SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 32 

A Monograph of the Lichen Genus 

BuZbothrix Hale (Parmeliaceae) 

Mason E. Hale, Jr. 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1976 
ABSTRACT 

Hale, Mason E., Jr. A Monograph of the Lichen Genus Bulbothrix Hale 

(Parmeliaceae). Smithsoniun Contributions to Botany, number 32, 29 pages, 7 

figures, 1976.-A world-level revision is given for the 29 species of Bulbothrix, 

a generic segregate of Parmelia characterized by marginal bulbate cilia and production of atranorin in the upper cortex. The genus is primarily tropical in 

distribution and best developed in secondary forests. The main center of speciation is Brazil with 14 species, and the genus is also well represented in Africa. 

The New World species have a high frequency of depsides and orcinol depsidones, whereas the Old World species usually contain p-orcinol depsidones. 

One new species, B. klementii Hale, and one new combination, B. goebelii 

(Zenker) Hale, are proposed. 

OFFICIAL I~~'I~C,IC,~TIOS D.yrI: is handstamped in a limited number of initial copics and is rccordetl 

in thc Institution's annual rcport, Suiithsoninn Yew. SI:RII:S COVER DMICS: Leaf clcaring from 

the katsura trcc Cercidiphyllirm jnponicum Siebold and Zuccaritii. 

Libra1 y of Congrcss Cataloging in Publication Data 

Hale, Mason E. 

A monograph of the lichcn gcnus Bulbothrix Halc (Parmcliaccac). 

(Smithsonian contributions to botany : no. 32) 

Bibliography: p. 

Includes index. 

Supt. of Docs. no.: SI 1.29:32 

1. Bulbothrix. I. Titlc. 11. Series: Smithsonian Institution. Smithsonian contributions to 

botany ; no. 32. 

QKlS2747 110. 32 [QK589.P2] 581'.08~ [589'.1] 75-619423 
Contents 

Pnge 

Introduction 1 

Morphology 1 

Chemistr) 3 

hforph Formation 1 

Phj togeography 5 

Key to the Species 7 

Classification of Bulbothrix 6 

Taxonomic Treatment 9 

Literature Cited 27 

Index 29 

... 

111 

A Monograph of the Lichen Genus 

Bulbothrix Hale (Parmeliaceae) 

Mason E. Hale, Jr. 

Introduction 

This world monograph of the 29 species of 

Bztlbothl-ix is part of my continuing effort to 

revise the parnielioid genera, utili~ing the most 

up-to-date analytical tools available, in particular 

tlie scanning-electron microscope and thin-layer 

chromatography. Genera already completed include Nypotl-nchyna (Vainio) Hale (Hale, 1975a) 

(the neotropical species), Relicinn (Hale and 

Kurokawa) Hale (Hale, 1975b), and Pseudopa?.- 

melia Lynge (Hale, 1976). Much of the introductory material in these revisions applies equall) 

well to Bulbothl-ix and will not be repeated here. 

I am especiallv thankful to curators of the various 

museums and herbaria, as listed Ly tlie standard 

herbarium acronyms under specimens examined, 

who so generously and promptly lent specimens for study. Special thanks for assistance in 

carrying out field studies are due Dr. M. LopezFigueiras of the Universidad de 10s Andes, hferida, 

Venezuela; Dr. P. G. Patwardhan, Maharashtra 

Association for tlie Cultivation of Science, Poona, 

India; and Dr. Flora Uyenco, University of the 

Philippines, Quezon City. Dr. S. Kurokawa a\- 

shed with descriptions and preliminary identifications of many of the species in 1961. 

The following grants received since 1960 are 

acknowledged with thanks: Sational Science 

Foundation (Systematic Biology and tlie U.S.- 

Japan Cooperative Science Program), MordenMason E. Hnlr, Jr., Departnient of Botany, Xutionnl Zflrcrttni 

of .Vatuinl History, Smithsonian Institution, Washinqton, 

D.C. 20560. 

Smithsonian Expedition to Dominica, Sational 

Geographic Society, and the Smithsonian liesearch Foundation. 

Morphology 

THE THALr.us.-species of b’iilboth rix have 

small, adnate to closely adnate tlialli. The lobes 

are suhlinear to more rarely sul,irregular, clepentling on the width. In these respects Rlrlbothrix is 

closest to Relicinn and Pai.nielinn. The most cliaracteristic feature, however, is the development of’ 

marginal bulbate cilia, which are essentially 

identical with those of Relirinn antl regularly dispersed around the lobe tips antl margins. Tl’lien 

rhizines are dichotomously Ixinchetl, the tips of 

these bulbae may also be branched. 

