ShIITHSOSI.4N CONTRIBLTIONS TO ROTAXI’ 0 XUAIBER 31 

A Monograph of the Lichen Genus 

Pseudoparmelia Lynge (Parmeliaceae) 

Mason E. Hale, Jr. 

S5f ITHSOSIAS IKSTITUTIOK PRESS 

City of IVashington 

1976 
ABSTRACT 

Hale, Mason E., Jr. A Alonograpli of the Lichen Genus Pse~idopai~melia Lynge 

(Parmeliaceae). Smifhsonian Contl-ibzitions to Botany, number 31, 62 pages, 18 

figures, 197(i.--A ~vorld monograph ot“ the genus Pseiidopai-melia is presented. 

There are 76 species, most occurring in dry subtemperate to tropical areas and 

especially concentrated in the arid scrub lands of South Africa, Australia, and 

Brazil. The genus is characterized 13) an unusually high number of species with 

di1,aricatic acid and related orcinol depsitles. The following new species are 

described: P. callicliroa KnrokaMa, P. concornitnns Hale, P. conlabiosa Hale, 

P. dahlii Hale, P. ncoquintal-ia Hale, P. siibarnbigzia Hale, P. .subamplexa Hale, 

and P. ixnczolnnu Hale. The following new combinations are made: P. L‘OV 

rugati-im (Kurokawa and Filson) Hale, P. cxomnta (Zahlbruckner) Hale, P. 

geilachci (Zahlbruckner) Hale, P. papillo.sa (Lynge ex Gyelnik) Hale, P. 

Totlrigiiesiana (Hue) H:ile, P. scliistncca (Kurokawa arid Filson) Hale, P. scotopliylla (Kurokawa) Hale, and P. spodochroa (Kui-okaxva and Filson) Hale. 

OFFICIAL l’IBL.ICATIO\ 1)Ai I: is hantlstampetl in a lilnitetl tiutiihcr of initi;tl copies and is recorded 

in rhc Institution’s annual report. Snzitiisoizicin l’cnr. SLKIFS co\ ER DESIG~: Leaf clearing from thc 

Katsnra trce Cei.cidiphylluin jnfmnicuin Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Hale, hlason E. 

.4 monograph of the lichen genus Pseudopar??~clk~ Llnge 

(Parmeliaceae) 

(Smithsonian Contributions to botany ; no. 31) 

15ibliography: p. 

Includes index 

Supt. of Docs. no.: SI 1.29:31 

I. Pseudoparmclia. I. Title. 11. Series: Sn1itl1sonian Institution. Smithsonian contributions to botzGi) , no. 31. 

QKlS2747 no. 31 [Qh585.P2] 381’.08s [389’.1] 73-619375 
Contents 

Page Introduction 1 ........................... .................... 

Morphology 2 

Chemistry 5 

Ecology and Habitats ........................................... 

Formation of Morphs and Chemical Populations ................ 

Phytogeography 8 

Taxonomic Treatment ...................................... 10 

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

................................................. 

....................................................... 

6 

7 .............................................. 

56 

Literature Cited ................................................ 58 

Index ........................................................... 61 

iii 

A Monograph of the Lichen Genus 

Pseudoparmelia Lynge (Parmeliaceae) 

Mason E. 

Introduction 

This world monograph of the lichen genus 

P,srutlopnrnielia is part of my continuing effort to 

rvvise the parmelioid genera. Although these genera 

are usually lumped under the collective name Pariii cliri, recent studies with the ’scanning-electron 

microscope have demonstrated that species previously classified under Parmelia fall into two distinct 

gi~oups, one with a paraplectenchymatous upper 

cortex antl very frequently with pseudocyphellae, 

t! pifietl by Pnrmelia saxatilis (L.) Acharius and including the P. boi-reri group, and another with a 

palisade plectenchymatous upper cortex and an 

ot.erlying pored epicortex hut never with pseudocyphellae (Hale, 1973a). 

The epicorticate group includes several heterogeneous elements differing, for example, in marginal 

Ornamentation (cilia, bulbate cilia), rhizine branching patterns (simple or dichotomously branched), 

arid lobe configuration and width (broad apically 

rotund to narrow obtuse lobes). These elements are, 

in my opinion, biologically isolated and represent 

good genera, as follows: Bzilbothrix Hale with bulbate cilia antl no cortical pigments (Hale, 1974), 

Hypotmcliynn (Vainio) Hale with narrow eciliate 

lobes antl dichotomously branched rhizines (Hale, 

1 Wja), Pnrmelinn Hale with narrow ciliate lobes 

(Hale, 1954), Paimotrerna Massalongo with broad 

rotund lobes (Hale, 1965a), Pseudopal-melia Lynge 

Ivith nai-row eciliate lobes, Relicina (Hale and Kurokatua) Hale with bulbate cilia and usnic acid in 

,ifason E, Hale, Jr,, Depart11ient of Botany, lYationnl dlust’uin of ‘Vl’ntuml History, Sftiit/isoniaii Institution, W’ashington, D.C. 20560 

Hale 3r. 

the cortex (Hale, 1975b), and Xnnlhoparmckz 

(Vainio) Hale with narrow eciliate lobes and usnic 

acid in the cortex (Hale, 1974). 

As with Hypotmchyna (Hale, 1975a) and Relicina 

(Hale, 1955b), much of the initial work on the 

chemistry of Pseiidoparmelin was completed before 

thin-layer chromatography came into use in 1965. 

Thus it has been necessary to reinvestigate the 

chemistry of many specimens. Even with the application of the most recent techniques, however, this 

monograph can be considered definitive only in the 

broadest sense. Fresh lichen collections are now being made every year in exotic regions, especially in 

Africa, Australia, and South America, by botanists 

and more significantly by professional lichenologists, 

who had not previously collected outside temperate 

areas and who choose to identify the specimens 

themselves rather than send them to a monographer. 

Under these circumstances, any monograph will be 

incomplete in two respects. For one, information on 

new records not available for study will be omitted, 

records that could profoundly affect interpretations 

of phytogeography. The older specimens now preserved in herbaria are a pitifully inadequate representation of lichen floras. Secondly, unpublished 

new species which are sure to be found in a genus as 

large as Pseiidoparmelia obviously cannot be included in the keys, and this lack makes species 

identification more difficult for lichen students. This 

monograph is offered, then, with full realization of 

its limitations but with the hope that it will provide a base for later workers. 

ACKNOWLEDGMENTS.-~ am much indebted to 

curators who sent specimens on loan and often 

allowed ’considerable extensions of loan periods. 

1 
2 SMITHSONIAK CONTRIBUTIONS TO BOTANY 

These include Mr. Peter James (BM), Dr. A. J. 

Kostermans (BO), Dr. A. Robyns (BR), Mr. M. S. 

Christiansen (C), Dr. M'. A. Weber (COLO), Dr. W. 

L. Culberson (DUKE), Dr. L. Williams (F), Dr. I. 

M. Lamb (FH), Dr. C. E. Bonner (G), Mr. C. E. 

Palmar (GLAM), Dr. T. Ahti (H), Dr. H. Schindler 

(KR), Dr. J. Poelt (M), Dr. H. A. Imshaug (MSC), 

Dr. H. Osorio (MVM), Dr. C. T. Rogerson (NY), Dr. 

Hildur Krog (0), Dr. E. F. Warburg (OXF), Mme. 

Jovet-Ast (PC), Dr. C. E. Smith (PH), Dr. S. Ahlner 

(S), Dr. S. Kurokawa (TNS), Dr. R. Alava (TUR), 

Dr. I. Tavares (UC), Dr. R. Santesson (UPS), Dr. H. 

Riedl (W), Dr. J. W. Thomson (WIS), and Dr. G. 

Cufodontis (TNU). Dr. Ove Almborn (LD) lent his 

own extensive collections in addition to materials 

from BOL, L, PRE, and TRH which he had on 

hand for study. Dr. Z. Cernohorskj kindly arranged 

for loans from BP at a time when travel in Hungary 

was not permitted. The late Dr. H. des Abbayes 

(REN) was extremely generous in sending loans as 

well as duplicates of type-collections of the new 

species he was describing. For allowing me to examine collections in their private herbaria, I am 

especially indebted to Dr. D. D. Awasthi, Dr. Gunnar Degelius, Dr. M. Mahu, Dr. T. D. V. SB' 'inscow, 

and Dr. Rolf Santesson. 

The specimens were photographed by the Smithsonian Office of Printing and Photographic Services. Scanning-electron microscope photographs 

were prepared by the Smithsonian Scanning-Electron Microscope Laboratory. 

Dr. S. Kurokawa assisted with species identifications, chemical tests, and descriptions prior to 1961, 

and his help is gratefully acknowledged. Field 

studies have been supported at various times by the 

National Science Foundation, National Geographic 

Society, Smithsonian Research Foundation, and the 

Morden-Smithsonian Fund. Cooperative field 

studies were conducted in India with the Maharashtra Association for the Cultivation of Science, working with Dr. P. G. Patwardhan, and in Venezuela 

with Dr. M. Lopez-Figueiras of the Universidad de 

10s Andes. Finally, I wish to thank Dr. Ove Almborn for reading the entire manuscript and checking the bibliography and spelling of place names. 

Morphology 

THALLw-The thallus is typically foliose, orbicular, 3-15 cm in diameter, and uniformly adnate, 

even at the margin. The degree of adnation varies 

from nearly subcrustose in P. xanthomelaena (Figure 18e) to loosely adnate in P. caperata. Comparable eciliate species of Parmotrema have wider, 

more or less marginally erect lobes. Lobe configuration is variable. While some species have sublinear, 

apically obtuse lobes 0.5-2.0 mm wide (Figures 6b,d, 

10e, 12d), an even larger number have subirregular, 

apically rotund lobes 2-6 mm wide (Figures 6a,c, 

9a, IOc, 12c). Lobes in both types are often contiguous and crowded. There is no marginal ornamentation in Pseudoparmelia; the margins are smooth and 

sometimes black rimmed. 

The lower surface of the thallus is either black 

or pale brown, and this color difference, although 

variable in some of the saxicolous species, can be 

an important taxonomic character. Broader lobed 

species with a black lower surface, such as the wellknown P. caperata, usually have a narrow brown 

zone at the lobe tips. While most species have a 

shiny paraplectenchymatous lower surface (Figure 

Ic), species in the P. amplexa group (P. amplexa, P. 

pachydactyla, and P. subamplexa) have a minutely 

grained, velvety appearing black lower surface to 

the margin (Figure la,b). This is caused by an irregularly thickened, minutely papillose cell layer 

sloughing from the lower cortex, a phenomenon 

which I have not seen in any other parmelioid 

genera. 

Rhizines are uniformly simple and produced in 

moderate abundance (Figure Id). The only exception to this rule is P. intertexta, which usually has 

pale branched rhizines, not unlike those of some 

species of Relicina but not as agglutinated (Hale, 

19758). Broad-lobed species usually lack rhizines 

along the margins at lobe tips or at most have a 

zone of short papillae. The rhizines are often blunt 

with pale, penicillate tips (Reznik et al., 1968). 

The internal structure of Pseudoparmelia presents no unusual features (Figure Za,c,f). The upper 

cortex has a basic palisade structure (Figure 2d) and 

is overlain by a thin pored epicortex (Figure le,f), 

as I had previously discovered by using the 

scanning-electron microscope (Hale, 1973a). The 

medullary hyphae are often conspicuously encrusted 

with lichen substances. The lower cortex is paraplectenchymatous (Figure 2e). In these respects it 

is identical with the other epicorticate genera in the 

family. 

VEGETATIVE PRoPAGuLES.-The isidia in Pseudo- 
NUMBER 31 3 

FIGURE 1.-Morphology of Pseudoparrnelia: a, lower surface of P. arzplexa (Degelius s. n.) 

(X 160); b, enlargement of surface in a (X 1600) with arrow pointing to a papilla; c, lower 

surface of P. rutidota (Degelius A-37) ( x 160); d, cross section of P. rnalaccensis to show simple 

rhizines (Hale 30353) ( x 140); e, surface of P. rahengensts showing pored epicortex (Kurokaum 

1602) (X 400); f, enlargement of surface in e showing pores and aggregations of crystals of usnic 

acid (x 1600). (All photographs taken with the scanning-electron microscope.) 
4 SMITHSONIAS COSTRIBUTIOSS TO BOTANY 

FIGURE 2.-Morphology of Pseudoparmelia: a, longitudinal cross section of P. uenezolana with 

arrow in upper right pointing to the epicortes (Hale 43326a) (x 390); 6, enlargement of epicortex in a (upper arrow) and upper cells of the cortex (lower arrow) (x 4000); c, longitudinal 

cross section of P. spodochroa (Weber 47220) (x 370); d, enlargement of upper cortex in c 

(x 800); e, longitudinal cross section of P. sphaerospora (Culberson 10324) (X 240); f, enlargement of lower cortex in e (x 800); g, isidia of P. dnlilii (isotype in US) (x 160). 
SUXIBER 31 5 

pavmelia are quite uniform. Twenty species (P. 

adspersa, P. amazonica, P. annexa, P. arcana, P. 

basutoensis, P. caroliniana, P. cinerascens, P. concrescens, P. conlabiosa, P. cyphellata, P. dahlii (Figure Zg), P. ecapernta, P. ischnoides, P. malaccensis, 

P. martinicana, P. neoquintaria, P. salacinifera, P. 

scoiophylln, P. siibtovtzila, and P. venezolann) have 

normal, cylindrical, simple to sparsely branched 

isidia. Two additional species (P. pachydactyla and 

P. pn@illosa) have abnormally thickened but otherwise normal isidia (Figure 14d). No species have 

lobulate, procumbent, or ciliate isidia. 

Pustules have been described and illustrated for 

Hypotrachynci (Hale, 1975a), and they are more or 

less the same in Pseudoparmelia (Figure 7b), very 

thickened, fragile, hollow isidioid structures which 

usually break open apically. Some then produce 

soredia, while in other cases no soredia form. Seven 

species (P. baltimorensis, P. eruptens, P. geesterani, 

P. owariensis, P. pustulescens, P. schistacea, and P. 

zim babzuensis) have largely nonsorediate pustules. 

One species (P. rnunkiaeri) has heavily sorediate 

pustules. 

Ssralia occur in 14 species: P. alabamensis, P. 

nptata, P. caperata, P. carneopruinata, P. crozalsiana, P. cryptochlorophaea, P. epilezica, P. gerlachei, 

P. labrosa, P. leucoxantha, P. soredians, P. subambigiin, P. subamplexa, and P. texana. They are usuall) orbicular and discrete except in P. capevata, 

which has coalescent, more diffuse soralia. All are 

strictly laminal except for P. leucoxantha (Figure 

13b), where the soralia are largely submarginal. It is 

perhaps noteworthy that all of the sorediate species, 

excepting P. alnbnmensis, are corticolous; sorediate, 

obliqately saxicolous species are very rare in the 

genus. 

The remaining 32 species lack any vegetative 

propagules. The older lobes, however, may become 

lobulate or heavily rugose, although, as pointed out 

above, no species have true laminal lobulae (phyllidia) or lobulate isidia. Virtually all of these species 

are fertile. 

APoTHECIA.-Apothecia, found so far in 47 of 

the 76 species, are extremely uniform. They are 

sessile to subpedicellate and 1-3 mm (rarely to 6 

nim) in diameter. The disc is always imperforate. 

The asci contain eight spores, and the bulk of the 

species have spores in the range of 6-10 pm wide 

and 7-13 Ipm long. Psezidoparmelia caperata and 

P. concomitaizs have the largest spores, up to 24 pm 

long. 

Chemistry 

The chemical constitutents of Pseudoparmelia 

were first summarized by Hale and Kurokawa (1964) 

on the basis of microcrystal tests. I have retested 

most of the species with thin-layer chromatography, 

using solvent sjstems A and B of C. Culberson 

(1972) and in large part confirming the earlier results. The lichen substances in the genus are listed 

below under the presently accepted classification 

with number of species containing each substance 

in parentheses. 

A\LIPHATIC ACIDS 

Caperatic acid (5) 

Protolichesterinic acid (3) 

Others (2) 

AROMATIC CO\lPOUNDS 

Orcinol Series 

para-Depsides 

DiLaricatic acid (9) 

Erernic acid (1) 

Gyrophoric acid (4) 

Lecanoric acid (4) 

Obtusatic acid (1) 

Perlatolic acid (2) 

Cr) tochlorophaeic acid (1) 

Sekikaic acid (1) 

Neoloxodic acid (2) 

Soilobaridone (3) 

ineta-Depsides 

Depsidones 

p-Orcinol Series 

para-Depsides 

Atranorin (55) 

Barbatic acid (1) 

4-0-Demethylbarbatic acid (1) 

Constictic acid (5) 

Fumarprotocetraric acid (1) 

Norstictic acid (2) 

Physodalic acid (2) 

Protocetraric acid (25) 

Salazinic acid (5) 

Stictic acid (8) 

Succinprotocetraric acid (1) 

Depsidones 

DIUESZOFURAXES 

Usnis acid (20) 
6 S\IITHSOSI.AS COSTRIBUTIOSS TO BOTASY 

ASTIlR iQLIXOSES 

Skyrin (2) 

Untletermined (3) 

Unidentified substances presumed to be depsidones are known in P. chaly beizans (“chalybeizans” 

unknown), P. chapadensis, P. exornata and P. papillosa (“conformata” unknown), and P. neoqiiintaria 

(“quintaria” unknowns). Unidentified depsides or 

orcinol depsidones are probably present in P. vanderbylii. unidentified yellow pigments have been 

reported for P. arcana, P. baltirnorensis, P. condyloirles, P. cor?xgativa, P. cyphellata, P. riitidota, and 

P. .rphaerospora, but none of these are accompanied 

by triterpenoids. Details on these will be found in 

“Taxonomic Treatment.” 

Divaricatic acid, perlatolic acid, and sekikaic acid 

are all accompanied by 1 to 3 weaker, as yet unidentified spots falling below the main acid. Perlatolic 

acid may occur mixed with (or even be replaced by) 

stenosporic acid, but these two substances are not 

resolved in the solvent systems employed here. 

Gyrophoric acid always occurs as an accessory with 

protocetraric acid, never alone, and norstictic is accompanied by salazinic acid or stictic acid. 

TYhile I hope to discuss the chemical features of 

Pseudoparrnelia in relation to the other parmelioid 

genera in a separate article, several chemical traits 

stand out. The abundance of orcinol series depsides, 

especially divaricatic acid, is noteworthy and in fact 

characterizes the genus. The absence of the commoner orcinol series depsidones such as alectoronic 

acid is significant, as is also the absence of echinocarpic acid, galbinic acid, lichexanthone, and triterpenoids, and the rarity of barbatic acid and 

norstictic acid. 

Ecology and Habitats 

Knowledge of the ecology of any lichen group can 

enhance our understanding of difficult species complexes, and it is best if this knowledge is gained 

through firsthand experience in the field. Herbarium labels often omit important data on general 

vegetation, substratum, and elevation. IVhile I have 

collected specimens of Pseudoparnzelia in various 

parts of the world, I have never visited Africa or 

Australia, two very important centers for the genus. 

It is still possible, however, to draw a general picture of the ecological requirements of the genus. 

It is evident, for example, that Pseudoparrnelia is 

best developed in temperate, subtropical, and arid 

tropical regions at low to mid elevations. This is in 

contrast to the strong boreal requirements of Parrnelia, and the highland tropical preferences of 

Hypofmchyna (Hale, 1975aj. Only two species, P. 

baltiniorensis and P. caperata, grow as far north as 

the southern edge of the boreal forest, although in 

the Southern Hemisphere P. gerlachei occurs in 

subantarctic localities. At the same time only four 

species occur with any frequency in humid lowland 

tropical forests: P. sphaerospora in the Americas 

and Africa and P. dahlii, P. intertexta, and P. malaccensis in Southeast Asia. 

Pseudopamzelia occurs most abundantly in the 

arid desert or sclerophyll scrublands of Africa and 

Australia. In these two regions alone nearly 30 

obligately saxicolous species are known and most 

of them are endemic. The approximately 40 obligately corticolous species in the genus occur 

mostly in open deciduous secondary forest or in 

disturbed sites such as pastures, parks, and botanical 

gardens. About 8 species occur on either rocks or 

trees, the most typical being P. texana. These are 

generally habitats of great environmental stress with 

intense insolation and prolonged drought. 

Psezidoparinelia competes with two other parmelioid genera: Parmotrema, which is mainly corticolous, and Xanthoparmelia, a saxicolous group. In 

South Africa Parmelia subgenus Melanoparmelia 

is also well devolped on rocks, and as a result Psezidoparmelia, Xanthoparmelia, and the Melanoparrneliae often have similar lobe configuration and 

close thallus adnation. They all have simple rhizines, lack marginal cilia, and rarely produce soredia. It may represent an example of convergent 

evolution for three genera in these stressed environments. 

Another possible example of. convergent evolution is represented by the three usnic-acid containing species in the humid lowland forests, P. dahlii, 

P. intertexta, and P. malaccensis. They occupy the 

same habitats (in particular canopy branches in 

dipterocarp forests) and have the same narrow, appressed, sublinear lobes and yellow pigmentation 

(usnic acid) as Relicina (Hale, 197513) but lack the 

marginal bulbate cilia. 
NUMBER 31 

Formation of Morphs and Chemical Populations 

7 

Parent (16) (lacking) 

A detailed discussion of morph formation and 

speciation in Hypotrachyna (Hale, 1975a), which 

applies equally well to Pseudoparmelia, has already 

been presented. Briefly, one assumes that nonfertile 

isidiate and sorediate morphs have evolved from 

fertile nonisidiate or nonsorediate parent species 

and that lacking any means of sexual reproduction 

the morphs tend to conserve chemical traits. A parent. morph may be traced rather easily, may already 

be modified through the usual genetic changes, or 

may have become extinct and now be represented 

only by the vegetative morph. While application 

of the morph concept may be somewhat arbitrary in 

actual practice, it has offered a reasonable explanation of species evolution in several foliose genera. 

There are seven theoretically possible combinations of parents and vegetative morphs if we consider only soredia and isidia (Figure 3). Pustules, 

which in any event are not common, would be 

considered as an offshoot of isidia. In Pseudoparmelia the largest category is the 16 fertile species 

which, as far as we can determine from herbarium 

collections now available, have not produced any 

vegetative morphs. Some will undoubtedly be found 

eventually, but the presumption is that these species (P. callichroa, P. caribaea, P. chapadensis, P. 

concomitans, P. condyloides, P. corrugativa, P. 

Iry$)ornilta, P. inornata, P. lecanoracea, P. rodriguesiana, P. rupicola, P. rutidota, P. somaliensis, P. subtiliacea, P. vanderbylii, and P. violacea), most rather 

rare, have not yet evolved successful vegetative 

morphs. 

Referring again to Figure 3, the next largest 

category is 12 isidiate mo.rphs for which no corresponding parents have been discovered. These include P. adspersa, P. amazonica, P. arcana, P. caroliniana, P. cineraseens, P. dahlii, P. geesterani, P. 

neoquintai-ia, P. owariensis, P. rahengensis, P. salacinifera, and P. venezolana. IZ'e might assume in this 

instance that the morphs evolved from a nonisidiate 

parent which has not been able to survive and propagate or which has not yet been discovered. 

Coexisting parents and isidiate morphs (but with 

no sorediate morph) are represented by the following nine species pairs: P. exornata - P. papillosa, 

P. intertexta - P. malaccensis (tentative), P. molybdiza - P. annexa, P. prolata - P. basutoensis, P. 

/\ /\ 

(lacking) (lacking) Isidlate (3) Sorehiate (3 ) 

Parent (9) (lacking) 

Parent (4) (lacking) 

Parent (2 

Isidlate (2) Soredim (2 1 

FIGURE 3.-Patterns of inorph formation in Pseudoparmelia 

with number of species in each categor) in parentheses. 

spodochroa - P. Scotophylla, P. tortula - P. subtortula, P. xanthomelaena - P. ischnoides, and P. 

zam biensis - P. ecapevata. 

Soredia are comparatively rare in Pseudoparmelia. There are four parent species with corresponding sorediate morphs: P. epileuca - P. schelpei, P. 

ferax - P. gerlachei, P. inhaminensis - P. crozalsiana 

(tentative), and P. scrobicularis - P. cavneopruinata. 

Six sorediate species, P. alabamensis, P. aptata, P. 

cryptochlorophaea, P. leucoxantha, P. soredians, 

and P. subambigua, appear to have no nonsorediate 

parent. 

Two groups of species hace the full complement 

of morphs: P. amplexa - P. pachydactyla - P. subamplexa and P. nairobiensis - P. concrescens - P. 

texana (including as well P. eruptens). A third 

series, P. rutidota - P. baltiniorensis (coarsely pustulate) - P. caperata (sorediate), is another possibility 

but the species here all diLerge rather widely from 

each other in some details. 

A final group of four species, P. martinicana - P. 

raunkiaeri and P. conlabrosa - P. labrosa, represent 

isidiate-sorediate morph pairs but have no fertile 

parent extant. 

Of the remaining species in the genus, P. geesterani, P. pustulescens, P. schistacea, and P. zini- 
8 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

babwensis are pustulate and have no obvious 

relatives. 

There is a particularly interesting group of saxicolous species in South Africa, including P. molybdiza (lecanoric acid), P. spodochroa (salazinic acid), 

P. tortula (norlobaridone), and P. vanderbylii (unknown depsides). They are so similar externally 

that chemical tests are needed to separate them. 

IVhile this group may have evolved from a common 

parent, the chemical evolution has followed very 

different pathways. The presumptive isidiate 

morphs of three of these (P. antzexa, P. scotophylla, 

and P. subtortula) are also sufficiently similar that 

chemical tests are required for positive identification. 

If one subscribes to the concept of vegetative 

morphs, it then follows that morphologically identical but chemically different vegetative morphs have 

evolved from different parents through parallel evolution and are therefore polyphyletic (W. Culberson, 

1973). Pseudoparmeliu texana (divaricatic acid) and 

P. uptutu (perlatolic acid), for example, differ in 

distribution, P. texana being pan-temperate whereas 

P. aptatu occurs only in the Old World. The two 

acids involved are para-depsides, divaricatic acid 

having a C,H, side chain and perlatolic acid C5Hll 

side chains (Figure 4). Another chemical pair, P. 

owariensis (divaricatic acid) and P. pustulescens 

(sekikaic acid), is confined to the Old World and 

R 

A ,,--CO - 0 -0-OH 

CH~O-V-OH -COOH 

I 

R 

FIGURE 4.-Comparison of molecular structures of acids in 

Pseudopnrmelia: Divaricatic acid (R = C,H,) and perlatolic 

acid (K= C,H,,) (above) and sekikaic acid (below). 

appears to be sympatric. The acids are particularly 

interesting since sekikaic acid is a meta-depside 

and divaricatic acid the homologous para-depside 

(Figure 4). 

Rarer species with chemical differences used as 

taxonomic characters include P. dahlii (lecanoric 

acid) and P. rahengensis (barbatic acid group) in 

Southeast Asia. Pseiidoparmelia rutidota, a common 

species in Australia and southwestern United States 

into tropical America, contains protocetraric acid; 

a virtually indistinguishable population, P. exornata, contains protocetraric acid with the “conformata” unknown and occurs principally in Uruguay. 

Other close relatives include P. ferax, which contains physodalic acid, and P. callichroa, which contains a fatty acid. 

The presence or absence of usnic acid is the main 

criterion for recognizing P. nairobiensis (usnic acid 

absent in the cortex) and P. zambiensis (usnic acid 

present) and their corresponding isidiate morphs, 

P. concrescens and P. ecaperata. 

Phytogeography 

The Psezidoparmelia floras of various geopolitical 

units are listed below. Many countries are so poorly 

collected, of course, that no meaningful floristic 

comparisons can be made. However, the United 

States, West Indies, Venezuela, South Africa, and 

perhaps Brazil and Australia are all rather well 

known. Several lichenologists are now working in 

East Africa and Australia, and their studies, when 

completed, will fill in many gaps. 

NORTH AMERICA 

Canada: P. baltimorensis, P. caperata. 

United States: P. alabamensis, P. amazonica, P. baltirnorensis, P. caperata, P. caroliniana, P. crozalsiana, P. cryptochlorophaea, P. martinicana, P. rutidota, P. salacinifera, P. 

sphaerospora, P. texana. 

MEXICO AND CENTRAL AMERICA 

Mexico: P. amazonica, P. carneopruinata, P. caroliniana, P. 

crozalsiana, P. leucoxantha, P. martinicana, P. raunkiaeri, 

P. rutidota, P. salacinifera, P. sphaerospora, P. texana. 

Guatemala: P. caperata, P. sphaerospora. 

