SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 10 
Fine Structure of the 
Cortex in the Lichen Family 
Parmeliaceae Viewed with the 
Scanning-electron Microscope 
Mason E. Hale, Jr. 
SMITHSONIAN INSTITUTION PRESS 
City of Washington 
1973 
ABSTRACT 
Hale, Mason E., Jr. Fine Structure of the Cortex in the Lichen Family Parme- 
liaceae Viewed with the Scanning-electron Microscope. Smithsonian Contributions 
to Botany, number 10, 92 pages, 150 figures, 1971.-The scanning-electron micro- 
scope was used to examine the cortical surface of 123 species of lichens in 12 
genera of the lichen family Parmelinceae. Two general types of cortex were found, 
one consisting of exposed hyphae and one with the hyphae covered by a thin 
polysaccharide epicortex. The epicorticate species fell into two groups, one with 
a tightly appressed continuous epicortex and one with a more loosely associated 
pored epicortex. Type of epicortex is a constant character at the genus and section 
level and appears to have considerable usefulness in the taxonomy of the family. 
OFFICIAL PUBLICATION DATE is handstam ed in a limited number of initial copies and is recorded 
in the Institution?s annual report, SrnitEsonian Year. SI PRESS NUMBER 4781. SERIFS COVER DESIGN: 
Leaf clearing from the katsura tree Cercidiphyllurn japonicum, Siebold and Zuccarini. 
Library of Congress Cataloging in Publication Data 
Hale, Mason E. 
Fine structure of the cortex in the lichen family Parmeliaceae viewed with the scanningelectron 
microscope. 
(Smithsonian contributions to botany, no. 10) 
Bibliography: p. 
1. Parmeliaceae. 2. Lichens-Anatomy. Ultrastructure (Biology). I. Title. 11. Series: Smith- 
QKl.SZ747 no. 10 [QK585.P2] 581?.08s [589?.1] 72-8039 
sonian Institution. Smithsonian contributions to botany, no. 10. 
For de by the Superintendent of Documents, U.S. Government Printing Office 
Washington, D.C. 40402 - Price $2 (paper cover) 
Contents 
Page 
Introduction ...................................................... 1 
The Scanning-electron Microscope ................................... 2 
Samples .......................................................... 2 
Structure of the Thallus 2 
Vertical Structure 2 
The Epicortex 3 
Nonepicorticate Type ............................................ 3 
Epicorticate Type ............................................... 3 
Fine Structure of the Lower Surface ............................... 4 
Variation in Epicortical Structure ................................... 4 
Value of Cortical Fine Structure in Systematics.. .................... 5 
Anzia Stizenberger ................................................ 6 
Asahinea Culberson ................................................ 6 
Cauernularia Degelius .............................................. 6 
Cetraria Acharius ................................................. 6 
Cetrelia Culberson and Culberson ................................... 7 
Heterodea Muiller-Argau ........................................... 7 
Hypogymnia (Nylander) W. Watson ................................ 7 
Menegazzia Massalongo ............................................ 7 
Pannoparmelia Darbishire ......................................... 7 
Parmelia Acharius ................................................. 7 
Subgenus Melanoparmelia Hue 7 
Subgenus Xanthoparmelia (Vainio) Hale. ......................... 8 
Subgenus Parmelia .............................................. 8 
............................................ 
................................................ 
.................................................... 
................................... 
Section Parmelia ............................................... 8 
8 
8 
8 
Section Simplices (Hale and Kurokawa) Hale .................... 
Section Irregulares (Vainio) Vainio ............................. 
Section Relicinae (Hale and Kurokawa) Hale .................... 
Section Bicornuta Lynge ....................................... 8 
Section Imbricaria (Schreber) Fries ............................. 9 
Section Cyclocheila (Vainio) Rasaanen 9 
Section Hypotrachyna Vainio ................................... 9 
Subgenus Everniiformes (Hue) Hale and Wirth .................... 
Subgenus Amphigymnia Vainio 9 
............................ 
9 
Parmelia Species of Uncertain Position .............................. 10 
Parmel iopsis N ylander ............................................. 10 
Platismatia Culberson and Culberson ................................ 10 
Pseudevernia Zopf ................................................. 10 
Atlas of SEM Photographs .......................................... 10 
Literature Cited ................................................... 11 
Figures ........................................................... IS 
................................... 
iii 

Fine Structure of the 
Cortex in the Lichen Family 
Parmeliaceae Viewed with the 
S canning-elec tron Microscope 
Mason E. Hale, Jr. 
Introduction He identified several orientations of cortical hyphae 
The scanning-electron microscope (SEM) is one of 
the newest tools available to biologists. Electron 
beams reflected from the surfaces of objects produce 
images with a remarkable three-dimensional quality 
because of the great depth of focus. The range of 
magnification, from 20 to 100,000 diameters, is 
very great, although photographs become fuzzy 
over 10,000 diameters. The electron microscope 
(EM), which is often used to study the internal 
structure of hyphal and algal cells, also magnifies 
at 10,000 diameters or more (see Jacobs and Ah- 
madjian, 1969), but the specimens can be examined 
only as two-dimensional sections. Thus the SEM 
bridges the gap between the EM and the low power 
light-reflecting stereomicroscope. 
The first use of the SEM in lichen research, as far 
as I am aware, was reported briefly by Peveling and 
Vahl (1968). The next study (Peveling, 1970, a 
report of a symposium held in 1969) contained 
more detailed descriptions, showing, for example, 
crystal formations on hyphae and certain surface 
details. Hawksworth in the same year (1969) pub- 
lished the first comprehensive investigation of a 
lichen family (Alectoriaceae) with the intent of 
applying the information to taxonomic problems. 
Mason A. Ha?e, Jr., Department of Botany, National Museum 
of Natural History, Smithsonian Institution, Washington, 
D.C. 20560. 
.- 
that were useful in delimiting genera. More re- 
cently, Tibell (1971) has shown the value of the 
SEM in studying ascospore structure in the Cyphe- 
liaceae. These first efforts have demonstrated con- 
vincingly that the SEM has real potential for 
opening up new and previously unexplored areas 
of study in lichen morphology and systematics. 
The purpose of this report is to describe in de- 
tail the cortical surface of representative species in 
the Parmeliaceae and to ascertain the value of fine 
structure in the taxonomy of the family. Interpre- 
tation of some of these structures is at best tentative 
and may indeed prove erroneous on future more 
intensive studies using data from the SEM in con- 
junction with EM sections and chemical analyses. 
It is hoped, nevertheless, that the photographs 
presented here will enable other lichenologists to 
make their own interpretations. 
ACKNOWLEDGMENTS.-~ have a number of col- 
leagues to thank for constructive discussions on the 
nature of cortical fine structure. They include Dr. 
Vernon Ahmadjian, Dr. William L. Culberson, Dr. 
D. L. Hawksworth, and Dr. Harold Robinson. Dr. 
Jerome Jacobs prepared EM sections of Parmelia 
croceopustulata and provided an interpretation of 
the micrograph that I have used. All of the photo- 
graphs except for this one (Figure 14) were re- 
produced by the Smithsonian Photographic Services 
Division. 
1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
The Scanningelectron Microscope 
The samples were photographed by Mr. Walter 
Brown of the Smithsonian Scanningelectron Lab- 
oratory with a Cambridge Stereoscan Mark 2A. Volt- 
age was uniformly 20KV since this was found to be 
most satisfactory with biological materials. The 
electron beams were perpendicular to the plane of 
the slide, giving the effect of viewing the specimens 
from directly above. In a few instances the samples 
were tilted as much as 69" to afford an oblique 
view. Separation of stereo pairs when two photo- 
graphs were taken was 7". Samples were plated with 
gold on a specially made biased rotating disk. The 
gold layer is estimated to be 200-400 angstroms 
thick. 
Samples 
All of the samples were taken from herbarium 
specimens preserved in the United States National 
Herbarium, unless otherwise stated. It would have 
been ideal to examine only type-specimens at first 
in order to establish a standard for the fine struc- 
ture of each species. I did, in fact, use types when- 
ever possible, but most specimens were recent col- 
lections identified by morphological and chemical 
comparisons with types. Older material from the 
19th century, especially that collected in the tropics, 
was often so densely covered with fine hyphae of 
molding fungi or with debris and unidentified par- 
ticles (Figure I) that surface details could some- 
times not be recognized. These artifacts are appar- 
ently caused by poor preservation. 
Preparation of the samples is very simple com- 
pared with the EM or other techniques where fixing 
and sectioning are required. One can take a dried 
lichen thallus directly from the herbarium and cut 
out sections as large as 10 mm (the maximum size 
accommodated by the Stereoscan SEM), although 1-2 
mm is adequate since, as will be shown below, the 
area of cortex examined will usually be less than 
0.20 mm. The sample is affixed to a cover glass with 
glue (in my case water-soluble Elmer's glue), air 
dried, and plated with gold in a vacuum. The en- 
tire surface abutting the cover glass must be firmly 
grounded or electrical discharge will give streaked 
photographs. Tall specimens, 1 mm or more high, 
are also prone to discharge. 
