rhizines black, simple, 1 mm<br />
or more long. Apo<strong>the</strong>cia rare, substipitate, <strong>the</strong><br />
amphi<strong>the</strong>cium isidiate, 2-3 mm in diameter; spores<br />
8,6 X 12pm.<br />
REMARKS.-PaYmelina usambarensis is a ra<strong>the</strong>r<br />
large, loosely attached, saxicolous lichen. The isidia<br />
are <strong>of</strong>ten quite sparse. It does not conform perfectly<br />
to <strong>the</strong> concept <strong>of</strong> Paymelina as I define <strong>the</strong> genus,<br />
but <strong>the</strong> simple rhizines and coarse, long marginal
NUMBER 33 51<br />
cilia at least remove it from Hypotrachyna (Vainio)<br />
<strong>Hale</strong>. On <strong>the</strong> o<strong>the</strong>r hand, it cannot be accommo-<br />
dated in Parmotrema LIassalongo because <strong>of</strong> <strong>the</strong><br />
sublinear subdivaricate lobes. There are no close<br />
relatives, except, very superficially, <strong>the</strong> African en-<br />
demic <strong>Parmelina</strong> enormis.<br />
SPECIMFSS Ex \vIsm.-Guinea: Baldruin 9849a (BM) . Ivory<br />
Coast: Man, tles .4bbnyes (LD, REN), Santesson 10632 (S,<br />
US). Uganda: Su’inscow 2U 15/6 (BM, US), Thornas 1678,<br />
3029 (BM). Thailand: Kuroknwa 1804, 1813, 1939, 1956 (TXS),<br />
1958 (TSS, US), Kurokawa in 1-icheria Rariores et Criiici<br />
Exsiccati 41 (US).<br />
45. <strong>Parmelina</strong> versiformis<br />
FIGURE 2Od<br />
Parmeliiza versiformis (Krempelhnber) <strong>Hale</strong>, 1964:483.<br />
Pnvr!ielin uersiformis Krempelhul)er, 1878:464 [type collection:<br />
Argentina, Lorentz crnd Hieronymuy (M, lectotype:<br />
isolectotypes in G, PC)].<br />
Parmeiia inuta/a Vainio, 1890.3‘) [t)pe collection: Sitio,<br />
Minas Gerais, Brazil, Vainio in <strong>Lichen</strong>es Brasilienses<br />
Exsiccati 539 (TUR, 1ectot)pe: isolectotypes in Bhl, FH,<br />
UPS)].<br />
Parmelia catliari?ie,!sis Miiller Argoviensis, 1891:239 [type<br />
colleclion: Near Santa Catarina, Brazil, Uie 1891 (G, lectotype;<br />
isolectotjpe in \V)].<br />
Parmelia weltsteinii Zahlbrnckner, 1909:173 [type collection:<br />
Near Taipas, Sao l’aulo, Brazil, Schiffner and Wettsteiii<br />
(W, lectotype, isolectotype in G)].<br />
DE~CRIPTION.-T~~~~US adnate to loosely adnate<br />
on bark, turning deep olive-buff in <strong>the</strong> herbarium,<br />
about 8 cm in diameter; lobes irregularly branched,<br />
sublinear-elongate, more or less imbricate, 2-8 mm<br />
wide, <strong>the</strong> margins crenate, ciliate mostly in lobe<br />
axils, <strong>the</strong> cilia <strong>of</strong>ten sparse, to 0.5 mm long; upper<br />
surface more or less shiny, not maculate, wrinkled<br />
and irregularly cracked on older lobes, soredia and<br />
isidia lacking; medulla white; lower surface dark<br />
brown or blackening, more or less wrinkled, sparsely<br />
rhizinate, <strong>the</strong> rhizines pale brown, simple, 0.1-0.3<br />
mm long. Xpo<strong>the</strong>cia common, substipitate, 2-10<br />
mm in diameter, <strong>the</strong> rim crenate and undulate, <strong>the</strong><br />
amphi<strong>the</strong>cium rugose, <strong>the</strong> disc blackish brown;<br />
spores 8, 10-14 X 18-28 pm, <strong>the</strong> episporium about<br />
3 pm thick.<br />
CHEmsmY.-Cortex K + yellow (atranorin); medulla<br />
K+ yellow turning red, C-, KC-, P+<br />
orange (salazinic acid).<br />
DISTRIBUTION.-~OUth America.<br />
REMARKS.--This is <strong>the</strong> only species <strong>of</strong> <strong>Parmelina</strong><br />
in <strong>the</strong> New World with salazinic acid, excepting<br />
rare P. min.rcowii. It is also unusual in having very<br />
large spores with a thick episporium as well as a<br />
lower surface that is dark brown to black ra<strong>the</strong>r<br />
than entirely black. The closest relative appears to<br />
be <strong>the</strong> Asian P. expallida, an isidiate species with<br />
smaller spores (Kurokawa, 19G8a: 192). Although P.<br />
veisiformis occurs in a botanically well-known re-<br />
gion, it has not been collected since 1901.<br />
Si~ec~si~,ss Ex.iwAFD.-Brazil: hhas Gerais, r’C’flTr7lifig 283,<br />
289, 286 (M); Rio de Janeiro, B~lrclzell 2409 (BM), Glaziou<br />
1822, 1823 (M).<br />
46. <strong>Parmelina</strong> wallichiana<br />
FIGURES 20e, 21<br />
Parmelinn wallichiniia (Taylor) <strong>Hale</strong>, 1974:483.<br />
Parmelia wallichiana Taylor, 1847:l 76 [type collection: Sepal,<br />
Wnlliciz (FH-TayI, lectotype)].<br />
Parinelia tiliacea rar. fximfa Steiner, 1888: 138 [type collection:<br />
Usambara, Tanzania, Meyer (G, lectotype)].<br />
Parmelia juflodii Steiner, 19073640 [type collection: Sanatorium,<br />
Cape Proiince, Union <strong>of</strong> South Africa, Junod 978<br />
(G, lectot!pe)].<br />