Honduras: P. amazonica, P. carneopruinata, P. caroliniana, 

Kicaragua: P. caroliniana. 

Costa Rica: P. carneopruinata, P. caroliniana, P. sphaerospora, 

Panama: P. caroliniana, P. sphaerospora, P. texana. 

P. cryptochlorophaea, P. salacinifera, P. texana. 

P. texana. 
NUMBER 31 

WEST ISDIES 

Bahamas: P. sphaerospora. 

Cuba: P. anlazonica, P. caribaea, P. cnroliniana, P. salacinifera, P. sphaerospora. 

Jamaica: P. carneopruinata, P. caroliniana, P. cryptochlorophaea, P. raunkiaeri. 

Hispaniola: P. cnperata, P. caroliniana, P. cryptochlorophaea, 

P. inornata, P. martinicana, P. raunkiaeri, P. sphaerospora, 

P. texana. 

Puerto Rico: P. nmazonica, P. niartinicana. 

Lesser Antilles and Virgin Islands: P. caribaea, P. cryptochlorophaea, P. inornata, P. martinicana, P. raunkiaeri, P. 

sphaerospora. 

Trinidad: P. nninzonica 

SOUTH AMERICA 

Colombia: P. arnazonica, P. carneopruinata, P. salacinifera. 

Venezuela: P. ainazonica, P. caperata, P. carnepruinata, P. 

P. caroliniana, P. cryptochlorophaea, P. martinicana, P. 

.ralacinifera, P. scrobicularis, P. sphaerospora, P. texana, 

P. venezolana. 

Ecuador: P. caroliniana. 

Peru: P. caperata, P. rutidota, P. sphaerospora, P. texana. 

Bolivia: P. rutidota. 

French Guyana: P. caribaea, P. sphaerospora. 

Brazil: P. aniazonica, P. caperata, P. carneopruinata, P. caroliniana, P. chapadensis, P. cinerascens, P. crozalsiana, P. 

cyphellata, P. exornata, P. hypolnilta, P. leucoxantha, P. 

papillosa, P. rupicola, P. rutidota, P. salacinifera, P. scrobicularis, P. sphaerospora, P. texana. 

Uruguay: P. carneopruinata, P. crozalsiana, P. exornata, P. 

papillosa, P. rutidota, P. texana. 

Paraguay: P. cinerascens, P. scrobicularis. 

Argentina: P. caperata, P. carneopruinata, P. crozalsiana, P. 

gerlnchei, P. papillosa, P. rutidota, P. scrobicularis, P. soredians. 

Chile: P. caperata, p. ferax, P. gerlachei, P. labrosa, P. rutidota, P. soredians, P. subnmbigua, P. texana. 

EUROPE AND AFRICA 

Europe: P. caperata, P. caroliniana, P. carneopruinata, P. 

Tunisia: P. cnperata. 

Guinea: P. ecaperata, P. pustulescens, P. rodriguesiana, P. 

Ivory Coast: P. caroliniana, P. ecaperata, P. inalaccensis, P. 

Cameroon: P. sphaerospora. 

Ethiopia: P. nptata. 

Somalia: P. soinalie~zsis. 

Uganda: P. nnnexa, P. arcana, P. caroliniana, P. concrescens, 

P. Jiiolybdiza, P. nairobiensis, P. owariensis, P. pustulescens, 

P. soinaliensis, P. subtortula, P. texana. 

crozalsiana, P. soredians. 

sphaerospora. 

owariensis, P. rodriguesiann, P. texana. 

Urundi: P. somaliensis. 

Kenya: P. caperata, P. epileuca, P. nairobiensis, P. pachydacZaire: P. concresceiis, P. crozalsiana, P. sphaerospora. 

Rhodesia: P. subamplexa, P. tortula. 

Angola: P. ainplexa, P. annexa, P. arcana, P. caroliniana, P. 

tyla, P. soredians, P. subtortula, P. texana. 

concrescens, P. ecaperata, P. inhaminensis, P. nairobiensis, 

P. pustulescens, P. sphaerospora, P. texana. 

Malawi: P. ecaperata. 

Zambia: P. somaliensis, P. zainbiensis. 

Tanzania: P. aptata, P. inolybdiza, P. nairobiensis, P. SOmaliensis, P. sphaerospora, P. texnna. 

hloqanibique: P. concrescens, P. epileuca, P. eruptens, P. 

nialaccensis, P. schelpei. 

Union of South Africa: P. ainplexa, P. annexa, P. aptata, 

P. arcana, P. basutoensis, P. concrescens, P. condyloides, P. 

crozalsiana, P. erufitens, P. geesterani, P. ischnoides, P. 

lecanoracea, P. niolybdiza, p. prolata, P. ptistulescens, 

P. rodriguesiann, P. soredians, P. spodochroa, P. texana, 

P. tortula, P. vanderbylii, P. uiolacea, P. xanthoinelaena. 

Madagascar: P. rodriguesiana, P. sonialiensis, P. sphaerospora, 

P. texana. 

ASIA 

India: P. aptata, P. caperata, P. crozalsiana, P. ecaperata, P. 

Sri Lanka: P. dahlii, P. texana. 

Thailand: P. adspersa, P. ecaperata, P. oujariensis, P. rahenLaos: P. caroliniana 

China: P. caperata. 

Japan: P. aptata, P. caperata, P. owariensis, P. texana. 

Hong Kong: P. oa'ariensis. 

Taiwan: P. aiiinzonica. 

Philippines: P. adspkrsa, P. intertexta, P. inalaccensis. 

Malaysia: P. intertexta, P. nialaccensis. 

Indonesia: P. nptata, P. intertesta, P. inalaccensis, P. texana. 

Sew Guinea: P. intertexta. 

malaccensis, P. pustulescens, P. texana. 

gensis, P. salacinifera, P. texana. 

AUSTRALIA AND NEW ZEALAND 

Australia: P. aptata, P. caperata, P. conlabrosa, P. corrugativa, 

P. ferax, P. intertexta, P. labrosa, P. neoquintaria, P. rutidota, P. schistacea, P. scotophylla, P. soredians, P. spodochroa, P. subtiliacea, P. texciiia, P. xcinthoinelaena. 

New Zealand: P. caperata, P. labrosa, P. rutidotn, P. soredians, 

P. subtiliacea. 

PACIFIC REGION 

New Caledonia: P. concomitans. 

Hawaii: P. caperata, P. texana. 

The greatest concentration of species is found in 

South Africa. This region with adjacent countries 

has a flora of 35 species, nearly half of the species 

in the genus (Figure 5). At least 15 of these are 

endemic and most saxicolous. Australia, a region 

of similar climate and yegetation, has 16 species, of 

which 5 {P. aptatn, P. soi~tlians, P. spodochroa, P. 

texana, and P. xanthomelaena) also occur in Africa. 

In the New It'orld, Brazil has the most species, 

18, of which 6 are endemic. North America has 12 

species but only 2 {P. nlabamensis and P. baltimorensis) are endemic there. 

Temperate and tropical eastern Asia (India to 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 5.--Number of species of Pseudoparmelia by major geopolitical regions. 

Japan) has a rather small Pseudoparmelia flora of 

only 14 species, 4 of them endemic and the remainder pantemperate or also widespread in Africa. 

In general the Old World and New World floras 

are quite distinct. The Old World, however, is 

much richer with about 55 species in contrast to 

the 30 species so far collected in North and South 

America. 

Some idea of the abundance of the various species can be gained by tabulating the number of 

countries (as listed above) where each species has 

been reported. This crude method of comparison 

does not, of course, take into consideration the 

relative sizes of the countries or the intensity of 

collecting, but it is unlikely that the order of 

abundance will be greatly changed by future collecting. The following species occiir in five or more 

countries: P. texana (24), P. caperata (17), P. sphaerospoi'a (16), P. caroliniana (13), P. amazonica (9), 

P. canzeopminata (9), P. crozalsiana (Y), P. rutidota 

(9), P. salacinifera (B), P. ecaperata (7), P. aptata (7), 

P. concrescens (6), P. cryptochlorophaea (6), P. 

malaccensis (6), P. martinicana (6), P. soredians (6), 

P. intertexta, (5), and P. owariensis (5). Within its 

range P. caperata is undoubtedly the the most commonly collected foliose lichen, followed by P. caroliniana and in Asia P. ecaperata. 

Twenty-nine species, nearly half the genus total, 

have been found so far in only one country, as can 

be determined from the lists of specimens examined 

in the species treatment, and while many of these 

will be found in other countries with more intensive 

collecting, this great a number reflects the high degree of endemism characteristic of the genus. 

Taxonomic Treatment 

The 76 species of Pseudoparmelia are arranged 

below in alphabetic order. All specimens collected 

by Hale are deposited in the Smithsonian Institution (US), and the herbarium acronym is not cited 

for these. 

The following key is divided into three sections: 

isidiate species, sorediate species, and nonisidiate, 

nonsorediate species. 
KUMBER 31 11 

Keys to the Species of Pseudoparmelia 

ISIDIATE 

I. Thallus greenish yellow (usnic acid present). 

2. Thallus black below with at most a narrow brown zone at the lobe tips. 

3. 

3. Isidia thick and inHated, 0.5-1 nim thick; mzdulla P+ retl. 

4. 1,owrr surface shiny black with a narrow hrown ?one at the lobe tips. 

5. Isidia irregularly scattered, often br.xking open apically. .. 

Isidia fine, no more than 0.2 mm thick; medulla P-. ........ P. ecaperata 

4. Lower surface velvety black to the margin. ....... P. pachydactyla 

5. Isidia dense, cylindrical. .................. ........... P. papillosa 

P. baltirnorensis 

2. Thallus iiniformly pale brown hrlow. 

6. Medulla P+ retl (protocetraric acid). . . P. malaccensis 

6. Medulla P- . 7. Medulla C + red (Iccanoric acid). ...... ..... ..... P. dahlii 

i. Medulla C- or C+ orange (barbatic acid). ............. P. rahengensis 

1. Thallus whitish to greenish mineral gray (usnic acid lacking). 

8. Collected on trees. 

9. Lower surface uniformly pale brown. 

10. 

10. 

9. Lower surface black toward the ccnter and black at the margin or with a narrow 

brown rone at the lobe tips. 

11. Isidia not inflated or pustulate, to 1 mm high. 

12. 

12. Upper surface continuous, not white-reticulate. 

13. Medulla P-. 

Isitlia moderate to dense, 1 mm high: medulla K+ red (salazinic acid). .......... 

.. ........................................ P. salacirtifera 

Isidia sparse, 2-3 mm high; medulla K- or K+ yellow (stictic acid) .......... 

................................................... P. cyphellata 

11. Isidia inHated, pustulate, 2-3 mm high ....................... P. eruptens 

Upper surface finely reticulate!y cracked and strongly white-reticulate ...... 

..................................................... P. caroliniana 

14. Medulla C+ red (lecanoric acid). ............... P. conlabrosa 

14. Medulla C - (divaricatic acid). .............. P. concrescens 

13. Medulla P+ red or orange. 

15. Medulla K+ (salazinic acid) ....................... .P. cinerascens 

15. Medulla K-. 

16. Fumarprotocetraric acid present. .................. P. adspersa 

16. Protocetraric acid present. 

17. Isidia fragile, crumbling; lobe tips white-reticulate. ...... 

........................................ P. rnartinicana 

17. lsidia normal; lobes not white-reticulate. ... P. arnazonica 

8. Collected on rocks. 

18. Isidia coarse, pustulate, usually erupting apically. 

19. Medulla K+ red (salazinic acid). ..................... .P. geesterani 

19. Medulla K-. 

20. Medulla P + red (protocetraric acid). ............. P. simbabwensis 

20. Medulla P - . 

21. Divaricatic acid present. ............................ .P. owariensis 

21. Divaricatic acid absent. 

22. Sekikaic acid present. ... ............ P. pustulescens 

22. Caperatic acid present. ......................... P. schistacea 

18. Isidia normal, thin and cylindrical, not pustulate. 

23. Medulla P+ red or orange. 

24. Lobes 2-3 mm wide, apically subrotund. ................ P. scotophylla 

24. Lobes about 1 mm wide. 

25. Medulla K-; collected in South America. ..... .... P. venesolana 

25. Medulla K+ ycllow (stictic acid); collected in Africa. ... P. ischnoides 

26. Lobes 2-4 mm wide, apically subrotund. 

23. Medulla P-. 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

27. Upper surface finely reticulately cracked. ................ P. cardiniana 

27. Upper surface continuous and smooth. ............... P. subtortula 

26. Lobes 1-2 mm wide, apically obtuse. 

28. Lower surface black. ................................... P. annexa 

28. Lower surface uniformly brown or tan. 

29. Thallus very closely appressed. .................... P. arcana 

29. Thallus adnate to loosely adnate. 

30. “Quintaria” unknowns present; collected in Australia. ........ 

................................. .P. neoquintaria 

30. “Quintaria” unknowns absent; collected in Africa. 

31. Norlobaridone present. .P. subtortula 

31. Norlobaridone absent. ................. .P. basutoensis 

....... 

............ ..... 

SOREDIATE SPECIES 

1. Thallus greenish yellow (usnic acid present). 

2. Lower surface uniformly velvety black below to the margin. .......... .P. subamplexa 

2. Lower surface shiny, black at the center and dark brown in a narrow zone at the lobe tips. 

3. Medulla K+ yellow turning red (salazinic acid). ................... .P. soredians 

3. Medulla K-. 

4. Medulla C+ red (lecanoric acid), P-. ..................... .P. subambigua 

4. Medulla C-, P+ red or orange. 

5. Soralia mostly apical or marginal. 

6. Soralia generally orbicular; collected in Antarctic regions or the high Andes. 

P. gerlachei 

6. Soralia elongate along lobe margins; collected in mid-elevation tropical re5. Soralia mostly laminal, becoming diffuse. .................... .P. caperata 

...... ........................................... 

gions. ................................................ .P. leucoxantha 

1. Thallus whitish to greenish mineral gray (usnic acid absent). 

7. Medulla P+ orange or red. 

8. Upper surface broadly reticulately ridged and foveolate (without lens); stictic acid 

present. 

9. Lobes subirregular, 2-5 mm wide. ............................ P. crozalsiana 

9. Lobes sublinear, 1-2.5 mm wide. .......................... .P. carneopruinata 

8. Upper surface smooth and plane; protocetraric acid present. 

10. Soredia coarse, densely produced over the upper surface in coalescing soralia. , . . , 

...................................................... .. .P. raunkiaeri 

10. Soredia farinose, produced in discrete orbicular soralia. 

11. Collected on rocks ..................................... .P. aiabamensis 

11. Collected on trees ......................................... .P. epileuca 

12. Medulla C+ deep red (lecanoric acid). ..................... .P. labrosa 

12. Medulla C- or C+ fleeting pink. 

13. Soralia strongly capitate along lobe margins. ....... P. cryptochlorophaea 

13. Soralia not capitate, entirely laminal. 

14. Divaricatic acid present. .............................. .P. texana 

7. Medulla P-. 

14. Perlatolic acid present. ................................. P. aptata 

NONISIDIATE, NONSOREDIATE SPECIES 

1. Thallus greenish yellow (usnic acid present; P. chapadensis, P. hgpotnilta, and P. sphaerosporu 

appear to be greenish yellow but lack usnic acid). 

2. Lower surface brown or tan throughout. 

3. Collected on trees. ........................................... .P. intertexta 

3. Collected on rocks. .............. .................. .P. chapadensis 

2. Lower surface black. 

4. Lower surface velvety black to the margin. ......................... .P. amplexa 

4. Lower surface shiny black and often with a narrow brown zone at the lobe tips. 

5. Medulla P-. 

6. Collected in Africa. .................................... P. zambiensis 

6. Collected in South America. .............................. .P. callichroa 
NUMBER 31 13 

3. Medulla P+ red. 

7. Physodalic acid present. .................................. .P. ferax 

Protocetraric acid present. .............................. .P. rutidota 

7. Physodalic acid absent. 

8. 

8. Protocetraric acid and the “conformata” unknown present. ..... .P. exornata 

1. Thallus whitish to greenish mineral gray (usnic acid absent; P. chapadensis, P. hypomilta, 

P. sphaerospora, and sometimes P. violacea appear to be yellowish green but lack usnic acid). 

9. Medulla pigmented purple to yellow. 

10. Medulla yellow or orange. 

10. Medulla completely deep purple. ........................... P. uiolacea 

tains no pigment.) ....................................... .P. sphaerospora 

12. Collected on rocks. ........................... .P. lecanoracea 

11. Medulla completely pale orange-yellow (P. condyloides appears yellow but con11. Medulla pigmented only in the lower half. 

12. Collected on trees. 

13. Lower surface brown; collected in South America. .... P. hrpomilta 

13. Lower surface hlack; collected in Australia. .......... .P. corrugatiua 

9. Medulla white. 

14. Collected on trees. 

15. Lower surface uniformly brown. 

16. Medulla P+ red (protocetraric acid). .P. somaliensis 

16. 

15. Lower surface black (often with a narrow brown zone at the lobe tips). 

17. Medulla P - . 

Medulla P- or P+ faint orange (stictic acid).. ............ .P. sphaerospora 

18. Collected in Africa; divaricatic acid present. .......... P. nairobiensis 

18. Collected in Australia and New Zealand; caperatic acid present. 

....................................................... P. subtiliacea 

19. Medulla K+ yellow (stictic acid); surface of’ lobes reticulately ridged 

and foveolate (without lens). 

20. Lobes 1-2 mm wide; collected in South America. ... P. scrobicularis 

20. Lobes 2-4 mm wide; collected in Africa. ......... .P. inhaminensis 

...... 

17. Medulla P+ red or orange. 

19. Medulla K- (protocetraric acid). 

21. Lobes 1-2 mm wide. .............................. .P. schelpei 

21. Lobes 3-6 mm wide. 

22. Medulla C-; collected in the West Indies ..... .P. inornata 

22. Medulla C+ rose; collected in New Caledonia. .............. 

....... ............................... , . .P. concomitans 

14. Collected on rocks. 

23. Lower surface uniformly brown. 

24. Medulla P+ orange (salazinic acid). 

25. “Chalybeizans” unknown present. .................... P. condyloides 

25. “Chalybeizans” unknown absent. ................... P. spodochroa 

26. Medulla C+ red (lecanoric acid). ................... P. molybdiza 

26. Medulla C-. 

27. Thallus rather loosely attached on rocks and soil; lobes subelongate. 

27. Thallus tightly adnate on rocks; lobes shorter, apically rotund. 

24. Medulla P-. 

...................................................... P. prolata 

28. Norlobaridone present. .......................... .P. tortula 

28. Unknown substances present. ................. P. uanderbylii 

23. Lower surface black (brown only in a narrow zone at the lobe tips). 

29. Medulla P+ orange or red. 

30. Lobes less than 1 mm wide. .................... .P. xanthomelaena 

30. Lobes 2-4 mm wide. 

31. Medulla K-; collected in the Caribbean region ..... .P. caribaea 

31. Medulla K+ yellow or red (salazinic acid); collected in Australia 

and Africa. ...................................... .p. spodochroa 

29. Medulla P-. 

32. Collected in South America. , . , . , , . , . . , , , , , , . , , , . . , , , , , .p. rupicola 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANI 

32. Collected in Africa. 

33. Medulla C+ red (lecanoric acid). . . . . . . . . . . . . . . . . . , P. molybdiza 

33. Medulla C - (divaricatic acid). . . . . . . . , , . . . . . P. rodriguesiana 

Pseudoparmelia 

Pseudoparmelia Lynge, 1914: 15. 

TYPE-SPECIES.-Pseudoparmelia cyphellata Lynge, 

1914:15. 

Lynge erected this genus on the basis of a single 

collection from Brazil, citing the presence of “pseudocyphellae” on the lower surface as the main 

difference from Parmelia. Santesson (1942) demonstrated conclusively that these are in reality artifacts 

caused by rhizines that had been torn off, revealing 

the medulla in irregular patches. He did not reject 

the genus, as implied in the title of his article, but 

regarded it as a synonym of Parmelia. I do not believe that one could invoke Article 71 of the Code 

of Botanical Nomenclature here to reject Pseudoparmelia. The gashes of dislodged rhizines on the 

lower cortex are not monstrosities in the accepted 

sense. Lynge simply described a new genus on a 

mistaken interpretation of the morphology. In any 

event, Article 71 was deleted from the Code at the 

sessions of the ‘International Botanical Congress 

in Leningrad in 1975. The only lichenologist to use 

this genus name in practice was Gyelnik (1933), 

whose species (Pseudoparmelia aradensis and P. 

pseudofallax) have been placed in Cetrelia (Culberson and Culberson, 1968). 

Pseudo par me lia ads persa 

FIGURE 6a 

Pseudoparmelia adspersa (Vainio) Hale, 1974: 189. 

Parmelin adspersa Vainio, 1907: 168 [type collection: Lem 

Sgob, Sam, Sclzinidt XV (TUR, lectotype; C, isolectotype)]. 

Parinelia filipina Hale, 1972a:99 [type collection: Tagaytay, 

Cavite, Philippine, Hale 26809 (US, holotype; TNS, UPS, 

isotypes)]. 

DEscRIPTIoN.-Thal1us adnate on trunks, whitish 

ashy gray, 5-8 cm broad; lobes subrotund, 3-6 mm 

wide, the upper surface plane, continuous or finely 

cracked with age, densely isidiate, the isidia mostly 

simple, cylindrical, 0.1-0.15 mm high, 0.05 mm in 

diameter; lower surface black except for a narrow 

dark brown zone along the margins, sparsely to 

moderately rhizinate. Apothecia (from Hale 26976) 

rare, substipitate, 2-4 mm in diameter, the amphithecium isidiate; spores 8, 5 X 10-12 pm, 

C -, KC -, P + red; atranorin, fumarprotocetraric 

acid, and succinprotocetraric acid. 

CHEMISTRY.-cOrteX K -k yellow, medulla K -, 

DIsTRIBuTIoN.-Phi~ippines, Thailand. 

HABITAT.-Planted Casuarina trees at low to mid 

elevation (200-2000 m). 

REMARKS.-The type collections (Schmidt XV 

and XXXIV) are fragmentary. When I finally resolved the chemistry with TLC, I soon discovered 

that the species is the same as Parmelia filipina. Des 

Abbayes (1958) compared P. adspersa with Parmelia 

inactiva (Zahlbruckner) Vainio, a much larger amphigymnioid lichen containing protocetraric acid. 

I had compared it with Pseudoparmelia carolinana 

(Nylander) Hale, which contains perlatolic acid 

and has a more consistently and conspicuously 

cracked cortex. 

SrEcIhiEws ExAiwNEo.-Philippines: Quezan Province, Hale 

26976; Mountain Province, Santos 88 (PHU, US). 

Pseudoparmelia alabamensis 

FIGURE 6b 

Pseudoparmelia alabainensis (Hale and McCullough) Hale, 

1974:189. 

Parmelia alabamensis Hale and McCullough, 1968:44 [type 

collection: St. Clair County, Alabama, Hale and McCuZlough 24072 (US; isotypes in DUKE, UPS)]. 

DEScRIPTIoN.-Thahs closely adnate on rock, 

ashy white, 2-4 cm broad; lobes narrow and sublinear, contiguous, 0.8-1.2 mm wide; upper surface 

transversely cracked with age, sorediate, the soralia 

laminal, capitate; lower surface black, moderately 

rhizinate, the rhizines black. Apothecia unknown. 

C-, KC- or rose, P+ red; atranorin and protocetraric acid. 

DISTRIBUTION.-United States (Tennessee and 

Alabama). 

HABITAT.-oII large sandstone rocks in open deciduous forests at about 300 m elevation. 

REMARKS.-This species is unique among the 

North American Pseudoparmeliae. An African speCHEMISTRY.-cOrteX K+ yellow, medulla K-, 
NIJhlRER 31 15 

FIGURE 6.-Species of Pseudoparmelia: a, P. adspersa (Hale 26809, holotype of Parmelia filipina 

Hale); b, P. alabamensis (McCullough 1842 in US); c, P. amazonica (Hale 19389); d, P. amplexa 

(Gossweiler 3256e in US); e, P. annexa (Almborn 5683, holotype in LD); f, P. aptata (Almborn 

6548 in LD). (Scale in mm.) 
16 SMITHSONIAX CONTRIBUTIONS TO BOTANY 

cies, P. epileuca (Hale) Hale, also has soredia and 

protocetraric acid, but it is corticolous and has 

broader lobes. 

SpEcrhrms ExAar1sED.-United States: Tennessee, Hale 

36927. 

Pseudoparmelia amazonica 

FIGURE 6c 

Pseudoparmelia amazonica (Nylander) Hale, 1974: 189. 

Parrnelia amaronica Nylander, 1885:611 [type collection: 

Santarem, Brazil, Spruce 11 1 (H, Nylander herbarium number 39111, lectotype; BM, G, NY, [V, PC, isolectotypes)]. 

DEscRIPTroN.-Thalhs closely adnate on bark, 

5-10 cm broad, light mineral gray and turning buff 

in the herbarium; lobes subirregular, apically rotund, 2-6 mm wide, contiguous; upper surface 

plane, continuous, moderately isidiate, the isidia 

rarely branched, to 2 mm high; lower surface black, 

densely rhizinate except for a narrow naked or 

papillate zone along the margins. Apothecia rare, 

1-3 mm in diameter; spores 8, 8-10 X 13-16 pm. 

CHErvmmY.-Cortex K + yellow, medulla K -, 

C-, KC + rose, Pf red; atranorin and protocetraric 

acid. 

DrsTR1BuTIoN.-United States (Florida), Central 

America, West Indies, Colombia, Venezuela, Brazil, 

Guinea, and Taiwan. 

HABITAT.-& trees (conifers, palm trees, deciduous trees), Opzintia, and on rocks in open forests 

at 100-1500 m elevation. 

REMARKs.-SUperficially this species could be confused with P. salacinifera (Hale) Hale, which has a 

pale brown lower surface and salazinic acid. The 

two occupy very similar habitats (trees in open 

secondary forests) and geographic ranges. 

SPECIXIESS ExAxr1NED.-United States: Florida, Hale 34139, 

16859, 17668, 17696, Moore 3357 (DUKE, US), Rapp 311, 662 

(US). Mexico: Veracruz, Hale 19389; Chiapas, Hale 19972 

(DUKE, LD, US), Hale 20187 (Bhl, US), 20607a. Honduras: El 

Paraiso, Standley 560 (F): Morazan, Standley and Williams 

1643 (F). Cuba: Pinar del Rio, Imshaug 25289, 25310, 25333 

(MSC); Oriente, Hioram 5963 (US). Puerto Rico: Howe 1162 

(XU). Trinidad: Wedertnann 2305 (8). Colombia: Santander, 

Killil, 15212, Nee and Mori 3768 (US), Venezuela: Merida, 

Sash 2025 (US). Bolivia: El Beni, Wedernzann 2305 (S). Brazil: 

Minas Gerais, Vainio 588 (BM, TUR), 603 (BM, TUR); Rio de 

Janeiro Burchell 946 (BM); Mato Grosso, Malme s.n., 20506b 

(S), 2369, 2408 (LD, S, US). Guinea: N'Zerekore, Santesson 

10573a (UPS). Taiwan: Asahina (TNSL 

Pseudoparmelia amplexa 

FIGURE 6d 

Pseudoparmelia amplexa (Stirton) Hale, 1974: 189. 

Parmelia amplexa Stirton, 1877:212 [type collection: Somerset East, Union of South Africa, MacOu'an (BM, lectotype; 

GLAM, isolectotype)]. 

Parmelia caperata var. glaucopis Muller ilrgoviensis, 18943258 

[type collection: Matangiri, Seen Region, Africa, Stuhlrnann 

359 (559 on type label) (G, lectotype; BM, isolectotype)]. 

Parmelia glaucopis (Muller Argoviensis) Vainio, 1900:4. 

Parmelia subconspersa var. benguellensis Vainio in TVelwitsch, 

1901:401 [type collection: Mt. hlorro de Lopollo, Huilla, 

Angola, Welwitsclz 31 (BM, lectotype)]. 

Parmelia benguellensis (T'ainio) Dodge, 1959:70. 

Pseudoparmelia benguellensis (Vainio) Hale, 1974:189. 

DEscRiPTIoN.-Thallus rather closely adnate on 

bark, coriaceous, greenish yellow, 3-5 cm broad; 

lobes sublinear, crowded, partly black rimmed, 1-2 

mm wide; upper surface plane, dull, rugose and 

becoming lobulate with age and heavily pycnidiate; 

lower surface black and minutely velvety to the 

margin, sparcely to moderately rhizinate. Apothecia 

numerous, sessile, 1-2 mm in diameter; spores 8, 

7-8 X 16-18 pm. 