A dried herbarium specimen obviously has low 
water content, and the hyphae and algae are shriv- 
eled to some extent. Peveling and Vahl (1968) 
made a special study of possible distortion in dried 
thalli compared with thalli that had been wetted 
and freeze-dried. They found relatively little, and I 
did not pursue this problem myself. Without 
freere-drying, incidentally, the water in a wet speci- 
men would evaporate off in the vacuum used in 
gold plating and probably cause severe degassing 
and cracking. Freeze-drying would be the method of 
choice for certain kinds of studies where shrinkage 
must be avoided and there is no reason why it 
could not be used with lichens. 
A specimen can be scanned rapidly at low power 
(20-100 diameters) in order to select a standard 
area for enlargement. I usually took an area in the 
center of a lobe tip 0.5-2.0 mm back from the mar- 
gin, depending on lobe width. It is difficult at first 
to gain perspective in SEM photography because the 
range of magnification is so great. The best working 
magnification, used by both Peveling and Hawks- 
worth, is between 500 and 2500 diameters. An 
example of how much of the thallus surface is 
included in such photographs is illustrated in 
Figure 2. At 500 diameters, for example, the stand- 
ard 4x5 negative (print area about 10x11 cm) 
encompasses an area about 0.20 mm wide and 0.22 
mm long. At 1000 diameters this would be reduced 
to 0.09x0.11 mm and at 2000 about 0.05X0.06 mm. 
For this study I used two standardized enlarge- 
ments, 500 and 2000 diameters. 
Structure of the Thallus 
VERTICAL STRUCTURE 
I prepared cross-sections of a few species to get a 
better idea of the general internal organization of 
the thallus and its relation to the surface layers. Ver- 
tical or oblique free-hand sections were made with 
a razor blade and thin strips about 0.5-1.0 mm wide 
mounted on their side. These were scanned and 
photographed at different magnifications. The fol- 
lowing layers were easily identified (see Figures 3,4). 
LOWER CORTEx.-This is a discrete layer consist- 
ing of densely conglutinated hyphae with lumina 
still intact. Medullary hyphae come off from the 
top of this layer. Rhizines originate as thick strands 
(Figure 5) which are composed of vertically ar- 
ranged conglutinated hyphae in cross-section, not 
unlike those in the lower cortex (Figure 6). 
NUMBER 10 
3 
MEDULLA.-The medulla occupies the bulk of the 
thallus and has an appearance little different from 
that in microscopic preparations. Degree of packing 
and conglutination differs widely from species to 
species (Figure 7) and even from lobe to lobe in the 
same plant (cf. Figures 3,4). Encrusting crystals of 
lichen substances may be conspicuous and dense to 
absent on the medullary hyphae (Figures 7,15,18). 
ALGAL LAYER.-The algae are much more diffi- 
cult to spot than in a stained cross-section. Viewed 
stereoscopically they stand out as round though 
shriveled cells scattered within a palisade-like layer 
(Figures 3,7,8) or lying just below the upper cortex 
(Figure 10). Some may break away when sections 
are cut with this technique. In any event the algae 
seem to have a very loose association with the 
hyphae. 
UPPER CoRTEx.-The lower part of the upper cor- 
tex is similar to the lower cortex in consisting of 
densely conglutinated hyphae, though with smaller 
more compressed lumina and perhaps not as clearly 
separated from the hyphae in the algal layer. The 
layer in total appears to be somewhat thinner, two 
to three cells thick (12~-18p), although this varies 
according to the species. The structure of the upper 
part of the cortex showed considerable differentia- 
tion. This is the layer viewed with the SEM and will 
be discussed in more detail below. 
The Epicortex 
The various species in the Parmeliaceae exam- 
ined could be divided into two major groups 
depending on the presence or absence of a thin 
polysaccharide layer above the cortical hyphae. 
D. L. Hawksworth (in litt.) proposed the name 
?epicortex? for this layer, and it seems to be com- 
parable in certain respects to Peveling?s ?Kittsub- 
stanz? (cementing substance). One group of 
lichens lacks any epicortical layer, the cortical 
hyphae being exposed directly at the surface, and 
they will be called ?nonepicorticate.? The other 
major group has an epicortal layer covering the 
cortical hyphae; these, then, are epicorticate. 
NONEPICORTICATE TYPE 
The horizontally arranged hyphae in the lower 
part of the upper cortex turn upward to form a 
forest of loosely packed stubby hyphal tips (cf. 
Figures 146-150). These stand out especially well 
in a stereo view (Figure 9). This type of cortex 
is characteristic of the genus Pseudevernia and has 
been lound in some species of the Graphidaceae and 
Thelotremataceae (see Figure 11 for Graphina con- 
fluens), but it is by no means typical of crustose 
iichens. This kind of structure is apparently the one 
called fastigiate by the morphologists of the 19th 
century. An interesting consequence of this type of 
organization is that gas exchange between the algal 
layer and the exterior is completely uninhibited 
since there are many gaps between the hyphae 
providing access to the algae below. 
EPICORTICATE TYPE 
The upper cortical hyphae in this type are com- 
pletely or partially covered by a thin amorphous 
epicortex 0.6~1 .OF thick. This noncellular layer is 
secreted by the hyphae and is apparently similar in 
composition to the thick outer polysaccharide layer 
that has been found enveloping hyphae in lichens. 
There are two kinds of epicortex which may or may 
not be morpliogenetically related, one without pores 
and one with pores. 
THE NONPORED EPIc0RTEx.-Here the entire up- 
per surface is covered by the thin epicortical mem- 
brane (Figure 12a). This layer, about ly thick, 
is so closely joined to the thick polysaccharide 
walls of underlying hyphae that outlines and con- I 
volutions of the hyphae are easily distinguished. As 
a resiilt, the surface of nonpored epicorticate lichens 
viewed from above almost always appears nodular 
or mammilate (Figure 13a,b) and in an oblique 
view undulating or sinuous (Figure 1%). 
A distinctive feature of the nonpored epicortex 
is its reaction to electron beams. Photographs of 
lichens with this structure tend to be fuzzy or 
blurred, even at 500 diameters but especially so at 
2000 (see, for example, Figures 42, 49, 66, 144). It 
would appear that some of the electrons penetrate 
the gold and are reflected back from a lower layer. 
This double reflection would account for the fuzzi- 
ness. Another feature is that nonpored epicorticate 
lichens tend to degass, shrink, and crack excessively 
when exposed to intense electron beams used in 
focusing at 10,000 to 50,000 diameters (cf. Figures 
30, 64, 80). Cracking can actually be seen develop- 
ing within a few seconds. 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
THE PORED EPIcoRTEx.-The basic structure here 
is a noncellular amorphous layer 0.6p-0.8p thick 
lying on and in part free from the hyphal layer be- 
low (Figure 12b). The three-dimensional nature of 
this layer stands out vividly in stereo photographs 
(cf. Figure 8) which show free edges of the layer 
as flaps curling up (also see Figures 53, 81, 88, 105). 
A section through the cortex of Parmelia croceopus- 
tulata viewed with an electron microscope (Figure 
14) shows an electron-dense polysaccharide layer 
that stains only with lead, barely distinguishable 
from the hyphal cell walls below. The reaction of 
the polysaccharides in the pored epicortex to elec- 
tron beams is also different on the average from 
that of the nonpored epicortex. Photographs are 
almost always sharper, indicating very little electron 
penetration below the epicortex. Degassing and 
cracking at high electron intensity are also en- 
countered much less often than in the nonpored 
epicortex. 
The underlying hyphae are basically arranged 
as in the nonepicorticate type with the outermost 
ones becoming short erect stubs (Figure 15; also 
Figures 91, 105), also seen more clearly in stereo 
photographs (Figure 16). This epicortex may be 
an extremely smooth and flat sheet, giving no hint 
of the presence of underlying hyphae (Figures 15, 
18a, 78), or an undulating layer where contours of 
individual hyphae can be recognized (Figures 18b, 
119, 130). 
The most remarkable feature of this type of epi- 
cortex is the formation of pores, actual perforations 
of the polysaccharide layer exposing the underlying 
cortical hyphae (Figures 15, 91, 105) and providing 
passageways to the algal layer for gas exchange 
(Figures 3, 8) . These pores cannot be distinquished 
with a stereo binocular even under highest magni- 
fication. They are produced more or less uniformly 
over the upper cortex (Figure 17) but in some 
groups, in particular Parmelia subgenus Parmelia 
section Irregulares and subgenus Everniiformes, 
they occur in groups (Figures 70, 71, 123). 