Parmelia nimandaira,la Zahlbrnckner, 1934a:55 [type collection:<br />
Mt. rlrisan, Taiivan, Asahimi 63 (W, lectotype; isolectotype<br />
in TSS)].<br />
Parrnelia nimandairana f. sirblaeui.~ Asahina, 19523131) [type<br />
collection: hlt. Buko, Prov. hfusashi, Japan, Asahiiza 35<br />
(TNS, lectotype)].<br />
DEscRIPTioN.-Thallus adnate to loosely adnate<br />
on bark or rock, yellowish glaucous to pale<br />
glaucous-green, 5-20 cm in diameter; lobes irregu-<br />
larly branched, apically rotund, 3-10 mm wide, <strong>the</strong><br />
margins more or less crenate, lobulate with age,<br />
short ciliate, especially in <strong>the</strong> axils, <strong>the</strong> cilia about<br />
0.5 mm long; upper surface shiny, not maculate,<br />
irregularly cracked on older lobes, sparsely isidiate,<br />
isidia cylindrical, mostly Simply, less than 1 mm<br />
high; medulla white; lower surface black, mod-<br />
erately to sparsely short rhizinate, dark brown and<br />
naked or papillate in a ra<strong>the</strong>r wide Lone near <strong>the</strong><br />
tips, <strong>the</strong> rhizines black, simple. Apo<strong>the</strong>cia rare,<br />
adnate, 2-7 mm in diameter; spores 8, 8-10 x<br />
14-18 Fm.<br />
CHEnmTRu.-Cortex K + yellow (atranorin); me-<br />
dulla K+ yellow turning red, C-, KC-, P+<br />
orange (salazinic acid).<br />
DISTRIBuTIoN.-Africa and Asia (Figure 2 1).<br />
REMARKS.-This is <strong>the</strong> most widespread and com-
52 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
monly collected Paimelina in <strong>the</strong> Old World. It is<br />
easily recognized by <strong>the</strong> adnate, moderately isidiate<br />
thallus that grows in a variety <strong>of</strong> habitats in <strong>the</strong><br />
highlands <strong>of</strong> Africa, India, and eastern Asia, rang-<br />
ing from temperate Japan to both higher elevations<br />
in <strong>the</strong> tropics and to ra<strong>the</strong>r low elevations near<br />
<strong>the</strong> equator in Guinea and <strong>the</strong> Ivory Coast. The<br />
parent morph is no longer extant nor are <strong>the</strong><br />
parallel sorediate or pustulate morphs.<br />
SPECIMENS EXAMINED.-GUinea: N’Zerekore, Santesson 10568b<br />
(UPS, US). Ivory Coast: Man, Santesson 10650 (UPS). Uganda:<br />
Dummer 5064 (US), Swinscow 2U 24/20 (BM, US). Kenya:<br />
Nyanza Province, Maas Geesteratius 10935 (L, LD, US), 4950,<br />
11164 (L); Rift \’alley Province, hfaas Geesteranus 4832 (L).<br />
Angola: Cuanza Sul, Degelius (Degelius herbarium, US).<br />
Rhodesia: Hoeg (TRH). Tanzania: Arusha Province, Santesson<br />
21339 (UPS, US); Kiliuiarijaro Province, Santesson<br />
20961, 20989, 20990 (UPS, US); Moshi District, Burnet T122<br />
(BM, US). Swaziland: Almborn 7901 (LD), K<strong>of</strong>ler (LD). Union<br />
<strong>of</strong> South Africa: Natal, Almborn 8494 (LD), 8683, 9732, 9871<br />
(LD, US), Hoeg (TRH), K<strong>of</strong>ler (LD); Transvaal, Almborn<br />
6430 (LD). Madagascar: des Abbayes in Licheiies Madagascarienses<br />
et Borbonici Selecti Exsiccati 10 (UPS, US, dis-<br />
tributed as Parmelia isidiza). Nepal: Awasthi 2156, 2157,<br />
2198, 2203 (Arvasthi herbarium), i\Torkett 5533 (BM), Togashi<br />
(TNS, US). Sikkih: Bose 136 (Awasthi herbarium). India:<br />
Uttar Pradesh, Awasthi 3175, 3509, 3814, 3980, 3986 (Awasthi<br />
herbarium), Hara (TNS, US); Tamil Nadu, Foreau 43<br />
(Awasthi herbarium), <strong>Hale</strong> 40220, 40274, 43620, 43740, 43870.<br />
Thailand: Kerr L4 (BM). Malaya: Pahang: <strong>Hale</strong> 30094,<br />
30217, 30218, 30219. Taiwan: Kurokawa 29, 530, 626, 2926<br />
(TNS). China: Kwantung Province, Petig 12578 (US); Man-<br />
churia, Asahina 25 (TNS). Hong Kong: Thrower 1586 (US).<br />
Japan: Prov. Aki, <strong>Hale</strong> 29508; Prov. Chikuzen, Kurokawa<br />
62545 (TNS); Prov. Higo Kiirokawn 63076 (TNS); Prov. Hizen,<br />
Kurokawa 63157 (TXS); I’rov. Hyuga, <strong>Hale</strong> 29645; Prov. Iyo,<br />
Kurokau~a 60098 (TNS); Prov. Iru, Asahina 94 (Th’S); Prov.<br />
Kii, Asahina (TNS), Kzirokawa 60237 (TNS, US), Kurokawa<br />
in <strong>Lichen</strong>es Rariores et Critici Selecti 42 (US); Prov. Musashi,<br />
Asahina 11 (TNS), Ktirokawa 59155 (TNS, US); Prov. Ohmi,<br />
<strong>Hale</strong> 29460; Prov. Satsuma, Kurokawa 63043 (TNS); Tosa,<br />
Fujikawa (TNS). Yoshimirra 1157 (US). Philippines: Benguet,<br />
Degelius As-787 (Degelius herbarium), <strong>Hale</strong> 26760, <strong>Hale</strong><br />
in <strong>Lichen</strong>es Selecti Exsiccati 672 (US); Mountain Province,<br />
Haze 26413. Sabah: <strong>Hale</strong> 28707, 28807, 29063, 29071. Indonesia:<br />
Java, Groenhart 3238 (L, US), 4358, 5881, 5913, 5917 (L), 5948<br />
(L, US), Schiflner 3294 (US), Voogd 2079 (L).