KC-, or KC+ rose, P+ red; usnic acid and protocetraric acid with or without a trace of atranorin. 

DIsTRisuTIo~.-Southern Africa. 

HABITAT.-on trunks and branches of trees in 

open rocky areas. 

REMARKS.-I had at first identified all African 

specimens as Pseudoparmelia rutidota (Hooker and 

Taylor) Hale, but close examination showed that 

they have consistently narrow lobes and a jet black 

velvety lower surface (Figure la, b). By comparison, P. rzitidota, which is known from Australia and 

the New World, often has quite broad lobes (to 

5 mm wide) and a shiny surface below and brown 

bare zone near the tips. The sorediate morph of 

P. amplexa is P. subamplexa Hale and the isidiate 

morph is P. pachydactyla (Hale) Hale. All three 

appear to lack any fatty acids, in contrast to the 

P. rutidota group, which usually produces one or 

more of the common fatty acids. 

CHEMISTRY.-cOrteX K-, medulla K-, C-, 

SPECIMENS ExAxr1mD.-Rhodesia: KoPer (LD, US), Schiitte 

(LD, US), Sheppard 2 (US). Angola: Bie, Gossweiler 3256e 

(BM, US), Degelius (Degelius herbarium, US); Huila, Degelius (Degelius herbarium); hfoxico, Degelius (Degelius 

herbarium. US). 
NUMBER 31 17 

Pseudoparmelia annexa 

FIGURE 6e 

Pseudoparmelia annexa (Kurokawa) Hale, 1974: 189. 

Parmelia annexa Kurokawa, in Hale and Kurokawa, 1964:151 

[type collection: Caledon, Cape Province, Union of South 

Africa, Almborn 5683 (LD, holotype; isotype in US)]. 

DESCRIPTIoN.-Tha11us tightly adnate on rock, 

ashy mineral gray, 3-7 cm broad; lobes subirregular 

to sublinear, 0.7-3.5 mm wide; upper surface plane, 

shiny, continuous or irregularly cracked with age, 

moderately isidiate, the isidia mostly simple, about 

0.2 mm high, darkening at the tips; lower surface 

black, sparsely rhizinate. Apothecia adnate, 1.5-5 

mm in diameter, the amphithecium isidiate; spores 

8, 5 X 7-8 pm. 

CHEMISTRY.-cOrteX K+ yellow, medulla K-, 

C + , KCf red, P- ; atranorin and lecanoric acid. 

DIsTRIBuTIoN.-Uganda, Angola, and Union of 

South Africa. 

level to 1200 m. 

REkfARKS.-This rare saxicolous species is the 

isidiate morph or close relative of P. molybdiza 

(Nylander) Hale, which lacks isidia and has generally broader lobes. Both are African endemics. 

HABITAT.-on exposed rock outcrops from sea 

SPECIMENS EXAMINED.-uganda: Pian, Swinscow 6/ 1970 

(BM, US), Other records from Angola and South Africa are 

listed in Hale and Kurokawa (1964:151). 

Pseudoparmelia aptata 

FIGURE 6f 

Pseudoparmelia aptata (Krempelhuber) Hale, 1974:189. 

Parnzelia aptata Krempelhuber in Nylander, 1869:291 [type 

collection: Kotgurh, Himalayas, India, Skolicrka 509 (M, 

lectotype; BM, H, K, US, isolectotypes)]. 

Parmelia asmarana Vainio, 1898b:37 [type collection: Asmara, 

Ethiopia, Levander (TUR, Vainio herbarium number 

2664, lectotype)]. 

Parmelia nipponica Zahlbruckner, 1927:353 [type collection: 

Hachinoche, Japan, Faurie 1239 (W, lectotype)]. 

DEScRIPTIoN.-Thallus adnate on bark, whitish 

mineral gray, 6-12 cm broad; lobes sublinear to 

subirregular, more or less apically rotund, 2-5 mm 

wide; upper surface plane, continuous or cracked 

with age, laminally sorediate, the soralia capitate; 

medulla white except for pale orange pigmentation 

below the soralia; lower surface black, sparsely 

rhizinate except for a narrow marginal papillate 

zone. Apothecia not seen. 

CHEMIsmY.-Cortex K + yellow, medulla K- , 

C -, KC + rose, P - ; atranorin, perlatolic acid, and 

an unidentified pigment. 

DIsTRIBuTI0N.-Africa, India, Indonesia, Australia, and Japan. 

HABITAT.-oII trees (Pinus, deciduous trees), 

secondary forests at low to mid elevations. 

REMARKS.-This Old World lichen is externally 

similar to P. texana (Tuckerman) Hale, which contains divaricatic acid and is pantropical-pantemperate. It apparently evolved in the Old World as 

a chemical variant from a now extinct fertile perlatolic acid-containing progenitor, related perhaps 

to P. nairobiensis (Steiner and Zahlbruckner) Hale. 

It is basically sympatric with P. texana in most of 

Asia and in Japan comprises about two-thirds of 

the specimens in this group. Chemical tests must 

be made to separate the species. 

SPECIMENS ExAarINEo.-Tanzania: Hoeg (TRH). Union of 

South Africa: Natal, Almborn 6548, 8538, 8697, 9643 (LD), 

Hoeg (TRH). India: .Sikkim, Hara (TNS). Japan: Province 

Aki, Hale 29517, 29542; Province Ohmi, Hale 29473. Indonesia: Java, Groenhart 8853, 8854 (BOR), 2197, 5322, 5324 

(L, US), Neeruoort 753 (BOR). Australia: Queensland, Bailey 

(BM). 

Pseudoparmelia arcana 

FIGURE 7a 

Pseudoparmelia arcana (Kurokawa) Hale, 1974: 189. 

Parmelia arcana Kurokawa in Hale and Kurokawa, 1964:151 

[type collection: Baba, Mocamedes, Angola, Degelius (Degelius herbarium; isotype in US)]. 

DESCRIPTIoN.-Thallus tightly appressed on rock, 

ashy white, 3-7 cm in diameter; lobes sublinear, 

crowded and appearing subareolate toward the 

center, 0.3-1 mm wide; upper surface plane to convex, continuous to tangentially cracked, moderately 

isidiate, the isidia mostly simple, about 0.2 mm 

high; medulla white or in part pale yellow; lower 

surface brown to pale brown, sparsely rhizinate. 

Apothecia not seen. 

C-, KC-, P-; atranorin, fatty acids, and unidentified pigments. 

DIsTRIBUTION.-Angola and Union of South 

Africa. 

CHEMISTRY.-cOrteX Kf yellow, medulla K-, 
18 SMITHSONIAN CONTRIBUTIOXS TO BOTANY 

HABITAT.-on rocks in dry open areas at about 

1000 m elevation. 

REMARKS.-This rarely collected species seems to 

be related to P. annexa (Kurokawa) Hale, which 

contains lecanoric acid. The yellow pigmentation 

does not seem to be a consistent character to judge 

from the few specimens available. 

SPECIMENS EXAXINED.-uniOn of South Africa: Basutoland, 

KoPer (LD); Cape Province, Hiieg (TRH); Orange Free 

State, Plank (PRE). 

Pseudoparmelia baltimorensis 

FIGURE 7b 

Pseudoparmelia baltirnorensas (Gyelnik and Foriss) Hale, 

1974:189. 

Parmelia baltimorensis Gyelnik and F6riss in Gyelnik, 1931a: 

167 [type collection: Gunpowder RiFer, Baltimore County, 

Maryland, Plitt (BP, holotype; US, isotype)]. 

DESCRIPTIoN.-Thal1us closely to loosely adnate 

on rocks, yellowish green, up to 15 cm broad but 

colonies of ten coalescing and covering extensive 

areas of rock; lobes sublinear to irregular, apically 

subrotund, 3-5 mm wide; upper surface plane, becoming covered toward the center with large, thick, 

simple or branched isidioid pustules, fragile and 

breaking open apically but not becoming sorediate; 

lower surface black except for a narrow, smooth 

to rugose, papillate, brown zone along the margins, 

moderately rhizinate. Apothecia rare, substipitate, 

1-3 mm in diameter, the amphithecium pustulate; 

spores 8, 6-7 X 13-15 pm. 

C+ rose, KC+ rose, P+ red; usnic acid, protocetraric acid, and very frequently gyrophoric acid. 

DIsTRIBuTroN.-Eastern United States and adjacent Canada. 

HABITAT.-sandstone and granite rocks in open 

deciduous forests. 

REhfARKs.-This species had been confused with 

the equally common but usually corticolous P. 

caperata (L.) Hale. The distinction between the 

coarse lamina1 pustules of P. baltimorensis and the 

more or less diffuse soralia of P. caperata, as well 

as the presence of gyrophoric acid as an accessory 

substance in P. baltimorensis, was recently discovered by Dr. W. L. Culberson (in litt.), who 

kindly showed me his preliminary results. He is 

now preparing a paper on the biology of these two 

CHEMISTRY.-cOrteX K-, medulla K--, c- or 

species that will present much more detailed information. 

REPRESENTATII E SPECI\IEhS EXAMINED (all in US).-Canada: 

Ontario, Hale. United States: Massachusetts, Hutchinson 

1744; Connecticut, Hale 19076; New Jersey, Hale 17266; 

Pennsylvania, Becking 57070315, Hale 16292, Heller, White 

210; Maryland, Hermann 13768, Leonard 2933, 3086, h'orden 

27, Plitt; West Virginia, Hale 10734, 13016; Kentucky, Allen 

545, Hale 13115; Illinois, Hale 13926, 14029, Skorepa 943; 

Wisconsin, Hale 23542, Thomson 919; Minnesota, Hale 23479; 

Missouri, Greeninan 4763, Ireland 3238, Schoop 254; Virginia, 

Hale 15107, 17317, Luttrell 120, 4876; Xorth Carolina, Culberson in Lichenes Selecti Exsiccati 915, Hale 16371; Tennessee, Degelius, Hale 31118, Phillips 321; South Carolina, Hale 

7747; Georgia, Hale 30877; Alabama, Hale 7125, 33841, McCullough 344, 2032; Mississippi, Hale 7846; Arkansas, Hale 

in Lichenes Ainericani Exsiccati 2, Hale 3644, 3813, Moore 

259; South Dakota, Vischer 4; Kansas, Hale 2934, 4534; Oklahoma, Hale 5053; Texas, Hale 34317. A number of records 

in other herbaria would have been identified as Parmelia 

caperata by me before 1970; it was impractical to borrow 

these specimens again and correct the names. 

Pseudoparmelia basutoensis 

FIGURE 7c 

Pseudoparmelia basutoensis (Hale) Hale, 1974:189. 

Parmelia basutoensis Hale, 1972b:342 [type collection: Roma, 

Basutoland, Union of South Africa, KoPer (LD, holotype; 

isotype in US)]. 

DEscRIPTroN.-Thallus adnate on rocks, pale 

greenish white, 3-5 cm broad; lobes sublinear to 

subirregular, contiguous, 2-3 mm wide; upper surface plane, isidiate, the isidia cylindrical, simple; 

lower surface pale brown, sparsely rhizinate. Apothecia sparse, 1-3 mm in diameter, spores not found. 

CHEMIsmY.-Cortex K + yellow, medulla negative with all reagents; atranorin and an unknown 

substance. 

DISTRIBuTIoN.-~nion of South Africa. 

HABITAT.-RockS in dry regions. 

REMARKS.-AS I pointed out in the original description, this species has the general aspect of a 

Xanthoparmelia but lacks usnic acid. The nonisidiate morph and presumptive parent is P. prolata 

(Hale) Hale (see below), but neither species has 

been collected in sufficient quantity for more definitive statements on biology and possible evolution of the group. The unknown substance appears 

to be a new fatty acid that reacts faintly yellow with 

sulfuric acid. The same substance has been discovered in a brown Parmelia, P. scabrella Esslinger 

(Esslinger, 1976). 
NUMBER 31 

FIGURE 7.-Species of Pseudoparmelia: a, P. arcana (Degelius, isotype in US); b, baltimorensis (Luttrell 3369 in US); c, P. basutoensis (Hewitt, holotype in TRH); d, P. caperata (Plitt in 

US); e, P. caribaea (Gallo 494, holotype in US); f, P. carneopruinata (Hale 42912). (Scale in mm.) 

19 
SMITHSONIAN CONTRIBUTIONS TO BOTAX'Y 

Pseudoparmelia callichroa Kurokawa, new species 

DESCRIPTIoN.-Thallus adnatus, ramulicola, 

viridi-flavicans, 3-6 cm latus, lobis subirregularibus, 

congestis, apice subrotundatis; superne planus vel 

mox irregulariter rugosus, isidiis sorediisque destitutus; cortex superior 10-13 Fm crassus, epicortex 

perforatus, stratum gonidiale 12 pm crassum, medulla alba, 100-120 pm crassa, cortex inferior 12-13 

pm crassus; subtus niger, nitidus, modice rhizinosus, 

rhizinis simplicibus, nigris. Apothecia numerosa, 

amphithecio ruguloso, margine crenato, 1 .O-4.5 mm 

diametro; hymenium 70-85 pm altum; sporae 8:nae, 

6-8 X 13-16 pm. 

CHmmmY.-Cortex K -, medulla negative with 

all reagents; usnic acid and protolichesterinic acid. 

HoLoTYPE.-Chile: Colchagua, Hacienda de Conquenes, P. Duse'n 91, 22 August 1896 (S; US, isotype). 

DISTRIBUTIoN.-Chi~e. 

REMARKs.-This species would probably be 

identified as P. rutidota (Hooker and Taylor) Hale 

without a chemical test (P. rutidota is Pf red with 

protocetraric acid). The thallus is smaller but we 

need many more collections to determine whether 

any morphological characters correlate with the 

chemical difference. 

HABITAT.-On twigs. 

Pseudoparmelia caperata 

FIGURE 7d 

Pseudopartnelia caperata (L.) Hale, 1974: 189. 

Lichen caperatus L. 1753:1147 [type collection: England, 

Dillenius, Historia Muscoruin 193, specimen represented 

by $1. 25: fig. 97 (OXF, lectotype)]. 

Parmelia caperata (L.) Acharius, 1803:216. 

Parmelia caperata b. cyliphora Acharius, 1814:196 [type collection: North America, Ltfulilenberg (H, lectotype; UPS, 

isolectotype): listed in error as "cylisphora."] 

Parinelia perlata var. flauicans Tuckerman, 1866: 13 [type 

collection: California, flolaizder TO (FH-Tuck, holotype)]. 

Parlnelia euplecta Stirton, 1877-78:299 [type collection: Brisbane, .4ustralia, Bailey (BM, lectotype)]. 

Parnielia flauicans (Tuckerman) Tuckerman, 1882:53 [not 

P. flauicans (Swartz) Acharius, 1803:268 (= Teloschistes)]. 

Partnelia ochrolewa f. sorediosa Muller Argoviensis, 1883b: 

77 [type collection: Mt. Dromedary, Australia, Reader (G, 

lectotype)]. 

Parmelia subglauca Nylander in Gasilien, 1894:126 [type 

collection: Saint Omer, France (H, lectotype: BM, PC isotypes)]. 

Parmelia cylisphora [sic] (Acharius) Vainio, 1896b:7. 

Parmelia cuperata var. subglauca (Nylander in Gasilien) Nylander, 1896:35. 

Partnelia caperata f. laeuissima Gyelnik, 1928:587 [type collection: Meleghegy, Fehkr, Hungary, GyeInik (BP, holoPnrmelia herreana Zahlbruckner, 1929:239 [based on P. fiauiPartnelia pseudosorediosa Gyelnik, 1931b:288 [based on ParParmelia negatiua Gyelnik 1934:301 [type collection: Salisbury 

Pseudoparmelia euplecta (Stirton) Hale, 1974: 190. 

Pseudoparmelia pseudosorediosa (Gyelnik) Hale, 1974: 190. 

type)]. 

cans (Tuckerman) Tuckerman]. 

nzelia ochroleuca f. sorediosa &fuller Argoviensis]. 

Cove, Maine, Plitt (BP, holotype)]. 

DEscRIPTrox.-Thallus adnate to loosely attached, 

growing on bark or more rarely on rocks, yellowish 

green, 5-20 cm in diameter, coalescing to form 

larger colonies; lobes subirregular, apically rotund, 

3-8 mm wide; upper surface plane to undulate or 

rugulose, continuous, sorediate, the soralia laminal, 

diffuse and coalescing; medulla rarely orange near 

the lower cortex; lower surface black, moderately to 

sparsely rhizinate except for a naked brown rugose 

zone along the margins. Apothecia very rare, 2-5 

mm in diameter; spores 8, 8-13 X 17-24 pm. 

KC+ rose, Pf red, atranorin, caperatic, protocetraric, and usnic acids; if pigmented, skyrin present. 

DrsTRIsuTronl.-Pantemperate on all major continents (Figure 8). 

HABITAT.-on conifers and deciduous trees in 

temperate forests, common on rocks only at the 

northern part of its range, from sea level to over 

3000 m elevation. 

RE&fARKs.-This is by far the most widespread 

and commonly collected foliose lichen in all temperate areas of the world. The range in morphological variation is very small and the chemistry 

remarkably uniform. In eastern North America it 

can be confused with the primarily saxicolous P. 

baltimorensis (see discussion above). The presumptive nonsorediate parent seems to be P. rutidota 

(Hooker and Taylor) Hale (see below). 

Older authors consistently placed this species in 

Parmelia section Amphigymnia (= Parmotrema). 

The lobes, however, are clearly adnate at the margins, the bare rim below is very narrow, the tips of 

rhizines near the margin are frayed, and the apothecia are adnate and nonperforate. This combination of characters relates the species much m6re 

closely to Pseudoparmelia than to Parmotrema. 

The species is apparently present in Australia 

CHESIISTRY.-COrteX K--, medulla K-, C-, 
NUMBER 31 21 

FIGURE 8.-World distribution of Pseudoparmelia caperata. (Dots may represent more than one 

collection.) 

and New Zealand, represented by the anonomously 

pustulate P. euplecta. Parmelia ochroleuca f. sorediosa, also described from Australia, is atypical in 

having rather discrete orbicular soralia. Future 

studies in this region may show that these populations are distinct from P. caperata. 

SPECIMENS EXAMINED.-%e Figure 8 for localities of specimens annotated in the major herbaria. Some specimens from 

eastern North America which I annotated before about 1972, 

especially those on rock, will be P. baltimorensis. 

Pseudoparmelia caribaea 

FIGURE 7e 

Pseudoparmelia caribaea (Hale) Hale, 1974:189. 

Parmelia caribaea Hale in Hale and Kurokawa, 1964:152 

[type collection: St. Barthdemy, West Indies, Le Gallo 494 

(US, holotype)]. 

DEscRxPTIoN.-Thahs adnate on rocks, ashy 

white, rather coriaceous, 6-10 cm broad; lobes irregularly sublinear, often densely imbricate, 3-6 

mm wide; upper surface plane or rugulose, cracked 

with age; lower surface black, moderately rhizinate. 

Apothecia subpedicellate, at first globose, urceolate 

at maturity, up to 5 mm in diameter; spores 8, 4-5 

X 10-13 pm. 

C -, KC + rose, P + red; atranorin and protocetraric 

acid. 

DISTRIBUTION.-weSt Indies and French Guiana. 

HABITAT.-oII schist and volcanic rocks in open 

areas up to 275 m elevation. 

REMARKS.-Except for the two specimens from 

French Guiana and the Isle of Pines, P. caribaea 

is confined to St. Barthblemy, where it has been collected many times. There are no close relatives in 

the genus. 

CHEMISTRY.-cOrteX K+ yellow, medulla K--, 

SPECIMENS EXAMINED.-cUbP: Isle of Pines, Rritton et al. 

14436, 15237 (US). St. Barthelemy: Le Gallo 403, 424, 527, 

536, 546 (US), 2604 (MSC, US). French Guyana: Degelius 

(Degelius herbarium). 
22 SMITHSONIAN COXTRIBUTIONS TO BOTANY 

Pseudoparmelia carneopruinata 

FIGURE 7f 

Pseudoparrrielia carneopruinata (Zahlbruckner) Hale, 1974: 

189. 

Parmelia carneopruinata Zahlbruckner, 1902:419 [type collection: Rio de Janeiro, Brazil, HBhnel 164 (W, lectotype)]. 

Parrnelia sbnrbaronis Bouly de Lesdain, 1923:278 [type collection: Catalupe, Varezze, Liguria, Italy, Gresino 11467 (F, 

lectotype)] . 

DEsCRIPTIor\;.-Thallus closely adnate, corticolous, greenish mineral gray or in the herbarium 

buff, 5-9 cm in diameter; lobes sublinear, 1-2.5 mm 

wide; upper surface strongly reticulately ridged and 

rugulose, becoming pruinose near the lobe tips, 

sometimes densely lobulate, sorediate, soralia about 

1 mm wide, often coalescing; lower surface black, 

sparsely rhizinate. Apothecia rare, adnate, 1-4 mm 

in diameter, the disc pruinose or naked, the amphithecium sorediate; spores 8, 6-9 X 9-1 3 pm. 

yellow, C-, KC-, Pf pale orange; atranorin, 

stictic acid, and constictic acid with associated unknowns. 

DIsTRIBuTIoN.-h!feXiCo, Central America, West 

Indies, Colombia, Venezuela, Brazil, Uruguay, Argentina, and southern Europe. 

HABITAT.-on shade trees in banana and Coffee 

plantations and on deciduous trees in open pastures 

at 300-2000 m elevation. 

REbfARKs.-This species is closely related to P. 

scrobicularis (Krempelhuber) Hale, which is or at 

least may be the nonsorediate progenitor. Another 

very close sorediate species, P. crozalsiana (Bouly de 

Lesdain) Hale, has much broader, subrotund lobes 

and a wider, more temperate distribution. The two 

species, however, probably intergrade and are not 

always easily distinguished. 

SPECIMENS ExAhrr?iEo.-Mexico: Xayarit, Weber 33609 

(COLO); Veracruz, Hale 19398, 19432 (DUKE, US), 19452, 

19654, 19672, 19794, 21081, 21195. Honduras: Morazln, 

Standley 11655 (F, US), 11657 (F). Costa Rica: San Josk, 

Standley 34811 (US). Jamaica: Orcutt 5603 (US), Plitt (Us). 

Colombia: Lindig 2590 (G). Venezuela: Mkrida, Hale 42785, 

42912. Brazil: Minas Gerais, Mexia 5322 (BM); SPo Paulo, 

Schindler 4512, 4525 (KR, US). Uruguay: Canelones, Osorio 

6236 (MVM). Argentina: Jujuy, Fries 43b (S). Europe: Italy, 

Sbarbaro (TUR). 

CHEMISTRY.-cOrteX K + yellow, medulla K f 

Pseudoparmelia caroliniana 

FIGURE 9a 

Pseudoparmelia caroliniana (Nylander) Hale, 1974: 189. 

Parmelia caroliniana Nylander, 1885:614 [type collection: 

South Carolina, Ravenel 404 (H, lectotype; FH-Tuck, isolectotype)]. 

Parrnelia perlata var. subrevoluta Miiller Argoviensis, 1880:267 

[type collection: Petropolis, Brazil, Deventer 45 (G, lectoParmelia isidiophora Zahlbruckner, 1902:420 [type collection: 

Botanical Garden, Rio de Janeiro, Brazil, Hohnel 169, (W, 

lectotype)]. 

Partnelia luteola Zahlbruckner, 1909:170 [type collection: 

Sear Barra Mansa, SPo Paulo, Brazil, Wettstein and Schifliier (W, lectotype)]. 

Parinelin zuainoana Lynge, 1914:87 [type collection: Santa 

Anna da Chapada, Mato Grosso, Brazil, Malme 2435C (S, 

lectotype)]. 

Pyxine azorea Nylander, 1895:lOO [type collection: Azores, 

Michel 66 (H, lectotype)]. 

DEsCRIPTION.-Thal~us closely adnate to bark, 

rarely on rock, 5-10 cm broad; lobes subirregular, 

rotund, 2-5 mm wide; upper surface plane, reticulately white-maculate and finely cracked, densely 

isidiate, the isidia 1-3 mm high, often branched; 

lower surface black, very rarely uniformly dark 

brown, moderately rhizinate except for a narrow 

naked or papillate zone along the margins. Apothecia rare, adnate, 1-3 mm in diameter, the amphithecium isidiate; spores 8, 6-8 X 13-15 pm. 

C -, KC + faint rose or purple, P - ; atranorin and 

perlatolic acid with associated unknowns. 

DIsmrnuTroN.-Southeastern United States, 

Mexico, Central America, West Indies, Venezuela, 

Ecuador, Brazil, Azores, Africa, and Thailand. 

HABITAT.-on trees (conifers and deciduous 

trees) in coffee plantations, secondary forest, and 

pastures and rarely on rocks from sea level to 2800 

m elevation. 

RE&aARKs.-The first study of this species was 

done by Culberson (1957), who clarified its relation and frequent confusion with Parmelia 

rudecta Acharius, another widespread species which 

differs in having pseudocyphellae and lecanoric acid 

as well as a paraplectenchymatous upper cortex. 

Pseudoparmelia caroliniana is the only isidiate species in the genus with perlatolic acid; P. aptata 

(Krempelhuber) Hale also has perlatolic acid but 

is sorediate and lacks the fine reticulation so characteristic of P. caroliniana. I wish to thank Dr. T. D. 

V. Swinscow for pointing out the synoymy of 

Pyxine azorea. 

type)]. 

CHEMISTRY.-cOTteX K + yellow, medulla K -, 

SPEcrarENs ExAMrNm.-United States: See Culberson (1957) 

for localities over the range in the United States and Moore 

(1968:223) for specimens from Florida. Mexico: Hidalgo, 
NUMBllR 31 23 

FIGURE 9.-Species of Pseudoparinelia: a, P. caroliana (Hale 20144); b, P. chapadensis (Malme 

2297B, lectotype in S); c, P. cinerascens (Xauier 713 in US); d, P. concomitans (Hill 12091, holotype in BM); e, P. concrescens (Swinscow 2U 24/21 in BM); f, P. condyloides (Almborn 4882, 

holotype in LD). (Scale in mm.) 
24 SMITHSONIAN CONTRIBUTIONS TO BO

Robinson (US); Veracruz, Hale 19471, 19488, 21131, 21135; 

Chiapas, Hale 19900, 20104, 20144, 20215, 21310, 20333. Honduras: MoraaBn, Standley 238, 11603 (F). Nicaragua: Standley 

8432 (F). Costa Rica: Guanacaste, Standley 44370 (US). 

Panama: Chiriqdi, Hale 38804. Cuba: Wright 71 (FH, M, PC, 

UPS, US); Oriente, Imshaug 24763, 24858 (MSC). Jamaica: . Imshaug 13762, 14200 (MSC), Plitt (US). Haiti: Ouest, Imshaug 

22542, 22859, 22944, 23061 (MSC, US), Wetmore 2882, 3076, 

3149, 3207 (MSC, US), Leonard 3709b (US). Dominican Republic: Cordillera Central, Imshaug 23714 (MSC, US, WIS), 

23718 (Bhf, G, MSC); La Vega, Allard 18033 (US). Venezuela: 

Distrito Federal, Dennis 1503 (BM). Santesson 6672, 6681 (S); 

Mbrida, Hale 42039, 42196, 43318, 44081, 44225; Tachira, 

Hale 45632, 45649. Ecuador: Galapagos, Johannsen 10 (WIS), 

Weber 40031 (COLO, UPS). Brazil: Rio de Janeiro, Eiten 

7373 (US); Sao Paulo, Eiten 2978, (US) Osorio 4925 (MVM); 

Santa Catarina, Reitz and Klein 13145, 15398 (US). Europe: 

Portugal, Persson (UPS). Ivory Coast: Man, Santesson 10494 

(UPS, US). Uganda: Burahiya County, Swinscow 2U 18 (BM); 

Kyadondo County, Swinscou! 2U 1/7 (BM). Angola: Huila, 

Degelius (Degelius herbarium). Laos: Tsuyama 19 (TNS). 

Pseudoparmelia chapadensis 

FIGURE 9b 

Pseudoparmelia chapadensis (Lynge) Hale, 1974: 189. 

Parmelia chapadensis L~nge, 1914: 153 [t) pe collection: near 

Bocca da Serra, Serra da Chapada, Mato Grosso, Brazil, 

Malme 2297B (S, lectotjpe)]. 