The origin of these pores is quite unknown at 
this time. Their regularity would seem to rule out 
random tearing or imperfections in the epicortex. 
They form very early in thallus development if the 
example of Parmelia gigas may be considered 
typical. A series of panoramic photographs of the 
area from the lobe tip to a point 0.6 mm from the 
tip in this species shows a very narrow marginal 
zone at the growing point where no pores are 
present. This intergrades abruptly into a disorgan- 
ized zone which may actually be nonepicorticate. An 
epicortex gradually forms and a mature zone of 
regularly formed pores become visible (Figure 19). 
These pores differ significantly from the better 
known pseudocyphellae in size and internal struc- 
ture. Pseudocyphellae (Figures 68, 69, 138), much 
larger than epicortical pores, actually interrupt and 
penetrate both the epicortex and the underlying 
hyphnl layer, and are closed off with a loosely 
packed irregular layer of shriveled, weakly con- 
glutinated hyphae (Figure 69). The fine structural 
pores described here, rarely more than 25p in 
diameter, are formed by perforations in the epi- 
cortex which do not disturb the cortical hyphae. 
The epicortex acts as a site for deposition of 
lichen substances. This layer contains electron- 
light extracted profiles that apparently represent 
crystals of atranorin originating at its interface 
with the hyphae (Figure 14). Crystals of typical 
cortical substances, atranorin (Figures 19a, 85, 97, 
107), lichexanthone (Figures 102, 111) , and usnic 
acid (Figures 19b, c, 96, 103, 121, 135), are ulti- 
mately deposited on the surface of the epicortex 
as well as in it and are also abundant around and 
inside the pores (Figure 19b) . They readily dissolve 
out in acetone leaving a polysaccharide skeleton 
bare of acids (Figure 19b). Although the crystals 
are secreted extracellularly by the cortical hyphae, 
they soon migrate through the polysaccharide 
epicortex intact. 
FINE STRUCTURE OF THE LOWER SURFACE 
The lower surface of three epicorticate species, 
two of them pored and one nonpored, was exam- 
ined to compare them with the upper cortex. All 
were uniformly epicorticate and nonpored, even 
when the upper cortex was strongly pored (Figure 
21). As one would surmise, gas exchange through 
the lower cortex is probably negligible. 
Variation in Epicortical Structure 
We must determine the constancy and range of 
variation in fine structure of the cortex before we 
can use it as a taxonomic character. The high cost 
of SEM work and the limited availability of the 
machine, however, easily discourage one from 
achieving this goal. Although I usually examined 
NUMBER 10 
5 
only one specimen of each species, 1 prepared 
samples of two or more specimens or lobes of the 
same specimen for a number of species, especially 
when a discrepancy or point of interest was found. 
In one instance I selected five specimens of the 
pan-temperate Parmelia caperata from several 
countries, All had distinct poring and three were 
virtually indistinguishable (Figures 22a,b, 91). 
The other two had fewer and smaller pores (Figure 
22c, d) . These differences were unrelated to 
geography. We should not rule out a correlation 
with ecological conditions, but this kind of informa- 
tion is only rarely given in sufficient detail on 
herbarium labels. 
Other examples of multiple samples can be 
illustrated for Parmelia conspersa (Figures 53,54) , 
P. croceopustulata (Figures 15, 105) P. crozalsiana 
(Figures 13, 93), P. dissecta (Figures 1, 83), P. 
fissicarpa (Figures 18b,l 10) , P. formosana (Figures 
17c,ll l), P. imbricatula (Figures 17d,l 13), P. 
physcioides Figures 1,118) , P. rigida (Figures 132, 
133), and P. xanthina (Figures 204135, lobes 
from the same specimen), I feel satisfied, in any 
event, that specimen-to-specimen variation for most 
species is quite small and that as exceptions are 
found they can be subjected to more exhaustive 
study as funding and machine time permit. Hawks- 
worth (1969) had earlier found an acceptably 
small range of variation in the orientation of 
hyphae in Alectoria. 
Another source of variation could be the age 
of the lobe (cf. Hawksworth, 1969). The center of 
the lobe where I took most samples is at most 2-3 
years old, assuming growth rates of 1-4 mm per 
year from these primarily tropical foliose species. 
To make comparisons with older lobes, I examined 
one long linear lobe of Parmelia gigas, photograph- 
ing the center of the tip (Figure 112), a point 5 cm 
back from the tip which I estimated to be about 
10 years old, and a point 10 cm back from the 
same tip (Figure 23) which may be as much as 20 
years old. Pores were best developed at the tip but 
were easily distinguished at 5 cm back and less so 
at 10 cm. Perhaps the pores in older areas gradually 
fill in as the need for gas exchange diminishes. 
VALUE OF CORTICAL FINE STRUCTURE IN SYSTEMATICS 
The family Panneliaceae was chosen for this 
study because it is a large, varied, and taxonomi- 
cally well-known group of at least 11 genera, one of 
which (Parmelia) is divided into 13 major sub- 
genera and sections. As I had hoped and as will be 
seen in the photographs, cortical fine structure 
proves to be a remarkably constant character at 
the generic and sectional levels. A summary of the 
results is given in Table 1 and details of individual 
groups are discussed more fully below. The most 
interesting discovery was that lichens with pseudo- 
cyphellae (or even nonpseudocyphellate lichens in 
genera that may produce them) never form a pored 
epicortex. Conversely, lichen species with a pored 
epicortex never produce pseudocyphellae. 
I can cite several examples where epicortical 
structure has helped in assigning species to the 
correct section in a genus. Parmelia uanderbylii 
Zahlbruckner was described from South Africa as 
TABLE 1.-Numerical and structural survey of 
species in Parmeliaceae 
Taxa 
Anzia .................................. 
Asahinea ............... 
Cnvernularia ......................... 
Cetraria ........................... 
Cetrelia ...................... 
Heterodea ..................... 
Nppogy mnia .................. 
Men egazzia ............... 
Pannoparmelia ................... 
Pa rnielia ................... 
Subgenus Melanopharm 
Subgenus Xanthoparmelia 
Subgenus Parmelia 
Section Parmelia ....... 
Section Simplices , . , , , , , . , 
Section Irregulares , , , . , , . , 
Section Relicinae , ,., ,,. 
Section Ricornutae ...... 
Section Im bricaria , , . . , , , 
Section Hypotrachyna . , 
Subgenus Everniiformes ,. 
Subgenus Amphigymnia . 
Parmeliopsis ............... 
Platismatia ......................... 
Pseudevern iac .................. 
Section Cyclocheila ..... 
80 8 + (+I 
80 7 + 
25 6 + + 
25 3 + + 
73 + 
25 6 + 
30 4 + 
38 8 + 
49 11 (+) + 
90 22 + 
14 4 + 
125 9 + 
52 + 
10 3 + + 
65 
* = Nonepicorticate. ?=Uncertain (?) =Rare occurrence. 
6 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
a member of section Melanoparmelia. It is indeed 
a small dark species, superficially a typical brown 
Parmelia. The SEM photograph, however, shows a 
pored epicortex more typical of species in section 
Cyclocheila (Figure 99), not of Melanoparmelia 
which always has a nonpored epicortex. Another 
example is Parmelia jamesii Hale, a species from 
New Zealand which seemed to belong in section 
Parmelia (the P. saxatilis group) because of the 
faint white reticulation at the lobe tips. The epi- 
cortex is pored (Figure 85) and this species can 
therefore be tentatively placed in section Zmbri- 
curia since species in section Parmelia are nonpored. 
A final example is the common Parmelia ulophyl- 
lodes (Vainio) Savicz, which appears to belong to 
section Cyclocheila but which, except for lack of 
distinct pseudocyphellae, also falls near P. flaven- 
tior and definitely does not belong in section 
Cyclocheila, although its ultimate disposition has 
not yet been decided. 
While the structure of the epicortex is a valuable 
character at the sectional level, too few data are 
available to evaluate its use as a species character. 
I did find one case, Parmelia formosana Zahlbruck- 
ner vs. P. pustulifera Hale (Figures 111, 119), 
where pore size and evenness of the surface were 
sufficiently different to be used with other characters 
to distinguish two species (Hale, 1972) . Preliminary 
study also suggests that pore size and density are 
correlated with chemistry in the isidiate species 
of the P. laevigata complex, including P. addita 
Hale (Figure loo), P. dentella (Figure 106), P. 
orientalis Hale (Figure 117), and P. imbricatula 
(Figure 113). Many more specimens will have to 
be examined, however, to make a conclusive judg- 
ment in this case. 
The essential features of each genus are discussed 
briefly below. Further elaboration on the classifica- 
tion and characters used in the taxonomy may be 
found in Culberson and Culberson (1968a), Hale 
and Kurokawa (1964), and Hale (1965). 