LOWER SURFACE AND RHmNr;.s.-The lower surface is black in some of tlie species, dark or pale 

brown in others. Rhizines are either simple to 

sparsely furcate, as in B. nfixn, A. chorc~oct7.ti.s. B. 

deciwtatn, B. fungicola, B. hypori~nea, R. imshcizrgii 

(Figure 1 b), R. isidiza, B. nieizo.rpom, B. pir,~tiila t a, B . .re n si b i 1 i.r, B . .re t sc h 7~ 1 n n c n .r i,s, R . .s i i b c omnata, B. siibinflatn, B. tnbncinn, and R. 7~entl-ico,sn, 

or more or less richly dichotomously In-anched, as 

in B. apophysata, B. ntrichelln, R. bicoi~~rtn, B. 

biilbochaeta, B. confoedercltn, R. coi-niitn, R. 

goebelii, B. klenientii, R. lncuignfiiln (Figure lc), 

B. schiffnel-i, I?. semilzrnatn, B. ,sztfixn, antl R. 

viiidescens. Branching is a useful and consistent 

taxonomic character. 

There is a close correlation bettveen chemistry 

and btanching pattern of the rhizines. All P+ 

1 
2 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 1.-Morphology of Rnlhothrix: n, cross section of 13. lnerjigntuln showing epicortex (arrow) 

(Allard 15992a in US) (X 500); b, simple rhizines of B. irnshaugii (Mnhu 3545 in US) (X 80); 

c, dichotomously branched rhizines of n. lnevigatiiln (Allard 15992a in US) (X 130); d, soralia of 

B. imsharigii (Rtmdel 7308 in US) (X 10); e, pustules of B. piistdata (holotype) (X 10); 

f, lobules of B. sufiixa (Hale 42753) (X 10); g, pycnidiate apothecia of B. uiridescens (Imshaug 

42530 in US) (X 10). (Photographs Q-c taken with scanning-electron microscope.) 
NUMBER 32 3 

species, that is, those containing norstictic acid, 

protocetraric acid, or salazinic acid, have simple 

unbranched rhizines. Most of these are Old World 

species. On the other hand, the remaining species, all of which are P- and contain gyrophoric 

acid, lecanoric acid, similar orcinol depsides and 

depsidones, or only atranorin, have richly branched 

rhizines and occur predominantly in the Nelv 

World. The only exceptions are B. afixa and B. 

fungicola, which have mostly simple rhizines 

although they contain gyrophoric acid. As far as 

I know, no other parmelioid genus shows such a 

clear-cut correlation between chemistry and morphology except the Pnrrnelza borrerz group (Hale, 

1965a), which shows a direct correlation between 

a pale lower surface and the presence of lecanoric 

acid and a black lower surface with gyrophoric 

acid. 

VFGETATII F PRoP.uuLEs.--Isidia are regularly 

produced in B apophysata, B. decititata, B. fltngtcola, B. goebelii, B. isidiza, B. klementii, B. lneviqntiiln, B. pigmentacen, B. szibinpatn, B. tnbacina, 

and B. uentrirosa. They are cylindrical and usuall\ 

unbranched but become procumbent and nearlv 

dorsiventral in B. fungicola. Dense lamina1 lobules 

without isidia occur in B. sitfixa (Figure If). 

These are ti ue lobules since they are corticate and 

may produce marginal bulbate cilia. 

Pustules are found in only one species, B. pustitlata. These large, inflated structures (Figui e 

le) tend to break open apically and seem identical 

with those described in the genus Hypot)achyna 

(Hale, 1975a:6). Thev do not form soredia. 

Soredia occur only in B. irnshairgii, a Chilean 

endemic. The soralia are rather diffuse over the 

upper surface toward the lobe tips, producing. 

large patches of farinose soredia (Figure 1 d). 

A rather large percentage of the species in the 

gent15 produce no special vegetative propagules 

other than small adventitious lobules so common 

in all paimelioid genera. This group includes B 

afixn, B at?ichella, B bicorniita, B. biilbochnetn, 

B. cho7ooenrir, B. confoederata, B. coronntn, B. 

hypocram, B. mezzospora, B. sztbcoronata, B 

schiffneri, B. yemilunata, B. yensibiliy, B. setsch7cwnensis, and B. viridescen y. ,411 have abundant 

a po theci a. 

APOTHECIAI CHARACTER$ -The apothecia of Bitlbothiix are uniformlJ small (rarely more than 4 

mm in diameter) and sessile or adnate. In a few 

species they are almost substipitate because of a 

constricted base. Pycnidia (coronate apothecia) 

are always produced around the rim in B. afixa, 

B. bulbochaeta, B. CoTonnta, B. fungicola, B. schiffneri, B. .semilzinnta, B. subcoronato, B. ventricosa, 

and B. viridescens. The thalli of these species (and 

in the case of B. biilbochnetn, B. coronata, and B. 

viridescens (Figure 1 g) the amphithecium) are 

sometimes heavily pycnidiate with normal immersed pycnidia. These apothecial pycnidia are 

apparently the same as those I described for Relirina (Hale, 19751,:8), the only other parmelioitl 

genus with this trait. 