DEscRIPTIoN.-Thallus closely adnate on rock, 

yellowish green, 1-1.5 cm broad; lobes sublinear, 

short, 0.5 mm wide, convex and expanded at the 

tips; medulla yellowish; lower surface pale brown, 

moderately rhizinate. Apothecia common, sessile, 

0.5-1.0 mm in diameter; spores 8, 4-5 X 8-10 p.m. 

C+, KC+ yellowish, P+ red; unidentified substances present. 

DIsTRIBuTION.-Brazil. 

HABITAT.-Rocks in open areas. 

REMARKS.-This peculiar species is tentatively 

placed in Pseudoparmelia because the lobes are 

eciliate and the rhizines simple. Contrary to my 

earlier findings (Hale, 1960), it does not contain 

usnic acid or protocetraric acid. The yellow coloration is caused by unidentified pigments in both the 

cortex and the medulla. Perhaps additional collections will clarify the status of the species. 

CHEMISTRY.-cOrteX K -k yellow, medulla K f , 

Pseudoparmelia cinerascens 

FIGURE 9c 

Pseudoparmelia ciwrascens (Lynge) Hale, 1974: 189. 

Parmelia cinerascens Lynge, 1914: 104 [type collection: Paraguari, Paraguay, Afalme 1498 (S, lectotype)]. 

DEscRIPTIoN.-Thallus adnate, fragile, ashy 

eral gray, 4-5 cm broad; lobes short, beco

imbricate, 1-3 mm wide; upper surface plan

regularly cracked with age, densely isidiate, i

cylindrical, simple or branched; lower surface b

rhizines moderate to sparse. Apothecia abun

adnate; spores 8, 6 X 12 pm. 

yellow turning red, C-, KC-, P+ orange

ranorin and salazinic acid. 

CHEMISTRY.-cOrteX K f yellow, medulla 

DIsTRIBunoN.-Brazil and Paraguay. 

HABITAT.-On trees in open forest. 

REMARKS.-This rare species is superficially 

P. salacinifera (Hale) Hale, which differs in ha

a pale brown lower surface. 

SPECIMESS ExAI\fINED.-Brazi1: Pernambuco, Xavier 713 

Pseudoparmelia concomitans, new species 

FIGURE 9d 

DEscRIPTIoN.-Thallus laxe adnatus, cort

bubalino-albidus, coriaceus, 4-6 cm latus, lobis 

irregularibus, apice rotundatis, 3-5 mm 

superne planus, nitidus albo-maculatusque, so

et isidiis destitutus; cortex superior 10-14 pm 

sus, epicortex sparse perforatus, stratum goni

10-12 pm crassum, medulla 90-110 pm c

Cortex inferior 8-10 pm crassus; subtus n

sparse rhizinosus, ambitu nudus, castaneus, nit

Apothecia numerosa, substipitata, 1-2 mm 

metro; sporae 8:nae, 12 X 20-23 pm. 

C +, KC + red, P + red, atranorin, gyrophoric 

and protocetraric acid. 

HoLoTYPE.-New Caledonia: Baie des Crabe

de Pins, on fallen branches of Araucaria cookii

Hill 12124 (BM; US, isotype) . 

CHEMISTRY.-cOrteX K + yellow, medulla 

DISTRIBUTION.-New Caledonia. 

HABITAT.-on branches of Araucaria. 

REMARKS.-This species is known only from 

type collection and seems unrelated to any 

Pseudoparmeliae. The lobes are coriaceous 

strongly white maculate. Only P. martinicana 

duces this combination of medullary substance

it is an isidiate, fragile lichen confined to the 

Indies. 

Pseudoparmelia concrescens 

FIGURE 9e 

Pseudoparmelia concrescens (Vainio) Hale 

Parmelia concrescens Vainio, 1901 :400 [type collection
NUMBER 31 25 

da Xella, Huila, Angola, Welwitsch 30 pro parte (TUR, 

lectotype; BM, isolectotype)]. 

Parmelia capensis Nylander, 1885:613 [type collection: Cape 

of Good Hope, Union of South Africa, Drege (H, Nylander 

herbarium number 35174, lectotype; PC, isotype); not 

P. capensis (Acharius) Sprengel, 1827:280 (= Teloschistes)]. 

Parmelia austroafricana Zahlbruckner, 1929:152 [type: based 

on P. capensis Nylander]. 

Parmelia caffrorum Zahlbruckner, 1932a:555 [type: based on 

P. capensis Nylander]. 

DEscRwTIoN.-Thallus adnate on bark, ashy 

mineral gray, 4-10 cm in diameter; lobes irregularly 

sublinear, 1-3 mm wide; upper surface plane, continuous or cracked on older lobes, densely isidiate, 

the isidia short, cylindrical, mostly simple; lower 

surface black, sparsely rhizinate. Apothecia rare, 

adnate, 2-3 mm in diameter; spores 8, 5 X 7-8 pm. 

C--, KCf faint wine red, P-; atranorin and divaricatic acid with associated unknowns. 

CHEMISTRY.-cOrteX K + yellow, medulla K - , 

DISTRIBUTION.-SOUthern Africa. 

HABITAT.-On trees or rocks in open woodland 

at 1000-2300 m elevation. 

REMARKS.-ThiS is one of the commoner, more 

widespread lichens in Africa. It is presumed to be 

an isidiate morph of P. nairobiensis (Steiner and 

Zahlbruckner) Hale, with which it is sympatric. 

It is also close to the more widespread P. ecaperata 

(Muller Argoviensis) Hale, which differs in containing usnic acid in the cortex. 

SPECIMENS ExA%rINED.-uganda: Dummer 602 (US); Busiro 

County, Swinscou! 2U 24/21 (BM, US); Kinkizi County, Dale 

L46 (BM, US). Zaire (Congo): Degelius (Degelius herbarium, 

US), Hoeg (TRH), Louis 7494 (BR), de Witte 2675 (BR). 

Angola: Cuanza-Sul, Degelius; Bie, Degelius; Huila, Degelius; 

Moxico, Degelius (all Degelius herbarium). Rhodesia: Arne11 

1282a (LD), Schutte 46c (LD, US). Moqambique: Alrnborn 

6888, 7015 (LD). Union of South Africa: Natal, Almborn 

8637 (LD). 

Pseudoparmelia condyloides 

FIGURE 9f 

Pseudoparmelia condyloides (Kurokawa) Hale, 1974: 189. 

Parmelia condyloides Kurokawa in Hale, 1972:343 [type collection: Kamieskroon, h’amaqualand, Union of South 

Africa, Almborn 4882 (LD, holotype; TNS, US, isotypes)]. 

DEScRIPTIoN.-Thallus closely adnate on rock, 

dark mineral gray with a yellowish cast, 3-5 cm 

broad; lobes subirregular to sublinear, crowded, 

1.5-2.5 mm wide; upper surface plane to rugosebullate, the cortex breaking away easily; lower surface pale brown, moderately rhizinate, the rhizines 

pale. Apothecia numerous, adnate, 1-2 mm in diameter; spores 8, 4 X 5 pm. 

yellow turning red, C-, KC-, Pf orange; atranorin, “chalybeizans” unknown, salazinic acid, and 

a pale yellow unidentified pigment. 

CHEMISTRY.-cOrteX K f yellow, medulla Kf 

DISTRIBUTION.-~outh Africa. 

HABITAT.-On rocks in open dry areas. 

REMARKS.-This is another species of Pseudoparmelia that bears a close resemblance to Xanthoparmelia. The “chalybeizans” unknown, in fact, was 

previously known only in Xanthoparmelia (Hale, 

1972b). This specimen apparently endemic to the 

arid regions of South Africa. 

Africa is listed in Hale (1972b3343). 

SPECIMENS ExAhrlNED.-Material from the Union of South 

Pseudoparmelia conlabrosa, new species 

FIGURE 10a 

DEscRIPTIoN.-Thallus arcte adnatus, corticola, 

viridi-albidus, 3-5 cm latus, lobis sublinearibus, 

apice plus minusve subrotundatis, congestis, 2-2.5 

mm latis; superne undulatus, nitidus, apice alboreticulatus, dense isidiatus, isidiis cylindricis, pro 

maxima parte simplicibus, usque ad 0.5 mm altis; 

cortex superior 10-12 pm crassus, epicorticatus, epicortice perforato, stratum gonidiale ca 10 pm 

crassum, medulla alba, ca 100 pm crassa, cortex inferior paraplectenchymatus, 8-10 pm crassus; subtus 

niger, modice rhizinosus, rhizinis nigris, simplicibus. Apothecia ignota. 

C +, KC + red, P - ; atranorin and lecanoric acid. 

HoLoTYPE.-Australia: New South Wales, between Majors Creek and Araluen, Weber and 

McVean L-47102, 18 October 1967 (US; COLO, 

isotype). 

REMARKs.-This species is obviously related to 

Pseudoparmelia labrosa (Zahlbruckner) Hale, a 

sorediate species with lecanoric acid also known 

from Australia. No parent morph has yet been discovered for these two species. 

CHEMISTRY.-cOrteX K + yellow, medulla K -, 

Pseudoparmelia corrugativa, new combination 

Parmelia corrugativa Kurokawa and Filson, 1975:38 [type 

collection: Near Bulhunnah, South Australia, Rogers 553 

(MEL. holotype) (not seen)]. 
26 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 10.-Species of Pseudoparmeliu: a, P. conlabrosa (Weber and McVean L-47102, holotype in US); b, P. croralsiataa (Culbersota 63S6 in US); c, P. cryptochlorophaea (Hale 19771); 

d, P. cyphellata (Maline 2532B, isolectotype in US); e, P. dahtii (Dahl, isotype in US); f, P. 

ecaperata (Tsuyama 5 in US). (Scale in min.) 
NUMBER 31 27 

DESCRIPTIoN.-Thallus closely adnate on twigs, 

whitish mineral gray, about 6 cm broad; lobes 

irregularly elongate, apically rotund, 1.5-3 mm 

wide; upper surface rugulose; medulla white in the 

upper part and deep yellow in the lower half; lower 

surface black, sparsely rhizinate. Apothecia numerous, 1-4 mm in diameter; spores 8, 7 X 10 pm. 

CHEMIsTRY.-Cortex K + yellow, medulla negative with color reagents except for the pigment 

which is K+ purple, atranorin, and unidentified 

substances. 

DISTRIBUTION.-~outh Australia. 

HABITAT.-on twigs of tree in open area. 

REMARKs.-ThiS species is very similar to P. 

subtiliacea (Nylander) Hale, which differs in having a white medulla without pigments but with 

fatty acids. One additional specimen, which I have 

not seen, was collected by Rogers (95) in ParaWirra national Park, South Australia. The species 

is well illustrated in Kurokawa and Filson (1975) 

(PI. 1: fig. 3). 

Pseudoparrnelia crozalsiana 

FIGURE 10b 

Pseudoparmelia crozalsiana (Bouly de Lesdain) Hale, 1974: 

189. 

Parmelia crozalsiana Bouly de Lesdain ex Harmand, 1909: 

555 [type collection: Agde, Herault, France, De Crozals, 

May 1909 (US, lectotype)]. 

DESCRIPTIoN.-Thalhs adnate to bark, greenish 

mineral gray, 5-10 cm broad; lobes subirregular, 3-6 

mm wide; upper surface strongly reticulately ridged 

and wrinkled, sometimes white-pruinose, sorediate, 

the soralia often produced along the ridges, coalescing; lower surface black, moderately rhizinate 

except for a narrow brown marginal zone. Apothecia not seen. 

yellow, C-, KC-, Pf pale orange, atranorin, 

stictic acid, constictic acid, and associated unknowns. 

DIsTRIBuTroN.-Eastern United States, Mexico, 

Brazil, Uruguay, Argentina, France, Italy, Zaire, 

Union of South Africa, and India. 

HABITAT.-on trees in open secondary forests 

at 100-2000 m elevation. 

REMARKS.-I first identified this species in North 

America in 1960 (Hale, 1960) where it had been 

misidentified as Parmelina aurulenta (Tuckerman) 

CHEMISTRY .-cortex K + yellow, medulla K $. 

Hale, a ciliate species. Since then it has been recognized from many localities. The lobes are usually 

quite broad and apically subrotund and the reticulate wrinkles very distinct without magnification. 

Once the chemistry is recognized, it could only be 

confused with Parmelia carneopruinata (see below), which has lobes less than half as wide on 

the average. There seems to be no nonsorediate 

progenitor extant, unless we consider it to be the 

rare African species P. inhaminensis (Dodge) Hale. 

When I first examined the surface of this species 

with the scanning-electron microscope (Hale, 

1973a), I noted a strongly nodular surface without 

pores, an anamolous condition for the genus. However, examination of other species has shown that 

pores do occur rarely but that even though the 

species does not seem to have a paraplectenchymatus upper cortex, as one would expect in a 

nonepicorticate Parmelia, the cortical structure of 

this species is rather atypical. 

SPECIMENS EXAnfINED.-United States: Illinois, Hale 13980, 

Skorepa 1757 (US); Indiana, Hale 14097; Kentucky, Hale 

13691; Ohio, Hale 13576; Virginia, Hale 15232, 15761, 18455; 

Arkansas, Keck 1216 (US): Alabama, Hale 33829. Mexico: 

Mexico, Hinton 7724 bis (BM); Veracruz, Hale 19494. Venezuela: Merida, Hale 42288, 42312, 43049a: Tachira, Hale 

45704. Brazil: Mato Grosso, Malme 2243C (S); Parana, Montes 

10121 (DUKE); Pernambuco, Xavier 700 (US). Uruguay: 

Lavalleja, Osorio 3667 (DUKE); Maldonado, Osorio 4879 

(US): Rivera, Osorio 1075 (US). Argentina: Tucumain, Venturi 336 (DAR). Europe: Italy, Davies (BM), Sbarbaro (US), 

Sbarbaro in Lichenotheca Pama 32 (M), Sbarbaro in Lichenes 

Selecti Exsiccati 96 (H, LD, US). Uganda: Mawakota County, 

Swinscow 2U 41/2 (BM, US). Zaire: Louis 8150 (BR). Union 

of South Africa: Natal, Almborn 8636 (LD), Hoeg (TRH). 

India: Tamil Nadu, Awasthi 4276 (Awasthi herbarium), Hale 

43976. 

Pseudoparmelia c yptochlorophaea 

FIGURE 10c 

Pseudoparinelia cryptochlorophaea (Hale) Hale, 1974: 189. 

Parinelia cryptochlorophaea Hale, 1959: 18 [type collection: 

Ciudad Trujillo, Dominican Republic, Allard 15715a (US, 

holotype)]. 

DESCRIPTION.-Thal~Us adnate to appressed on 

large branches, 5-10 cm in diameter, light mineral 

gray but soon turning buff in the herbarium; lobes 

3-5 mm wide, apically subrotund, crowded; upper 

surface rugulose and shiny, more or less reticulate 

white-madate at the tips, becoming cracked in 

older parts; margins sorediate, the soredia in dense 
28 ShlITHSONIAN CONTRIBUTIOSS TO BOT.4NY 

erect soralia, 0.6-1.0 mm thick, 1-1.5 mm high: 

lower surface black and rhizinate at the center, 

brown and naked to papillose in a narrow zone at 

the margins. Apothecia rare, 1-3 mm in diameter, 

spores 8, 4-5 X 6-9 pm. 

C-, KCS. rose, P-; atranorin, caperatic acid (C. 

Culberson, 1965) , and cryptochlorophaeic acid. 

DISTRIBUTION.-SOUtherrI United States, ~~exico, 

Central America, Tt7est Indies, Venezuela, and 

Brazil. 

HABITAT.-OII trees (cashew, Xylopia, bamboo, 

etc.) in mature forests at sea level to 500 m elevation. 

RE&fARKS.-The most unusual feature of this 

New Tliorld species is the chemistry, the only reported occurrence of cryptochlorophaeic acid in the 

Parmeliaceae. The capitate soralia are also unique. 

The white reticulation is similar to that in the 

Parinelia texana group, with which it is obviously 

closely allied. 

CHEMISTRY.-cOrteX Kf yellow, medulla K-, 

SPECIXIESS ExAarIxED.-United States: Georgia, Hale 16797, 

21985; Florida, Hale 16989, 17584, 17609, 21674, 21815, 21839, 

Rapp (US) (for furthcr records from Florida in DUKE see 

Moore, 19683225). Slexico: \’eracruz, Hale 19771, 19811. Honduras: Comayagua, Standley 5462, 5930 (F). Jamaica: Imshaug 

15967, 15969 (US), Plitt s.n. (US). Dominican Republic: La 

Vega, Allard 16833, 16845, 16834, 16861, 18028; Santiago, 

Znishaug 23884 (XISC, US); Santo Domingo, Allard 15715, 

16182, 16187, 16191a (US). Trinidad: Degelius s.n. (Degelius 

herbarium); Lassen (C); Lewis 347 (PH). Venezuela: Barinas, 

Steyerniark and Rube 96533 (US); Mkrida, Hale 42305a. 

Brazil: Ceara, Cutler 8072c (F); Rio de Janeiro, Glaziou 1848 

(M). 

Pseudoparmelia cyphellata 

FICCRE 10d 

Pseudopnrmelm c)phel[ata Llnge, 1914.15 [type collection. 

Santa Anna da Chapada, Mato Grosso, Brazil, 21\lnln2e 

2532B (S, 1ectot)pe; LD, UPS, CS, isolectot)pes)] 

Parnzelra cTphellata (Lynge) Santesson, 1942.473. 

DEsCRIPTIoN.-Thallus as in P. sphaerospora (see 

below) except the upper surface isidiate, isidia 

cylindrical, rather sparse, up to 1 mm tall. Apothecia abundant, adnate to substipitate, 2-3 mm 

in diameter; spores 8, about 5 X 7 pm. 

lowish to orange with color reagents; atranorin, 

stictic acid, and an unidentified pigment. 

CHEhlISTRY.-cOrteX K f yellowish, medulla yelDIsTRIsuTIoN.-Brazil. 

HABITAT.-On trunks of trees in forest. 

REMARKs.-This species is clearly the isidiate 

morph of P. sphaeyospora (Nylander) Hale. Unfortunately it is still known only from the type 

locality. 

Pseudoparmelia dahlii, new species 

FIGURE 10e 

Thallus adnatus, corticola, viridi-flavicans, 3-4 

cm latus, lobis sublinearibus, contiguis, 1.5-2.0 

mm latis; superne convexus, opacus, dense isidiatus, 

isidiis cylindricis, simplicibus, usque ad 0.5 mm altis; 

cortex superior 12-14 pm crassus, epicorticatus; 

stratum gonidiale 15-20 pm crassum; medulla alba, 

140-160 pm crassa; cortex inferior paraplectenchymatus, 12-14 p~i crassus; subtus pallide castaneus, 

dense rhizinosus, rhizinis pallidis, simplicibus, elongatis. Apothecia adnata, 1 mm diametro: hymenium 

40-45 pm altum; sporae S:nae, 6 X 7-9 pm. 

KC+ red, P-, usnic acid, caperatic acid and lecanoric acid. 

HoLoTYPE.-sri Lanka: Polonnaruwa, E. Dahl, 6 

January 19‘72 (0; US isotype). 

DISTRIBUTION.-.!%”i Lanka. 

HABITAT.-On trees in lowland forest. 

RE~rARKs.-Superficially this lichen resembles P. 

malaccensis (Nylander) Hale yery closely, but the 

rhizines are denser and longer, the lobes thicker 

and dull without alboreticulate patterning, and 

the chemical components different. It is, however, 

clearly a member of the P. intertexta-P. malaccensis 

lowland rain forest complex. Another closely related 

species, P. rakengensis (Vainio) Hale, which occurs 

in the higher elevation monsoon forest in Thailand, 

is also externally very close. Chemical tests are 

needed to identify the species. I wish to thank Dr. 

Hildur Krog for allowing me to see the specimens 

and to describe them. 

CHEhfISTRY.-cOrteX K -, medulla K-, c +, 

SPECI\fENS EXA\rlhED.-sri Lanka: Dambulla, DahZ (0, us). 

Pseudoparmelia ecaperata 

FIGURE lOf 

Pseudoparrnelia ecapernta (Muller Argoviensis) Hale, 1974: 

190. 

Parrnelia acaperata Muller Argoviensis, 1891:378 [type collection: Shire River, “Zambesica,” Africa, Kirk (G, lectotype; BM, isolectotype)]. 
NUMBER 31 29 

Parmelia nialaccensis var. laetepavens Vainio, 1921:38 [typecollection: Doi Sutep, Thailand, Hosseus, 1904 (TUR, 

Vainio herbarium number 2764, lectotype)]. 

Parmelia laetepavens (Vainio) Gyelnik, 1938a:32. 

Parmelia djalonensis des Abbayes, 1951:966 [type collection: 

Fouta-Djalon, Dalaba, Guinea, des Abbayes (REN, lectotype)] 

DEScRIPTIoN.-Tha1Ius as in P. concrescens (see 

above) except yellowish green. Apothecia rare, 1-4 

mm in diameter, the amphithecium isidiate; spores 

8, 6 X 11-14 .pm. 

KC+ faint wine red, P-; atranorin, divaricatic 

acid, and usnic acid, rarely with protolichesterinic 

acid. 

DISTRIBUTION.-Africa, Nepal, India, and Thailand. 

HABITAT.-On trees or more rarely rocks in open 

forest up to 2000 m elevation. 

REMARKs.-This species represents the isidiate 

morph of P. zambiensis (Hale) Hale. It also represents the usnic acid-containing counterpart of P. 

concrescens, a species which does not occur outside 

of Africa. It has no relationship to P. caperata, as 

the name might be construed to imply, which contains protocetraric acid. 

SPECIMENS EXAI\lINED.-IVOry Coast: Seguela, Santesson 

10698a, 10702b (UPS, US). Guinea: N’Zerekore, Santesson 

10568a (UPS, US). Zaire: Hoeg (TRH), Schmitt 3048 (BR). 

Angola: Bie, Degelius (Degelius herbarium); Cuanza Sul, 

Degelius (Degelius herbarium); Huila, Degelius (Degelius 

herbarium); Moxico, Degelius (Degelius herbarium). Rhodesia: Angus (BM), Hoeg (TRH), Kofler (LD), Schutte 61b 

(LD, US). hfalawi: Jellicoe 55 (BM). Nepal: Poelt 107, 114 

(M, US), Noordyte (L), Togashi (TNS). India: Tamil Nadu, 

Awasthi 4431, 4432 (Awasthi herbarium, US), Degelius As256 (Degelius herbarium), Foreau 54 (Awasthi herbarium, 

US), Hale 43673, 43713, 43866, 43948; Uttar Pradesh, Awasthi 

3496, 3972 (Awasthi herbarium). Thailand: Kurokawa 1670, 

1954 (TNS, US), Tsuyama (TNS, US). 

CHEMISTRY.-cOrteX K--, medulla K--, c-, 

Pseudoparmelia epileuca 

FIGURE Ila 

Pseudoparmelia epileuca (Hale) Hale, 1974:190. 

Parmelia epileuca Hale, 1972:343 [type collection: District 

Inhambane, MoGambique, Schelpe 4461 (BOL, holotype; 

isotypes in LD, US)]. 

DEscRIPTIoNs.-Thallus closely adnate on bark, 

ashy white, 3-5 cm broad; lobes sublinear, contiguous, with smooth dark rimmed margins, 1.5-2.0 mm 

wide; upper surface plane, sorediate, the soralia 

orbicular, confluent with age; lower surface black, 

moderately rhizinate. Apothecia unknown. 

C-, KC-, P+ red; atranorin and protocetraric 

acid. 

CHEMISTRY .-cortex K 4- yellow, medulla K - , 

DIsTRIBuTI0N.-Kenya and Mocambique. 

HABITAT.-On coconut and other trees in open 

forest or savanna up to 300 m elevation. 

REMARKS.-This rare lichen appears to be the 

sorediate morph of P. schelpei (Hale) Hale, which 

also occurs in Mocambique. Two superficially related sorediate species with protocetraric acid occur 

in the New World: P. alabamensis (Hale and McCullough) Hale, which is saxicolous and has generally narrower lobes, and P. raukiaeri (Vainio) 

Hale, which has irregular pustular soredia. 

SPECIMENS ExAxfINED.-Kenya: Coast Province, Santesson 

20898 (UPS, US), Mocambique: Sul do Save, Schelpe 4461a 

(BOL, US), 4460a, 4461b (BOL). 

Pseudoparmelia eruptens 

FIGURE llb 

Pseudoparmelia eruptens (Kurokawa) Hale, 1974: 190. 

Parmelia eruptens Kurokawa in Hale and Kurokawa, 1964: 

153 [type collection: Lydenburg, Transiaal, Union of South 

Africa, Alm born 7498 (LD, holotype; US, isotype)]. 

DESCRIPTION.-Tha11us adnate, corticolous, whitish mineral gray to buff in the herbarium, 5-8 cm 

broad; lobes subirregular, rotund, 2-8 mm wide; 

upper surface plane, continuous, moderately isidiatepustulate, the isidia irregularly inflated, basally 

constricted, bursting apically; lower surface black, 

sparsely rhizinate except for a narrow naked zone 

at the tips. Apothecia rare, adnate, 1-3 mm in 

diameter, the amphithecium coarsely isidiate; spores 

8, 5-7 X 19-12 pm. 

C-, KC- or KC+ purple violet, P-; atranorin 

and divaricatic acid with associated unknowns. 

DIsTRIBUTIoN.-Mocambique and Union of South 

Africa. 

HABITAT.-on trees (and rocks?) in open forest. 

REMARKS.-This rare species is probably most 

closely related to sorediate P. texana (Tuckerman) 

Hale. The large pustules of P. eruptens do not become sorediate. 

CHEMISTRY.-COrteX K+ yellow, medulla K-, 

SPECIMENS ExAM1NED.-Mocambique: Mitchell 332 (us). 

Union pf South Africa: Transvaal, hlaas Geesteranus 6453 

(L, US), 6455 (L). 
SMITHSONIAN CONTRIBUTIONS TO BOTANT 

* I a'"- 

FIGURE 11.-Species of Pseudoparvielin: a, P. epileuca (Schelpe 4461b, isotype in BOL); b, P. 

eruptens (Almborn i498, holotype in LD); c, P. exoriiata (Lamb 1101 in US); d, P. ferax (Rogers 

1326 in US); e, P. geesterani (Maas Geesteranus 6405, holotype in L); f, P. gerlachei (Santesson 

6499 in US). (Scale in mm.) 
NUMBER 31 31 

Pseudoparntelia exornata, new combination 

FIGURE llc 

Parmelia caperata var. exornata Zahlhruckner, 1912:379 [type 

collection: Cerillos Canelones, Uruguay, Felippone 431 (W, 

lectotype; G, isolectotype)]. 

Parmelia rutidota f. filizans Lynge, 1914:153 [type collection: 

Quinta, Rio Grande do Sul, Brazil, Malme 727 (S, lectotype)]. 

DEsCRIPTIoN.-Thallus adnate to closely adnate 

on bark, yellowish green, 4-10 cm broad; lobes subirregular, contiguous, apically subrotund, 2-4 mm 

wide, sometimes marginally dissected, becoming 

lobulate and crowded toward the center, with numerous pycnidia; lower surface black and sparsely 

rhizinate but with a rugose bare brown zone at the 

margin near the tips. Apothecia common, adnate 

to substipitate, 2-5 mm in diameter, the rim crenate; spores 8, 8-10 X 15-18 pm. 

KC- or KC+ rose, P+ orange; usnic acid, protocetraric acid, and the “conformata” unknown. 

DIsTRIBUTIoN.-Southeastern Brazil and Uruguay. 

HABITAT.-& trees in open forest at 300-600 m 

elevation. 

REMARKs.-This species is closely related to P. 

rutidota in chemistry and morphology. It produces 

an unidentified substance, the “conformata” unknown, just below protocetraric acid in the hexane 

solvent, the same chemistry known for its presump 

tive isidiate morph, the saxicolous P. papillosa. The 

lobes tend to be thinner and more filiform and 

finely lacinate than typical P. rutidota. It occupies 

a restricted range in Brazil and Uruguay, whereas 

P. rutidota occurs sporadically in the Andean chain 

and in northern Brazil, as well as in North America 

and Australia. 

CHEMISTRY .-cortex K - , medulla K - , c - , 

SPECIMENS EXAhfINED.-BraZil: Rio Grande do Sul, Malme 

715 (S). Uruguay: Lavalleja, Lamb 1101 (FH, US); San Jok, 

Osorio 877, 6154, 6195 (MVM); Tacuarembo, OsOriO 1106, 

1165 (MVM); Treinta y tres, Osorio 5979 (MVM). 

Pseudoparmelia ferax 

FIGURE lld 

Pseudoparmelia ferax (Muller Argoviensis) Hale, 1974: 190. 