NUMBER 33 53<br />
47. <strong>Parmelina</strong> xantholepis<br />
FIGURE 20f<br />
Parmeliria sun tholepis (Montagne and van den Bosch) <strong>Hale</strong>,<br />
1974:438.<br />
Parrne/ia xantholepis hIontagne and van den Bosch,<br />
1855:428 [type collection: Gode, Java, Teydom (L,<br />
lectotype)].<br />
Parrnelia hiformis T’ainio f. dataensis Vainio [= f. biformis],<br />
1909:660 [type collection: Mt. Data, Luzon, Philippines,<br />
Alerrill 4987 (TUR, T’ainio herbarium number 2607,<br />
lectotype)].<br />
Parmelia biformis f. pauniensis T’ainio, 1909:660 [type collec-<br />
tion: Pauai, Benguet, Luzon, Philippines, Mearns 4434<br />
(TUR, T’ainio herbarium number 2605, lectotype)].<br />
DEscRIP’rIos.-Thallus adnate on bark or rock,<br />
yellowish mineral gray, very fragile, 4-15 cm broad;<br />
lobes sublinear to subirregular, 1-3 mm wide,<br />
crowded, dissected, <strong>the</strong> marginal cilia coarse, 0.2-0.8<br />
mm long; upper surface plane, maculate, becoming<br />
densely lobulate toward <strong>the</strong> lobe margins, isidia<br />
and soredia lacking; medulla barium yellow; lower<br />
surface black, densely rhizinate, <strong>the</strong> rhizines black,<br />
long, simple or squarrose. Apo<strong>the</strong>cia adnate, 1-3<br />
mm in diameter; spores 6-8 X 9-13 pm.<br />
CHEXIISTRY.-cOl.teX K + yellow (atranorin); me-<br />
dulla more intensively yellow with K and C, P-<br />
(zeorin, leucotylin and related terpenes, and seca-<br />
lonic acid A.)<br />
DISTRImTIoN.-hdia and Nepal to <strong>the</strong> Philip-<br />
pines.<br />
REM.mKs.-This species is easily identified by <strong>the</strong><br />
fragile, lobulate thallus with a pale yellow-orange<br />
medulla. It is closely related to <strong>the</strong> P. amagiensis-P.<br />
clenegans series, definitely in chemistry but less so in<br />
thallus texture. It is usually collected at <strong>the</strong> base <strong>of</strong><br />
trees or on rocks, <strong>of</strong>ten growing among mosses, at<br />
higher elevations (1000-2300 m) in evergreen hard-<br />
wood cloud forests.<br />
SPECIMESS Ex \MIsIm-h’epaI: Poelt 101, 111 (hI) , Norkett<br />
9227 (BM). India: Tamil Nadu, Foreau 4118 (ALvasthi herbarium,<br />
US), <strong>Hale</strong> 43862; Uttar Pradesh, Awasthi 3839<br />
(Awasthi herbarium). Thailand: Kziroknwa 1669 (TNS, US).<br />
Philippines: Mountain Province, <strong>Hale</strong> 25993, 26096, 26187,<br />
26344, 26554. Indonesia: Java, Groenhart 2841, 5939 (L),<br />
5941, 6004 (L, US).<br />
Doubtful and Rejected Names<br />
Parmelia coilocarpa<br />
Parriwlia coilocnrpn Stirton, 1877-78:202 [t)pe collection:<br />
Feinando Po, West Africa, G. Thornson (Bhf, lectotype;<br />
CLAM, isolectotype)].<br />
The type material is too fragmentary for ade-<br />
quate study. It is a nonsorediate, nonisidiate, sub-<br />
irregularly lobed species with a black lower surface,<br />
simple rhizines, and numerous marginal cilia. The<br />
spores are large, 15 X 28 pm. Atranorin and sala-<br />
zinic acid are present. It is almost certainly a<br />
Pnrmelina, and its status will only be clarified as<br />
more collections are made in TYest Africa.<br />
ParmeIin orchidophlia<br />
Pnrnielin orcliidoljhila Dodge, 1953:374 [type collection:<br />
Nyinabitsa, TVestern Pro\-ince, Uganda, Omastin 118.1<br />
(Bhf, lectot! pe)]<br />
The type collection is too Eragmentary for study.<br />
Salazinic acid was proved niicrochemically. It may<br />
be <strong>Parmelina</strong> zunllichiann (Taylor) <strong>Hale</strong>.<br />
Parmelia tiliacea var. lezicina<br />
Parmelin tilicea Tar. leucintr Miillrr Argoviensis, 1880:267<br />
[type collection: Sear Petropolis, Brazil, Deventer (G,<br />
lec totype)].<br />
This variety is equal to Hypotrachyna dactylifera<br />
(Vainio) <strong>Hale</strong> (<strong>Hale</strong>, 1975a:30).<br />
Pal-melia tilacea var. rimzilosa<br />
Pnrrrielin tilicea tar, rriricilosa Muller Aigoiiensis, 1882:458<br />
[tJpe collection: Socotra, h’nlfoirr (C, lectotipe)].<br />
No type specimen could be located at Geneva.<br />
One <strong>of</strong> two specimens so labeled by hluller is Par-<br />
motrema reticiilatum (Taylor) Choisy, and <strong>the</strong><br />
o<strong>the</strong>r is Pseudoparmelia carneopruinata (Zahl-<br />
bruckner) <strong>Hale</strong>.<br />
Pal-melina bagziioensis<br />
Panneliizu bnguioensis (<strong>Hale</strong>) <strong>Hale</strong>, 1974:482 [type collection:<br />
Baguio, Mountain Province, Philippines, Ha/e 26768 (US,<br />
holotype)].<br />
Closer examination <strong>of</strong> this sorediate species con-<br />
vinced me that it is a member <strong>of</strong> <strong>the</strong> genus Hypo-<br />
tl-achyna (H. bagtrioensis (<strong>Hale</strong>) <strong>Hale</strong>, new combi-<br />
nation). The rhizines are ra<strong>the</strong>r sparse but clearly
54 SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
dichotomously branched at maturity. In any event,<br />
<strong>the</strong> chemistry-fumarprotocetraric acid-was ano-<br />
malous for <strong>Parmelina</strong>, and this was <strong>the</strong> first clue to<br />
its correct generic status.<br />
<strong>Parmelina</strong> carporrhizans<br />
<strong>Parmelina</strong> carporrhizans (Taylor) <strong>Hale</strong>, 1974:482.<br />
I now consider this to be a synonym <strong>of</strong> Parme-<br />
lina quercina.<br />
<strong>Parmelina</strong> crystallorum<br />
<strong>Parmelina</strong> crystallorum (Lynge) <strong>Hale</strong>, 1974:482<br />
This species is synonymized under <strong>Parmelina</strong><br />
damaziana.<br />
Parme 1 ina h om ogen es<br />
<strong>Parmelina</strong> homogenes (Nylander) <strong>Hale</strong>, 1974:482.<br />
I have placed this species in synonymy under<br />
Pa rm e 1 in a szi bau ru 1 en t a.<br />
<strong>Parmelina</strong> nylanderi<br />
<strong>Parmelina</strong> nylanderi (Lynge) <strong>Hale</strong>, 1974:483.<br />
Parmelia nylanderi Lynge, 1914:82 [type collection: Sear $450<br />
Jeronyno, Serra da Chapada, Mato Grosso, Brazil, Malme<br />
2747 (S, lectotype)].<br />
I originally considered this loosely attached saxi-<br />
colous lichen to be a member <strong>of</strong> <strong>Parmelina</strong>, but<br />
after examining a second collection (Eiten 3429<br />
(US) from S5o Paulo, Brazil), I now feel that <strong>the</strong><br />
chemistry (usnic acid in <strong>the</strong> cortex and salazinic<br />
and gyroplioric acid in <strong>the</strong> medulla) and general<br />
habit, in spite <strong>of</strong> <strong>the</strong> narrow ciliate lobes, place it<br />
more appropriately in Parmotrema (P. nylanderi<br />
(Lynge) <strong>Hale</strong>, new combination). It is related to<br />
<strong>the</strong> much broader lobed P. delicatulzim (Vainio)<br />
<strong>Hale</strong> group so common on sandstone in South<br />
America.