Anzia Stizenberger 
FIGURB 24-26 
SPECIES EXAMINED.-Anria colpodes, A. japonica, 
and A. ornata. 
While this genus is better placed in the Anziaceae 
(Culberson and Culberson, 1968a), it is included 
here because of its long association with the Par- 
meliaceae. The two characters that distinguish the 
genus are numerous spores in each ascus (rather 
than the normal eight) and a unique mesh of 
black hyphae on the lower surface. There is a 
well-developed epicortex which appears to be non- 
pored or irregularly pored. The unusual sculpturing 
in A. colpodes (Figure 24) is similar to that in 
Heterodea muelleri (Figure 36) , The epicortical 
structure, then, seems to support the logic of a 
separdte family, but a clearer definition of the 
cortex here will require examination 
specimens. 
of many more 
Asahinea Culberson 
FIGURES 27, 28 
SPECIES EXAMINED.-Asahinea chrysantha and A. 
scholanderi. 
This is one of the segregates of Cetraria, sensu 
lato, proposed by Culberson and Culberson 
(1968b). Both species investigated have a smooth 
to undulating nonpored epicortex similar to that 
in other cetrarioid species. 
Cavemularia Degelius 
(not illustrated) 
This small genus of two species (C. hultenii 
Degelius and C. lophyrea (Acharius) Degelius) is 
superficially similar to Hypogymnia but has unique 
invaginations of the lower cortex into the medulla. 
Both species have a typical nonpored epicortex. 
Cetraria Acharius 
FIGURES 29-35 
SPECIES EXAMINED.-ct?tTa?"ia commixta, c. halei, 
C. nivalis, C. oakesiana, and C. pinastri. 
This large temperate-boreal genus has great 
morphological diversity, ranging from tiny foliose 
species like Cetraria fendleri (Tuckerman) Ny- 
lander to the fruticose Iceland Moss (C. islandica 
(L.) Acharius) . I examined five species and found 
all to have a nonpored epicortex and a nodular 
surface. This would seem to be the characteristic 
cortical structure for the genus. 
NUMBER 10 7 
Cctrelia Culberson and Culberson 
FIGURES 34, 35 
SPECIES EXAMINED.-cetl.elia chicitae and c. olive- 
torum. 
This genus includes broad-lobed pseudocyphel- 
late parmelioid species formerly classified in either 
Parmelia or Cetraria (Culberson and Culberson, 
1968b). As one would expect, the epicortex is non- 
pored and the surface nodular. 
Heterodea Muller-Argau 
FIGURE 36 
SPECIES ExAMlNED.-Hetel-odea mue/le?.i. 
This genus is endemic to Australia and has been 
classified in the Parmeliaceae more for want of a 
more appropriate family than from any real affinity. 
The cortical structure is unusual, a nonpored epi- 
cortex with very deep sculpturing, much deeper 
than in Anzia or Hypogymnia. The genus should 
probably be removed from this family. 
This primary austral genus, a segregate of Par- 
melia, has hollow lobes as in Hypogymnia, large 
perforations in the upper cortex, and no rhizines. 
Both species examined have a distinctly epicorticate 
nonpored layer and a nodular surface, not unlike 
that in Hypogymnia. 
Pannoparmelia Darbishire 
FIG- 45, 46 
SPECIES EXAMINED.-PannOpa7meEia angustata and 
P. anzioides. 
This genus is very similar to Anzia in having a 
dense network of black hyphae below but differs 
in having eight spores and, in the two species seen, 
usnic acid. The upper layer of hyphae are arranged 
mostly horizontally in an irregular configuration 
and at least some pores appear to originate from 
gaps between hyphae, although the epicortex is 
clearly developed (Figure 8b). Interestingly it 
differs in this respect from Anzia (Figures 24-26) 
which is basically nonpored and nodular. 
Hypogymnia (Nylander) W. Watson 
Parmelia Acharius 
FIGURES 3742  FIGURE^ 47-141 
SPECIES ExAMINED.-Hypogymnia bittel-iana, H. 
enteromorpha, H. metaphysodes, and H. physodes. 
This is a well-established segregate of Parmelia 
with hollow lobes and no rhizines. Nine specimens 
of four species were examined, but it is obvious that 
much more study will be needed to find a pattern 
in the genus. Most specimens appear to be typically 
nonpored and epicorticate. Hypogymnia bitteriana 
has very sparce pores while one specimen of H. 
enteromorpha had what seemed to be distinct pores. 
Eastern United States specimens of H. enterornor- 
pha, long suspected of being specifically distinct 
from the western population, have greater or lesser 
development of the pecularly sculptured furrowing 
seen in Anzia and Heterodea. 
Menegarzia Massalongo 
FIGUW 43, 44 
SPECIES ExAMINED.-Menegazzia cincinnata and 
M. terebrata. 
Subgenus Melanoparmelia Hue 
FIGURFS 47-52 
SPECIES ExAMINED.-Parmelia acetabulum, P. dis- 
juncta, P. olivacea, P. ryssolea, P. septentrionalis, 
and P. subolivacea. 
The systematic position of this subgenus is not 
firmly established. The main character for the 
group is the brown color attributed to an uniden- 
tified apparently noncrystalline pigment, ?Parmelia 
brown.? The. thallus morphology, short often 
rotund lobes, simple rhizines, and lack of cilia, 
places it superficially near section Cyclocheila of 
subgenus Parmelia, which has a pored epicortex. 
The infrequent Occurence of small pseudocyphellae 
might even relate it to section Simplices. All six 
species (eight specimens) investigated have a non- 
pored epicortex and a more or less strongly nodular 
surface. Variation between species is very small. 
All evidence suggests that Melanoparmelia should 
have at least subgeneric status in Parmelia. 
8 ShlITHSONIAN COSTRIBUTIOSS TO BOTANY 
Subgenus Xanthoparmelia (Vainio) Hale 
FIGURES 53-60 
SPECIES ExAMINED.-Paimelia conspersa, P. cum- 
berlandia, P. hyporhytida, P. ioanis-stmae, P. plittii, 
P. taractica, and P. tasmanica. 
All species in this subgenus are saxicolous 
(rarely on soil), contain usnic acid, and have 
simple rhizines without marginal cilia. Section 
Cyclocheila, except for lack of usnic acid in most 
species, is very close in morphology. Seven species 
and nine specimens in subgenus Xanthoparmelia 
were examined. All have a pored epicortex with 
abundant deposition of usnic acid crystals. The 
well-known P. conspersa (Figure 54) has very large 
pores but the largest occur in P. hyporhytida 
(Figure 56). Two closely related species, P. tar- 
actica (Figure 59) and P. tasmanica (Figure 60), 
which differ primarily in color of the lower surface, 
have sparse development of pores, a character that 
may well separate them from the ?conspersa? 
group. The SEM may, indeed, prove to be a useful 
way to delineate the numerous otherwise poorly 
differentiated species in this subgenus. 
Subgenus Parmelia 
FIGURES 61-121 
Section Parmelia 
FIGURES 61-66 
SPECIES EXAMINED.-PUimelia cunninghamii, P. 
fraudans, P. omphalodes, P. saxatilis, P. squarrosa, 
and P. sulcata. 
The arctic-boreal to temperate species in this 
section have similar lobe configuration, pseudo- 
cyphellae or pseudocyphellae-like white reticulation, 
and simple chemistry (salacinic or rarely proto- 
cetraric acid). Six species and eight specimens 
studied consistently had a nonpored epicortex and 
nodular surface. The cortex is virtually indistin- 
guishable from that in section Simplices. 
Section Simplices (Hale and Kurokawa) Hale 
FIGURES 67-69 
SPECIES EXAMINED.-Parmelia bolliana, P. borreri, 
and P. rudecta. 
This small group of temperate pseudocyphellate 
Parmeliae has a uniformly nonpored epicortex and 
a nodular surface. It is closely related to section 
Parmelia above in fine structure but the chemical 
characters are completely different, the chief lichen 
substances being gyrophoric and lecanoric acids. 
Section Zrregulares (Vainio) Vainio 
FIGURES 70, 71 
SPECIES ExAMINED.-Parmelia cetrata, P. maura, 
and P. simulans (not illustrated). 
This small section is characterized by a finely 
reticulately cracked cortex. The cracks show up 
well in the SEM photographs (Figure 71). Three 
specimens and three species have a very similar 
pored epicortex and a smooth surface. The pores 
tend to be grouped with broad intervening areas 
of noppored cortex. This section forms a link 
between subgenus Amphigymnia and the narrow- 
lobed subgenus Parmelia (especially section Im bri- 
caiia) but the epicortical structure does not help 
us decide to which group it is better assigned. 