Parmelia ferax Muller Argoviensis, 1886:257 [type collection: 

Australia: New South Wales, Gutawang, Hamilton 2 (G, 

lectotype)]. 

Parmelia obversa Stirton, 1899:76 [type collection: Australia, 

Paton (GLAM, lectotype; BM, isolectotype)]. 

Parmelia citrinescens Gyelnik, 1938b:271 [type collection: 

Lago Nahuel, Puerto Blest, Patagonia, Argentina, Duste 

163 (BP, holotype; S, US, isotypes)]. 

DEScRIPTIoN.-Thallus closely adnate on bark, 

greenish yellow, 5-8 cm broad; lobes subirregular, 

crowded, apically rotund, 3-4 mm wide; upper surface soon wrinkled and rugose, in part warty and 

lobulate; lower surface black and coarsely rhizinate 

except for a narrow brown zone at the margins. 

Apothecia common, sessile, plane to almost urceolate, 2-4 mm in diameter; spores 8, 7-8 X 13-16 pm. 

CHEMISTRY.-cOrteX K + yellowish, medulla K -, 

C-, KC-, P+ red; atranorin, usnic acid, and 

physodalic acid. 

DIsTRIBUTIoN.-Australia and Chile. 

HABITAT.-on branches and trunks of shrubs and 

trees in arid habitats up to 1200 m elevation. 

REiMARKs.-This species was almost always identified as “Parmelia rutidota” until the chemistry 

was clarified by Kurokawa (1967). Pseudoparmelia 

rutidota contains protocetraric acid and generally 

has a more expanded, thallus. While described from 

Australia, P. ferax seems to be most common in 

Chile. 

SPECIMENS ExAh%INEo.-Australia: South Australia, Rogers 

1320, 1326 (US); Victoria, Filsm 6597 (US). Chile: Aconcagua, 

Follrnann 11784-L (US): Nuhle, Mahu 3538 (US); Santiago, 

Mahu 1117, 2034, 3334 (US), Santesson 7124 (S, US); Valparaiso, Imshaug 36657 (MSC), Rundel 7335 (US). 

Pseudoparmelia geesterani 

FIGURE 11 e 

Pseudoparmelia geesterani (Hale) Hale, 1974: 190. 

Parmelia geesterani Hale, 1972h:344 [type collection: Transvaal, Union of South Africa, Maas Geesteranus 6405 (L, 

holotype; LD, US, isotypes)]. 

DESCRIPTIoN.-Tha11us closely adnate on rock, 

dark olive greenish to whitish, 2-4 cm broad; lobes 

sublinear, crowded, 0.5-1 .O mm wide; upper surface 

plane, isidiate, isidia coarse and irregularly inflated; 

lower surface brown or blackening, sparsely rhizinate. Apothecia unknown. 

yellow turning red, C -, KC -, P + orange; atranorin and salazinic acid. 

CHEMISTRY.-cOrteX KS yellow, medulla Kf 

DISTRIBUTION.-union of South Africa. 

HABITAT.-On exposed rocks at 1500-1800 m eleREMARKS.-This lichen is still known only from 

vation. 
32 S;ZIITHSOSIAS CONTRIBUTIONS TO BOTANY 

the type collection. It is not closely related to any 

other Psezidoparmeliae although it might be confused with isidiate-pustulate P. owariensis (4sahina) 

Hale or P. imperfecta (Kurokawa) Hale. 

Pseudoparmelia gerlachei 

FIGURE llf 

Parinelia gerlachei Zahlbruckner, 1929:137 [type collection: 

based on Painzelia antarctica Vainio]. 

Partnelia antarctica Vainio, 1903: 13 [t)pe collection: Cap van 

Beneden, Terre de Danco, Gerlaclze 226 (TUR, Vainio 

herbarium number 2839, lectotype); not P. antarctica Bitter, 1901:248 (= Hypogymnia)]. 

Xanthofiarrnelia gerlachei (ZahlbrucLner) Hale, 1974:487. 

DESCRIPTION.-Tha11us closely adnate on rock, 

rather coriaceous, pale greenish yellow, 3-5 cm 

broad; lobes subirregular, crowded, apically rotund, 

2-4 mm wide; upper surface dull, sometimes lightly 

pruinose, broadly rugose or foveolate, appearing 

somewhat inflated, soredia developing in orbicular 

to irregular laminal, capitate soralia 2-4 mm in 

diameter; lower surface black and moderately 

rhizinate, papillate in a narrow brown marginal 

zone. Apothecia not found. 

CHEMISTRY.-cOrteX K - , medulla K -, c - , 

KC-, P+ red; usnic acid and either physodalic 

acid or protocetraric acid or a mixture of both. 

DIsTRIBuTIoN.-Andean mountain chain and 

Antarctica. 

HABITAT.-on rocks in exposed paramo or tundra 

at 2900-3800 m in the Andes and to sea level in 

Antarctica. 

REMARKS.-I had previously considered this to be 

a Xanthoparnzelia because of the saxicolous habitat 

and presence of usnic acid. The apically rotund lobe 

configuration and penicillate rhizines and papillae 

at the margin, as well as physodalic acid, are more 

characteristic of Pseudoparmelia, as suggested by 

Kurokawa (1967). Pseudoparmelia gerlachei has 

evolved in extremely harsh environments where 

trees are completely absent. The most closely related 

species and the probable progenitor is P. ferax 

(Miiller Argoviensis) Hale, which occurs on trees in 

Australia and Chile, Both species contain physodalic 

acid in the “typical” state, but P. gerlachei in Venezuela contains only protocetraric acid whereas 

Chilean and Argentinian specimens produce both 

acids. This combination is similar to that in Hypotrachyna physodalica (Hale) Hale (Hale, 1975a). 

SPECIMENS EXAa~txED.-VeneZUela: Merida, Hale 42706, 

42843, 44667, 44676. Chile: Magallanes, Santesson 1924 (S, US). 

Argentina: Santa Cruz, Santesson 7079 (S, US). Antarctica: 

Graham Land, Lamb 2644 (FH, US); Cap Fan Beneden, Gerlache (TUR, Vainio herbarium numbers 2840, 2841, syntypes 

of P. antarctica). 

Pseudoparmelia hypomilta 

FIGURE 12a 

Pseudoparmelia hypomilta (Fee) Hale, 1974: 190. 

Parmelia hypomilta Fee, 1837:123 [type collection: Peru (G, 

lectotype; H, isolectotype)]. 

Parmelia regnellii Lynge, 1914:40 [type collection: Sao Joao

d’el Rey, Minas Gerais, Brazil, Malme 308 (S, lectotype; 

BhI, MICH, isolectotypes)]. 

Parmelia regnellii f. arida Lpge, 1914: 141 [type collection: 

Bocca da Serra, hiato Grosso, Brazil, Malme 2240 (S, lectotype; UPS, US, isolectotypes)]. 

DESCRIPTION.-Thallus closely adnate on bark, 

rather coriaceous, buff to straw colored, 1-3 cm 

broad; lobes sublinear 1.0-1.5 mm wide, often black 

rimmed; upper surface plane to’ convex, densely 

white maculate; medulla white in upper half, dull 

red in the lower half; lower surface brown, moderately rhizinate, the rhizines brown. Apothecia (description from Malme 171) common, adnate, 1-2 

mm in diameter; spores 8, more or less uniseriate, 

5 X6pm. 

CHEMISTRY.-cOrteX K f yellowish, medulla K + , 

Cf, KC+ yellowish (pigment K+ purple), P-; 

atranorin and unidentified substances. 

DIsTRrBuTIoN.-Brazi~. 

HABITAT.-On trees and rocks in open forests. 

~EnlARKs.-~seildopar?ne~ia hypomilta is a variable species in terms of lobe width. Lynge’s f. arida 

has quite narrow lobes (1 mm or less) but the types 

of P. hypomilta and Parmelia regnellii are comparable in size. While it is difficult to generalize on 

a species so poorly represented in herbaria, it seems 

closely related to P. congruens but is readily differentiated by the medullary pigment and, as far as we 

can determine, common occurrence on rock. 

SPECIMENS ExAariNED.-Brazil: hlinas Gerais, Gardner 

(BM), Malme 171 (LD, UC, US), Warming 302 (M); Mato 

Grosso, klaltne 2240 (S,US), Malnze in Lichenes Austroainericani 92 (G, LD, S, UPS). 

Pseudoparmelia inhaminensis 

FIGURE 12b 

Pseudoparrnelia inhaminensis (Dodge) Hale, 1974:190. 

Parmelia inhaininensis Dodge, 1959:130 [type collection: Inhambane, Inhatnine, Angola, Sousa (BM, holotype)]. 
NUMBER 31 

FIGURE 12.-Species of Pseudoparmelia: a, P. hypomilta (Malme 2745B in US); b. P. inhaminensis (Degelius in US); c, P. inornata (Imshaug 24454, isotype in US); d, P. intertexta (Pentype, 

lectotype of P. gracilis Muller Argoviensis in G); e, P. .ischnoides (Almborn 1698, holotype in 

LD); f, P. labrosa (James 577 in US). (Scale in mm.) 

33 
34 SMITHSOXIAIU CONTRIBUTIONS TO BOTANY 

DEsCRIPTIoN.-Thallus adnate on twigs, soft and 

fragile, buff mineral gray, 3-5 cm broad; lobes subirregular, apically rotund, 1.5-2 mm wide; upper 

surface more or less regularly rugose and wrinkled, 

the cortex easily breaking away, heavily pycnidiate; 

lower surface black and moderately rhizinate except 

for a narrow brown, bare zone at the tips. Apothecia 

substipitate, 2 mm in diameter; spores 8, 5-8 X 

10-12 pm. 

yellow, C-, KC-, P+ orange; atranorin, stictic 

acid, and constictic acid. 

REMARKS.-The chemical constituents and rugose 

upper surface place P. inhaminensis close to P. CTOzalsiana (Bouly de Lesdain) Hale, which is sorediate, 

and P. sc.r.obicuZaris (Krempelhuber) Hale, which 

is nonsorediate and has a pruinose apothecial disc, 

a smaller, very rugose thallus, and unusually large 

conidiospores (about 20 pm long). The conidiospores of P. inhaminensis are about 12pm long. It is 

a possible candidate as nonsorediate progenitor of 

P. crozalsiana. 

CHEMISTRY.-cOrteX K + yellow, medulla K $- 

SPECIhrEhS EXAMINED.-.hgOla: Mocamedes, L)egeliUS (Degelius herbarium, US). 

Pseudoparmelia inornata 

FIGURE 12c 

Pseudoparmelia inornata (Hale) Hale, 1974: 190. 

Parrnelia inornata Hale, 197la:32 [type collection: Grand 

Cayman, Imshaug 24454 (MSC, holotype; US, isotype)]. 

DEscRIPTIoN.-Thallus adnate on bark, pale 

greenish mineral gray, 5-10 cm broad; lobes subirregular, apically rotund, 3-7 mm wide; upper 

surface plane, continuous to cracked with age, usually densely pycnidiate, white-reticulate at the lobe 

tips; lower surface black and sparsely rhizinate except for a narrow bare, brown zone at the margins. 

Apothecia common, substipitate, 2-4 mm in diameter; spores 8, 7-8 X 16-18 pm. 

C-, KC+ rose, Pf red; atranorin and protocetraric acid. 

 HEMI IS TRY .-cortex K -k yellow, medulla K - , 

DISTRIBUTION.-\~eSt Indies. 

HABITAT.-oD. trees in shaded woods near sea 

level. 

REhfARKS.-This rather broad-lobed species might 

be considered as a possible progenitor of isidiate P. 

martinicana (Nylander) Hale, but it is more coriaceous and has a different aspect. It appears to be best 

developed in dry sea-level forests of smaller islands. 

SmcnrExs ExA\cmm.-Records from the Bahamas, Grand 

Cayman, and Haiti are listed in Hale (1971332). 

Pseudoparmelia intertexta 

FIGURE 12d 

Pseudoparmelia i7itertexta (Montagne and van den Bosch) 

Hale, 1974:190. 

Parnielia intertexta Montagne and van den Bosch in Montagne, 1896:327 [type collection: Java, Junghuhn (L, lectotype; PC, isolectotype)]. 

Parrnelia ecoronata Nylander, 187354 [type collection: PuloPenang, Malaya, Collingham [Cunningham in publication] 

(H, Nylander herbarium number 32999, lectotype)]. 

Parrnelia subrupta NJlander in Xylander and Crombie, 1883: 

51 [type collection: Allagajah, near Malacca, Malaya, 

hlaingay (BM, lectotype; H, isolectotype)]. 

Parmelia gracilis hliiller Argoviensis, 1887:317 [type collection: Daintree River, Australia, Pentzke (G, lectotype); not 

Parrnelin gracilis (Persoon) Sprengel, 18273277 (= Usnea)]. 

Parrnelia relicina var. ecoronata (Nylander) Muller Argoviensis, 1891:378. 

Parnzelia gracilenta Vainio, 1900:6 [type collection: Based on 

Parrnelia gracilis Muller Argoviensis]. 

DEscRIPTIoN.-Thallus closely adnate, corticolous, 

marguerite yellow, 3-10 cm in diameter; lobes sublinear-elongate, 0.5-2 mm wide; upper surface more 

or less convex, faintly maculate; lower surface pale 

brown to tan, densely rhizinate, the rhizines simple 

to densely branched, pale. Apothecia numerous, 

adnate, 0.7-2 mm in diameter; spores 8, 3-5 X 5-7 

Pm. 

CHEMISTRY.-cOrteX K 4- yellowish, medulla K -, 

C-, KCf rose, Pf orange red; atranorin, protocetraric acid, protolichesterinic acid, and usnic acid. 

DISTRIBuTroN.-Andaman Islands, Thailand, Malaysia, Philippines, Indonesia, New Guinea, and 

Australia. 

HABITAT.-on canopy branches of trees (dipterocarps and Quercus) in rain forest at 150-1600 xn 

elevation. 

REMARKs.-This is one of two Pseudoparmeliae 

(the other being P. malaccensis (Nylander) Hale) 

that have evolved in the Southeast Asian rain forests, primarily on dipterocarps. The only other common parmelioid genus there is Relicina (Hale, 

1975b), which is also characterized by a closely 

adnate habit and presence of usnic acid. Pseudoparmelia intertexta is a presumptive nonisidiate 

progenitor of P. malaccensis (Nylander) Hale (see 
NUMBER 31 35 

discussion under that species), although it is anomalous in having branched rhizines. 

SPECIMENS ExAhzlNED.-Andaman Islands: KUrZ 3 (M, UPS, 

W). Thailand: Chang, Schmidt XVI (TUR). Malaya: Pahang, 

Hale 30184, 30225, 30482, 30498; Selangor, Hale 30073, 30260, 

30261, 30264, 30299, 31 188. Philippines: Mountain Province, 

Hale 25811; Negros Occidental Province, Hale 26422, 26511, 

26523. Malaysia: Sabah, Hale 28206, 28813, 29065, 30363, 

30364. New Guinea: Versteegh (L, US). 

Pseudoparmelia ischnoides 

FIGURE 12e 

Pseudoparmelia ischnoides (Kurokawa) Hale, 1974:190. 

Parmelia ischnoides Kurokawa in Hale and Kurokawa, 1964: 

155 [type collection: Window Stream, Kirstenbosch, Wynberg, Cape Province, Union of South Africa, Almborn 

1698 (LD, holotype; TNS, US, isotypes)]. 

DEscRIPTIoN.-Thallus closely adnate on rock, 

fragile, whitish ashy gray, 4-10 cm in diameter; 

lobes sublinear-elongate, 0.5-2 mm wide; upper surface plane to convex, continuous, isidiate, the isidia 

short, simple, darkening at the tips; lower surface 

black, sparsely rhizinate. Apothecia adnate, 1-2 mm 

in diameter, the amphithecium isidiate; spores 8, 

5-6 X 7-8 pm. 

yellow, C-, KC-, P+ pale orange; atranorin, stictic acid, and constictic acid. 

CHEMISTRY.-cOrteX K+ yellow, medulla K+ 

DIsTRIBuTIoN.--union of South Africa. 

HABITAT.-on rocks in open areas. 

REMARKs.-This species has a very limited range 

in South Africa and probably represents a typical 

Cape endemic. It bears a superficial resemblance to 

two other small saxicolous species, P. annexa (Kurokawa) Hale (lecanoric acid present) and P. arcana 

(Kurokawa) Hale (fatty acids, pale below). 

for records in the Union of South Africa. 

SPECIMENS EXAhfIXED.-~ee Hale and Kurokawa (1964: 156) 

Pseudoparmelia labrosa 

FIGURE 12f 

Pseudoparnielia labrosa (Zahlbruckner) Hale, 1974: 190. 

Parmelia tenuirirna var. labrosa Zahlbruckner, 1941 :lo8 [type 

collection: Saddle Hill, Dunedin, New Zealand, Thomson 

V34 (W, lectotype)]. 

DESCRIPTIoN.-Tha11us adnate on bark, light buff 

mineral gray, 4-8 cm broad; lobes subirregular, 

Parrnelia labrosa (Zahlbruckner) Hale, 1968:325. 

apically subrotund, 1.5-3 mm wide; upper surface 

plane to rugulose, shiny, sorediate, the soralia originating from coarse pustular ridges, becoming irregular to diffuse; lower surface black and moderately 

rhizinate. Apothecia adnate, 3-5 mm in diameter; 

the rim sorediate; spores 8, 5 X 10-12 pm. 

CHEMISTRY.-cOrteX Kf yellow, medulla K-, 

C + , KC + red, P - ; atranorin and lecanoric acid. 

DIsTRIBuTIoN.-Austra~ia, New Zealand, and 

Chile. 

HABITAT.-On trees (Drachophy llum, Myrsine, 

Hymenanthera, and Betula) in open woods at low 

elevations. 

RElhlARKs.-The lobe configuration places this 

austral species near P. texana (Tuckerman) Hale, 

but the soralia are more diffuse and the chemistry is 

distinct. The parallel isidiate morph is probably 

P. conlabrosa Hale; there seems to be no fertile 

progenitor. The syntype of P. tenuirima var. labrosa 

(Thomson 2A 683 in W) is Parmotrema reticulatum 

(Taylor) Choisy. 

SPECIMENS ExAmNED.-chile: Chiloe, Santesson 2262 (s). 

Other records from Australia and New Zealand are listed in 

Hale (1968:325). 

Pseudoparmelia lecanoracea 

FIGURE 13a 

Pseudoparmelia lecanoracea (Muller Argoviensis) Hale, 1974: 

190. 

Parmelia lecanoracea Muller Argoviensis, 1888:529 [type collection: Arisdrift, Oranje River, Namaqualand, Union of 

South Africa, Schenck 543 (G, lectotype)]. 

DEscRIPTIoN.-Thahs closely adnate, appearing 

areolate at the center, pruinose whitish buff, 2-3 cm 

broad; lobes sublinear, 0.6-1.0 mm wide, black 

rimmed; upper surface convex, rugose with age, 

roughened; medulla pigmented reddish yellow in 

the lower half; lower surface tan or darkening, moderately rhizinate, the rhizines brown. Apothecia 

rare, adnate, 1 rnm in diameter; spores 8, 7 X 8-10 

CHEMISTRY.-cOrteX K+ yellow, medulla K-, 

Cf, KCf rose; atranorin and evernic acid; pigmented medulla K+ purple, skyrin present. 

DIsTRIBuTroN.-~nion of South Africa. 

REMARKS.-ThiS species is still only known from 

the rather fragmentary type collection. As Muller 

noted, it could be confused with a Lecanora. The 

HABITAT.-On rocks. 
36 SMITHSOSIAK CONTRIBUTIONS TO BOTASY 

FIGURE 13.-Species of Pseudoparmelia: a, P. lecanoracea (Schenck 543, lectotype in G); b, 

P. leucoxantha (Eiten 449TA in US); c, P. trialaccensis (Hale 24859); d, P. martinkana (Hale 

35662); e, P. molybdiza (Degelius SA-39 in US); f, P. nnirobiensis (Swinscow 6/19TO n US). 

(Scale in mm.) 
NUMBER 31 37 

chemistry is anomalous for the genus. It is included 

here in the hope that future workers will be in a 

better position to assess its exact relationship. 

Pseudoparmelia leucoxantha 

FIGURE 13b 

Pseudoparmelia leucoxantha (Muller Argoviensis) Hale, 1974: 

190. 

Parmelia leucoxantha Muller Argoviensis, 1881:85 [type 

collection: Brazil, Sao Paulo, Puiggari 1050 (G, lectotype; 

W, isolectotype)]. 

DEscRIPTIoN.-Thallus adnate to loosely attached 

on rock or bark, dull greenish yellow, 3-6 cm broad; 

lobes subirregular, apically rotund, 3-5 mm wide; 

upper surface plane, rimose with age, sorediate 

along the margins and in part on the surface, 

soralia irregular in capitate or elongate masses, the 

soredia coarse; lower surface black and sparsely 

rhizinate except for a brown, bare or papillate zone 

along the margins. Apothecia rare, sessile, up to 2 

mm in diameter, the amphithecium sorediate; 

spores not developed. 

CHEiwsTRY.-cortex K+ yellowish, medulla K -, 

C--, KC- or KC+ rose, P+ red; atranorin (trace), 

usnic acid, and protocetraric acid. 

DIsTRIBuTroN.-Mexico and Brazil. 

HABITAT.-oII sandstone boulders, more rarely 

on trees, in dry scrubby areas (chapada vegetation 

in Brazil) at 300-1 100 m elevation. 

REhfARKS.-There is considerable resemblance to 

Pseudoparmelia caperata (L.) Hale in lobe configuration, but the soralia are more discrete and distinctly marginal and lateral. It may well have 

evolved from nonsorediate P. rutidota (Hooker 

and Taylor) Hale but differs in substratum. 

Brazil: Maranhao, Eiten and Eiten 4497A (US); Mato Grosso, 

Malme (S); Sao Paulo, Schindler 4564b (KR, US). 

SPECIMENS EXAhiIIVED.-Mexico: Oaxaca, Hale 20643, 20648. 

Parmelia malaccensis 

FIGURE 13c 

Pseudoparmelia malaccensis (Nylander) Hale, 1974:190. 

Parmelia malaccensis Nylander in Nylander and Crombie, 

1883:52 [type collection: St. Johns Hill, Malacca, Malaya, 

Maingay 21 (BM, lectotype; FH, H, Nylander herbarium 

number 34984, isolectotypes)]. 

DESCRIPTION.-ThallUS closely adnate on bark, 

yellowish green, 2-8 cm in diameter; lobes short, 

sublinear, 0.5-1.5 mm wide; upper surface plane 

to convex, shiny, distinctly white reticulate at the 

lobe tips, tangentially cracked with age, isidiate, the 

isidia simple, 0.2-0.5 mm high; lower surface pale 

brown, densely rhizinate, the rhizines simple to 

sparsely branched, becoming dark brown. Apothecia rare, adnate, 1-2 mm in diameter; spores 8, 

4-5 X 6-7 pm. 

CHEMIsTRY.-cortex K+ yellowish, medulla K-, 

C-, KC+ rose, Pf red; protocetraric acid and 

usnic acid. 

DISTRIBUTION.-Africa, India, Sri Lanka, Indonesia, Malaysia, and the Philippines. 

HABITAT.-On trunks and canopy branches of 

trees in lowland rain forest from sea level to 150 m 

elevation. 

REMARKS.-while 1 tentatively consider this as 

the isidiate morph of P. intertexta, mostly because 

of the chemical characters and similar habitat in the 

lowland rain forest of Southeast Asia, the two 

species have diverged considerably in rhizine structure. The rhizines of P. malaccensis are simple and 

often turn darker brown that the lower cortex; they 

are pale and rather richly branched in P. intertexta. 

The upper cortex is strongly white-reticulate in P. 

malaccensis, continuous in P. intertexta. 

SPECIMENS ExAhlINED.-IVOry Coast: Santesson 10397a (UPS, 

US). India: Tamil Nadu, Hoeg (Awasthi herbarium, US). 

Sri Lanka: Fosberg 51032 (US). Philippines: Basilan, Hale 

24941, 25331; Zamboanga del Norte, Hale 24727; Zamboanga 

del Sur, Hale 24859. Malaya: Selangor, Hale 30055, 30068, 

30262, 30266, 30284, 30292, 30295, 31187. Sarawak: Hale 29992, 

29994, 29998, 30000. Sabah: Hale 30353, 30362, 30368. Indonesia: Java, Spanjaard 6102 (L). 

Parmelia martinicana 

FIGURE 13d 

Pseudoparmelia martinicana (Nylander) Hale, 1974:190. 

Parmelia martinicana Nylander, 1885:609 [type collection: 

Martinique, Tardin (H, lectotype)]. 

DEScRIPTIoN.-Thallus adnate, 3-9 cm broad, 

pale tan mineral gray; lobes irregular, subrotund, 

short, 2-5 mm wide; upper surface becoming rugose 

toward the center, densely isidiate, isidia initially 

papillate, cylindrical to irregularly thickened, simple or branched, rarely turning granular at the tips; 

lower surface black and sparsely rhizinate at the 

center, rugose, brown, and naked in a narrow zone 

at the margin. Apothecia not seen. 
38 SMITHSONIAN CONTRIBUTIOSS TO BOTANY 

CHEMISTRY.-cOrteX K + yellowish, medulla K-, 

CS, KC+ rose, P+ orange red; atranorin, gyrophoric acid, and protocetraric acid (rarely with 

norlobaridone). 

DrsTRIBuT1oN.-Southeastern United States, Mexico, West Indies, and Venezuela. 

HABsrAT.-On tree trunks and branches in pastures a?d secondary forests from sea level to 1000 

m elevation. 

REMARKS.-ThiS is one on the commonest foliose 

lichens in scrub forest and dry savanna in the West 

Indies. It may be related to nonisidiate P. inornata 

(Hale) Hale but has a thinner, more fragile thallus. It is most closely related to P. raunhiaeri 

(Vainio) Hale, which is distinctly coarsely pustulate-sorediate. One specimen (Allard 17325) contained norlobaridone in addition to protocetraric 

acid, but further collections should be examined 

before deciding whether this population represents 

a distinct species or not. 

SPECIAIE%S ExA\rINED.-United States: Florida, Hale 36813, 

36864. Mexico: Veracruz, Hale 19743. Bahamas: illerrill (US), 

Britton 942, 1080 (FH). Cuba: Pinar del Rio, Inishaug 25289 

(MSC); Guantanamo, Hzoram 2701 (US); Oriente, Hioram 

5477, 5706 (US). Dominican Republic: Allard 17325 (US), 

Wetmore 3922 (MSC), 3947 (MSC, US). Puerto Rico: Britton 

1691 (FH, NY). St. Croix: Britton 75 (NY, US), Boergessen 

(C), Raunkiaer 547 (C), Paulsen (C). St. Martin: Le Gallo 

478b, 571 (US). St. Barthelemy: Le Gallo 415, 497, 515, 521, 

542a, 545, 563 (US). Guadeloupe: Culberson 14536, 14544 

(DUKE), Degelius (Degelius herbarium) Duss (C), 489 (NY), 

Le Gallo 465, 471, 489, 574, 575, 580 (US), 2729 (MSC), 2741 

(BM). Dominica: Imshaug 33159, 33282 (MSC) (other records 

listed in Hale, 1971:18), Martinique: Culberson 14732 (DUKE), 

Degelius (Degelius herbarium), Jardin (H). St. Lucia: Imshaug 

29701, 29968, 30144 (MSC), Evans 75 (FH, NY, YU), 101 

(US). St. Vincent: EIliott (TUR), Itnshaug 30369, 30692 

(MSC). Tobago: Imshaug 31565 (MSC). Trinidad: Broadway 

8099 (BM). Venezuela: Magdalena, firoadway 805 (US). 

Pseudoparmelia molybdiza 

FIGURE 13e 

Pseudoparmelia mol) bdiza (Nylander) Hale, 1974:190. 

Parrtzelia rnolybdiza Nylander in Crombie, 1876a: 19 [type 

collection: Table Mountain, Cape of Good Hope, Union of 

South Africa, Eaton (BM, lectotype; H, Nylander herbarium 

number 35234, isolectotype)]. 

Partnelia atricliotdes Nylander in Crombie, 187613: 167 [type 

collection: Cape of Good Hope, Union of South Africa, 

Eaton (BM, lectotype; H, isolectotype)]. 

Parwelia brachyphylla Muiller Argoviensis, 1886:256 [type 

collection: Near Lydenburg, Transvaal, Union of South 

Africa, Wi[tns 2752 (BM, lectotype). 

Parmelia perfissa Steiner and Zahlbruckner, 1926:519 [type 

collection: Port Elizabeth, Cape Province, Union of South 

Africa., Brutznthaler PV, lectotype; WU, isolectotype)]. 