des Ahhayes, H.<br />
1951. <strong>Lichen</strong>s recoltes en Guice francaise et en Cote<br />
d’Ivoire. Bulletin de l’lnstitut Franpis d’Afrique<br />
Noire, 13 (4):965-977.<br />
Acharius, E.<br />
1794. Forsok ti1 en forbattrad lafvarnes indelning<br />
(Dianome <strong>Lichen</strong>um). Konglige Vetenskaps-Academiens<br />
Handlingar, 12:237-254.<br />
1798. <strong>Lichen</strong>ographiae Suecicae Prodromus. 264 pages.<br />
Lincopiae.<br />
1803. Methodus qua omnes detectos lickenes. 394 pages.<br />
Stockholmiae.<br />
1814. Synopsis methodica lichenuni. 392 pages. Lundae.<br />
Asahina, Y.<br />
1951a. <strong>Lichen</strong>es Japoniae novae vel minus cognitae (4).<br />
Journal <strong>of</strong> Japanese Botany, 263225-228.<br />
19511). <strong>Lichen</strong>es Japoniae novae vel minus cognitae (5).<br />
Journal <strong>of</strong> Japanese Botany, 263257-261,<br />
1951~. <strong>Lichen</strong>es Japoniae novae vel minus cognitae (6).<br />
Journal <strong>of</strong> Japanese Botany, 26:289-293.<br />
1952. <strong>Lichen</strong>s <strong>of</strong> Japan, 11: <strong>Genus</strong> Parmelia. 162 pages.<br />
Tokyo.<br />
1960. <strong>Lichen</strong>ologische Sotizen (160-163). Journal <strong>of</strong> Japanese<br />
Botany, 35:97-102.<br />
1963. <strong>Lichen</strong>ologische Notizen (191). Journal <strong>of</strong> Japanese<br />
Botany, 38:225-228.<br />
Berry, E. C.<br />
1941. A <strong>Monograph</strong> <strong>of</strong> <strong>the</strong> <strong>Genus</strong> Parmelia in North<br />
America, Sorth <strong>of</strong> Mexico. Annals <strong>of</strong> <strong>the</strong> Missouri<br />
Botanical Garden, 28:31-146.<br />
Bouly de Lesdain, M.<br />
1914. <strong>Lichen</strong>s de Mexique. 31 pages. Mexico.<br />
Burnham, S. H.<br />
1922. <strong>Lichen</strong>s <strong>of</strong> <strong>the</strong> Lake George Region. Bryologist,<br />
25:72-80.<br />
Cengia Samho, M.<br />
1938. <strong>Lichen</strong>i del Kenia e del Tanganica raccolti dai Rev.<br />
Padri Della Consolata. Nuouo Giornale Botanico<br />
Italiano, 45:364-387.<br />
Chao, Chi-ding<br />
1964. [A Preliminary Study on Chinese Parmelia.] Acta<br />
Phytotaxoninzica Sinica, 9: 139-166. [In Chinese.]<br />
Culberson, C.<br />
1972. Improved Conditions and New Data for <strong>the</strong> Identification<br />
<strong>of</strong> <strong>Lichen</strong> Products by a Standardized Thin-<br />
Layer Chromatographic Method. Journal <strong>of</strong> Chromatography,<br />
7211 13-125.<br />
Culherson, W.<br />
1955. The Corticolous Communities <strong>of</strong> <strong>Lichen</strong>s and<br />
Literature Cited<br />
55<br />
Bryophytes in <strong>the</strong> Upland Forests <strong>of</strong> Nor<strong>the</strong>rn Wisconsin.<br />
Ecological <strong>Monograph</strong>s, 25: 2 15-23 1.<br />
1958. Variation in <strong>the</strong> Pine-inhabiting L’egetation <strong>of</strong><br />
North Carolina. Ecology, 39:23-28.<br />
1961. The Parmelia quercina Group in h’orth America.<br />
American Journal <strong>of</strong> Botany, 48: 168-174.<br />
1972. Disjunctive Distributions in <strong>the</strong> <strong>Lichen</strong>-forming<br />
Fungi. Annals <strong>of</strong> <strong>the</strong> Missouri Botanical Garden,<br />
59:165-173.<br />
Dahl, E., and H. Krog<br />
1973. Macrolichens <strong>of</strong> Denmark, Finland, Sorway, and<br />
Sweden. 185 pages. Oslo.<br />
Degelius, G.<br />
1940. contributions to <strong>the</strong> <strong>Lichen</strong> Flora <strong>of</strong> North America,<br />
I: <strong>Lichen</strong>s from Maine. Arkiv fur Botanik, 30A<br />
(1) : 1-62.<br />
1941. Contributions to <strong>the</strong> <strong>Lichen</strong> Flora <strong>of</strong> North America,<br />
11: The <strong>Lichen</strong> Flora <strong>of</strong> <strong>the</strong> Great Smoky<br />
Mountains. Arkiv for Botanik, 30A (3): 1-80.<br />
Dey, J.<br />
1975. The Fruticose and Foliose <strong>Lichen</strong>s <strong>of</strong> <strong>the</strong> High<br />
Mountain Areas <strong>of</strong> <strong>the</strong> Sou<strong>the</strong>rn Appalachians. 603<br />
pages. Unpublished Ph.D. dissertation, Duke Universi<br />
ty.<br />
Dobson, F. S., and D. L. Hawksworth<br />
1976. Parmelia pastillifera (Harm.) Schuh. & Klem. and<br />
P. tiliacea (H<strong>of</strong>fm.) Ach. in <strong>the</strong> British Isles. <strong>Lichen</strong>ologist,<br />
8:47-59.<br />
Dodge, C. TV.<br />
1953. Some <strong>Lichen</strong>s <strong>of</strong> Tropical Africa. Annals <strong>of</strong> <strong>the</strong><br />
Missouri Botanical Garden, 40:271-412.<br />
1959. Some <strong>Lichen</strong>s <strong>of</strong> Tropical Africa, 111: Parmeliaceae.<br />
Annals <strong>of</strong> <strong>the</strong> Missouri Botanical Garden, 46:39-193.<br />