Section Relicinae (Hale and Kurokawa) Hale 
FIGURES 72-76 
SPECIES EXAMINED.-P. fluorescens, P. malesiana, 
P. planiuscula, P. ramosissima, and P. subabstrusa. 
Species in this section have inflated marginal 
bulbate cilia and always produce usnic acid, al- 
though the crystals are not easily seen in the SEM 
photographs. All of the five species and eight 
specimens examined have a pored epicortex. The 
pores are generally small and relatively sparce, a 
correlation perhaps with the humid environment 
of the tropical rainforest where most of the species 
have evolved. 
Section Bicornuta Lynge 
FIGURES 77-80 
SPECIES EXAMINED.-P. confoederata, P. coronata, 
P. pigmentacea, and P. subinflata. 
This section has the same bulbate cilia as section 
Relicinae but lacks usnic acid. Few of the species 
occur in the humid lowland tropics; most are 
montane-tropical or temperate. Four species and 
NUMBER 10 9 
five specimens all have a distinct pored epicortex, 
the pores in P. coronata being unusually well de- 
veloped (Figure 78). 
Section Zmbricaria (Schreber) Fries 
FIGURES 81-88 
SPECIES ExAMINEu.-Parmelia antillensis, P. aurul- 
enta, P. dissecta, P. horrescens, P. jamesii, P. quer- 
cina, P. scortea, and P. wallichiana. 
There is a great range of morphological variation 
in this rather heterogeneous assemblage of margin- 
ally ciliate species. Eight species and ten specimens 
studied have a pored epicortex and a smooth to 
undulating surface. Pore variation was rather great 
in P. dissecta. The holotype from France has very 
large elongate pores (Figure la,b,) whereas two 
other specimens, one from the United States and 
one from Japan (Figure 83) have uniformly smaller 
round pores. The curly objects on the type of P. 
dissecta (Figure la,b) and on P. wallichiana (Fig- 
ure 88) have not been identified. 
Section Cyclocheila (Vainio) Rasanen 
FIGURFS 89-99 
SPECIFS ExAntrivEu.-Parmelia aptata, P. baltimor- 
ensis, P. caperata, P. congruens, P. crozalsiana, P. 
rryptochloi.oplraea, P. mnlaccensis, P. rutidota, P. 
salacinifera, P. texana, and P. vanderbylii. 
This section is also rather heterogeneous and 
contains many narrow-lobed species with simple 
rhizines but lacking marginal cilia. Some of the 
species, e.g., Parmelia caperata, have rather broad 
rotund lobes, but the bare marginal zone below is 
much narrower than in subgenus Amphigymnia. 
Sixteen specimens of eleven species have a pored 
epicortex. Parmelia crozalsiana was represented by 
one specimen with small sparse pores (Figue 93) 
and two specimens with a typical, well-developed 
nonpored epicortex with a nodular surface, (Figure 
13a,b), much as in section Parmelia although there 
is no other justification for placing it there. Three 
other species that are closely related in lobe con- 
figuration and chemistry, P. aptata (Figure 89), P. 
cryptochlorophaea (Figure 94), and P. texana (Fig 
ure 98), also tend to have a strongly nodular sur- 
face, but pores, though small and sparce, are 
developed in the epicortex. 
Section Hypotrachyna Vainio 
FICUW 100-121 
SPECIES EXAMINED.-PaYmelia addita, P. adjuncta, 
P. brasiliana, P. caraccensis, P. chicitae, P. croceo- 
pustulata, P. dentella, P. endochlora, P. erythro- 
cardia, P. exsecta, P. fissicarpa, P. formosana, P. 
gigas, P. imbricatula, P. laevigata, P. livida, P. 
neodissecta, P. orientalis, P. physcioides, P. pustuli- 
fera, P. scytodes, and P. sinuosa. 
This is the largest single section in the subgenus. 
All species have dichotomously branched rhizines 
and sublinear lobes. Most have evolved in tropical 
montane regions. Chemical variation is very rich 
and indicates a high level of evolution. There was 
a considerable degree of uniformity in cortical 
structure among the 38 specimens of 22 species 
examined. All have a pored epicortex, and although 
variation in pore size and density between species 
may be great, specimen-to-specimen variation in the 
same species is not. 
Subgenus Everniifmmes (Hue) Hale and Wirth 
FIGURES 122-125 
SPECIES ExAMiNED.-Parmelia americana, P. neo- 
cirrhata, P. pseudonepalensis, and P. vexans. 
This small montane-tropical group has linear 
channeled lobes and long marginal cilia. Rhizines 
are either absent below or simple to branched. It 
seems most closely related to section Hypotrachyna 
although the rhizines are quite different. Four 
species (five specimens) examined all have a pored 
epicortex with some clumping of the pores. 
Subgenus Amphigymnia Vainio 
FIGURES 126135 
SPECIES ExAMINED.-Parmelia crist iferu, P. haitien- 
sis, P. hypoleucina, P. hypotropa, P. louisianae, P. 
perforata, P. rigida, P. tinctorum, and P. xanthina. 
This subgenus is characterized by broad rotund 
lobes 10-30 mm wide and a zone at the margin 
below free of rhizines. Both chemical and morpholo- 
gical diversity are great. Nine species and eleven 
specimens were examined. All except P. perforata 
have a well-developed pored epicortex, and no 
significant difference could be found between the 
two major sections, Amphigymnia and Perlatae. 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
Parmelia pei.forata (Figure 131) is anamolous in 
that much of the cortex appears to be nonepicorti- 
cate as in Pseudevernia. It is of interest that its 
presumptive chemical counterpart, P. rigida (Fig- 
ure 133) which contains alectoronic acid instead of 
norstictic acid, has a nmal pored epicortex, as 
does the presumed sorediate counterpart of P. 
perforata, P. hypotropa (Figure 129). 
Pannelia Species of Uncertain Position 
FIGURES 136-141 
Several species of Parmelia have not yet been 
assigned to any of the recognized sections because 
they are anomolous in one or more respects. Parm- 
elia Raventior, for example, seems amphigymnioid 
but differs sharply from species in subgenus Am- 
phigymnia in having pseudocyphellae. On the other 
hand, the chemistry (lecanoric acid and usnic acid) 
is totally different from that of Cetrelia where it 
might otherwise be placed. As would be expected 
with a pseudocyphellate species, the epicortex is 
nonporetl and nodular. Parmelia ulophyllodes 
(Figure 141) appears to be related to P. flaventior 
but lacks obvious pseudocyphellae and is smaller, 
much as the species in subgenus Parmelia section 
Cyclocheila. It is, however, nonpored and very 
similar to P. paventior. Another relative of P. 
flaventior, P. praesignis (Figure 138), lacks soredia 
but has identical epicortical structure. Another 
amphigymnioid species, P. thomsonii (Figure 140) , 
has a normal pored epicortex but because of the 
conspicuous erect marginal pycnidia does not fit 
well into subgenus Amphigymnia. 
Parmelia sphaerosporella has all of the character- 
istics of a species in section Cyclocheila, but it has 
a nonpored nodular epicortex (Figure 139). Fin- 
ally, P. meiophora, an endemic Asian species, ap 
pears to be in section Zmbricaria but the nonpored 
epicortex and probably pseudocyphellae place it 
closer to section Parmelia (Figure 137), 
Parmeliopsis Nylander 
 FIGURE^ 2, 142 
SPECIES ExAMINED.-Parmeiiopsis am bigua and P. 
placorodia. 
This small genus resembles Parmelia rather 
closely but always has a pale lower surface and a 
different type of conidiophore. The SEM photo- 
graphs show a strongly pored epicortex similar to 
that in many sections of Parmelia, suggesting a 
close affinity. 
Platismatia Culberson and Culberson 
FIGURE 143-145 
SPECIES ExAMINED.-Platismatia glauca, P. steno- 
phylla, and P. tuckermanii. 
This small segregate of Cetraria includes the 
widespread boreal P. glauca. The epicortical struc- 
ture is quite uniform, nonpored and nodular. It 
falls very close to both Cetrelia and Asahinea. 
Pseudevernia Zopf 
FIGURES 146-149 
SPECIES EXAMINED.-Pseudevernia cladonia, P. 
ronsocians, P. ful-furacea, P. olivetorina, and P. in- 
tensa. 
Although only recently generally accepted as a 
segregate of Parmelia, this genus is amply distinct 
in morphology (Hale, 1968). Rhizines are lacking 
and the lower surface is strongly channeled. The 
surface structure is unusual. There is no epicortex 
at all, the crowded erect hyphal tips forming a kind 
of pseudocortex with various degrees of conglutin- 
ation. 
Atlas of SEM Photographs 
The following series of photographs (Figures 24- 
150) illustrates the cortical surface of 123 species 
taken at two standardized enlargements, 500 
diameters (upper photograph, 1 mm = 2~) and 
2000 diameters (lower photograph, 2 mm = 1~). 