DESCRIPTION.-Tha~~Us adnate to appressed on 

rock, whitish mineral gray, 4-8 cm broad; lobes subirregular, apically rotund, crowded toward the 

thallus center, 2-3 mm wide; upper surface plane, 

rimose with age; lower surface dark brown and 

blackening, moderately rhizinate. Apothecia numerous, adnate, 1-2.5 mm in diameter; spores 8, 5 X 6 

Pm. 

CmmsTRY.-Cortex K + yellow, medulla K -, 

Ct, KC+ red, P--; atranorin and lecanoric acid. 

DIsTRIBuTIoN.-Uganda and Union of South 

Africa. 

HABITAT.-on rocks in open or partly shaded 

habitats from sea level to about 1200 m elevation. 

REMARKS.-PSeUdo~al-melia moly bdiza is especially common in Cape Province. The isidiate morph 

appears to be P. annexa (Kurokawa) Hale. The 

brilliant Cf red test identifies it immediately. Pseudoparmelia spodochroa (Kurokawa and Filson) 

Hale, P. tortula (Kurokawa) Hale, and even P. 

uanderby Zii (Zahlbruckner) Hale are very similar 

externally and occur in the same localities where 

P. molybdiza is found. They would be differentiated by a negative C test. 

SrEciarEivs ExAMlSED.-uganda: Pian County, Sroinscow 

2U 31/10A (BM, US). Union of South Africa: Natal, Almborn 8645, Hoeg (TRH); Transvaal, Maas Geesteranus (L); 

Basutoland, Kofler (LD); Orange Free State, Ahborn 5818, 

5831, 5834 (LD); Cape Province, Almborn 1803, 2065, 2066, 

2M7, 4837, 4948, 4980, 5690, 11115 (LD), Degelius SA-39 

(US), H6eg (LD, TRH), Maas Geesteranus 6706, 6707, 6730 

(I>). 

Pseudoparmelia nairobiensis 

FIGURE 13f 

Pseudoparmelia nnirobiensis (Steiner and Zahlbruckner) Hale, 

1974: 190. 

Partnelia nairobiensis (“neirobiensis’? Steiner and Zahlbruckner, 1926:517 [type collection: Nairobi, Kenya, Schroder 

287 (W, lectotype)]. 

Parmelia gracifescens var. angolensis Vainio in Welwitsch, 

1901:401 [type collection: Serra da Xella, Huila, Angola, 

Welujitsch 30 pro parte (TUR, Vainio herbarium number 

3059, lectotype)]. 

Parmelia angolensis (Vainio) Dodge, 1959:103. 

Parmelia ganguelIensis Dodge, 1959: 109 [type collection: 

Ganguelas and Ambuelas, Benguela, Angola, Gossweiler 

(BM, lectotype)]. 
NUMBER 31 39 

Parnielia hansfordii Dodge, 1959:127 [type collection: Kampala, Uganda, Hansford 1455 (BM, lectotype)]. 

DEscRiPTIoN.-Thallus loosely adnate to appressed on bark, rather coriaceous, greenish to buff 

mineral gray, 5-10 cm broad; lobes sublinear to subirregular, usually apically subrotund, 2-5 mm wide, 

often becoming marginally lobulate with age; upper 

surface plane, shiny, usually conspicuously pycnidiate, reticulately rimose with age; lower surface 

black except for a narrow marginal brown zone, 

sparsely to moderately rhizinate. Apothecia common, adnate, 2-5 mm in diameter; spores 8, 6-8 X 

8-13 pm. 

C - , KC+ faint violet or KC -, P - ; atranorin and 

divaricatic acid with associated unknowns. 

DISTRIBUTION.-Kenya, Uganda, Angola, Zaire, 

Rhodesia, and Tanzania. 

HABITAT.-on trees and rocks in open or secondary forest at 1000-1700 m elevation. 

REMARKs.-This is one of the more common foliose lichens in central Africa. It appears to be (or 

at least would be similar to) the progenitor for 

isidiate P. concrescens (Vainio) Hale and sorediate 

P. texana (Tuckerman) Hale. 

SPECIMENS EXAMINED.-Uganda: Buddu County, Lye L167 

(BM, US); Burahiya County, Swinscow 2U 17/21 (BM, US). 

Kenya: Central Province, Maas Geesteranus 10272 in Lichenes 

Africani 29 (L, LD, US). Zaire: Degelius (Degelius herbarium), Louis 4857E, 4858 (BR). Tanzania: Kilimanjaro Prov., 

Santesson 20959 (UPS). Angola: BiC, Degelius (Degelius herbarium); Huila, Degelius (Degelius herbarium); Moxico, 

Degelius (Degelius herbarium). Rhodesia: Bankart (BM), 

Bullock 2105 pars, Hoeg (TRH). Tanzania: Proctor 1023 B 

CHEMISTRY.-cOrteX K+ yellow, medulla K--, 

(BM). 

Pseudoparmelia neoquintaria, new species 

FIGURE 14a 

DEscRIPTIoN.-Thalhs laxe adnatus, saxicola, 

cinereo-albidus, 3-7 cm latus, lobis sublinearibus, 

1-2 mm latis, margine laciniatis; superne planus, 

continuus vel aetate rimosus, isidiatus, isidiis cylindricis, simplicibus vel ramosis, usque ad 0.5 mm 

altis; cortex superior 10-15 pm crassus, epicorticatus, 

epicortice perforato, stratum gonidiale 10-1 5 pm 

crassum, medulla alba, ca 200 pm crassa, cortex inferior paraplectenchymatus, 10-12 pm crassus; subtus castaneus, modice rhizinosus, rhizinis elongatis, 

simplicibus, castaneis. Apothecia ignota. 

CHEMISTRY.-cOrteX K + yellow, medulla Kor K+ reddish, C-, KC-, P-; atranorin and the 

“quintaria” unknowns. 

HoLoTYPE.-Australia: New South Wales, Mount 

Wilson, Blue Mountains, G.E and G. Du Rietz 781, 

12 November 1926 (UPS; US, isotype). 

DIsTRIBUTIoN.-Australia. 

HABITAT.-on exposed sandstone rocks. 

REMARKs.-The thallus of this unique Species is 

rather loosely attached with a tendency for the lobes 

to become revolute, a kind of growth form often 

seen with soil-inhabiting Xanthoparmeliae growing 

under harsh conditions. The “quintaria” unknowns 

have been discovered in the brown Parmeliae and 

in Xanthoparmelia quintaria (Hale) Hale, which 

has quite different lobe configuration and a black 

lower surface. 

Pseudoparmelia owariensis 

FIGURE 14b 

Pseudoparmelia owariensis (Asahina) Hale, 1974: 190. 

Parrnelia owariensis Asahina, 1953: 135 [type collection: 

Inuyama, Province Owari, Japan, Asahina (TNS, lectotype)]. 

DEsCRIPTroN.-Thallus closely adnate on rocks, 

whitish mineral gray, 2-5 cm broad; lobes sublinear, 0.5-2 mm wide; upper surface plane, continuous 

or cracked with age, isidiate-pustulate, the isidia 

coarse, short, cylindrical to irregularly inflated, 

bursting open apically; lower surface black, sparsely 

rhizinate. Apothecia unknown. 

C--, KC+ faint purple violet, P--, atranorin, and 

divaricatic acid with associated unknowns. 

DIsTRIBUTioN.-Africa, Thailand, Hong Kong, 

and Japan. 

HABITAT.-On rocks in open areas at lower elevations. 

REMARKS.-Pseudoparmelia owariensis is a member of a saxicolous pustulate complex that also includes P. pustulescens. These two species cannot be 

distinguished except by chemical tests. Many more 

specimens will have to be examined to comprehend 

more fully their relationships. 

CHEMISTRY.-cOrteX Kf yellow, medulla K-, 

SPECIMENS EXANINED .--Ivory Coast: Seguela, Santesson 

10713a (UPS). Uganda: Nyabushozi County, Swinsco7u 2U 

22/2A;22 22/4 (BM, US). Thailand: Kurokawa 1718,1719, 1720, 

1723, 1724, 1814, 1874 (TNS). Hong Kong: Thrower 1189 (BM). 
40 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

FIGURE 14.-Species of Pseudoparrnelia: a, P. neoquintaria (Du Rietz 781, holotype in US); 

0, P. owariensis (Kurokawa 1874 in US); c, P. pachydactyla (Kofler in US); d, P. papillosa 

(Zorron 2334 in US); e, P. prolata (Hoeg, holotype in TRH); f, P. pustulescens (Santesson 

10691b in US). (Scale in mm.) 
NUMBER 31 41 

Pseudoparmelia pachydactyla 

FIGURE 14c 

Pseudoparmelia pachydactyla (Hale) Hale, 1974:191. 

Parmelia caperata var. isidiophora Steiner, 18973215 [type 

collection: Athi Plains, Kenya, Liechtenstein (WU, lectoParmelia pachydact)la Hale, 1972b:345 [type collection: 

based on P. caperata lar. isidiophora Steiner]. 

Parmelia steineri Dodge, 19593125. [t)pe collection: based on 

P. caperata bar. zsidiophora Steiner: not P. steineri Gyelnik, 1938b:289 (= Xantho$arinelia molliuscula (Acharius) 

Hale)]. 

DEsCRIPTIoN.-Thallus closely adnate on rock, 

light greenish yellow, 4-6 cm broad; lobes sublinear, 

crowded toward the center of the thallus, 1-1.5 mm 

wide; upper surface plane, dull, isidiate, the isidia 

scattered, thick, about 0.3 mm wide and to 0.5 mm 

high; lower surface black and velvety to the margin, 

sparsely rhizinate, the rhizines coarse, dull. Apothecia unknown. 

KC- or KC+ rose, Pf red; usnic acid and protocetraric acid. 

type)]. 

CHEMISTRY.-cOrteX K-, medulla K-, c--, 

DIsmrBuTIoN.-Kenya and Rhodesia. 

HABITAT.-oII rocks in semiarid regions at mid 

elevation. 

REMARKS.-The black velvety lower surface is 

identical to that of Pseudoparmelia amplexa (Stirton) Hale, the presumptive parent species. The 

sorediate morph is P. subamplexa Hale. Pseudoparmelia pachydactyla is different from both of these 

species in being saxicolous. 

ler (LD, US). 

SPECllfENS ExAwNED.-Rhodesia: Division Victoria, KofPseudoparmelia papillosa, new combination 

FIGURE 14d 

Parmelia papilkxa Lynge ex Gyelnik, 1939343 [type collection: Canelones, La Paz, Uruguay, F. Felippone 752 (W, 

lectot) pe)]. 

Xanthopartnelia phpillosa (Lynge ex Gyelnik) Hale, 1974: 

488. 

DESCRIPTION.-Thal1us adnate on rock, greenish 

yellow, 4-6 cm broad; lobes subirregular, apically 

rotund, 24 mm wide; upper surface plane and 

shiny, somewhat white-reticulate at the tips, becoming rugose and densely isidiate toward the center, 

the isidia irregularly inflated, to 0.4 mm thick and 

up to 1 mm high, solid but the tips easily eroding 

away, not becoming sorediate; lower surface black 

and moderately rhizinate except for a bare or papillate zone at the tips. Apothecia common, substipitate and often appearing immersed among the 

isidia, 1.5-3 mm in diameter, the amphithecium and 

rim densely isidiate; spores 8, 7-8 X 16-18 pm. 

KC-, or KC+ reddish, P+ red; usnic acid, protocetraric acid, and the “conformata” unknown. 

DIsTRIBuTIoN.-Brazi~, Uruguay, and Argentina. 

HABITAT.-On acidic rocks in open areas from 

sea level to 1000 m elevation. 

REMARKS.--While I had recently transferred this 

species to Xanthoparmelia, partly because of its saxicolous habit, it displays features characteristic of 

Pseudoparmelia: broad, apically rotund lobes, a 

bare or papillate brown zone at the tips below, rhizines with brown, fibrous tips, and presence of protocetraric acid which is rare in Xanthoparmelia, 

especially in New World species. As so delimited, 

P. papillosa is another offshoot from the P. rutidota complex differentiated by the very large almost 

pustular isidia. ,4 domparable saxicolous species containing protocetraric acid, without the “conformata” unknown, P. baltimorensis (Gyelnik and 

F6riss) Hale, has even larger more typically pustular 

outgrowths. 

SITCIMEYS EXAhrINED.-BraZil: Santa Catarina, Reitz and 

Klein 15977, 16120 (US). Uruguay: Lavalleja, Lamb 1107 

(FH, US); Maldonado, 050riO 5026 (MVM, US); Rocha, 

Herter 826a (H, S, US), Zorron 2334 (US): Treinta y Tres, 

Heiter 2 (H). Argentina: Buenos Aires, Eyerdam 2369b (F). 

CHEMISTRY.-COrteX K--, medulla K-, c-, 

Pseudoparmelia prolata 

FIGURE 14e 

Pseudoparmelia prolata (Hale) Hale, 1974:191. 

Parmelia prolata Hale, 1972:344 [type collection: Cape 

Province, Union of South Africa, Hoeg (TRH, holotype; 

LD, US, isotypes)]. 

DEscRrPTIoN.-Thallus adnate to loosely attached 

on rock or rarely on soil, pale olivaceous mineral 

gray, 4-7 cm broad; lobes sublinear, extended, almost divaricate, 2-3 mm wide; upper surface plane, 

shiny; lower surface pale brown, moderately rhizinate, the rhizines pale brown. Apothecia not seen. 

CHEMISTRY.-cOrteX K + yellow, medulla negative with all reagents; atranorin and an unknown 

spot (aliphatic compound?). 
42 SMITHS0NI.W COSTRIBUTIONS TO BOTANY 

DIsTRIBuTIox.-union of South Africa. 

HABITAT.-on rocks, more rarely on loose soil, 

in open areas. 

REMARKs.-The general configuration of this 

species is that of a Xanthoparmelia. In many respects it resembles P. neoqziintah Hale, which 

occurs in similar habitats in Australia but has a 

different chemistry. The isidiate morph is P. basutoensis (Hale) Hale. 

SPECIMEYS Ex.iVIhED.-see Hale (1972b:344) for records 

from the Union of South Africa. 

Pseudoparmelia pustuleseem 

FIGURE 14f 

Pseudoparmelia pustulescens (Kurokawa) Hale, 1974: 191. 

Parrnelia pustulescens Kurokawa in Hale and Kurokawa, 

196.1: 156 [type collection: Vila Flor, Humbo, Angola, 

Degeliiis (Degelius herbarium, holotype; TNS, US, isotypes)]. 

Parmelia iinperfecta Kurokawa in Hale and Kurokawa, 1964: 

155 [tjpe collection: Bcrgville, Natal, Union of South 

Africa, Ahborn 8815 (LD, holotype; US, isotype)]. 

DEscRIPTIoN.-Tha~~us as in P. owmiensis (See 

above). Apothecia adnate, 1-2 mm in diameter, the 

amphithecium pustulate; spores 8, 4-5 X 7-9 ium. 

norin, sekikaic acid, fatty acids, and unknown substances. 

CHEMISTRY:-?VledUlla K - , c -, KC -, P - ; atraDIsTRIBUTIoN.-Africa and India. 

HABITAT.-On rocks in open areas up to 2000 m 

elevation. 

RE&fARKs.-This species is essentially indistinguishable from P. owariensis (Asahina) Hale and 

a chemical test is needed to separate them. They 

appear to be sympatric although collected together only in the Ivory Coast. By using thin-layer 

chromatography I had earlier synonymized P. imperfecla (Hale, 1972b). 

SI’ECI 11 cxs ExA >I Is ED.-I\ 01 y Coast: Seguela, Santesson

10691 b (UPS, US). Guinea: Zerekorce, Santesson 10584. Union 

of South Africa: Natal, Alrnborn 6110 (LD), Hoeg (TRH). 

India: Tamil Nadu, Hale 43674. 

Pseudoparmelia rahengensis 

FIGURE 15, 

Psezidopnriicelia ia/ieiigeiesi$ (Vainio) Hale, 1954:191. 

Pai itcelia ralcei~geiisis Vainio, 1921:39 [type collection: Nong 

Boa, near Raheng, Thailand, Hosseus 320 (TUR, lectotype)]. 

DEsCRIPTIo~..-ThallLis closely adnate on bark or 

rock, yellowish green, 3-6 cm broad; lobes sublinear, 0.5-2 mm wide; upper surface convex, continuous, densely isidiate, the isidia mostly simple, to 

0.2 mm high; lower surface brown to tan, moderately rhizinate, the rhizines tan. Apothecia adnate, 

1-3 mm in diameter, the amphithecium sparsely 

isidiate; spores 8, 3 X 5 pm. 

CHEMISTRY.-cOrteX K f yellowish, medulla K - , 

C-, or C+ yellow, KCf orange, P- or P+ faint; 

barbatic acid, obtusatic acid, 4-0-demethyl-barbatic 

acid, unidentified substance, rarely with an unidentified pigment. 

DIsTRIsUTIoN.-Thailand. 

HABITAT.-on tree bark or rocks in open deciduous forests at 300-1350 m. 

REMARKs.-This species is unique in producing 

barbatic acid. Is is superficially similar to two other 

narrow lobed, usnic acid-containing Asian species, 

P. dahlii Hale (lecanoric acid), and P. malaccensis 

(Nylander) Hale (protocetraric acid). Chemical 

tests should be made to separate them. 

SPECIMENS ExA?rrxm.-Thailand: Kerr L28, (BM, US), Kurokarc*a 1602, 1815 (TSS, US). 

Pseudoparmelia raunkiaeri 

FIGURE l5b 

Pseudoparmelia raunkiaeri (Vainio) Hale, 1974: 191. 

Parrrielia 7aunkiaeri Vainio, 1915:19 [type collection: Cane 

Bay, St. Croix, Raunkiaer 461 (TUR, Vainio herbarium 

number 2552, lectotype; C, FH, isotypes)]. 

Parjnelia scnbrosa Vainio, 1896a:33 [type collection: Chateau 

Belair, St. Vincent, Elliott 249 (TUR, lcctotype; BM, isolectotjpr); not Paririelia scabrosa Ta)lor, 1847:162 (= Xanthopi-i1zelia ~cnbroin (Taylor) Hale)]. 

Pni-?reelin 7~incercfina Zahlbruekner, 1929:222 [type collection: 

based on Parrnelia scabTosa l’ainio]. 

DEscRrPTIoN.-Thallus closely adnate on bark, 

light buff mineral gray, 5-8 cm broad; lobes subirregular, apically rotund, 2-4 mm wide; upper surface plane, becoming irregularly rugose toward the 

center, pustulate-sorediate, the pustules coalescing 

in a coarsely sorediate mass in older portions; lower 

surface black and sparsely rhizinate except for a 

narrow bare, brown zone at the tips. Apothecia unknown. 

C-, KC-, or C+ KCf rose; atranorin and protocetraric acid with or without gyrophoric acid and 

unidentified fatty acids. 

CHEMISTRY.-cOrteX Kf yellow, medulla K-, 
NUMBER 31 43 

FIGURE 15. Species of Pseudoparmelia: a, P. rahengensis (Kurokawa 1602 in US); b, P. raunkiaeri (Evans in US); c, P. rodriguesiana (Santesson 105 60a in US); d, P. rupicola (Reitz and 

Klein 16113 in US); e, P. rutidota (Heller 260 in US); f, P. salacinifera (Moore 4406 in US). 

(Scale in mm except for P. rahengensis which is x3.) 
44 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

DIsTRrBvTIoN.-iZiest Indies and Mexico. 

HABITAT.-on trees in dry, open or secondary 

forest at sea level to 500 m elevation. 

REMARKS.-The soredia of this species are rather 

coarse and originate from small pustulate-isidiate 

growths on the upper surface. These burst open 

and coalesce so that the major part of the thallus 

is sorediate. The relationship to another West Indian species, isidiate P. martinicana (Nylander) 

Hale, is very close. The chemistry is variable. Specimens from St. Croix (including the lectotype) and 

Grand Cayman lack gyrophoric acid. All others contain both protocetraric and gyrophoric acids, as does 

P. martinicana. 

SPECIJIENS Ex~~ll~f~.-hfexico: Tamaulipas, Purse11 5197 

(US). Cuba: Oriente, Hiorant 5545 (US). Grand Cayman: 

1JilShaUg 24480, 24449 (MSC, US), 24508 (MSC). Jamaica: 

Zinsliaug 15000, 15571 (MSC), 19653, 16044 (MSC, US). 

Haiti: Sud, Ij?is/iaug 231 19, 23140 (MSC). Dominican Republic: Santiago, 1Vetinore 3910 (MSC). St. Croix: Britton 77 

(XY, US), Evans (US, YU), Raunkiaer 461 (C). Tortola: Fishlock 493 (FH, NY). St. Barthklemy: Le Gallo 426, 520, 547, 

584 (US), 2626 (MSC). hlonserrat: Evans 46 (US, YU) Guadeloupe: Degelius (Degelius herbarium). 

Pseudoparmelia rodriguesianu 

FIGURE 15c 

Pa?rizelia lodtiguesiana Hue, 1899:167 [type collection: Ambosi tra, Madagascar, Rodriguez (PC, lectot) pes)]. 

DESCRIPTIoN.-Thallus closely adnate on rock, 

whitish mineral gray, 5-10 cm in diameter; lobes 

sublinear to subirregular, 2-3 mm wide; upper surface plane to rugulose, cracked with age; lower surface black, sparsely rhizinate except for a narrow 

naked zone near the tips, Apothecia common, sessile, 2-8 mm in diameter; spores 8, 5-7 X 8-13 pm. 

C-, KC+ faint purple violet or KC-; P-; atranorin and divaricatic acid with associated unknowns. 

DIsTRrBuTro;u.-Guinea, Ivory Coast, Angola, 

Union of South Africa, and Madagascar. 

HABITAT.-on open or exposed rocks at 1000- 

1700 m elevation. 

REMARKS.-This rarely collected lichen would 

seem to be closely related to P. iiairobiensis (Steiner 

and Zahlbruckner) Hale, which is normally corticolous and somewhat larger and more loosely 

adnate. Further field studies will be needed to determine the exact relationship between these two speCHEMISTRY.-cOrteX Kf yellow, medulla K--, 

cies. Pseudoparmelia rupicola (Lynge) Hale in 

South America is also saxicolous and contains divaricatic acid, but it has smaller, closely appressed 

lobes. 

SPECIMENS ExAhmED.-Guinea: N'Zerekore, Santesson 

10560a (UPS, US). Ivory Coast: Man, Santesson 10635 bis 

(UPS, US). Angola: Huambo, Degelius (Degelius herbarium, 

US). Union of South Africa: Natal, Almborn 8594 (US); 

Transvaal, A'ofier (LD). 

Pseudoparmelia rupicola 

FrciR~ 15d 

Pseudopariiielia 1 upicola (L)nge) Hale, 1974:191. 

Parmelza ruprcola LJnge, 1914:132 [type collection: Porto 

Alegre, Rio Grande do Sul, Brazil, Malme 1339 (S, lectot,pe)I. 

DESCRIPTIoiv.-Thal~us closely adnate on rock, 

whitish to ivory mineral gray, 4-8 cm broad; lobes 

sublinear, contiguous, 1-2 mm wide; upper surface 

plane to rugulose, shiny, transversely rimose with 

age, becoming lobulate toward the center; lower 

surface black except for a dark brown zone at the 

tips, moderately rhizinate, the rhizines black. Apothecia common, adnate, 1-1.5 mm in diameter; 

spores 8, 6 X 7-8 pm. 

C-, KC- or KC+ wine colored, P-; atranorin 

and divaricatic acid with associated unknowns. 

CHEMISTRY.-COrteX Kf yellow, medulla K--, 

DIsTRIBUTIoN.-Brazil and Uruguay. 

HABITAT.-oII sandstone outcrops in open areas 

at about 1000 m elevation. 

REMARKs.-Pseudoparmelia rupicola closely resembles sorediate P. alabamensis (Hale and McCullough) Hale in lobe configuration and adnation 

on sandstone, and they may be remotely related. 

Strictly saxicolous species of Pseudoparmelia are 

rare in New World. 

Srrci~r~vs Ex4un~~.-Brazil Catarina, Reitz and Klein 

16111, 16113, 16115, 16121 (US). Uruguay: Lavalleja, Osorio 

6425 (MVM); Rocha, Hosseur (H). 

Pseudoparmelia rutidota 

FIGURE 13, 

Pseudopartrielia rutidota (Hooker and Taylor) Hale, 1974: 

Partnelia i utidota Hooker and Taylor, 1844:645 [type collecParvielia caperata var. caperatula Nylander, 1860:377 [type 

191. 

tion: Van Diemen's Land, Australia (FH, lectotype)]. 
NUMBER 31 45 

collection: Australia (H, Wylander herbarium number 

35731, lectotppe; FH, isolectotype)]. 

Parmelia caperata f. rafnealis Nylander, 1861 :373 [type collection: Andes, Bolivia, Mandon (H, Nylander herbarium 

number 35692, lectotype; FH, S, isolectotypes)]. 

Parmelia jelinekii Krempelhuber, 1868:321 [type collection: 

Australia, Jelinek 27 (M, lectotype; W, isolectotype)]. 

Parmelia ochroleuca Muller Arguviensis, 1882:306 [type collection: Near Illawarra, Australia, Kirton 1 (G, lectotype); 

not Parnaelia ochroleuca Taylor, 1848:24 (= Sticta)]. 

Parmelia splendidula Delise ex Nylander, 18853605 [type 

collection: Peru (a, Nylander herbarium number 35733, 

lectotype)]. 

Parrnelia caperatula (Kylander) Nylander, 1885:606. 

Parmelia subcaperatula Nylander, 1885:606 [type collection: 

Derwent River, Tasmania, Brown (H, Nylander herbarium 

number 35730, lectotype; BM, isolectotype)]. 

Parmelia confertula Stirton, 1899:77 [type collection: Brisbane, Australia, Bailey (BM, lectotype)]. 

DEsCRIPTIoN.-Thahs adnate to appressed on 

bark, light greenish yellow, 3-8 cm broad; lobes subirregular, apically rotund, contiguous, 24 mm 

wide; upper surface plane or becoming rugulose 

and cracked on older lobes; medulla white but 

sometimes with a reddish pigment near the lower 

cortex; lower surface black, shiny, sparsely rhizinate, 

usually with a bare dark brown naked zone at the 

tips. Apothecia common, sessile, the rim crenate, 

1-3 mm in diameter; spores 8, 7-10 X 14-20 pm. 

dulla K--, C-, KC- or KC+ rose, Pf red; usnic 

acid, rarely atranorin, protocetraric acid with or 

without associated unknowns, and with or without 

caperatic acid (and protolichesterinic acid?) and 

unidentified K + purple pigments. 

DIsTRIBuTI0N.-United States, Mexico, South 

America, and Australia. 

HABITAT.-on trunks and branches of trees in 

semiarid regions at 100-2000 m elevation. 

REMARKs.-Pseudoparme~ia rutidota is a rather 

variable species, as one might assume from the long 

list of synonyms. It occurs very commonly in certain 

semiarid regions, especially Texas in the United 

States and New South Wales in Australia. Outside 

of this it is relatively rare. The comparable African 

populations appear to consist entirely of P. amplexa (Stirton) Hale, a smaller, more congested 

species with a black velvety lower surface. On the 

other hand, some specimens from Mexico and South 

America are quite a bit larger than the average. 

Obviously environmental modification plays a role 

in this variation. Basically P. rutidota could be, or 

closely resemble, a now extinct progenitor (or proCHEMISTRY.-cOrteX K- or K+ yellowish, megenitors?) of the sorediate morph P. caperata (L.) 

Hale, the pustulate morph P. baltimorensis (Gyelnik and F6riss) Hale, and the isidiate morph P. 

papillosa (Gyelnik) Hale. None of these fit their 

respective roles perfectly but together they form a 

coherent species group. 

Chemical variation centers around the presence 

or absence of fatty acids, in most cases caperatic 

acid, and a medullary pigment. There seems to be 

no geographic pattern here, but assuredly many 

more field studies and collections are needed to 

determine this. For the present a rather broad species concept seems unavoidable. 

SPECIMENS EXAblINED.-United States: Texas, Darrow 4863 

(US), Hale 5316, 5504, Heller in Lichenes boreali-americani 

197 and Decades of North American Lichens 260 (US), 

Hubricht B1891, B1927, Jerrny (US), Jones (US), Orchard 4 

(US), Slater 3 (US), Whitehouse 2283, 2289, 2290 (US). Mexico: Tamaulipas, Nakanishi 184 (US), Purcell 5564, 5601 (US). 