Ecoronate apothecia are characteristic of the 

remaining 15 fertile species. Bzilbothrix npopkysata, B. decirrtntci, B. klementii, and B. piginentncea have never been collected with apothecia. 

There is no obvious correlation between chemistry and presence of coronate apothecia, but the 

geographical correlation is strong. All coronate 

species, excepting B. afixa from Angola and A. 

bitlbochaeta from India, are endemic to or more 

common in the New TZrorltl. Most species endemic 

to the Old T\'orlcl are ecoronate. 

The spores of Ritlbothi.ix are colorless, onecelled, and either ovoid or bicornute. Ovoid 

spores are rather small, 5-8 ym wide and 6-12 pm 

long on the average. Only B. bulbochnftn, R. 

ch07~~oen.sis, B. confoederntn, and 23. Iat4gatitla 

have spores no more than 5-7 ym long, the typical range for species of Relicinn (Hale, 197511). 

As a rule, species with salazinic acid, these being 

largely endemic to the Old \l'orld, have the largest 

spores, 8-20 pm long. Bicornute spores are known 

for three apparently closely related Brazilian species, B. bicornirtn, B. .tchifj'neri, and B. .rrmilirnnta. 

These spores are quite variable in shape, from 

nearly ovoid ivitli two short but distinct horns to 

long, narrow, and semi-lunate. This type of spore 

has recently heen discovered in trvo species of 

Relicinn (Hale, 1975b), but other than these five 

examples no other species in any parmelioid 

genera have bicornute spores. \Ye cannot, however, determine the evolutionary significance of 

unique spores until more is known of their cytology and ontogen?. 

Chemistry 

The chemistry of Biilbothrix is the least varied 
4 ShlITHSONIAS CONTRIBUTIONS TO BOTAXY 

of all the epicorticate parmelioid genera. Only 

twelve lichen substances, as enumerated below, 

have been detected. By comparison, the closely 

related bulbate-ciliate genus Relicina, with almost 

the same number of species, has at least 20 different substances, and of these, only atranorin, 

gyrophoric acid, norstictic acid, protocetraric acid, 

and salazinic acid are shared by Bztlbothrix. For 

further comparison, the completely unrelated 

genus Hypotrachyna has 39 lichen substances (Hale, 

1975a.) 

Atranorin: All species in the genus and the main component 

Gyrophoric acid: B. afixa, B. atrichella, B. coronata, B. funLecanoric acid: R. bicornrcta, B. confoederata, and B. laeuiLobaric acid: B. apophysata. 

h’orlobaridone: B. decurtata (as a iare accessory substance 

with salazinic acid). 

Sorstictic acid: B. subcoronata and B. ventricosa (probably 

accompanied by the unknown substance connorstictic acid). 

Protocetraric acid: B. chowoensis and B. subinpata. 

Salazinic acid: B. decztrtata, B. hypocraea, B. imshaugii, 

B. isidiza, B. meizospora, B. pustulata, B. sensibilis, B. setschwanensis, and B. tabacina (rarely accompanied by the 

unknown substance consalazinic acid, an H,SO, + reddish 

spot that remains at the point of origin in the usual 

chromatographic solvents). 

in B. bulbochaeta and B. viridescens. 

gicola, B. goebelii, B. sckniffneri, and B. sufixa. 

gatula. 

Skyrin (rhodophyscin): B. semdunata. 

Unidentified anthraquinone: B. fiigrnentacea. 

On the whole the genus exhibits a relatively 

low level of evolution in terms of chemistry (Hale, 

1966). This is reflected both in the abundance 

of gyrophoric and salazinic acids, relatively 

“primitive” substances, and in the lack of biogenetically more “advanced” substances such as 

alectoronic acid, barbatic acid, divaricatic acid, 

evernic acid, lichexanthone, perlatolic acid, and 

stictic acid, as well as the protocetraric acid derivatives fumarprotocetraric acid and succinprotocetraric acid. 

On further analysis we can see that the geographical distribution of the acids follows a definite trend. Salazinic acid, for example, predominates in the species endemic to the Old World, 

and the two species with protocetraric acid (B. 

chowoensis and B. subinflata) are also confined 

there. On the other hand, lecanoric acid, the two 

orcinol depsidones colensoinic acid and lobaric 

acid, and norstictic acid are all restricted to species 

endemic to the New World (North and South 

America). The only exception to this generalization is the occurrence of norlobaridone with salazinic acid in the African B. decurtata. 

Two pantropical species, B. isidira and B. 

tabacina, have salazinic acid and one other, B. 

goebelii, has gyrophoric acid. These differences 

suggest strongly that the Bulbothrix floras of the 

New and Old Worlds have been separated for a 

long period of time and deprived of opportunites 

for gene exchange that might increase chemical 

diversity. In any event, the New World species 

have been evolving more rapidly in terms of chemistry than the conservative Old World species. 