See Figure 2 for the relative areas included in the 
original photographs. The final prints as shown 
here were trimmed to 9 x 9 cm. The x 2000 en- 
largement was usually taken at or near the center 
of the x 500 photograph and included in it, but in 
a few cases it was selected outside this area to avoid 
accumulations of debris. Many kinds of artifacts are 
to be seen on the cortical surface, including pollen 
grains (Figures 66, 108, 123), crystals of lichen 
substances (Figures 85, 121, 135), pruina caused by 
calcium oxalate crystals (Figure 93), hyphae of 
saprophytic fungi (Figures 61, 74, 89, 95, 97, 108), 
and unidentified debris (Figures 25, 87). White 
NUMBER 10 
11 
areas in the photographs, especially around pores, 
are created by a thinner layer of gold. 
The photographs are arranged alphabetically by 
genus and within each genus by species. In Parmelia 
the sections are phylogenetically arranged but 
within sections the species are alphabetic. Data to 
identify each specimen include collector, collector?s 
number, and country or state of origin. 
Literature Cited 
Culberson, W. L., and C. F. Culberson. 
1968a. A Phylogenetic View of Chemical Evolution in the 
Lichens. Bryologist, 73:l-33 
1968b. The Lichen Genera Cetrelia and Platismatia (Par- 
meliaceae). Contributions from the United States 
National Herbarium, 34:449-558. 
A Monograph of Parmelia Subgenus Amphigymnia. 
Contributions from the United States National 
Herbarium, 36 193-358. 
A Revision of the Lichen Genus Pseudevernia. Bry- 
ologist, 71:l-11. 
Parmelia pustulifera, a New Lichen from South- 
eastern United States. Brittonia, 24:22-27. 
Hale, M. E. 
1965. 
1968. 
1972. 
Hale, M. E., and S. Kurokawa. 
1964. Studies on Parmelia Subgenus Parmelia. Contribu- 
tions from the United States National Herbarium, 
36:121-191. 
The Scanning Electron Microscope: An Aid to the 
Study of Cortical Hyphal Orientation in the 
Lichen Genera Alectoria and Cornicularia. Journal 
Microscopie, 8:753-760. 
Hawksworth, D. L. 
1969. 
Jacobs, J. B., and V. Ahmadjian. 
1969. The Ultrastructure of Lichens, I: A General Sur- 
vey. Journal of Phycology, 5:227-240. 
Peveling, E. 
1970. Die Darstellung der Oberflachenstrukturen von 
Flechten mit dem Raster-Elektronenmikroskop. 
Berichte der Deutschen Botanischen Gesellschaft, 
neue folge 4:89-101. 
Die Anwendung der Gefrieratzmethode fur Unter- 
suchungen mit Raster-Elektronenmikroskop im 
Vergleich mit herkommlichen Untersuchungsmeth- 
oden. Beitrag Elektronenrnikroskopiche Direktab- 
bildung von Oberpachen, 1:205-212. 
The Genus Cyphelium in Europe. Svensk Botanisk 
Tidskrift, 65: 138-164. 
Peveling, E., and J. Vahl. 
1968. 
Tibell, L. 
1971. 

NUMBER 10 
FIGURES 1-150 
FIGURE 1.-Artifacts on the cortical surface: a, unidentified curled artifacts on Parmelia dissccta 
Nylander (Nylander, France, holotype, P) (~500); b, same specimen enlarged (x2WO); c, 
presumed fungal infection of P. physcioides Nylander (Humboldt and Bonpland s.n., Venezuela, 
isotype, US) (~500); d, same specimen enlarged (xZO00). 
13 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
NUMBER 10 15 
FIGURE P.-Comparison of areas viewed with the SEM at dif- 
ferent magnifications using Parmeliopsis ambigua (Acharius) 
Nylander (Tuckerman, New Hampshire): a, x50, lobe about 
2 mm long showing approximate areas viewed at ~100 
(larger rectangular shown in b) and at x500 (smaller rec- 
tangular equivalent to area viewed in c); b, xl00 (showing 
area equivalent to that ir. c); c, ~500, print area about 0.20 
x 0.22 mm (rectangle indicating area enlarged in d); d, 
x2O00, print area about 0.05 x 0.06 mm. 
FIGURE 3.-Vertical cross-section of Parmelia xanthina (Muiller-Argau) Vainio (same plant shown 
in Figure 135), showing the upper cortex (epicortex too thin to see), a pore (upper arrow), 
palisadelike algal layer (algal cell identified with arrow), medulla, and lower cortex (lower 
arrow) (about ~1000, 1 mm=about lP). 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 5.-Lower surface of Parmelia saxatilis (L.) Acharius 
(Hale 36387, Canada) (~100). (Stump of broken rhizine in 
right center enlarged in Figure 6.) 
FIGURE 4.-Oblique cross-section of Parmelia xanthina (second 
lobe of plant shown in Figure 3) (about ~700). 
FIGURE 6.-Cross-section of a rhizine of Parmeh saxatilk 
(same specimen shown in Figure 5) 
(~2000). 
NUMBER 10 
17 
FIGURE 7.-Cross-sections of various species of Parrnelio: a, Parinelia saxatilis (same plant shown 
in Figure 64); b, P. fluorescens Hale (same plant shown in Figure 72) 
(this species has an 
unusual arrangement of the cortical hyphae in columns); c, P. neodissecta Hale (same plant 
shown in Figure 116) (blob at the top right is glue; arrow points of top of cortex); d, P. cro- 
ceopustulota Kuroknu~n (Hale 33269, North Carolina) n, b, andd, about ~500; c about ~600. 
FIGURE %-Stereo photographs of a cross-section of Parmelia croceopustulata (same plant shown 
in Figure 105) (xlO00). These can be viewed with a standard stereo map viewer at approxi- 
mately 12 an above the photograph. 
FIGURE 9.-Stereo view of a cross-section of Pseudevernia furfuracea (L.) Hale and Culberson 
showing arrangement of cortical hyphae, the cortical layer, and the upper medullary area (same 
plant shown in Figure 148) (x 1000). Compare Figure 148 for a perpendicular view of the same 
plant. 
FIGURE 10.-Close-up of the algal layer in Parmelia xanthina 
(same plant shown in Figure 4) (~2000). Arrow points to an 
algal cell. Part of the upper surface of the cortex, the cortical 
layer, and part of the medulla are also shown. 
FIGURE 11.-Surface of Crafihina confluens (Fee) Muiller -Argau 
(Hale 20169, Mexico) (~2000). 
FIGURE lZ.-Cross-sections of the cortical layer in (a) a non- 
pored epicorticate species, Parinelia bollinnu Muiller-Argau 
(Adam 34, Oklahoma) (xlO00) and (b) a pored epicorticate 
species, P. croceopustulata Kurokawa (same plant shown in 
Figure 74 (~5000). Arrow points to epicortex. 
20 
ShfITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 13.-Cortex surface of some nonpored epicorticate species: a, nodular surface of Parmelia 
croralsiana Bouly de Lesdain (Skorepa 1757, Ilinois) (~500); b, same plant enlarged ~2000; c, 
oblique view of the nodular surface of P. squarrosa Hale (same plant shown in Figure 65) 
(About ~2500) (Compare Figure 65 for a perpendicular view of this specimen). 
NUMBER 10 
FIGURE 14.-Section of the cortex and epicortex of Parmelia croceopustulata Kurokawa (same 
plant as in Figure 105) viewed with electron microscope (~24,000). The electron-light extracted 
areas at periphery of the hyphae and below the epicortex are apparently sites of lichen acid 
deposition. Photograph by Dr. Jerome Jacobs. 
21 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 15.-Cross-section of Parmelia croceofiustulata Kuro- 
kawa (same plant shown in Figure 105), showing the flat 
epicortex, one pore, hyphal stubs in the pore, cross-section of 
the cortical hyphae, and top of the medulla and crystal- 
encrusted hyphae (about x2OOO). 
FIGURE 16.-Stereo view of the sur- 
face of Parmelia croceopustulata 
Kurokawa (same plant shown in 
Figure 'id) (~1000) . For method of 
stereo viewing see Figure 8. 
FICURF. 17.-Pore configuration in se\ era1 species of Pnrtnelia: a, general arrangement of pores in 
part of a lobe of P. azwulenta Tockerman (same plant shown in Figure 82) (~50); b, P. 
fissicarpa Kurokawa (Esterhuysen 101 15, Cape Province, South Africa) (~500); c, P. formosana 
Zahlbruckner (Hale 21883, Mexico) (XjOO) : d, P. inibricalula Zahlbruckner (same plant shown 
in Figure 18a) (~500). 