Bolivia: Mandon (BM). Brazil: Pernambuco, Xavier 776, 

815, 816, 826. Chile: Valparaiso, Follrnann 13013 (US). Uruguay: Lorentz 740 (M); Minas, Heiter 90669 (H). Argentina: 

Lorentz (M). Australia: New South Wales, Degelius A-37, 

A-38 (Degelius herbarium, US), Doing (L, US), Du Rietz 683, 

693 (UPS, US), Hamildon L1812 (XSiV), Watts L1817 (NSW); 

South Australia, Rogers 1463, 1697 (US); Western Australia, 

Irvine (G); Tasmania, Pearcey (BM); Victoria, Filson 5419 

(US); Australian Capital Territory, Weber in Lichenes Exsiccati 268 (US). 

Pseudoparmelia salacini f era 

FIGURE 15f 

Pseudoparmelia salacinifera (Hale) Hale, 1974: 191. 

Parinelia salacinifera Hale in Hale and Kurokawa, 1964:157 

[type collection: Sanford, Seminole County, Florida, Ru@ 

(US, holotype; FLAS, isotppe)]. 

DEsCRrPTroN.-Thallus adnate on bark, light ashy 

buff, 6-12 cm broad; lobes subirregular, apically 

subrotund, 3-5 mm wide; upper surface plane to 

rugulose, fissured with age, moderately isidiate, the 

isidia simple, to 0.3 mm high; lower surface brown 

to tan, moderately rhizinate except for a narrow 

naked zone along the margins. Apothecia rare, 2-4 

mm in diameter, the amphithecium isidiate; spores 

8, 8-9 X 13-16 ,pm. 

yellow turning red, C-, KC-, P+ pale orange; 

atranorin and salazinic acid. 

DIsTRIBUTION.-~Outheastern United States, Mexico, Central America, West Indies, Colombia, Venezuela, Brazil, and Thailand. 

CHEMISTRY.-cOrteX K+ yellow, medulla Kf 
46 SMITHSOSIAN COSTRIBUTIOSS TO BOTANY 

HABITAT.-On trees (deciduous trees, palm, conifers) in open or secondary forest from sea level to 

1000 m elevation. 

REhfARKs.-The distinguishing features of this 

species are the isidia and pale brown lower surface. 

The only other comparable salazinic acid-containing 

species is saxicolous P. scotophylla Kurokawa from 

Australia. It has a smaller thallus and small spores. 

Pseudoparmelia cinerascens (Lynge) Hale has a 

black lower surface. The only other species that 

could be confused with it because of the similar 

habitat, range, thallus color, and lobe configuration 

is P. amazonica (Nylander) Hale, which has a 

black lower surface and contains protocetraric acid 

(K-h 

S[,Ecihims EXAMiNED.--Cnited States: Georgia, Hale 16843; 

Florida, Hale 7994, 16714, 17045, 17083, 17735, 21690, 21967, 

Moore 4406 (DUKE, US). Rapp (FLAS, US), Standiey 13084 

(US). See Maore (1968:225) for further records from Florida. 

Merico: Chiapas, Hale 20607. Honduras: Comayagua, Standley 

6522 (F, US); Cortes, Morton 5896 (US). Cuba: Pinar del Rio, 

Ztnshaug 25215 (MSC, US). Jamaica: Plitt (US). Colombia: 

Santander, A'ee and Mori 3736 (US). Venezuela: Maracay, 

Nasli 1946 (US). Brazil: Maranhao, Eiten 4198 (US); Mato 

Grosso, Afaltiie (UPS). Thailand: Kurokawa 1931 (TNS, US). 

Pseudoparmelia schelpei 

FIGLRE 16a 

Pseudoparttieha schelpei (Hale) Hale, 1974:191. 

Pa? inelia sclfelpei Hale, 1972b:344 [ty pe collection: Maxixe, 

Sul do Sate, hlocambique, Sclielpe 4460 (BOL, holotype; 

LD, US, isotypes)]. 

DESCRIPTIoN.-ThalluS closely adnate on bark, 

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

crowded, 1.5-2.0 mm wide; upper surface plane, 

dull, rugulose with age; lower surface black, 

sparsely rhizinate. Apothecia numerous, adnate, 1-2 

mm in diameter; spores 8, 4 X 8-10 pm. 

C-, K- or KC+ rose, P+ orange; atranorin and 

protocetraric acid. 

CHEMISTRY.-cOrteX K+ yellow, medulla K-, 

DIsTRrBuTroN.-h~ocambique. 

HABITAT.-on coconut palms near sea level. 

REMARKS.--NO other species in Pseudoparmelia 

has protocetraric acid and such narrow lobes. Pseudoparmelia caribaea (Hale) Hale, a saxicolous 

species in the West Indies, is much larger. The 

sorediate morph of P. schelpei is presumed to be 

P. epileuca (Hale) Hale, which also occurs in Mocambique as well as in Kenya. I suspect both species 

will be more frequently collected as lichenologists 

visit coastal localities in East Africa. 

SPECIMENS ExAar!sED.-hfopcbique: Afogg 2099 (PRE, US). 

Pseudoparmelia schistacea, new combination 

Pa, iirelia scliistacea Kurokawa and Filson, 1975:44 [type collection: 65.5 km west of Kingoonya, South Australia, Filson 

11921 (hIEL, holotjpe) (not seen)]. 

DESCRIPTIoN.-ThallUS closely adnate, pale olivaceous gray, 2-4 cm broad; lobes sublinear-elongate, 

crowded at the center of the thallus, 0.5-1.5 mm 

wide; upper surface plane to convex, shiny, becoming tangentially rimose, pustulate; medulla 

white; lower surface pale brown, sparsely rhizinate. 

Apothecia not seen. 

CmMIsrRY.-Cortex K + yellow, medulla K -, 

C-, KC-, P-; atranorin, a trace of usnic acid, 

caperatic acid, and an unidentified fatty acid. 

DISTRIBLIT~~N .-Australia. 

HABITAT.-on rocks in open areas. 

REMARKS.-The authors of this species compare 

it with P. arcana (Kurokawa) Hale, both having 

closely adnate, almost subcrustose thalli. They have 

different morphologies, however, P. arcana being 

typically isidiate. A trace of usnic acid is alleged to 

occur in P. schistacea. The species is illustrated in 

Kurokawa and Filson (1975, pl. 4: fig. 1). 

Pseudoparmelia scotophylla, new combination 

FIGURE 166 

Partrielin scotophl lla Kurokawa in Kurokawa and Filson, 

1955 45 [type collection. Ardglen Gap, Liberpool Range, 

Sew South TVales, Australia, Kuiokauw 5174 (TNS, holot\pe; MEL, isotjpe) (not seen)]. 

DESCRIPTIoN.-Thallus closely adnate, whitish to 

dark mineral gray, 4-12 cm broad; lobes sublinear, 

congested, apically subrotund, 1-2 mm wide; upper 

surface plane, shiny, becoming densely isidiate, the 

isidia cylindrical, simple, the tips blackened; lower 

surface dark brown or blackening, sparsely to moderately rhizinate, the rhizines brown or black, simple. Apothecia (from type description) substipitate, 

6 mm in diameter, the amphithecium isidiate; 

spores 8, 5 X 7-8 pm. 

yellow turning red, C-, KC-, P+ orange; atranorin and salazinic acid. 

CHEMISTRY.-cOrteX Kf yellow, medulla K+ 
NUMBER 31 

FIGURE 16.-Species of Pseudoparmelia: a, P. schelpei (Schelpe 4460, isotype in US); b. P. 

scotophylla (Weber L-47301 in US); c, P. scrobicularis (Malme 1949, isotype of Parmelia 

longiconida Lynge in US); d, P. somaliensis (Jellicoe 40 in US); e, P. soredians (Sampaio 247 

in US); f, P. sphaerospora (Hale 21970A). (Scale in mm.) 

47 
S>llTHSOSI.AN COKTRIBUTIOKS TO BOTANY 

DISTRIBuTIoN.-Austra~ia. 

HABITAT.-on rocks in dry shrubby sclerophyll 

woodlands. 

REMARKs.-This species is part of the Pseudoparmelia molybdiza (Nylander) Hale complex, characterized by isidia and salazinic acid. The color of 

the lower surface seems to be variable, dark brown 

and blackening or remaining rather pale. A closely 

related nonisidiate species and possible progenitor 

is P. spodochroa (Kurokawa and Filson) Hale, 

which usually contains both salazinic and norstictic 

acids. 

SrEcI\rms EXA\fINED.-.~UStralia: Xew South Wales, Du 

Rietz 389 (UPS, US). 

Pseudoparmelia scrobicularis 

FICURE 16c 

Pseudoparrnelia scrobicularis (Krempelhuber) Hale, 1974:191. 

Partnelia scrobicularis Krempelhuber, 1873:lO [type collection: 

Lagoa Santa, Brazil, Warming (M, lectotype)]. 

Partnelia longiconida Lpnge, 1914:130 [type collection: Rio 

Apa, Colonia Risso, Paraguay, Alalme 1949 (S, lectotype: LD, 

US, TV, isolectotypes)]. 

DEscRIPTIoN.-Tha11us closely adnate on bark, 

light buff mineral gray, 2-5 cm broad; lobes sublinear, contiguous, 1 .O-1.5 mm wide: upper surface 

finely foveolate, dull to shiny, lower surface black, 

moderately rhizinate. Apothecia common, adnate, 

1.5-2.5 mm in diameter, the disc white pruinose; 

spores 8,6-7 X 10-12 &m. 

yellow, C-, KC-, Pf orange; atranorin, stictic 

acid, and constictic acid. 

CHEMISTRY.-cOrteX Kf yellow, medulla K+ 

DISTRIBUTION.-sOUth America. 

HABITAT.-On trees in open or secondary forest. 

REMARKs.-The reticulate wrinkles are very 

strongly developed but are not visible without low 

power magnification. Two possibly related species, 

P. carneopruinata (Zahlbruckner) Hale, a presumptive sorediate morph, and P. crozalsiana 

(Bouly de Lesdain) Hale, have broad reticulately 

ridged lobes. 

SmcihiEiw ExAhrrNEo.-Venezuela: Distrito Federal, Ernst 

(G). Brazil: Minas Gerais, Henschen (UPS). Paraguay: Malme 

(UPS); Paraguara, Balansa (G). Argentina: Misiones, Montes 

10089 pro parte (LD). 

Pseudoparmelia somaliensis 

FIGURE 16d 

Pseudoparinelia sonzaliensis (Muller Argoviensis) Hale, 1974: 

191. 

Partrzelia sonalienxis Muller Argoviensis, 18853501 [type collection: Somalia, Hildebrandt (G, lectotype)]. 

Parnzelin scottii Vainio, 1898a:40 [type collection: Ruwenzori, 

Africa, Scott-Elliott (TUR, Vainio herbarium number 2693, 

lectatppe)]. 

Parnzelia rainulicokz Dodge, 1959: 172 [type collection: Madagascar, Hildebrandt (FH, holotype)]. 

DESCRIPTIoN.-Thallus adnate on twigs, whitish 

mineral gray, rather soft and appearing inflated, 

3-5 cm broad; lobes subirregular, 2-5 mm wide; 

upper surface plane to rugulose, dull, becoming 

white-pruinose; lower surface dark brown to black 

and moderately rhizinate except for a brown naked 

zone along the margins. Apothecia common, substipitate, almost urceolate, 2-5 mm in diameter, 

the amphithecium rugose, pruinose; spores 8, 5-7 

X 9-13 pm. 

C-, KC+ rose, P+ red; atranorin and protocetraric acid. 

DISTRIBUTION.-Centra1 Africa and Madagascar. 

HABITAT.-OII twigs and branches of trees in 

open areas up to 2300 m elevation. 

REMARKs-The most characteristic habitat of this 

species is small twigs. The thallus is thick and soft 

with large apothecia. There are no close relatives in 

the genus for this African endemic. 

CHEMISTRY.-cOrteX K + yellow, medulla K -, 

SPECIMENS ExAwxED.-Zambia: Jellicoe 40 (BM, US). Tanzania: Hoeg (TRH). 

Pseudoparmelia soredians 

FIGURE 16e 

Psetcdoparmelia soredians (Nylander) Hale, 1974:191. 

Parinelia soredians Nylander, 1872:426 [type collection: Forqa, 

Reale, Spain, Xylander (H, Kylander herbarium number 

34690, lectotype)]. 

Parmelia conspersa var. polyphylla f. sorediosa Muller Argoviensis, 1891 :378 [type collection: San Miguel, Azores, Godittan (G, lectotype)]. 

Parmelia san-rniguelii Gyelnik, 1931b3288 [type collection: 

based on P. conspersa var. polyphylla f. sorediosa Miiller 

Argoviensis]. 

Parmelia halmaiana Gyelnik, 1935:47 [type collection: Bergues, France, Bouly de Lesdain, in Zahlbruckner Lichenes 

Rariores Exsiccati 236 (BP, holotype; BM, BPI, MICH, 

UPS, isotypes)]. 
NUMBER 31 49 

DEscRIPTIoN.-Thahs adnate on branches and 

trunks of trees, rarely on rocks, yellowish green, 

6-1 5 cm broad; lobes subirregular, apically rotund, 

2-5 mm wide; upper surface plane, shiny, becoming 

rugose toward the center, sorediate, soralia orbicular, coalescing, often occurring along ridges and 

wrinkles; lower surface black except for a marginal 

brown zone, sparsely rhizinate, the rhizines black. 

Apothecia rare, adnate; spores not seen. 

CHEmsmY.-Cortex K - , medulla K + yellow 

turning red, C-, KC-, Pf orange; usnic acid and 

salazinic acid. 

D1sTRIBUTIoN.-Europe, Central and South Africa, Argentina, Chile, and New Zealand. 

HABITAT.-On trees and rarely on rocks in open 

forests. 

REMARKS.-This species is similar to P. caperata 

(L.) Hale except for the discrete, orbicular soralia 

and different chemistry (P. caperata is P+ red 

with protocetraric acid). It is totally absent from 

North America and eastern Asia, where P. caperata 

is very common. In England it behaves as a Mediterranean species, occurring mostly in the southeastern parts of the country where rainfall is low 

and there are no severe frosts (James and Rose, 

1973). 

SPECIMENS ExAiwnim-Europe: Ireland, Mitchell (M); England, Holmes (BM); France, Crozals (US), Dahl (WIS), des 

Abbayes in Lirhenes Armorici Spectabiles 13 (LD), Mangillon 

668 (US), Santesson 10077 (UPS); Portugal, Persoon (UPS), 

Sainpaio in Lichenes de Portugal 247 (LD, M, US), Tauares 

in Lichenes Lusitaniae selecti exsiccati 140 (H, LD, M, US, 

WIS); Italy, Sbarbaro (US), Sbarbaro in Lichenes Selecti Exsiccati 24 (H, LD, M, US). Kenya: Central Province, Maas Geesteranus (L). Union of South Africa: Transvaal, Almborn 6603, 

7501, 7505 (LD), Hiieg (TRH); Natal, Almborn 9641, 9763 

(LD), Hdeg (TRH); Basutoland, h’ofler (LD); Cape Province, 

Almborn 1604, 1997, 1998, 2030, 2099, 2100, 2153, 2121, 4108, 

4735, 5361, 10662, 10862 (LD), Hoeg (TRH), Kofler (LD), 

Maas Geesteranus 6623 (L), Penfield 220 (PRE). Argentina: 

Buenos Aires, Santesson 48 (S); Chubut, Santesson 195 (S). 

Chile: Santiago, Mahu 1117, 2034 (US); Concepci6n, Burros 

3, 193 (H). New Zealand: Scott 323 (BM). 

Pseudoparmelia sphaerospora, new combination 

FIGURE 16f 

Parinelia sphaerospora Nylander, 1859:254 [type collection: 

Parnielia leucochlora Tuckerman in Nylander, 1860:392. 

Parmelia cubensis Nylander, 1885:611. 

Parnielia uleana Miiller Argoviensis, 1889:506. 

Madagascar, (H, lectotype; PC, isolectotype)]. 

Parmelia Pavidoglauca Vainio, 1890:65. 

Parinelia endoxantha Merrill, 1909:73. 

Parinelia bipindensis Dodge, 1959:59. 

Parmelia zenkeri Dodge, 1959:74. 

[Full citations of the synonyms are given in Hale (1971b:10).] 

DEscRIPTroN.-Thahs closely adnate on bark, 

5-10 cm in diameter, pale greenish or yellowish 

mineral gray, often turning chamois in the herbarium; lobes sublinear-elongate, 2-4 mm wide; upper surface plane to convex, more or less maculate, 

rugulose to minutely pitted with age; medulla 

white to pale yellow; lower surface light tan to pale 

olive brown or darkening, rugose, moderately rhizinate, the rhizines simple, tan. Apothecia common, 

adnate, 2-3 mm in diameter; spores €!, 6-8 X 7-9 

Pm. 

dulla yellowish to orange with color reagents; atranorin, stictic acid, norstictic acid, gyrophoric acid, 

“quintaria” unknowns, and an unidentified yellow 

pigment in various combinations. The majority of 

specimens contained atranorin and the pigment; 

the next largest group (about 20% of the specimens) had atranorin, stictic acid, and the pigment. 

The other combinations (atranorin with the “quintaria” unknown, with stictic and norstictic acids, 

or with gyrophoric acid) occurred in only a few 

specimens. 

DISTRIBUTION.-Southeastern United States, Mexico, Central America, West Indies, South America, 

and Central Africa. 

HABITAT.-on shaded tree trunks in mature forests, Taxodium swamps, or orchards from sea level 

to 1100 m elevation. 

REMARKS.-I had previously discussed the occurrence of this species in Dominica under the name 

Parmelia congruens Acharius (Hale, 1971b: 11). 

After making as definitive a summary as possible of 

the chemistry, however, I discovered that the lectotype of P. congruens contains usnic and hypoprotocetraric acids and is better placed in the genus 

Xanthoparmelia, leaving P. sphaerospora as the 

earliest name for the Pseudoparmelia species. The 

medullary color often darkens in old herbarium 

specimens. The isidiate morph is P. cyphellata 

Lynge. 

SPECIMENS ExAMINED.-United States: South Carolina, Culberson 10329 (DUKE, US); Georgia, Egan El-3403, El-6744 

(US), Hale 16832; Alabama, Evans 93, 201, 345, 371 (US); 

Mississippi, McDaniel L-3 (US), Purse11 3083, 3153, 4181 (US); 

CHEMISTRY.-cOrteX K- or K+ yellowish, me- 
50 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Louisiana, Langlois 47 (US); Texas, Jenny (US); Florida, 

Calkins 2 (US), Culberson 10893 (DUKE, US), Hale 16728, 

16889, 17652, 17738, 21623, 34134, 36829, 36887, h’ash 2035 

(US), RaPP 19, 2086 (US). Rapp in Cryptogarnae Vindobonenses 3654 (US), Rapp in Lichenes Rariores Exsiccati 379 

(BM, UPS, US), Standley 73421 (US) (for further records from 

Florida see Moore 1968:224). Mexico: Jalisco, Wirth (US); 

Chiapas, Hale 20030, 20174. Guatemala: Baja Verapaz, Hale 

45998, Xellerrnan 8045 (US). Panama: Cocle, Hale 43571; 

Panama, Hale 38546. Costa Rica: Tonduz (US). Bahamas: 

Brace 6792 (FH, SY). Cuba: W’right 76 (BM, FH, G, XY, 

UPS, US); Pinar del Rio, Imshaug 25349, 25376 (MSC). Haiti: 

Nord, Leonard 8023 (S, US), 9002 (NY). Dominica: Hale 

35636, 35775, 35779. Venezuela: Distrito Federal, Dennis 1548 

(BM); Guarico, Goodland 2 (US), Nilson 1500 (UPS). Peru: 

San Martin, Allard 2294a, 22295a (US). French Guiana: 

Degelius (Degelius herbarium). Brazil: Spruce 130 (BM); 

Maranhao, Eiten 3887 (US); Goias, Iru’in 32073 (NY, US); 

Mato Grosso, Malrne in Lichenes Awtroamericani 96 (BM, 

LD, S, UC, US), 2642 (UC, UPS, US), Rainos 6545 (BM); SPO 

Paulo, Schindler 4586 (US). Cameroon: Zenker 4053 (BR). 

Zaire: Hoieg (TRH), Louis 6765, 7151, 8482 (BR, US). Guinea: 

Santessoti 10574 (UPS, US). Angola: Gossuqeiler 8013a (BM). 

Tanzania: Hoeg (TRH). Madagascar: des Abbayes in Lichenes 

Madagascarienses 5 (US). 

Pseudoparmelia spodochroa, new combination 

FICCRE l7a 

Parrrzelia spodochroa Kurokawa and Filson, 1975:46 [type 

collection: Warren Gorge, Flinders Range, South Australia, 

K. Filsorz 11976 (MEL, holotype; TNS, isotype) (not seen)]. 

DESCRIPTION.-Tha~~us adnate on rock, whitish to 

dark mineral gray, 3-6 cm broad; lobes sublinear, 

apically subrotund, contiguous, becoming lobulate 

with age, 1.5-2.5 mm wide; upper surface plane, 

transversely rimose with age; lower surface brown 

or blackening, moderately rhizinate, the rhizines 

brown or black. Apothecia (from Degelius SA-57) 

adnate, 2-3 mm in diameter; spores 8, 6 X 8-11 

P. 

yellow turning red, C-, KC-, P+ orange; atranorin, salazinic acid, and usually norstictic acid. 

DIsTRIBUTION.-AUStralia and Union of South 

Africa. 

HABITAT.-on more or less exposed rocks in dry 

woodlands. 

REMARKS.-This species is represented by only 

five collections, which diverge rather widely in several characters. The color of the lower surface, for 

example, varies from almost entirely black to pale 

brown. Chemistry is uniform except for Almborn 

4900 and the holotype, which lack norstictic. The 

CHEMISTRY.-cOrteX Kf yellow, medulla K+ 

lobe surface is smooth and shiny except for Degelius 

SA-57, which has a roughened, almost coarsely 

pruinose surface, a trait seen in specimens of other 

genera that grow in exposed desert regions. Further 

collecting in Africa and Australia will be needed to 

decide whether the limits of the species as described 

here are too broad. 

SPECIMENS ExA~~iv~D.-Australia: Australian Capital Territory, Weber L-47220. L-47282 (COLO, US). Union of South 

Africa: Cape Province, Alniborn 4900 (LD, US), Degelius 

SA-57 (Degelius hcrbarium, US). 

Pseudoparmelia subambigua, new species 

FIGURE 17b 

DESCRIPTIoN.-Thalhs arcte adnatus, mollis, 

viridi-flavicans, 5-10 cm latus, lobis subirregularibus, brevibus, apice subrotundatis, congestis, 1 .O3.0 mm latis; superne planus, centrum versus rugosus, dense sorediatus, soraliis orbicularibus, confluentibus; cortex superior 8-10 pm crassus, epicortex perforatus, stratum gonidiale 20 pm crassum, 

medulla alba, 110-120 pm crassa, cortex inferior 

7-10 pm crassus; subtus niger, sparse rhizinosus, 

rhizinis simplicibus, nigris. Apothecia ignota. 

KCf red, P-; usnic acid and lecanoric acid. 

HoLoTYPE.-chile: Antofagasta, Paposo Quebrada Guanillo, elevation 880 m, M. Mahu 3405, 

12 September 1972 (US; Mahu herbarium, isotype). 

DIsmmuTIoN.-Chile. 

HABITAT.-on shrubs and fence posts in open 

areas from sea level to 900 m elevation. 

REh.fARKs.-This lichen could easily be misidentified as Parmeliopsis ambigua (Acharius) Nylander. The lower surface is black, however, and the 

chemistry distinctive. Larger specimens may also 

resemble Pseudoparmelia soredians (Nylander) 

Hale, which contains salazinic acid and which may 

well be the most closely related species. It is known 

only from Chile. 

CHEMISTRY.-cOrteX K - , medulla K - , c -l-, 

SPEClhlENS EXA?mNED.-chile: Coquimbo, Runde2 7230 (Us), 

Santesson 2519 (S, US); Santiago, Rundel 7323 (US); Valparaiso, Santesson 2808 (S, US). 

Pseudoparmelia subamplexa, new species 

FIGURE 17c 

DEsCRIPTroN.-Thallus ut in Pseudoparmelia amplexa (Stirton) Hale sed superficie sorediatus, soraliis 
NUMBER 31 

FIGURE 17.-Species of Pseudoparmelia: a, P. spodochroa (Weber L-47220 in US); b, P. subambigua (Santesson 2519 in US); c, P. subamplexa (Schiitte 34k, isotype in US); d, P. subtiliacea 

(Knight 7, lectotype in H); e, P. subtortula (Lye LZOZA, holotype in BM); 1, P. texana (Hale 

42248). (Scale in mm.) 

51 
52 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

orbicularibus, discretis vel pro parte confluentibus; cortex superior 15-18 pm crassus, epicortatus, 

epicortice perforato, stratum gonidiale 15-20 ,pm 

crassum, medulla alba, 110-130 ,pm alta, cortex inferior 14-20 pm crassus; subtus niger, opacus, modice 

rhizinosus, rhizinis simplicibus. Apothecia ignota. 

rose, P+ red; usnic acid and protocetraric acid. 

HOLOTYPE.-on tree, Chishawasha, Salisbury District, S. Rhodesia, K. Schutte 34k, 30 December 1953 

(LD; US, isotype). 

CHEMISTRY.-cOrteX K-, c-, KC- or KC+ 

DrsTRIsuTroN.-Rhodesia. 

HABITAT.-on trees in arid scrub land. 

REMARKS.-ThiS is the sorediate morph of P. 

amplexa (Stirton) Hale. It has the same peculiar 

black velvety lower surface. As with the parent 

species, it is restricted to Africa. 

SPECIMENS ExAMINED.-Rhodesia, Hoeg (TRH, us), Schutte 

42 (LD, US). 

Pseudoparmelia subtiliacea 

FIGURE 17d 

Pseudoparnielia subtiliacea (Nylander) Hale, 1974:191. 

Parmelia subtiliaeea Nylander, 1885:614 [type collection: New 

Zealand, Knight (H, Nylander herbarium number 35176, 

lectotype; UPS, isolectotype)]. 

Parmelia tiliacea var. feracissima Miiller Argoviensis, 1886: 

256 [type collection: Guntawang, New South Wales, Australia, Hamilton 5 (G, lectotype)]. 

DESCRIPTION.-Thallus adnate on twigs, whitish 

mineral gray, 3-5 cm broad; lobes subirregular, 

black rimmed, 1.5-3 mm wide; upper surface plane 

to rugulose or foveolate; lower surface black, sparsely rhizinate. Apothecia common, substipitate, 1-5 

mm in diameter; spores 8, 6-7 X 12-14 pm. 

CHEMIsTRY.-Cortex K + yellow, medulla negative with all reagents; atranorin and caperatic acid. 

DIsTRIBuTIoN.-Australia and New Zealand. 

HABITAT.-on twigs of subalpine shrubs and in 

dry scrub land. 

REMARKs.-Pseudoparmelia subtiliacea is a very 

rare lichen and as a consequence it is poorly understood. The only close relative seems to be another 

Australian lichen, P. corrugativa (Filson and Kurokawa) Hale, which has a pigment in the lower half 

of the medulla. 

Pseudoparmeliu subtortula 

FIGURE 17e 

Pseudoparinelia subtortula (Hale) Hale, 1974:191. 

Parmelia subtortula Hale, 1973b:3 [type collection: E. Mengo 

District, Uganda, Lye L202A (BM, holotype; US, isotype)]. 

DESCRIPTION.-Thal~US as in P. tortula (Hale) 

Hale (see below) except the upper surface isidiate, the isidia cylindrical, simple. Apothecia not 

seen. 

CHEMxsmY.-Cortex K + yellow, medulla K - , 

C - , KC + rose, P - ; atranorin, norlobaridone, and 

neoloxodic acid. 

DIsTRIBuTIoN.-Uganda and Kenya. 

HABITAT.-On rocks in open areas at about 1000 

m elevation. 

REMARKs.-Pseudoparmelia subtortula is the presumptive isidiate morph of P. tortula, a species with 

identical chemistry, which occurs chiefly in South 

Africa. Both are saxicolous African endemics unique 

in producing norlobaridone and neoloxodic acid. 

Uganda and Kenya. 

SPECIMENS ExAMINED.-see Hale (1973b) for records from 

Pseudoparmelia 

Pseudoparmelia texana 

FIGURE 17f 

texana (Tuckerman) Hale, 1974:191. 