Fink, B.<br />
1935. The <strong>Lichen</strong> Flora <strong>of</strong> <strong>the</strong> United States. 426 pages.<br />
Ann Arbor.<br />
Fries, E.<br />
1825. Systenia orbis vegetabilis. Part 1, 374 pages. Lund.<br />
Gyelnik, V.<br />
1931. Additamenta ad cognitionem Parmeliarum, 11.<br />
Fedde Repertorium, 29:273-291.<br />
1932a. Enumeratio lichenum europaeorum novorum rariorumque.<br />
Annales Mycologici, 30:442-455.<br />
1932b. Additamenta ad cognitionem Parmeliarum, 111.<br />
Fedde Repertorizrni, 30:209-226.<br />
<strong>Hale</strong>, M. E., Jr.<br />
1955. Phytosociology <strong>of</strong> Corticolous Cryptogams in <strong>the</strong><br />
Upland Forests <strong>of</strong> Sou<strong>the</strong>rn Wisconsin. Ecology,<br />
36:45-63.
56<br />
SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
1958. Studies on <strong>the</strong> Chemistry and Distribution <strong>of</strong> North Hedrick, J.<br />
American <strong>Lichen</strong>s (10-13). Bqologist, 61:81-85.<br />
1934. New Genera and Species <strong>of</strong> <strong>Lichen</strong>s from <strong>the</strong> Her-<br />
1959. New or Interesting Parmelias from North and Tropical<br />
America. Bryologist, 62:123-132.<br />
barium <strong>of</strong> Bruce Fink, 11. Mycologia, 26:153-166.<br />
H<strong>of</strong>fmann, G. F.<br />
1960. A Revision <strong>of</strong> <strong>the</strong> South American Species <strong>of</strong> Par- 1784. Enumeratio <strong>Lichen</strong>um. 102 pages. Erlangae.<br />
melia Determined by Lynge. Contributions from <strong>the</strong> Hue, A. M.<br />
United States National Herbarium, 36: 141.<br />
1899. <strong>Lichen</strong>es extra-europaei. Nouvelles Archives du Mu-<br />
1965. A <strong>Monograph</strong> <strong>of</strong> Parmelia Subgenus Amphigymnia.<br />
seum Paris, series 3, 1: 1-250.<br />
Contributions from <strong>the</strong> United States National Her- James, P. W.<br />
barium, 36: 193-358.<br />
1965. A New Check-List <strong>of</strong> British <strong>Lichen</strong>s. <strong>Lichen</strong>ologist,<br />
1967. Chemistry and Evolution in <strong>Lichen</strong>s. Israel Journal<br />
3:95-153.<br />
<strong>of</strong> Botany, 15:150-157.<br />
Koerber, G. W.<br />
1971a. Morden-<strong>Smithsonian</strong> Expedition to Dominica: The 1855. Systenza <strong>Lichen</strong>um Gernianiae. 458 pages. Breslau.<br />
<strong>Lichen</strong>s (Parmeliaceae). <strong>Smithsonian</strong> Contributions Krempelhuber, A.<br />
to Botany, 4: 1-25.<br />
1971b. Two Species <strong>of</strong> Parmelia New to North America.<br />
1878. <strong>Lichen</strong>es collecti in republica Argentina a Doctoribus<br />
Lorentz et Hieronymus. Flora, 61:461-464.<br />
Bryologist, 74:44-47.<br />
1972a. Four New Species <strong>of</strong> Parmelia (<strong>Lichen</strong>es) from India<br />
Kurokawa, S.<br />
1968a. Parmelia expallida, a Xew <strong>Lichen</strong> Species from<br />
and <strong>the</strong> Philippines. Bryologist, 75:97-101.<br />
Eastern Asia. Bulletin <strong>of</strong> <strong>the</strong> National Science Mu-<br />
1972b. Pamielia jamesii, an Unusual Species in Section<br />
seum, 11:191-194.<br />
Imbricaria (<strong>Lichen</strong>es) from Australia and New Zea- 1968b. New or Noteworthy Species <strong>of</strong> Parmelia <strong>of</strong> Japan.<br />
land. Phytologia, 23: 179.<br />
Journal <strong>of</strong> Japanese Botany, 43:349-353.<br />
1972~. Six New Species <strong>of</strong> Parmelia (<strong>Lichen</strong>es) from Atrica. 1972. Probable Mode <strong>of</strong> Differentiation <strong>of</strong> <strong>Lichen</strong>s in<br />
Phytologia, 23:343-349.<br />
Japan and Eastern North America. Pages 139-146<br />
1973a. Fine Structure <strong>of</strong> <strong>the</strong> Cortex in <strong>the</strong> <strong>Lichen</strong> Family<br />
in A. Graham, Floristics and Pale<strong>of</strong>loristics <strong>of</strong> Asia<br />
Parmeliaceae Viewed with <strong>the</strong> Scanning-Electron<br />
and North America. Amsterdam.<br />
Microscope. <strong>Smithsonian</strong> Contributions to Botany, Kurokawa, S., and M. Mineta<br />
10: 1-92.<br />
1973. Enumeration <strong>of</strong> Parmeliae <strong>of</strong> Ceylon. Annual Re-<br />
1973b. New Parmeliae (<strong>Lichen</strong>es) from Africa. Phytologia,<br />
port <strong>of</strong> <strong>the</strong> Noto Marine Lnboratory, 13:71-76.<br />
27~1-6.<br />
de Lamarck, J., and A. P. de Candolle<br />
1974. Bulbothrix, <strong>Parmelina</strong>, Relicina, and Xanthopar- 1805. Flore Francaise. Volume 2, 600 pages. Paris.<br />
melia, Four New Genera in <strong>the</strong> Parmeliaceae (Lich- Lynge, B.<br />
enes). Phytologia, 28:47%490.<br />