FIGURE 18.-Oblique views of the cortex surface of pored epi- 
corticate species: a, smooth surface of P. imbricatula Zahl- 
bruckner (Kurokawa 2181, Java) (x 1OOO) (for perpendicular 
view of the same plant see Figure 18d) ; b, nodular surface of 
P. aurulenta Tuckerman (Haze 17391, Pennsylvania) (x 1OOO) 
(for perpendicular view of a different specimen see Figure 
82). 
FIGURE 19.-Panorama of photographs taken from the lobe 
tip to about O.GO mm from the tip of Parmelia gigas Kuro- 
kawa (Killip 18394, Columbia). Center of this lobe (about 
1.5 mm from the tip) is illustrated in Figure 112. 
NUMBER 10 
25 
FIGURE 20.-Deposition of crystals of lichen substances in the epicortex: a, atranorin in ParrneIia 
gigas Kurokawa (same plant shown in Figure 112) (~2000) ; b, usnic acid around a pore in P. 
sinuosa (Smith) Achariiis (same plant shown in Figure 121) (~10,000) : c, usnic acid in P. 
xanthina (Muller-Argau) Vainio (second lobe of plant shown in Figure 135) (~2000) : d, sample 
from same lobe of specimen in c but extracted with acetone (~2000). 
26 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 21.-Comparison of the upper and lower surfaces of 
Parmeh perforatu (Jacquin) Acharius: upper surface (above: 
see also Figure 131) and lower surface from a lobe on the 
same specimen (Hale 16921, Florida) (both x2OOO). 
NUMBER 10 
27 
FIGURE PP.-Comparison of pore development in four different specimens of Parrnelia caperata 
(L.) Acharius: a, Hale 20343, Mexico; b, Maas Geesteranus 14153, Netherlands; c, Culberson 
12000, France; d, Kurokawa 64462, Japan (all ~500). See Figure 91 for another specimen of 
Parmelia caperata. 
28 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 23.-Pore development in Parmelia gigas Kurokawa at an area 5 cm from the lobe tip 
(left) and 10 cm from the tip (right) (same plant illustrated in Figures 19 and 112) (~500). 
NUMBER 10 
FICURF 24.--Anzia colpodes (Acharius) Stizenberger 
(Willey 3, Massachusetts). 
29 
FIGURE 25.--Anxia japonica (Tuckerman) Muller-Argau 
(Wright s.n., Japan). 
30 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 26.-Anxia ornata (Zahlbruckner) Asahina (Kurokawa FIGURE 27.-Asahinea chrysantha (Tuckerman) Culberson and 
58239, Japan). Culberson (Potter 6103, Alaska). 
NUMBER 10 
31 
FIGURE ZE.-Asahinca scholanderi (Llano) Culberson and Cul- 
berson (Thomson 15715, Canada). 
FIGURE 29.4ctraria commixta (Nylander) T. Fries (Crombie 
m., England). 
32 
ShlITHSOPI'IAN CONTRIBUTIONS TO BOTANY 
FIGURE 30.--Cetraria haEei Culberson (Hale 34200, Michigan). FIGURE 31 .--Cetraria nivalis (L.) Acharius (Bird 17254, 
Canada). 
NUMBER 10 33 
FIWRE 32.--Cetraria oakesiana Tuckerman (Culberson 4556, FIGURE 33.-Cetraria pinastri (Scopali) S. F. Gray (Bourn- 
New Hampshire). gartner 874 in Kryptogamae Vindobonensis, Austria). 
34 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 
FIGURE 34.-Cetreliu chicitae (Culberson) Culberson and Cul- FICVRE 35.--Cetreliu olivetorun (Nylander) Culberson and 
berson (Hale 19212, Virginia). Culberson (Hale '33411, Virginia). 
NUMBER 10 
35 
FIGURE 36.-Heterodea muelleri (Hampe) Nylander (James FIGURE 37.-Hypogymnia bitteriana (Zahlbruckner) Krog 
2106a, Australia). (Kjellm'ert s.n., Sweden). 
36 ShfITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 38.-Hypogymnia enteromorpha (Acharius) Nylander FIGURE 39.-Hypogymnia enteromorpha (Acharius) Nylander 
(Hale 10972, West Virginia). (Hale 33698, California). 
NUMBER 10 37 
FIGURE 4O.-Hypogymnia enteromorpha (Acharius) Nylander FIGURE 41.-Hypogym,nia metaphysodes Asahina (Fujikawa 
(Hale 21520, Oregon). s.n:, Japan, isotype specimen). 
38 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 42.-Hypogymnia physodes (L.) Nylander (Culberson FIGURE 43.-Menegazzia cincinnata (Acharius) Bitter (Mahu 
11994, France). 1642, Chile). 
NUMBER 10 
39 
FIGURE 44.-Menegazzia terebrata (Hoffmann) Massalongo FIGURE 45.-Pannoparmelia angustata (Persoon) Zahlbruckner 
(Denison 836, Canada). (Imshaug 47661, New Zealand). 
40 SIIlITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 46.-Pannoparm'elia anzioides Darbishire (Imshaug FIGURE 47.-Parmelia acetabulum Acharius (Tomin s.n., 
45476, Chile). Russia). 
NUMBER 10 
FIGURE 48.-Parmelia disjuncta Erichsen ( Thomson 12466, 
Canada), 
41 
FIGURE 49.-Parmelia olivacea (L.) Acharius (Hale sen., 
Canada). 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 50.-ParmeZia ryssolea Acharius (Kraschennikov s.n., FIGURE 51 .-Parmelia septentrionalis (Lynge) Ahti (Hale 
Russia). 33909, Canada). 
FIGURE 52.-Parnelia su bolivacea Nylander (Shushan 15202, FIGURE 53.-Parmelia conspersa (Acharius) Acharius (Hale 
California). 34653, Canada). 
44 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 54.-Parmelia conspersa (Acharius) Acharius (Sampaio FIGURE BEi.-Parmelia cumberlandia (Gyelnik) Hale (Hale 
246, Portugal). 36560, Canada). 
FIGURE 52.-Parnelia su bolivacea Nylander (Shushan 15202, FIGURE 53.-Parmelia conspersa (Acharius) Acharius (Hale 
California). 34653, Canada). 
NUMBER 10 45 
FIGURE 56.-ParmeZia hyporhytida Hale (Kofler s.n., Basuto- FIGURE 5?i-Parmelia iounis-simae Gyelnik (NichoGas 5982, 
land, holotype in LD). Mexico, isotype specimen). 
46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
Ficwm 58.-Parmelia plittii Gyelnik (Denison 683, Canada). FIGURE 59.-Parmelia taractica Krempelhuber (Hakulinen 
sn., Finland). 
NUMBER 10 47 
FIGURE 60.-Parmelia tasmanica Hooker and Taylor (Hale FIGURE 61.-Parmelia cunninghamii Crombie (Cunningham 
23606, Minnesota). s.n., Falkland Islands, holotype in BM). 
48 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE BP.-ParmeZia fraudans Nylander (Hale 36414, FIGURE 63.-Parmelia omphalodes (L.) Acharius (Muller 
Canada). 1355, Alaska). 
NUMBER 10 49 
FIGURE 64.-Parmelia saxatilis (L.) Acharius (Hale sm., FIGURE 65.--ParmeZia squarrosa Hale (Hate 18922, Virginia). 
Canada). 
50 
FIGURE 66.-ParmeZia sulcata Taylor (Hale 14914, 
Connecticut). 
ShfITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 67.-Parmelia bolliana Muiller-Argau (Hale 23691, 
Minnesota). 
NUMBER 10 
51 
FIGURE 68.-PurmeZia borreri (Smith) Turner (Culberson FIGURE: 69.--ParmeZiu rudecta Acharius (Hale 14769, Conecti- 
cut), .4 large pseudocyphella is included in the photographs. 10099, Ohio). 
52 
SMITHSONIAN COETRIBUTIONS TO BOTANY 
FIGURE 70.--Parmelia cetrata Acharius (Hale 30821, Georgia). FIGURE 71.-ParmeZia maura Persoon [=P. reticdata Taylor] 
(PUtt s.n., Jamaica). 
NUMBER 10 53 
FIGURE i2.-Parmelia fluorescens Hale (Hale 28637, Sabah, FIGURE 73.-Parmelia malesiana Hale (Hale 25178, Philip- 
holotype). pines). 
54 SMITHSONIAN CONTRIBUTIOXS TO BOTANY 
FIGURE 74.-Parmelia planiuscula Kurokawa (Neewoort 427, FIGURE 15.-Parmelia ramosissima Kurokawa (Buwaldn 4583, 
Java, holotype). Thallus infested by a fungus. Java, holotype). 
NUMBER 10 
55 
FIGURE 76.-Parmelia subabstrusa Gyelnik (Hale 25141, FIGURE 77.-ParmeEia confoederata Culberson (Culberson 
Philippines). 9010, North Carolina, isotype). 