Parmelia texana Tuckerman, 1858:424 [type collection: 

Blanco, Texas, Wright (FH, lectotype; M, US, isolectoParmelia sublaevigata var. texana (Tuckerman) Nylander in 

Triana and Planchon, 1867:307. 

Parrnelia confluexens Nylander, 1875:725 [type collection: St. 

Paul (H, Nylander herbarium number 35178, lectotype; 

BM, PC, UPS, isolectotypes)]. 

Parmelia cingalensis Stirton, 1876: 159 [type collection: Galle, 

Ceylon (BM, lectotype); illegitimate name]. 

Parmelia leptophylla Miiller Argoviensis, 1891:377 [type collection: Baziya, South Africa, Baer 714 (BM, lectotype; G, 

isolectotype)]. 

Parmelia tenuirinia f. sorediata Muller Argoviensis, 1894:258 

[type collection: Usambara (Tanganyika), Africa, Holst 787 

pro parte (G, lectotype)]. 

Parmelia exoriens Stirton, 1899:76 [type collection: Brisbane, 

Australia, Bailey 215 (BM, lectotype; GLAM, isolectotype)]. 

Parinelia symrniga Hue, 1899:168 [type collection: Coonoor, 

Nilgherries Mountains, India, Gray (PC, lectotype)]. 

Partnelia subconfluescens Magnusson, 1942:5 [type collection: 

Puu Waawaa, north of Hualalai, Hawaii, Selling 5669 (S, 

holo type)]. 

types)]. 
NUMBER 31 53 

Parmelia mangenotii des Abbayes, 1951:969 [type collection: 

Mankono, Cercle of Seguela, Ivory Coast, des Abbayes 

(REN, lectotype; US, isolectotype)]. 

Parnielia pseudorutidota Asahina, 1952: 17 [type collection: 

Nagachi, Shinshu, Japan, Takahashi (TNS, lectotype)]. 

Parmelia tQnQkQe Asahina, 1954:371 [type collection: KUZUMura, Buzen, Prov. Yamato, Japan, Tanaka (TNS, lectoParmelia QlbQnienSlS Dodge, 1959:121 [type collection: forests 

of Albany, Cape of Good Hope, Union of South Africa, 

Zeyher 3 (FH-Tayl, holotype)]. 

DEscRrPTroN.-Thallus closely adnate on trees 

or rocks, ashy white, 6-12 cm broad; lobes sublinear 

to subirregular, apically rotund, 3-5 mm wide; upper surface plane to rugulose, usually deeply rimose, 

sorediate, the soralia laminal, punctiform to capitate, in part initially pustulate with formation of 

soredia; lower surface black and moderately rhizinate except for a narrow naked brown zone along 

the margins. Apothecia rare, 2-5 mm in diameter; 

spores 8,6-7 X 9-11 pm. 

C--, KCf pale violet or KC-, P--; atranorin, 

divaricatic acid, and associated unknowns. 

DIsTRrsuTIoN.-Pantemperate outside of Europe 

and pantropical at higher elevations. 

erus, Pinus, Quercus) and more rarely on rocks in 

open or secondary forest and in parks and gardens 

at 50-2400 m elevation. 

REMARKS.-This is one of the most widespread 

species in the genus and while not usually occurring in as great abundance, for example, as P. cuperuta (L.) Hale, it is well represented, if not 

frequently misidentified, in most large herbaria. 

The range of morphological variation is small and 

the chemistry uniform and distinctive. It is difficult 

to understand why so many lichenologists failed to 

recognize it. The nonsorediate progenitor may be 

represented by the African P. nairobiensis (Steiner 

and Zahlbruckner) Hale and the companion isidiate morph by P. concrescens (Vainio) Hale. 

REPRESENTATIVE SPECIMENS EXAhlINED.-united States: Wisconsin, Thomson 10921 (US, WIS); Tennessee, Hale 37032; 

West Virginia, Hale 10308; Illinois, Skorepa 934 (us); Virginia, Hale 12814 (US); South Carolina, Culberson 7515 

(DUKE); Alabama, Hale 34057; Georgia, Hale in Lichenes 

Selecti Exuccati 270 (US); Florida, Hale 36897; Arkansas, 

Hale 5567. Mexico: Chihuahua, Matthews 43794 (US); Hidalgo, Chase 7429 (F, US), Weber et d. 33690 (COLO); 

Michoacaan, Hale 20849; Oaxaca, Cain 27578 (TRT, US), 

Hale 20621, 20630; Veracruz, Hale 19407; Chiapas, Hale 

t,pe)l. 

CHEMISTRY.-cOrteX Kf yellow, medulla K-, 

HABITAT.-On trees (A CaCia, Eucalyptus, Junip20278. Honduras: Morazan, Standley 4105, 11623 (F). Costa 

Rica: San Jose, Q,uiros 1246, 1474 (US). Panama: Chiriqui, 

Hale 38914, Scholander (US). Haiti: Ouest, Imshaug 22537 

(MSC), Wetmore 2670 (MSC, US). Venezuela: Merida, Hale 

42121, 42920. Peru: Abancay, Iltis 2322 (WIS). Brazil: Minas 

Gerais, Eiten 6864, 6870 (US), Mosen 2309 (S, US): SaO Paulo, 

Eiten 2978, 8060 (US), Milne 32 (BM), Osorio 4919 (MVM); 

Santa Catarina, Reitz and Klein 15114 (US); Rio Grande do 

Sul, Lima 90 (US). Uruguay: Canelones, Zorron 1929 (US); 

Montevideo, Osorio 1849, 6824 (MVM). Chile: Antofagasta, 

Rundel 7205 (US). Uganda: Busiro County, Swinscow 2U 

24/4 (BM, US). Angola: Bie, Degelius (Degelius herbarium). 

Kenya: Rift Valley Province, MQQS Geesteranus 4551 (L). 

Rhodesia: Hoeg (TRH). Tanzania: Arusha Prov., Santesson 

21338 (UPS); Kilimanjaro Prov., Santesson 20948 (UPS); 

Union of South Africa: Natal, Hoeg (TRH); Transvaal, 

Alrnborn 6253 (LD); Cape Province, Almborn 10664 (LD). 

Madagascar: des Abbayes in Lichenes Madagascarienses et 

Borbonici Selecti EXS~CCQ~~ 11 (LD, US). India: Tamil Nadu. 

Foreau 4092 (Awasthi herbarium), Hale 40167, 43708. Thailand: Kurokawa 1948 (TNS). Japan: Province Ohmi, Hale 

29473. Indonesia: Java, Groenhart 1711 (L), Kurokawa in 

Lichenes Rariores et Critici EXS~CCQ~~ 39 (TNS, US); Sumatra, 

Groenhart 957 (L). Australia: New South Wales, Flockton 

724B (US). 

Pseudoparrnelia tortula 

FIGURE 18a 

Pseudoparrnelia tortula (Kurokawa) Hale, 1974:191. 

Parmelia tortula Kurokawa in Hale and Kurokawa, 1964:157 

[type collection: Namaqualand, Cape Province, Union of 

South Africa, Almborn 4805 (LD, holotype; TNS, US, iso- 

‘Y pes)l. 

DESCRIPTION.-Thallus closely adnate on rock, 

whitish mineral gray, 5-10 cm in diameter; lobes 

subirregular to sublinear, apically subrotund, 

crowded toward the center, 1.5-4 mm wide, often 

twisted and contorted when free from the substratum; upper surface plane to rugulose or undulate, 

continuous, sometimes faintly pruinose; lower surface pale brown, sparsely rhizinate, the rhizines 

brown. Apothecia adnate to substipitate, 1-5 mm 

in diameter; spores 5-6 X 7-9 ,ym. 

C -, KC - or KC + faint rose, P - ; atranorin, norlobaridone, and neoloxodic acid. 

CHEMISTRY.-cOrteX K f yellow, medulla K--, 

DISTRIBUTION.-uniOn of South Africa. 

HABITAT.-on rocks in open treeless or sparsely 

vegetated areas up to 1000 m elevation. 

REMARKs.-This rather undistinguished lichen is 

still known only from South Africa, where it seems 

to be quite common. The presumptive isidiate 
54 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 18.-Species of Pseudoparmelia: a, P. tortula (Almborn 4805, holotype in LD); b, 

P. vunder6ylii (AlmbOrn 5110 in US); c, P. venezolana (Hale 45526a, holotype in US); d, P. 

uiolacea (Almborn 1771, holotype in LD); e, P. xanthomelaena (Maas Geesteranus 12145 in US); 

f, P. zambiensis (Jellicoe 50, holotype in BM); g, P. zimbabwensis (Hceg, holotype in TRH). 

(Scale in mm.) 
NUMBER 31 55 

morph, P. subtortula (Hale) Hale, occurs farther 

north in Kenya and Uganda. 

SPECIMENS EXAMINED.-&e Hale and Kurokawa (1964: 158) 

for records from the Union of South Africa. 

Pseudoparmelia vanderbylii 

FIGURE 18b 

Pseudoparmelia vanderbylii (Zahlbruckner) Hale, 1974:191. 

Parmelia vanderbylii Zahlbruckner, 1932a:252 [type collection: Union of South Africa, Van Rhynsdorp, Cape Province, P. A. van der Byl (W, lectotype)]. 

DEscRIPTIoN.-Thallus closely adnate on rocks, 

brownish or dark mineral gray, 3-6 cm broad; lobes 

subirregular, apically subrotund, contiguous, 1-2 

mm wide; upper surface plane, shiny, rimose with 

age; lower surface pale brown, sparsely rhizinate, 

the rhizines pale brown. Apothecia common, sessile, 1-2 mm in diameter; spores 8, 6 X 8-10 pm. 

CHEMISTRY.-cortex Kf yellow, medulla Kor Kf very faint orange, C-, KC-, P-; atranorin and two or more unidentified substances. 

DrsTRIBuTroN.-~nion of South Africa. 

HABITAT.-oII exposed rocks in open areas. 

REMARKS.-This species falls near P. molybdiza 

(Nylander) Hale in general aspect and habitat but 

differs in chemistry. Two specimens (the lectotype 

and Almborn 4636) have identical chemistry (two 

spots in both the hexane and benzene solvent systems) and the third (Almborn 51 10) has distinctly 

different but still unidentified higher spots. All 

three share the same two lowermost faint spots. 

None matched any of the usual KC.- depsidones 

and depsides with which they were co-chromatographed. 

ince, Alvrborn 4636, 5110 (LD, US). 

SPEClhtENS EXAhllNED.-uniOn of South Africa: Cape PrOVPseudoparmelia venezolana, new species 

FIGURE 18c 

DEsCRIPTION.-Thallus adnatus, saxicola, cinereoalbus, 3-7 cm latus, lobis sublinearibus, congestis, 

0.6-1 .O mm latis; superne planus, nitidus, dense 

isidiatus, isidiis cylindricis, simplicibus vel ramosis, 

0.2-0.4 mm altis; cortex superior 15-18 pm crassus, 

epicorticatus, epicortice distincte perforato, stratum gonidiale ca 10 pm crassum, medulla alba, 100 

pm crassa, cortex inferior paraplectenchymatus, 8- 

10 pm crassus; subtus pallide brunneolo-albus, 

sparse rhizinosus, rhizinis simplicibus, pallidis. Apothecia ignota. 

CHEMISTRY.-cOrteX Kf yellow, medulla K-, 

C-, KC- or KC+ rose, P+ red; atranorin and 

protocetraric acid. 

HoLOTYPE.-Venezuela: Tachira, Pic0 Banderas, 

Pdramo de Tama elevation 3200 m, M. E. Hale 

45526a, 27 March 1975 (US). 

REMARKS.-ThiS inconspicuous species is easily 

overlooked in the high paramo areas where it appears to be confined. This is the highest elevation 

at which the genus is found. There seem to be no 

close relatives. It would more likely be confused 

with Parmeliopsis, which is characterized by the 

presence of thamnolic acid. 

SPECIMENS EXAMINED.-VeneZUeh: Taichira, Hale 45499a, 

45523, 45528a. 

Pseudoparmelia violacea 

FIGURE 18d 

Pseudoparmelia violacea (Kurokawa) Hale, 1974: 191. 

Parmelia violacea Kurokawa in Hale and Kurokawa, 1964: 

158 [type collection: Blinkwater Ravine, Table Mountain, 

Cape Province, Union of South Africa, Almborn 1771 (LD, 

hobtype)]. 

DEscRIPTIoN.-Thallus closely adnate on rock, 

greenish mineral gray, 3-6 cm in diameter; lobes 

sublinear, 1-3 mm wide; upper surface plane to 

distinctly rugulose, continuous; medulla vinaceous 

purple; lower surface pale brown, moderately rhizinate, the rhizines pale. Apothecia adnate, 1-1.5 

mm in diameter; spores not developed. 

CHEMISTRY.-cOrteX K + yellowish, medulla KSpurple, P - ; unidentified anthraquinones. 

DISTRIBUTION.-union of South Africa. 

HABITAT.-on rocks in open forest. 

REMARKS.-This remarkable species has been recollected once since the original description. It is 

another typical saxicolous African endemic verging 

close to Xanthoparmelia. The pigments are, in fact, 

very close to those of X. endomiltoides (Nylander) 

Hale, one being identical. No lichen substances are 

present in the cortex. 

SPECIMENS EXAMINED.-union of South Africa: Cape Province, Degelius SA-70 (Degelius herbarium, US). 
56 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

Pseudoparmelia xanthomelaena 

FIGURE 1% 

Pseudoparnielia xantfiornelaenn (Muller Argoviensis) Hale, 

1974: 191. 

Parnzelia xanthomelaena Mulller Argoviensis, 1883a:48 [type 

collection: Grampian Mountains, Australia, Sullivan 28 

(G, lectotype)]. 

DE~CRIPTI~N .-Thallus closely appressed on rock, 

buff mineral gray but darkening to brownish olive 

gray at the center, 3-5 cm broad; lobes sublinear, 

contiguous, lobulate, 0.5-1.5 mm wide; upper surface plane, shiny, transversely rimose with age; 

lower surface black, moderately rhizinate. Apothecia common, adnate, up to 1 mm in diameter; 

spores 8,6 X 7-10 pm. 

yellow, C-, KC-, P+ orange; atranorin, stictic 

acid, and constictic acid. 

DISTRIBUTION.-uniOn of South Africa and Australia. 

HABITAT.-Exposed rocks from sea level to 250 

m elevation. 

REMARKS.-This is one of the smallest species of 

Pseudoparmelia. It is probably difficult to collect 

since it cannot be removed free of the rock substratum. There are no obviously related species in 

the genus, unless we consider P. ischnoides (Kurokawa) Hale as a possible isidiate morph. 

CHEMISTRY.-cOrteX K + yellow, medulla K + 

SPECIMENS EXAbiINED.-~niOII of South Africa: Basutoland, 

KoPer (LD); Cape Proiince, Almborn 125, 275, 1136a. 1444, 

1546, 1503, 3023, Degelius 54-337 (Degelius herbarium), Maas 

Geesteranus 12145 (L). 

Psewloparmelia zambiensis 

FIGURE 1Sf 

Pseudoparmelia zambiensis (Hale) Hale, 1974:191. 

Parnielia zambiensis Hale, 1972b:346 [type collection: Kyika 

Plateau, Zambia, lellicoe 59 (BM, holotype; US, isotype)]. 

DEsCRIPTIoN.-Thallus adnate on bark, greenish 

yellow, 4-6 cm broad; lobes subirregular, crowded, 

2-3 mm wide, becoming lobulate toward the center; 

upper surface plane to rugulose, rimose with age; 

lower surface black, sparsely rhizinate, the rhizines 

thick, black. Apothecia common, adnate 2.5-3.0 

mm in diameter; spores 8, 6 X 12 pm. 

KC- or KC+ faint rose, P-; usnic acid and divaricatic acid with associated unknowns. 

CHEMISTRY.-cOrteX K--, medulla K--, c-, 

DIsTRIBUTIoN.-Centra1 Africa. 

HABITAT.-on trees in open areas at 2300 m. 

REMARKS.-This is another African species that 

will surely be collected more often as lichenologists 

visit the continent. It appears to be the nonisidiate 

progenitor of P. ecaperata (Muller Argoviensis) 

Hale, which has a wide range in Africa and Asia. 

It could also be considered as the usnic acid chemical counterpart of P. nairobiensis, which contains 

atranorin only in the cortex. A full understanding 

of the relations of these species can only be gained 

when more specimens are collected. 

Pseudoparmelia rimbabwensis 

FIGURE 18g 

Pseudoparnielia zinibabwensis (Hale) Hale, 1974: 191. 

Parmelia zim babzc~emis Hale, 1972b:346 [type collection: Zimbabwe, Rhodesia, Hoeg (TRH, holotype; LD, US, isotypes)]. 

DEscRIPTIoN.-Thallus closely adnate on rock, 

fragile, light mineral gray, 2-5 cm broad; lobes sublinear, contiguous, black-marginate, 1 .O-1.5 mm 

wide; upper surface plane to rugulose, in part 

pruinose, pustulate, the pustules thick, apically 

pruinose and finally erupting without soredial formation; lower surface black, moderately rhizinate, 

the rhizines black. Apothecia not seen. 

CHEMISTRY.-cOrteX K+ yellow, medulla K-, 

C-, KC-, P+ red; atranorin and protocetraric 

acid. 

DIsTRisuTIoN.-Rhodesia and Union of South 

Africa. 

HABITAT.-oII boulders in open areas. 

REMARKS.-The pustules of this species are similar to those of P. pusutlescens (Kurokawa) Hale, a 

somewhat larger species containing sekikaic acid 

P->* 

SrEcIirENs EXA\fINED.-uniOn of South Africa: Transvaal, 

Almborn 6108, 6491, 6496 (LD). 

Doubtful and Rejected Names 

Parmelia appressa Sprengel, 1987:58 [type collection: On Cinchona, tropical America]. 

The type collection could not be found. A specimen in Acharius’ herbarium labeled “Parmelia 

appressa” appears to be identical with Pseudoparmelia sphaerospora (Nylander) Hale. How- 
NUMBER 81 57 

ever, Sprengel mentioned affinity with Parmelia 

perforata and P. perlata, both large Parmotrema 

species. 

Parmelia caperata var. coerulea Bouly de Lesdain, 

1954: 222 [type collection: Rocamadour, France, 

Duval]. 

The type specimen is probably in PC but I was 

not able to locate it there. 

Parmelia protorevoluta Gyelnik, 1931b3288. 

Parinelio revoluta f. nuda Muller Argoviensis, 1891:378 [type 

collection: Santarem, Brazil, Spruce 135 (G, lectotype; BM, 

isolectotype)]. 

The fragmentary type collections contain divaricatic acid. This is almost certainly a Pseudoparmelia, perhaps P. nairobiensis (Steiner and 

Zahlbruckner) Hale, a presumed African endemic. 

Parmelia rutidota var. vestita Zahlbruckner, 1941: 

107. 

This variety is based on two collections, one 

Thomson ZA217 from Southland, New Zealand, 

which is Parmelia rudecta Acharius, and the 

other Cranwell ZA242 from Coromandel Peninsula, New Zealand, which can be identified as 

Parmotrema crinitum (Acharius) Choisy. Both 

specimens are preserved in W. 

Parmelia soredians f. farinosa (Vainio) Gyelnik, 

1938a:27. 

This is based on Parmelia farinosa Vainio (1890: 

62) (Lichenes Brasilienses Exsiccati 551 in 

TUR). It is a species of Xanthoparmelia (Vainio) 

Hale. 

Parmelia soredians f. lobulata Gyelnik, 1934:306 

[type collection: La Calera Sud, Argentina, Hosseus 22 (BP, holotype)]. 

This is a species of Xunthoparmelia (Vainio) 

Hale. 

Parmelia soredians f. muscicola Gyelnik, 1934:306 

[type collection: Capilla del Monte, Argentina, 

Hosseus 23 (BP, holotype)]. 

This is also a Xanthoparmelia. 

Culberson and Culberson (1969: 5 15) place this 

Pseudoparmelia aradensis Gyelnik, 1933:6. 

species in synonymy under Cetrelia olivetorum 

Pseudoparmelia euplecta (Stirton) Hale, 1974: 190. 

I have tentatively placed this species as a synonym 

of Pseudoparmelia caperata in the “Taxonomic 

Treatment.” 

Pseudoparmelia leucopis (Krempelhuber) Hale, 

1974: 190. 

(Nylander) W. Culberson and C. Culberson. 

Parmelia leucopis Krempelhuber, 1878:461 [type collection: 

Lorentz and Hieronytnus, Argentina (M. lectotype)]. 

After reexamining the type collection, I found 

some cilia in the lobe axils. These would exclude 

the species from Pseudoparmelia, and I hesitate 

to reassign it to another genus until better material is available. 

Pseudoparmelia meiosperma (Hue) Hale, 1974: 190. 

Parmelia internexa f. rneiosfierina Hue, 1899: 185 [type collection: Mafate, Reunion, Rodriguez (PC, lectotype)]. 

The type-specimen has rather broad lobes with 

dense isidia. The rhizines are dense and more or 

less branched. The chemical constituents include 

atranorin, evernic acid, and traces of lecanoric 

acid, a combination that is known in several species of Hypotrachyna (Hale, 1975a) but in no 

Pseudoparmeliae. It does not fit well in Pseudoparmelia and until additional material is available it is impossible to settle its generic position. 

Culberson and Culberson (1968:459) consider 

this to be a species of Cetrelia but not having 

examined the type collection they could not decide which one. 

Pseudoparmelia pseudofallax var. cretaceoides Gyelnik, 1933:7. 

This taxon also belongs in the Cetrelia cetrarioides complex. 

Pseudoparmelia pseudosoTediosa (Gyelnik) Hale, 

1974: 191. 

This has been reduced to synonymy under Pseudoparmelia caperata in the “Taxonomic Treatment” above. 

Pseudoparmelia pseudofallax Gyelnik, 1933:6. 
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Index 

(Synonyms in italics) 

Cetrelia olivetorum, 57 

Lichen caperatus, 20 

Parmelia adspersa, 14 

alabamensis, 14 

albaniensis, 53 

amazonica, 16 

amplexa, 16 

angolensis, 38 

annexa, 17 

an tarctica, 3 2 

appressa, 56 

aptata, 17 

arcana, 17 

asmarana, 17 

atrichoides, 38 

austroafricana, 25 

balt i morensis, 18 

basutoensis, 18 

benguellensis, 16 

bipindensis, 49 

brachyphylla, 38 

caflrorum, 25 

capensis, 25 

caperata, 20 

caperata var. caperatula, 44 

caperata var. coerulea, 51 

caperafa b. qliphora, 20 

caperata var. exornata, 31 

capera fa var. glaucopis, 16 

caperata var. isidiophora, 41 

caperata f. Eaevissima, 20 

caperata f. ramealis, 45 

caperata var. subglauca, 20 

caperatula, 45 

caribaea, 2 1 

carneopruinata, 22 

caroliniana, 22 

chapadensis, 24 

cinermcens, 24 

cingalensis, 52 

citrinescens, 31 

concrescens, 24 

condy loides, 25 

confertula, 45 

conpuescens, 52 

congruens, 49 

conspersa var. polyphylla f. sorediosa, 

corrugativa, 25 

48 

crozalsiana, 27 

cryptochlorophaea, 27 

cubensis, 49 

cylisphora, 20 

cyphellata, 28 

d ja lonensis, 29 

ecaperata, 28 

ecoronata, 34 

endoxantha, 49 

epileuca, 29 

eruptens, 29 

euplecta, 20, 21 

exoriens, 52 

ferax, 31 

filipina, 14 

pavicans, 20 

pavidoglauca, 49 

ganguellensis, 38 

geesterani, 31 

gerlachei, 32 

glaucopis, 16 

gracilenta, 34 

gracilescens var. angolensis, 38 

gracilis, 34 

halmaiana, 48 

hansfordii, 39 

herreana, 20 

hypomilta, 32 

imperfecta, 42 

inactiva, 14 

inhaminensis, 32 

inornata, 34 

internexa f. meiosperma, 57 

intertexta, 34 

ischnoides, 35 

isidioplzora, 22 

jelinekii, 45 

labrosa, 35 

laetepavens, 29 

lecanoracea, 35 

leptophylla, 52 

leucochlora, 49 

leucopis, 57 

leucoxantha, 37 

longiconida, 48 

malaccensis, 31 

malaccensis var. laetepavens, 29 

mangenotii, 53 

martinicana, 37 

molybdiza, 38 

nairobiensis, 38 

negativa, 20 

nipponica, 17 

obversa, 31 

ochroleuca, 45 

ochroleuca f. sorediosa, 20, 21 

owariensis, 39 

pachydactyla, 41 

papillosa, 41 

perfissa, 38 

perlata var. pavicans, 20 

perlata var. subrevoluta, 22 

prolata, 41 

protorevoluta, 57 

pseudorutidota, 53 

pseudosorediosa, 20 

pustulescens, 42 

rahengensis, 42 

ramulicola, 48 

raunkiaeri, 42 

regnellii, 32 

regnellii f. arida, 32 

relicina var. ecoronata, 34 

revoluta f. nuda, 51 

rodriguesiana, 44 

rupicola, 44 

rutidota, 44 

rutidota f. filizans, 31 

rutidota var. vestita, 57 

salacinifera, 45 

san-miguelii, 48 

sbarbaronis, 22 

scabrella, 18 

scabrosa, 42 

schelpei, 46 

schistacea, 46 

scotophylla, 46 

scottii, 48 

scrobicularis, 48 

somaliensis, 48 

soredians, 48 

soredians f. farinosa, 57 

soredians f. lobulata, 57 

soredians f. muscicola, 57 

sphaerospora, 49 

splendidula, 45 

spodochroa, 50 

steineri, 41 

61 
62 SMITHSONIAN CONTRIBUTIONS TO BOTAXY 

subcaperatula, 45 

subconpuescens, 52 

subconspersa var. benguellensis, 16 

subglauca, 20 

sublaeuigata Tar. texana, 52 

subrupta, 34 

subtiliacea, 52 

subtortula, 52 

sy m miga, 52 

tanakae, 53 

tenuiriina var. labrosa, 35 

tenuiriina f. sorediata, 52 

texana, 52 

tiliacea var. feracissinia, 52 

tortula, 53 

uleana, 49 

vanderbylii, 55 

uincentiana, 42 

uiolacea, 55 

wainioana, 22 

xanthornelaena, 56 

;am biensis, 56 

zenkeri, 49 

rimbabwensis, 56 

Parmotrema crinitum, 57 

Pseudoparmelia adspersa, 14 

alabamensis, 14 

amazonica, 16 

amplexa, 16 

annexa, 17 

aptata, 17 

aradensis, 57 

arcana, 17 

baltimorensis, 18 

basutoensis, 18 

Denguellensis, 16 

callichroa, 20 

caperata, 20 

caribaea, 21 

carneopruinata, 22 

caroliniana, 22 

chapadensis, 24 

cinerascens, 24 

concomitans, 24 

concrescens, 24 

condyloides, 25 

conlabrosa, 25 

corrugativa, 25 

crozalsiana, 27 

cryptochlorophaea, 27 

cyphellata, 28 

dahlii, 28 

ecaperata, 28 

epileuca, 29 

eruptens, 29 

euplecta, 20, 57 

exornata, 31 

ferax, 31 

geesterani, 31 

gerlachei, 32 

hypomilta, 32 

inhaminensis, 32 

inornata, 34 

intertexta, 34 

ischnoides, 35 

labrosa, 35 

lecanoracea, 35 

leucopis, 57 

leucoxantha, 37 

malaccensis, 37 

martinicana, 37 

meiosperma, 57 

molybdiza, 38 

nairobiensis, 38 

neoquintaria, 39 

owariensis, 39 

pachydactyla, 41 

papillosa, 41 

prolata, 41 

pseudofallax, 57 

pseudofallax var. cretaceoides, 57 

pseudosorediosa, 20, 57 

pustulescens, 42 

rahengensis, 42 

raunkiaeri, 42 

rodriguesiana, 44 

rupicola, 44 

rutidota, 44 

salacinifera, 45 

schelpei, 46 

schistacea, 46 

scotophylla, 46 

scrobicularis, 48 

somaliensis, 48 

soredians, 48 

sphaerospora, 49 

spodochroa, 50 

subambigua, 50 

subamplexa, 50 

subtiliacea, 52 

subtortula, 52 

texana, 50 

tortula, 53 

vanderbylii, 55 

venezolana, 55 

violacea, 55 

xanthomelaena, 56 

zambiensis, 56 

zimbabwensis, 56 

Pyxine azorea, 22 

Xanthoparmelia gerlachei, 32 

molliuscula, 41 

papillosa, 41 

scabrosa, 42 

* U.S. GOVERNMENT PRINTING OFFICE: 1976--211--559/18