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 Parmeliaceae Viewed with the Scanning-electron Microscope. Smithsonian Contributions 

to Botany, number 10, 92 pages, 150 figures, 1971.-The scanning-electron microscope 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: SmithQKl.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 Ahmadjian, 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) published 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 recently, Tibell (1971) has shown the value of the 

SEM in studying ascospore structure in the Cypheliaceae. These first efforts have demonstrated convincingly 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 detail the cortical surface of representative species in 

the Parmeliaceae and to ascertain the value of fine 

structure in the taxonomy of the family. Interpretation 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 conjunction 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 colleagues 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 photographs except for this one (Figure 14) were reproduced 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 Laboratory with a Cambridge Stereoscan Mark 2A. Voltage 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 photographs 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 structure of each species. I did, in fact, use types whenever possible, but most specimens were recent collections 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 particles (Figure I) that surface details could sometimes not be recognized. These artifacts are apparently caused by poor preservation. 

Preparation of the samples is very simple compared 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 entire 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 shriveled 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 specimen 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 margin, 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 Hawksworth, 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 standard 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 enlargements, 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. Vertical 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 following layers were easily identified (see Figures 3,4). 

LOWER CORTEx.-This is a discrete layer consisting 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 arranged 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 difficult 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 cortex 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 differentiation. This is the layer viewed with the SEM and will 

be discussed in more detail below. 

The Epicortex 

The various species in the Parmeliaceae examined 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 comparable in certain respects to Peveling’s “Kittsubstanz” (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 confluens), 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 completely 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 upper surface is covered by the thin epicortical membrane (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 fuzziness. 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 developing 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 below (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 croceopustulata 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 electron 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 encountered 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 epicortex 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 magnification. 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 disorganized 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 structure. 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 conglutinated hyphae (Figure 69). The fine structural 

pores described here, rarely more than 25p in 

diameter, are formed by perforations in the epicortex which do not disturb the cortical hyphae. 

The epicortex acts as a site for deposition of 

lichen substances. This layer contains electronlight 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 ultimately 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 examined 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 information 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. Hawksworth (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, photographing 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 taxonomically well-known group of at least 11 genera, one of 

which (Parmelia) is divided into 13 major subgenera 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 pseudocyphellae (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 epicortex is pored (Figure 85) and this species can 

therefore be tentatively placed in section Zmbricuria since species in section Parmelia are nonpored. 

A final example is the common Parmelia ulophyllodes (Vainio) Savicz, which appears to belong to 

section Cyclocheila but which, except for lack of 

distinct pseudocyphellae, also falls near P. flaventior 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 Zahlbruckner 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 judgment in this case. 

The essential features of each genus are discussed 

briefly below. Further elaboration on the classification 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 Parmeliaceae. 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 nonpored 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) Nylander 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. olivetorum. 

This genus includes broad-lobed pseudocyphellate parmelioid species formerly classified in either 

Parmelia or Cetraria (Culberson and Culberson, 

1968b). As one would expect, the epicortex is nonpored 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 epicortex 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 Parmelia, 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. enterornorpha, 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. disjuncta, 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 unidentified 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 nonpored 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. cumberlandia, 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. taractica (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, pseudocyphellae or pseudocyphellae-like white reticulation, 

and simple chemistry (salacinic or rarely protocetraric acid). Six species and eight specimens 

studied consistently had a nonpored epicortex and 

nodular surface. The cortex is virtually indistinguishable 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 narrowlobed subgenus Parmelia (especially section Im bricaiia) 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, although 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 developed (Figure 78). 

Section Zmbricaria (Schreber) Fries 

FIGURES 81-88 

SPECIES ExAMINEu.-Parmelia antillensis, P. aurulenta, P. dissecta, P. horrescens, P. jamesii, P. quercina, P. scortea, and P. wallichiana. 

There is a great range of morphological variation 

in this rather heterogeneous assemblage of marginally 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 (Figure 88) have not been identified. 

Section Cyclocheila (Vainio) Rasanen 

FIGURFS 89-99 

SPECIFS ExAntrivEu.-Parmelia aptata, P. baltimorensis, 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 configuration and chemistry, P. aptata (Figure 89), P. 

cryptochlorophaea (Figure 94), and P. texana (Fig 

ure 98), also tend to have a strongly nodular surface, 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. croceopustulata, P. dentella, P. endochlora, P. erythrocardia, P. exsecta, P. fissicarpa, P. formosana, P. 

gigas, P. imbricatula, P. laevigata, P. livida, P. 

neodissecta, P. orientalis, P. physcioides, P. pustulifera, 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. neocirrhata, 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. haitiensis, 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 morphological 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 nonepicorticate as in Pseudevernia. It is of interest that its 

presumptive chemical counterpart, P. rigida (Figure 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. Parmelia Raventior, for example, seems amphigymnioid 

but differs sharply from species in subgenus Amphigymnia 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 characteristics of a species in section Cyclocheila, but it has 

a nonpored nodular epicortex (Figure 139). Finally, 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 photographs 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. stenophylla, and P. tuckermanii. 

This small segregate of Cetraria includes the 

widespread boreal P. glauca. The epicortical structure 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. intensa. 

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 conglutination. 

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 enlargement 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 (Parmeliaceae). 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. Bryologist, 71:l-11. 

Parmelia pustulifera, a New Lichen from Southeastern United States. Brittonia, 24:22-27. 

Hale, M. E. 

1965. 

1968. 

1972. 

Hale, M. E., and S. Kurokawa. 

1964. Studies on Parmelia Subgenus Parmelia. Contributions 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 Survey. 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 Untersuchungen mit Raster-Elektronenmikroskop im 

Vergleich mit herkommlichen Untersuchungsmethoden. Beitrag Elektronenrnikroskopiche Direktabbildung 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 different 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 rectangular 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. croceopustulota Kuroknu~n (Hale 33269, North Carolina) n, b, andd, about ~500; cabout ~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 approximately 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 nonpored 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 Kurokawa (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 crystalencrusted hyphae (about x2OOO). 

FIGURE 16.-Stereo view of the surface 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 epicorticate species: a, smooth surface of P. imbricatula Zahlbruckner (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 Kurokawa (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 Culberson (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 (BournNew 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, Conecti10099, Ohio). cut), .4 large pseudocyphella is included in the photographs. 
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, Philipholotype). 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 

India, isotype). (Lichenes Selecti Exsiccati) , Austria) . 
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 (Martian00, 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 Culson (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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