1914. Die Flechten der ersten Regnellschen Expedition.<br />
1975a. A Revision <strong>of</strong> <strong>the</strong> <strong>Lichen</strong> <strong>Genus</strong> Hypotrachyna<br />
Die Gattungen Pseudoparmelia gen. nov. und Par-<br />
(Parmeliaceae) in Tropical America. <strong>Smithsonian</strong><br />
melia Ach. Arkiv for Botanik, 13 (13):l-172.<br />
Contributions to Botany, 25:l-73.<br />
Massalongo, A. B.<br />
1975b. A <strong>Monograph</strong> <strong>of</strong> <strong>the</strong> <strong>Lichen</strong> <strong>Genus</strong> Relicina 1856. Schedae Criticae. Part 10, number 328.<br />
(Parmeliaceae). <strong>Smithsonian</strong> Contributions to Bot- hfichaux, A.<br />
any, 26: 1-32.<br />
1803. Flora Boreali-Americana. Volume 2, 340 pages. Paris.<br />
1976a. A <strong>Monograph</strong> <strong>of</strong> <strong>the</strong> <strong>Lichen</strong> <strong>Genus</strong> Pseudoparmelia<br />
Lynge (Parmeliaceae). <strong>Smithsonian</strong> Contributions to<br />
Botany, 31:l-62.<br />
1976b. A <strong>Monograph</strong> <strong>of</strong> <strong>the</strong> <strong>Lichen</strong> <strong>Genus</strong> Bulbothrix <strong>Hale</strong><br />
(Parmeliaceae). <strong>Smithsonian</strong> Contributions to Botany,<br />
32: 1-29.<br />
H ale, M E., Jr., and S. Kurokawa<br />
1962. Parmelia Species First Described from <strong>the</strong> British<br />
Isles. <strong>Lichen</strong>dogist, 2: 1-5.<br />
1964. Studies on Parmelia Subgenus Parmelia. Contributions<br />
from <strong>the</strong> United States h’ational Herbarium,<br />
36~121-191.<br />
H armand, J.<br />
Montagne, J. K. C., and R. B. van den Bosch<br />
1855. <strong>Lichen</strong>es, Collemaceae. Pages 427-494 in F. Junghuhn,<br />
PIantae Junghuhnianae [sive Enumeratio<br />
Plantarum quas in insulis Java et Sumatra].<br />
Moore, B. J.<br />
1968. The Macrolichen Flora <strong>of</strong> Florida. Bryologist, 71:<br />
161-266.<br />
Muller Argoviensis, J.<br />
1877. <strong>Lichen</strong>ologische Beitrage, V: <strong>Lichen</strong>en aus Texas.<br />
Flora, 60:77-80.<br />
1880. Licheologische Beitrage, XI. Flora, 63:260-268.<br />
1882. Diagnoses <strong>Lichen</strong>um Socotrensium novorum. Proceedings<br />
<strong>of</strong> <strong>the</strong> Royal Society <strong>of</strong> Edinburgh, 11:457-<br />
472.<br />
1897. Catalogue descriptif des <strong>Lichen</strong>s observes dans la 1883. <strong>Lichen</strong>ologische Beitrage, XVII. Flora, 66:4548.<br />
Lorraine. Bulletin de la Socidtd des Sciences de 1887. <strong>Lichen</strong>ologische Beitrage, XXVI. Flora, 70:316-320.<br />
Nancy, series 2, 14:l-510.<br />
1888a. <strong>Lichen</strong>es Montevidenses quos legit et communicavit<br />
1910. <strong>Lichen</strong>s de France. Volume 4, pages 483-732. Paris.<br />
Pr<strong>of</strong>. Arechavaleta. Revue Mycologique, 10: 1-5.<br />
1928. <strong>Lichen</strong>s d’Indo-Chine recueillis par M. V. Demange. 1888b. <strong>Lichen</strong>ologische Beitrage, XXIX. Flora, 71 : 195-208.<br />
Annales Cryptogarnie Exotique, 1 :319-337.<br />
1888~. <strong>Lichen</strong>ologische Beitrage, XXX. Flora, 71 :528-552.
NUMBER 33<br />
1891. <strong>Lichen</strong>ologische Beitrage, XXXV. Flora, 74:371-382. 1847. New <strong>Lichen</strong>s, Principally from <strong>the</strong> Herbarium <strong>of</strong><br />
Nylander, W.<br />
W. J. Hooker. Hooker Journal oj Botany, 6:148-<br />
1856. Prodromus <strong>Lichen</strong>ographiae Galliae et Algeriae.<br />
197.<br />
Actes de la Societe Lznndenne de Bordeaux, 21:249- Tuckerman, E.<br />
467.<br />
1858. Supplement to an Enumeration <strong>of</strong> North American<br />
1868. Enumeration des lichen, recoltes par M. T. Husnot<br />
<strong>Lichen</strong>s: Part First, Containing Brief Diagnoses <strong>of</strong><br />
aux Antilles francaises. Bulletin de la Societe Lin-<br />
New Species. American Journal <strong>of</strong> Science and Arts,<br />
ndenne de Normandie, (2)3:259-280.<br />
series 2, 25:422-430.<br />
1873. Observata <strong>Lichen</strong>ologica in Pyrenaeis orientalibus. 1882. A Synopsis <strong>of</strong> <strong>the</strong> North American <strong>Lichen</strong>s. 261<br />
Bulletin de la Societe Linntenne de Normandie,<br />
pages. Boston.<br />
(2)6:256-328.<br />
Vainio, E. A.<br />
1882. Addenda nova ad <strong>Lichen</strong>ographiam europaeam. 1890. Etude sur la classification naturelle et la morpho-<br />
Flora, 653491458.<br />
logie des <strong>Lichen</strong>s du Bresil. Acta Societatis pro<br />
1885. Parmeliae exoticae novae. Flora, 68:605-615.<br />
Fauna et Flora Fennica, 7(7): 1-247.<br />
1890. <strong>Lichen</strong>es Japoniae. 122 pages. Paris.<br />
1896. <strong>Lichen</strong>es Antillarum a W. R. Elliott collecti. Jour-<br />