56 
SMMITHSONIAN COiYTRIBUTIONS TO BOTANY 
FIGURE 78.-Parmelia coronata Fee (Hale 20505, Mexico). FIGURE 'iS.-Parmelia pigmentacea Hale (Hale 26895, Philip. 
pines, holotype). 
NUMBER 10 
57 
FIGURE 80.-ParneZia subinflata Hale (Hale 26641, Philip- FIGURE 81 .-Parmelia antillensis Nylander (Hale, 35504, 
pines, holotype). Dominica). 
58 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 82.-Parmelia aurulenta Tuckerman (Thieret 24389. FIGURE 83.-Parmelia dissecta Nylander (Hale 29536, Japan). 
Louisiana). 
NUMBER 10 59 
FIGURE 84.-Parmelia horrescens Taylor (Hale 36970, FIGURE 85.-ParmeZia jamesii Hale (James 21 18, New Zealand, 
Tennessee). holotype). 
60 
FIGURE 86.-Parm'eZia quercina (Willdenow) Vainio (Tavares 
1082, Portugal). 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 87.-Parmelia scortea Acharius 
(Vea 762, Yugoslavia). 
NUMBER 10 61 
FIGURE 88.-ParmeZia wallichiana Taylor (Wallich s.n., India, FIGURE 89.-Parmetia aptata Krempelhuber (Togashi m., 
holotype in FH-Tayl). Nepal). 
62 
FIGURE 90.--ParneZia battitnorensis Foriss and Gyelnik (Fink 
and Fuson 340, Indiana). 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 91.-Parmelia caperata (L.) Acharius 
(Hale, 22229, Virginia). 
NUMBER 10 63 
FIGURE 92.-Parmelia congruens Acharius (Kelley 5713, FICURF. 93.-Parmelia crozalsiana Bouly de Lesdain (Sbarbaro 
Florida). 96, Italy). 
64 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 
FIGURE 94.-Parmelia cryptochlorophaea Hale (Allard 15715a, FIGURE 95.-Parmelia malaccensis Nylander (Maingay 21, 
Dominican Republic, holoype). Malaya, holotype in H). 
NUMBER 10 65 
FIGURE 96.-ParmaZia rutidota Hooker and Taylor (Weber FIGURE 97.-Parrnelia salacinifera Hale (Rapp s.n., Florida, 
268 (Lichenes Exsiccati), Australia). holotype). 
66 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 98.-Purmelia texanu Tuckerman (Wright, Texas, FIr.uRE W-Parmelia rianderbylii Zahlbruckner (van der By! 
isotype). s.n., South Africa, holotype in W). 
NUMBER 10 
FIGURE 100.-Parmelia addita Hale (Hale 28342, Sabah, 
holotype). 
67 
FIGURE 101.-Parmelia adjuncta Hale (Kurokawa 50110, 
Japan). 
68 
FIGURE 102.--PurmeZiu brasilianu Nylander (Eiten 2031, 
Brazil). 
SMITHSONIAN COKTRIBUTIONS TO BOTANY 
FIGURE 103.--Purmeliu caruccensis Taylor (Poelt 7512, 
Venezuela). 
NUMBER 10 
69 
FIGURE 104.-Parmelia ckicitae Hale (Culberson 13210, Costa FIGURE 105.-Parmelia croceopustulata Kurokawa (Imshaug 
Rim, holotype). 22275, North Carolina). 
70 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 106.-Parmelia dentella Hale and Kurokawa (Hale FIGURE 107.-Parmelia endochlora Leighton (Hale 31300, 
34176, Alabama, topotype). Hawaii). 
NUMBER 10 
FIGURE 108.--Parmelia erythrocardia Zahlbruckner (Damazio 
1761, Brazil, holotype in W). 
71 
FIGURE 109.--Parmelia exsecta Taylor (Hale 26103, 
Philippines). 
72 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 110.-Parmelia fcssicarpa Kurokawa (Almborn 8374, FIGURE 11 1.-Parmelia formosana Zahlbruckner (Hale 17503, 
South Africa, holotype). Florida). 
NUMBER 10 
73 
FIGURE 112.-Parmelia gigas Kurokawa (Killip 18394, FIGURE 113.-Parmelia imbricatula Zahlbruckner (Kurokawa 
Colombia). 2181, Java). 
74 
ShlITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 114.-Parmelia Zaeuigata (Smith) Acharius (Yoshinura 
650533, Mexico). 
FIGURE 115.-Parmelia livida Taylor (Hale 15995, Virginia). 
NUMBER 10 
75 
FIGURE 116.-ParmeZia neodissecta Hale (Santesson 10597d, FIGURE 117.--Parmelia orientalis Hale (Togashi sn., Nepal). 
Guinea, holotype). 
76 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 1 IS.-ParmeEia physcioides Nylander (King 472, FIGURE 119.-Parmelia pustulifera Hale (HaEe 30865, Georgia, 
Colombia). holot ype) . 
NUMBER 10 
77 
FIGURE 120.-Parmelia scytodes Kurokawa (Togashi s.n., FIGURE 121 .-Parmelia sinuosa (Smith) Acharius (Steiner 283 
(Lichenes Selecti Exsiccati) , Austria) . India, isotype). 
78 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 122.-Parmelia americana (Meyen and Flotow) Vainio FIGURE 123.-Parmelia neocirrhata Hale (Aredne 3726, 
(Mahu 2130, Chile). Mexico, holotype). 
NUMBER 10 
79 
FIGURE 124.-Parmelia pseudonepaknsis Hale (Pringle 10713, FIGURE 125.-Parmelia vexans Zahlbruckner (Asahina s.n., 
Mexico, holotype). Taiwan, holotype in TNS). 
80 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 126.-Parmelia cristifera Taylor (Hale 35800, FIGURE 127.-ParmeZia haitiensis Hale (Orcutt 2987, Jamaica, 
Dominica). holotype). 
NUMBER 10 81 
FIGURE 128.-Parmelia hypoleucina Steiner (Rondon 963 FIGURE 129.-Parmelia hypotropa Nylander (Nakanishi 72, 
(Lichenes Selecti Exsiccati), France). Tennessee). 
82 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 130.-Parmelia louisianae Hale (Hale 34013, FIGURE 131.-ParmeZia perforata (Jacquin) Acharius (Reesc 
Louisiana, holotype). 6407, Louisiana). 
NUMBER 10 
a3 
FIGURE 132.-Parmelia rigida Lynge (Moore 2456, Florida) FIGURE 133.-Parmelia rigida Lynge (Malme 827B, Brazil, 
holotype in S). 
84 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 134.-Parmelia tinctorum Nylander (Degelius P-376, FICURF 135.-Parmelia xanthina (Muiller-Argau) Vainio (Hale 
Fiji). 20594, Mexico). 
NUMBER 10 
85 
FIGURE 136.-Parmelia flaventior Stirton (Fotlmann 14875-H, FICURF 137.-Parmelia meiophora Nylander (Delaway s.n., 
Chile). China, isotype in US). 
86 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 138.-Parmelia praesignis Nylander (Bourgeau s.n., FICURF 139.-Partnelia sphaerosporella Muiller-Argau (Zmshaug 
Mexico, holotype in H). 379, Washington). 
NUMBER 10 
87 
FIGURE 140.--Pnrm,elia thornsonii (Stirton) Culberson (Sarma 
s.n., India). 
FIGURE 141.-Parmelia ulophytlodes (Vainio) Savicz (Martian- 
00, sen., U.S.S.R., holotype in TUR). 
88 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 142.-Parmeliopsis placorodia (Acharius) Nylander FIGURE 143.-Platism'c~tia glauca (L.) Culberson and Culber- 
(Hale 13051, Virginia). son (Hale 33265, Sweden). 
NUMBER 10 
89 
FIGURE 144.--Platismatia stenophylla (Tuckerman) Culber- FIGURE 145.--Platismatia tuckermanii (Oakes) Culberson and 
son and Culberson (Shushan 14737, California). Culberson (Hale 12634, Virginia). 
90 SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FIGURE 146.-Pseudevernia cladonia (Tuckerman) Hale and FIGURE 147.-Pseudevernia consocians (Vainio) Hale and 
Culberson (Merrill 73, Vermont). Culberson (Hale 14294, \Vest Virginia). 
NUMBER 10 
91 
FIGURE 148.--Pseudevemia furfuracea (L.) Hale and Culber- FIGURE 149.-Pseudevernia olivetorina (Zopfj Hale and Cul- 
son (Steiner 4135 in Kryptogamae Vindobonensis, Austria). berson (Vrang 6075, Sweden). 
92 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 
FSCURE 150.--Pseudevernia intensa (Nylander) Hale and 
Culberson (Wetmore 15182, New Mexico). 
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