Morph Formation 

As I explained in studies of Hypotrachyna 

(Hale, 1975a), Pseudoparmelia (Hale, 1976), and 

Relicina (Hale, 1975b), speciation in the parmelioid genera may be attributed in large degree to 

the formation of isidiate and/or sorediate morphs 

from fertile sexual parents. In comparison with 

other genera, however, Bulbothrix, with relatively 

fewer species, has not undergone extensive morph 

formation, and this mechanism has contributed 

relatively little to speciation in the genus. 

There are, for example, an unusually high number of fertile parent species that could give rise to 

vegetative morphs but have not done so, judging 

at least from the collections now available. About 

a third of the species (11) fall into this category: 

B. bicornuta, B. bulbochaeta, B. chowoensis, B. 

coronata, B. klementii, B. meirospora, B. schiflneri, 

B. semilunata, B. setschwanensis, B. subcoronata, 

and B. viridescens. The next largest group, five species, B. apophysata, B. decurtata, B. pigmentacea, B. 

subinflata, and B. ventricosa, are isidiate species 

which lack a corresponding nonisidiate parent. 

The assumption here is that such parents did exist 

at some stage in the evolutionary history of the 

genus but have not survived to this time. A similar 

situation holds for pustulate B. pustulata, sorediate B. imshaugii, and lobulate B. sufixa, none of 

which appears to have an extant parent. 

Finally, four species pairs represent nonisidiate 

parent-isidiate morph series: B. afiixa - B. fungicola, B. atrichella - B. goebelii, B. hypoel-aea - B. 

isidiza, and B. sensibilis - B. tabacina. Except for 

lobe width differences, B. confoederata and B. 

Zaevigatula also form a plausible parent-isidiate 
NUMBER 32 5 

morph pair. As I remarked in my monograph on 

Hypotrachyna (Hale, 1975a), matching parent 

morphs and their presumptive vegetative morphs 

is often quite subjective since minor morphological differences are bound to have arisen in the 

course of evolution because of adaptations to 

changing environments, modifications of the parent morph through genetic recombination, etc. 

Phytogeography 

Bulbothrix is not a commonly collected genus. 

Even where it is ideally developed, as in the semiarid scrub forests of Brazil, no species predominate 

in the vegetation as does, for example, Hypotruchyna in the cloud forests of the Andes (Hale, 

1975a) or Relicina in the dipterocarp forests of 

the Philippines (Hale, 1975b). One must search 

with great care to find any specimens of Bulbothrix at all. Another measure of its comparative 

rarity is the fact that few major herbaria have 

more than a dozen sheets on file. None of the species occurs in Europe, where collecting intensity 

has been high, and few lichenologists have had 

the opportunity to collect in the tropical regions 

where Bulbothrix does occur. At the same time, 

collections by some contemporary lichenologists, 

especially in Africa where I have not collected, 

were unavailable for this monograph. Consequently, many of the statements made here on 

species abundance and distribution will be tempered by the results of other workers in the future. 

Generalizing, then, from what cannot be considered wholly adequate collections, we find that 

Bulbothrix resembles Parmotrema and Pseudoparmelia (Hale 1976) in showing highest develop 

ment in semi-arid woodlands and secondary forests. It appears that the cerrado vegetation of 

South America and the upland scrub forests of 

south-central Africa are ideally suited for Bulbothrix. Unlike Pseudoparmelia, however, the greatest number of species is known from Brazil (Fig 

ure 2), less from South Africa, and only one from 

Australia. The noteworthy point to be made here 

FIGURE 2.-Number of species of Bdbothrix in major geopolit'ical regions (see also list of species 

in each country in text). 
SMITHSONIAN CONTRIBUTIOXS TO BOTANY 

is that the morphologically similar genus Relicina 

has an almost completely separate distribution in 

the rain forests of Southeast Asia where Bulbothrix is rare. 

The Bulbothrix floras of each geopolitical unit 

where the genus has been found are as follows. 

NORTH AND SOUTH AMERICA 

United States: B. confoederata, B. coronata, B. goebelii, and 

Mexico: B. coronata, B. goebelii, B. isidiza, B. laevigatzila, 

Guatemala: B. goebelii, 11. isidiza, and B. sufixa. 

Honduras: B. sufixa. 

Costa Rica: N. ventricosa and “Parmelia stenophylla” (see 

Panama: A. npophysata, B. goebelii, and B. ventricosa. 

Cuba: I1. goebelii, B. laevigatiila, B. sufixa, and B. tabacina. 

Jamaica: B. goebelii, B. laevigatula, B. sufixa, and B. tabaHispaniola: B. apophysata, B. goebelii, B. laevigatula, B. sufLesser Antilles and Trinidad: B. apophysata, B. goebelii, 

Colombia: B. atrichella, B. goebelii, and B. laevigatula. 