1900. <strong>Lichen</strong>es Ceylonenses. Acta Societatis Scientiarum<br />
nal <strong>of</strong> Botany British and Foteign, 34:31-36.<br />
Fennicae, 26(10): 791-821.<br />
1899. <strong>Lichen</strong>es in Caucaso et in Peninsula Taurica annis<br />
Poelt, J.<br />
1884-1885 ab H. Lojka et hi. a Dechy collecti.<br />
1961. Mitteleuropaische Flechten, VII. Mitteilungen der<br />
Terineszetrajzi Fuzetek, 22:269-343.<br />
Botanischen Staatssammlung Munchen, 4: 171-197. 1909. <strong>Lichen</strong>es Insularum Philippinarum, I. Philippine<br />
1969. Bestimmungsschliusel<br />
pages. Lehre.<br />
Schauer, .- 1.<br />
europascher Flechten. 757<br />
Journal <strong>of</strong> Science, 4:651-662.<br />
Willdenow, C. L.<br />
1787. Florae Berolinensis Prodromus. 440 pages. Berlin.<br />
1965. Oceanische Flechten im Nordalpenraum. Portu- Yosioka, I.<br />
galiae Acta Biologica, (B)8: 17-229.<br />
1966. The Structure <strong>of</strong> Leucotylic Acid, a New Triterpenic<br />
Schreber, J. C. D.<br />
Acid from a <strong>Lichen</strong>. Tetrahedron Letters, 6:607-<br />
1791. Linne, Genera Plantarum. Eighth edition, 872 pages.<br />
612.<br />
Frankfurt am Main.<br />
Yosioka, I., and T. Nakanishi<br />
Schubert, R., and 0. Klement<br />
1963. Structure <strong>of</strong> Leucotylin. Chemical Pharmaceutical<br />
1966. Beitrag zur Flechtenflora von Sord- und LMittel-<br />
Bulletin, 11: 1468-1470.<br />
indien. Nova Hedwigia, 11 : 1-73.<br />
1966. On <strong>the</strong> Triterpenic Constituents <strong>of</strong> a <strong>Lichen</strong>, Par-<br />
Sernander-Du Rietz, G.<br />
melia ento<strong>the</strong>iochroa Hue, Zeorin, Leucotylin,<br />
1926. Parnzelia tiliacea, en kustlav och marin inlandsrelikt<br />
Leucotylic Acid, and Five Sew Related Triterpenic<br />
i Skandinavien. Svensk Botanisk Tidskrift, 20:352-<br />
Acids. Chemical Pharmaceutical Bulletin, 14:804-<br />
365.<br />
807.<br />
1957. Om yttre faktores inverkan pa apo<strong>the</strong>ciebildningen<br />
hos Parmelia tiliacea. Svensk Botanisk Tidskrift,<br />
51:454-488.<br />
Shibata, S., S. Natori, and S. Udaga1r.a<br />
1964. List <strong>of</strong> Fungal Products. 170 pages. Tokjo.<br />
Steiner, J.<br />
1888. Ueber afrikanische Flechten. Jahres-Bericht der<br />
Schlesischen Gesellschaft jur vaterlundische Cultur,<br />
66: 133-150.<br />
1907. <strong>Lichen</strong>es austro-africani. Bulletin de l’Herbier Boissier,<br />
2(7) : 637-646.<br />
Stirton, J.<br />
1877-78. On Certain <strong>Lichen</strong>s Belonging to <strong>the</strong> <strong>Genus</strong><br />
Yosioka, I., T. Nakanishi, S. Izumi, and I. Kitagawa<br />
1968a. Structure <strong>of</strong> a <strong>Lichen</strong> Pigment Ento<strong>the</strong>in and its<br />
Identity with Secalonic Acid A, a Major Ergot Pigment.<br />
Chemical Pharmaceutical Bulletin, 16:2090,<br />
2091.<br />
Yosioka, I., T. Nakanishi, and I. Kitagawa<br />
3968b. On <strong>the</strong> Stereostructures <strong>of</strong> Zeorin and Leucotylin.<br />
Tetrahedron Letters, 12:1485-1490.<br />
Zahlbruckner, A.<br />
1905. <strong>Lichen</strong>es a cl. Damazio in Brasilia lecti. Bulletin de<br />
I’Herbim Boissier, series 2, 5:539-543.<br />
1908. Beitriige zur Flechtenflora Brasiliens. Bulletin de<br />
Parmelia. Scottish Naturalist, 4:200-203.<br />
1’Herbier Boissier, series 2, 8:459468.<br />
1899. <strong>Lichen</strong>es. In F. M. Bailey, Contributions to <strong>the</strong> 1909. <strong>Lichen</strong>es (Flechten). In R. I\’ettstein and 1’. Schiff-<br />
Flora <strong>of</strong> Queensland. Queensland Agricultural Jour-<br />
ner, Ergebnisse der botanischen Expedition der<br />
nal, 5:483-488.<br />
kaiserlichen Akademie der Wissenschaften nach<br />
Stizenberger, E.<br />
Sudhrasilien 1901. Denkschrift Akademie der Wis-<br />
1890. <strong>Lichen</strong>aea africana. Bericht uber die Taigkeit der<br />
senschaften in Wien. Ma<strong>the</strong>matisch-A’aturwissen-<br />
St. Gallischen Naturruissenschaftlic~ieii Gesellschaft,<br />
schajtliche Klasse, 83:87-211.<br />
1888-89: 105-209.<br />
1926. Afrikanische Flechten (<strong>Lichen</strong>es). Botanische Jahr-<br />
Ta)lor, T.<br />
bucher jur Systetnatik, 60:468-552.<br />
1836. <strong>Lichen</strong>s. In J. T. hlackay, Flora Hibernica. 279 1927. Additamenta ad <strong>Lichen</strong>ographiatn Japoniae. Bopages.<br />
Dublin.<br />
tanical Magazine (Tokyo), 41:313-364.<br />
57
58 S\iITHSOXI 4S CONTRIBUTIOSS TO BOTASY<br />
1929. Catalogus iichenum tcniiwsrriic. T.olume 6, 323 1934a. Flechten der Inrel Forniosa. FeddP Reperiorium<br />
pages. Leipiig. 33:22-C,F.<br />
1930. <strong>Lichen</strong>es. 171 H. Handel-hlazretti, Sjmboiae Sinica, 1934. Nachtrage zur Flechtenflora Chinas. Hedwigia,<br />
3:1-234. 74: 193-21 3.