Venezuela: B. goebelii, B. laevigntzila, B. sensibilis, B. sufixa, 

Peru: 11. atrichella, B. coronata, B. goebelii, and B. laeviEcuador: B. laevigntula. 

Guyana: B . la evigat uln , 

Brazil: B. bicornuta, B. coronnta, B. fungicola, B. goebelii, 

B. Iiypocraea, B. isidirn, B. klementii, B. laevigatula, B. 

schiffneri, 13. seniiltinata, B. sr~bcoronata, B. sicfixa, B. tabacinn, and B. uiridescens. 

B. laevigatiila. 

B. tabacina, and B. ventricosa. 

page 00). 

cinn. 

fixa, B. tabacina, and B. ventricosa. 

B. laevigatula, B. szifixa, and B. tabacina. 

B. tabacinn, and B. ventricosa. 

got ZL la. 

Uruguay: B. viridescens. 

Paraguay: B. coronata, B. fungicola, B. goebelii, R. isidiza, 

Chile: n. goebelii and B. irnsliaiigii. 

.4 rgen t ina : B . sii bcorona to. 

B. sitbcoronnta, and B. stifixa. 

AFRICA 

Sierra Leone: B. goebelii. 

Ivory Coast: 13. decitrtnta and B. isidiza. 

Guinea: B. sensibilis and B. tabacinn. 

Cameroon: n. ineizosporn. 

Uganda: 11. decurtatn, B. hypocraen, B. isidiza, and B. tabaUrundi: 13. pirstttlata. 

Kenya: B. isidizn and B. sensibilis. 

Zaire: n. Iiypocraea, 13. isidizn, and B. sensibilis. 

Zambia: B. cliorcoet~sis, B. Iiypocraea, B. isidizn, and 23. sensiMalawi: 11. isidizn and B. sensibilis. 

Angola: B. ofiixn, B. hypocraea, 13. isidiza, B. sensibilis, and 

Rhodesia: B. liypocraen and B. isidizn. 

cinn. 

bilis. 

B. tabncina. 

Tanzania: B. hypocraea, B. isidiza, and B. tabacina. 

Mozambique: B. chowoensis. 

Union of South Africa: B. coronata, B. decurtata, B. goebelii, 

Madagascar: I#. tabacina. 

Mauretius: 23. sufixa. 

B. isidiza, B. sufixa, B. tabacina, and B. ventricosa. 

ASIA 

Pakistan: B. meizospora. 

Nepal: 13. isidiza, B. meizospora, B. setschwanensis, and B. 

ta bacina. 

India: B. bulbothrix, B. isidiza, B. meizospora, B. setschwanensis, and B. tabacina. 

Thailand: B. goebelii. 

Indochina: B. isidiza. 

Malaysia: B. goebelii, B. isiddza, B. pigmentacea, B. subinIndonesia: B. goebelii, B. isidiza, and B. tabacina. 

Philippines: B. goebelii, B. isidiza, B. fiigmentacea, B. subinflatn, and B. tabacina. 

New Guinea: B. goebelii. 

Taiwan: B. goebelii, B. isidiza, and B. tabacina. 

China: B. setschwanensis. 

Japan: B. isidiza. 

Australia: B. tabacina. 

Hawaii: B. isidiza. 

Pacific Area: B. goebelii and B. isidiza. 

flatn, and B. tabacina. 

The overall breakdown may be summarized as 

follows: The New World (North and South 

America) has a total flora of 19 species, 11 of these 

endemics, 5 also occurring in Africa, and 3 pantropical. Africa has a total of 13 species, 4 of them 

endemic, 5 also occurring in the New World, 1 

shared with Asia, and 3 pantropical. All of Asia 

has a total flora of 8 species, 4 of them endemic, 

1 shared with Africa, and 3 pantropical. The 

three commonest species are B. goebelii, known 

from 23 countries, B. isidiza from 21, and B. tabacina from 16. These are the three pantropical 

species. Eight species are known from only their 

type localities or, at most, one other collection: 

B. afixa, B. atrichella, €3. bicornirta, B. bulbochaeta, B. chozuoensis, B. klementii, B. pwstulata, 

B. schiflneri, and B. semilzinata. 

Classification of Bulbothrix 

Species now recognized in Bzilbothiix were 

previously classified in the collective genus Parmelia. The first lichenologist to sense an affinity 

between some of them was Lynge (1914), who 

did so, however, on the basis of bicornute spores 

(as opposed to ovoid) in three species, B. bicoi- 
NUMBER 32 

nuta, B. schiflneri, and B. semilunata. He grouped 

these three species in section Bicornuta, coordinate with section Amphigymnia, but overlooked 

the marginal bulbate cilia characteristic of all 

three species. He also failed to recognize bulbate 

cilia in Brazilian specimens of Parmelia acariospora (= B. goebelii), P. coronata (= B. COSOnata), P. fungicola (= B. fungicola), and P. virtdescens (= B. viridescens), all of which he either 

identified or described as new. He did make 

note of coronate apothecia where they occurred. 