lmbricaria quercina, 42<br />
tiliacea, 48<br />
<strong>Lichen</strong> quercinus, 42<br />
tiliaceus, 48<br />
Par in elia a 1 b ido -s tra min ea, 19<br />
amagiensis, 18<br />
ainazonica var. hirsnotii, 26<br />
antillensis, 18<br />
atricha, 42<br />
aurulenta, 19<br />
halansae, 22<br />
balansae \ar. sorediata, 40<br />
biformis f. dataensis, 53<br />
biforinis f. pnuaiensis, 53<br />
blastica, 18<br />
brachpconidia, 23<br />
brachsconidia \ ar. chlorocarpa, 23<br />
budapestiiiensis, 42<br />
ca m t rchadalis i'ar. epiph y lla , 26<br />
carporrhizans, 42<br />
carporrhizans f. inalicola, 42<br />
catharinensis, 51<br />
coilocarpa, 53<br />
consors, 22<br />
conspiciia, 47<br />
con t in en talis, 22<br />
cryptochlora, 23<br />
crystalloruin, 23<br />
damaziana, 23<br />
deminuta, 30<br />
denegans, 25<br />
dissecta, 25<br />
endoleuca, 27<br />
enormis, 27<br />
ento<strong>the</strong>iochroa, 28<br />
expallida, 28<br />
fecunda, 47<br />
finkii, 38<br />
fraudans ssp. subfraudans, 35<br />
galbina, 30<br />
galbina var. rugosa, 30<br />
galbina 1-ar. sitbradiata, 30<br />
hayachinensis, 31<br />
heteroloba, 31<br />
homalotera, 47<br />
homogenes, 47<br />
homogenes f. minor, 47<br />
horrescens, 3 1<br />
Index<br />
(Synonyms in italics)<br />
hubrichtii, 26<br />
hrinanensis, 19<br />
immiscens, 32<br />
insinuata, 34<br />
insin itat ula, 34<br />
irrugans, 34<br />
jarnesii, 35<br />
junodii, 51<br />
laruignta ssp. estremi-orienialis, 30<br />
iaeuigata var. gracilis f. furfuracea, 25<br />
laevigatoides, 50<br />
lericotylita, 35<br />
leucotqlix f. rugulosa, 35<br />
leiico/ylizn f. siiblaevis, 35<br />
lin d ilia nii, 36<br />
mrlaiiochneta, 36<br />
nietnrevoiiita, 36<br />
inichoncnnensis, 32<br />
ni inn JIL m, 26<br />
riii/el/eri, 38<br />
uiiitata, 51<br />
myriocnrpa, 47<br />
11 ima nda i ra n a, 5 1<br />
iiimaiidairnna f. sublaevis, 51<br />
nylanderi, 54<br />
obsessa, 36<br />
orchidophila, 53<br />
pastillifera, 39<br />
perisidians, 39<br />
phlyctina, 40<br />
pilosa, 40<br />
pruinata, 42<br />
piiiggari, 26<br />
quercifolia var. scrotea f. inicrophylla,<br />
48<br />
quercina, 42<br />
querrina var. stclphurosa, 30<br />
revolnta i.ar. granulata, 42<br />
sampniana, 22<br />
.scortea var. papillosa, 48<br />
scortra var. pastillifera, 39<br />
scortea iar. ramifica, 48<br />
scortea var. visegradensis, 48<br />
silvestris, 19<br />
simplicior, 44<br />
sinuosa var. hypothrix, 42<br />
spathulata, 46<br />
sp umosa, 46<br />
59<br />
subaurulenta, 47<br />
siibaurulenta var. inyriocarpa, 47<br />
subbalansae, 40<br />
subcremea, 47<br />
sribfatisceiis, 47<br />
subquercifolia, 30<br />
subqiiercifolia f. Aiibradiata, 30<br />
subreuolitta, 19<br />
subsz~lphitrata, 39<br />
tilracea. 48<br />
tiliacea \ ar. afjixa, 42<br />
tilicicea Tar. efflorescens, 19<br />
tiiiacva var. exirnin, ,51<br />
tiliacen ~ar. hsmthri\, 42<br />
tiiiacra \ar. leuciiia, 53<br />
tiliacea var. minor, 30<br />
ti/iacen \ar. rimiilosn, 53<br />
t iliacea su bq ii r rci folio, 30<br />
tilinrea var. siiipliurouz,, 30<br />
tiliaren I ar. siilphurosa f. asperata, 36<br />
usariibarensis, 50<br />
versiforniis, 5 1<br />
wailichiaiia, 51<br />
wettsteinii, 51<br />
yalungana, 42<br />
vnnthorarpa, 34<br />
santholefiis, 53<br />
<strong>Parmelina</strong> amagiensis, 18<br />
antillensis, 18<br />
aurulenta, 19<br />
baguioensis, 53<br />
carporrhizans, 54<br />
consors, 22<br />
crassata. 22<br />
cr) ptochlora, 23<br />
crystallorum, 54<br />
tlamaziana, 23<br />
degelii, 25<br />
tlenegans, 25<br />
dissecta, 25<br />
endoleuca, 27<br />
enormis, 27<br />
ento<strong>the</strong>iochroa, 28<br />
expallida, 28<br />
galbina, 30<br />
hayachinensis, 31<br />
heteroloba, 31<br />
homogenes, 47, 54
horrescens, 31<br />
immiscens, 32<br />
indica, 34<br />
jamesii, 35<br />
leucotyliza, 35<br />
lindmanii, 36<br />
melanochaeta, 36<br />
metarevoluta, 36<br />
muelleri, 38<br />
nylanderi, 54<br />
obsessa, 38<br />
pastillifera, 39<br />
perisidians, 39<br />
phlyctina, 40<br />
pilosa, 40<br />
pruinata, 42<br />
quercina, 42<br />
rhytidodes, 43<br />
schindleri, 44<br />
simplicior, 44<br />
SMITHSONIAN CONTRIBUTIONS TO BOTANY<br />
spathulata, 46<br />
spumosa, 46<br />
suhaurulenta, 46<br />
suhfatiscens, 47<br />
swinscowii, 48<br />
tiliacea, 48<br />
usamharensis, 50<br />
versiformis, 51<br />
wallichiana, 51<br />
xantholepis, 53