If we go back to Fee’s descriptions (1824 antl 

1837) of the first two bulbate species to be described, we discover that he correctly clemibed 

coronate apothecia in Parmelia coronatn and P. 

glandzilifera and possibly marginal bulbate cilia 

in P. glandzilifera (= B. coronata). Zenker 

(1 827) noticed the richly branched rhirines of 

B. goebelii and their position on the lobe margins. 

Neither Muller Aargau nor Nylander recognized 

bulbate cilia in any of the species they described, 

although Nylander mentioned them for Parmelici 

circiimnodata (= Relicina ciscumnodata (NJlander) Hale). Vainio (1 890) meticulously described the morphology of Parmelia coronatu antl 

P. coronata var. isidiosa Muller Argoviensis (the 

latter, incidentally, a misidentified Parnzelinn), 

as well as marginally bulbate Parmelia abTtriiTa 

(= Relicina abstriisa (Vainio) Hale), but failed 

in all these instances to observe bulbate cilia on 

the lobe margins. 

While Zahlbruckner (1909) did not attempt to 

group marginally bulbate species, he compared 

Parmelia cchiffneri (= B. schiflneri) with Pal melia relicinella (= Relicinn relicinella (Sylander) 

Hale), a marginally bulbate species with usnic 

acid, Parmelia coronata (spores ovoid), and P. 

bicornuta (lobes twice as broad as in P. rchiflnen’ 

and medulla C+ red). In this case, too, Zahl7 

bruckner seems to have sensed a close affinity for 

these species without observing the marginal hdbate cilia. He did not mention these cilia for 

Parmelia acariosporn (= B. goebelii) either, 

when describing it as new. 

Although Dodge (1959:89) recognized bulbate 

(“conic”) cilia on the lobes of Pamelia .slrblnevigatoides (= B. tabacina), he did not see them on 

other species of Btilbothrix that lie described. 

Hale and Kurokawa (1964) were apparently 

the first lichenologists to recognize mai~ginally 

bulbate species as a natural group. I recently 

separated the usnic acid-containing species as ;I 

distinct genus, Relicina (Hale and Kurokawa) 

Hale (Hale, 1975b), leaving the usnic acid-free 

species as the nucleus of B1tlbotksi.l: (Hale, 1974). 

The chief features of B7tlbothri.x are the marginal bulbate cilia, small, adnate to appressetl 

thallus, adnate imperforate apothecia antl frequent occurrence of coronate apothecia, absence 

of usnic acid in the cortex (antl conversely prev 

ence of atranorin), variable color of the 1o.iver 

surface (pale brown or black), and simple or 

branched rhizines. Anatomically the genus is tlistinguishetl by a palisade upper cortex overlaid by 

a pored epicortex (Figure In) (Hnle, 1973). 

The unique chemical profile is discussed under 

“Chemistry” (above). 

Bulbothrix as circumscribed in this monograph 

is most closely related to Relirina in morphology, 

these being the only two parinelioid genera witli 

bulbate cilia, coronate apothecia, antl bicornute 

spores. However, as I emphasized in my study of 

Relicina (Hale, 3975b), the gener;i are almost 

totally different in ecological requirements and in 

geography. Most species of Relirina have evolved 

in the evergreen rain forests of Southeast Asia. 

Their chemical profiles are strikingly different, 

making highly unlikely any significant gene exKey to the Species 

SPECIES WITHOUT ISIDIA, SOREDIA, OR PUSTULES 

1. Apothecia present; spores bicornute. 

2. Medulla C negative B. semilunata 

2. Medulla C+ rose or red. 

B. bicornuta 

B. schiflneri 

3. Lecanoric acid present: lobes 1-2 mm wide; apothecial rim ecoronate 

3. Gyrophoric acid present; lobes less than 1 mm wide: apothecial rim coronate 

1. Apothecia, if present, with ovoid or elliptical spores. 

4. Rhizines richly branched. 
8 SMITHSONIAN COKTRIBUTIONS TO BOTANY 

5. Medulla C+ rose or red. 

6. Lobes 1 mm wide or less; lecanoric acid present B. confoederata 

6. Lobes 1-2 mm wide; gyrophoric acid present. 

7. Apothecia coronate .............................................................................. B. coronata 

7. Apothecia ecoronate ............................................................................. B. atrichella 

8. Collected in India .................................................................. B. bulbochaeta 

5. Medulla C negative. 

8. Collected in tropical America. 

9. Rhizines densely branched; tips of bulbate cilia often branched ...... B. klementii 

9. Rhizines simple to sparsely branched; tips of bulbate cilia simple . B. viridescens 

4. Rhizines simple or sparsely furcate. 

10. Apothecia coronate. 

11. Medulla Kf yellow turning red ........................................................... B. subcoronata 

12. Lobes 0.5-1 mm wide; collected in Africa ........................................... B. afixa 

Lobes 1-2 mm wide; collected in South America ................ B. viridescens 

13. Medulla K negative (protocetraric acid) ...................................... B. chowoensis 

15. Lobes sublinear; spores 7-9 pm long .......................................... B. sensibilis 

15. Lobes broader, subirregular; spores 14-20 km long ............ B. meizospora 

pm long ..................................................................................... B. hypocraea 

11. Medulla K negative. 

12. 

10. Apothecia, if present, ecoronate. 

13. Medulla K+ yellow turning red (salazinic acid). 

14. Lower surface black with at most a narrow marginal brown zone. 

14. Lower surface pale brown. 

16.Lobes more or less sublinear, often crowded, 1-3.5 mm wide; spores 8-14 

16. Lobes broader, 2-5 mm wide; spores 12-19 km long ......... B. setschwanensis 

SPECIES WITH ISIDIA, SOREDIA, OR PUSTULES 

B. imshaugii 

2. Thallus pustulate B. pustulata 

2. Thallus not pustulate. 

1. Thallus sorediate 

1. Thallus not sorediate. 

3. Thallus isidiate, the isidia becoming procumbent and dorsiventral. 

4. Isidia fine, intermixed with thin, often branched lobulate isidia (Figure 3d) 

4. Isidia entirely lobulate (Figure If) 

3. Thallus isidiate, the isidia erect, cylindrical. 

5. Medulla Kf yellow tuining red (salazinic or norstictic acids). 

B. fungicola 

B. sufixa 

6. Lower surface brown B. isidiza 

6. Lower surface black with at most a narrow brown zone at the margins. 

7. h’orstictic acid present B. ventricosa 

7. Salazinic acid present. 

B. tabacina 

B. decurtata 

8. Thallus usually corticolous; isidia normal 

8 Thallus saxicolons; isidia blackening at the tips 

5. Medulla K negative. 

9. Medulla C+ rose or red. 

10. Lecanoric acid present 

10. Gyrophoric acid present. 

B. laevigatula 

11. Rhizines richly branched, often pale; isidia uniformly erect and cylindrical B. goebelii 

11. Rhizines sparsely branched, black; isidia in part procumbent and dorsiventral B. fungicola 

B. subinflata 

9. Medulla C negative. 

12. Lower surface brown; medulla P+ red (protocetraric acid) 

12 Lower surface dark brown to black; medulla P negative. 

13. Lobes appressed, less than 1 mm wide; collected in Southeast Asia 

13. Lobes adnate, 1-2 mm wide; collected in tropical America 

B. pigmentacea 

B. apophysata 
NUMBER 32 9 

change in the past evolution of the two genera. 

If we look for a parmelioid group with as 

"primitive" an assemblage of lichen substances as 

in Bulbothrix, it would be Parmelina Hale, another genus common in secondary, though more 

temperate forests, with a comparably small thallus 

but with normal marginal cilia and simple rhizines. I doubt very much, however, that Bulbothrix and Parmelina have evolved from a common 

ancestor or undergone a significant amount of 

generic hybridization. 

Taxonomic Treatment 

Bulbothrix 

Bulbothrix Hale, 1974:480. 

Parmelia section Bicornuta Lynge, 1915:17 [type-species: 

Parmelia bicornuta Lynge (= Bulbothrix bicornuta 

(Lynge) Hale)]. 

Parmelia subgenus Bicornuta (Lynge) Gyelnik, 1932:219. 

Parmelia subgenus Parmelia section Imbricaria subsection 

Bicornutae series Bicornutae Hale and Kurokawa, 19643135. 

The 29 presently known species of Bulbothrix 

are listed below in alphabetic order. Chemistry 

was determined with thin-layer chromatography 

in two solvent systems, hexane-ether-formic acid 

and benzene-dioxane-acetic acid. The lichen substances reported are those for the type material, 

unless otherwise stated. Locations of specimens 

examined are cited by the standard herbarium 

acronyms; collections by Hale are all preserved in 

US and the acronym is not given. 

Bulbothrix afixa 

FIGURE 3a 

Bulbothrix afixa (Hale and Kurokawa) Hale, 1974:428. 

Parmelia afixa Hale and Kurokawa, 1964:137 [based on 

Parmelia coronata var. denudata Vainio]. 

Parmelia coronata Fee var. denudata Vainio in IVelwitsch, 

1901:401 [type collection: Morro de Lopollo, Huila, .4ngola, 

Welwitsch 33 (TUR, lectotype; BM, isolectotype)]. 

DEscRIPTIoN.-Thallus very closely adnate, corticolous, light mineral gray, 2-3 cm in diameter; 

lobes sublinear, 0.3-1 mm wide; bulbate cilia 

dense; upper surface plane, continuous or cracked 

on older lobes; lower surface black, moderately 

rhizinate, the rhizines mostly simple, black. Apothecia numerous, adnate, 0.5-1.5 mm in diameter, 

the exciple coronate, the base retrorsely rhizinate; 

spores 8, 4-5 X 8-10 pm. 

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

C - , KC + faint rose, P - , atranorin and gyrophoric 

acid. 

HABITAT.-oII trees in open areas. 

DI~TRIBUTI~N .-Angola. 

REMARKS.-This small species, the only coronate 

one endemic to Africa, is still known only from 

Angola. It probably occurs more commonly there 

and elsewhere in Africa but would be overlooked 

because of the small size. It may be considered as 

the presumptive parent morph of B. fungicola, a 

rare New World species. The other nonisidiate 

species with gyrophoric acid in Africa, B. coronata, 

is significantly larger and has branched rhizines. 

SPECIMENS ExAbriNED.-Angola: Huila, Degelius (Degelius 

herbarium, US). 

Bulbothrix apophysata 

FIGURE 3b 

Bulbothrix apophytata (Hale and Kurokawa) Hale, 1947:480. 

Parmelia apophysata Hale and Kurokawa, 1964:138 [type 

collection: Piedra Blanca, La Vega, Dominican Republic, 

Allard 16073 (US, holotype; TNS, isotype)]. 

~ESCRIPTION.-Tha~~us closely adnate, corticolous, light mineral gray, 3-6 cm broad; sublinearelongate, 0.7-1.5 mm wide; bulbate cilia distinct, 

numerous; upper surface plane, faintly maculate, 

irregularly cracked on older lobes, sparsely to 

moderately isidiate, the isidia simple, less than 

0.3 mm high; lower surface variable, usually 

blackening in the center, dark brown in a broad 

marginal zone, densely rhizinate, the rhizines pale 

brown to black, densely branched. Apothecia not 

seen. 

C-, KC+ rose, P-, atranorin and lobaric acid. 

HABITAT.-on trees in pastures and disturbed 

areas at 200-1500 m elevation. 

DrsTRIsuTIoN.-Dominican Republic, Trinidad, 

and Panama. 

REMARKS.-This species has so far been collected 

only in secondary forests. The chemistry is unique 

in the genus and no nonisidiate morph is extant. 

Three other isidiate species within the range of 

B. apophysata would be distinguished as follows: 

CHEMISTRY.-cOrteX K + yellow, medulla K- , 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 3.-Species of Bulbothrix: a, B. afixa (lectotype in BM) (X 4); b, B. apophysata (Hale 

38873); c, B. atrichella (Lindig 110 in H); d, B. bicornuta (lectotype in G) (X 1); e, B. bulbochaeta (holotype in US); j, B. chowoensis (isotype in US). (Scale for b, c, e, f in mm.) 

B. laevigatula is somewhat smaller and more Bulbothrix atrichella 

appressed and contains lecanoric acid; B. goebelii 

has a more fragile thallus with a darker, sometimes 

olivaceous color, and produces gyrophoric acid; Bulbothrix atrichella (Nylander) Hale, 1974:480. 

FIGURE 3c 

and B. tabacina has simDle black rhizines and Parmelia atrichella Nylander, 1885:614 [type collection: CO- I 

contains salazinic acid. lombia, Lindig 110 (H, Nylander herbarium 35233, lectotype; FH, BM, M, isolectotypes)]. -. 

SPECIMENS EXAMINED.-Dominican Republic: La Vega, 

Allard 18032 (US). Trinidad: Zmshaug 32318 (MSC). Panama: 

Chiriqui, Hale 38873; Veraguas, Hale 45941. 

DEscRIPTIoN.-Thallus closely adnate on bark, 

light mineral gray, turning olive-buff in the her- 
SUMBER 32 11 

barium, 3-6 cm in diameter; lobes sublinearelongate, 0.5-2 mm wide, the margins densely bulbate, sometimes lobulate; upper surface plane, 

shiny, maculate; lower surface black, densely 

rhizinate, the rhizines densely branched. Apothecia adnate, 1-2.5 mm in diameter, the disc carob 

brown, ecoronate; spores 8, 5 X 7-8 pm. 

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

C+ rose, KC+ red, P--, atranorin and gyrophoric 

acid. 

HABITAT.-on trees in open forest at about 2000 

m elevation. 

DIsTRisuTioh..-Colombia and Peru. 

REMARKS.-This species is known from only two 

collections. It is very close to B. coronata but lacks 

coronate apothecia. The upper surface is rather 

distinctly maculate and the thallus color more 

olivaceous than in B. coronata. Bulbothrix goebelii may well be the isidiate morph. 

(US). 

SPECIMENS EXAMINED.-PerU: San Martin, Allard 21472 bis 

Bulbothrix bicornuta 

FIGURE 3d 

Bzilbothrix bicornuta (Muller Argoviensis) Hale, 1974:480. 

Parmelia bicornuta Muller Argoviensis, 1891:377 [type collection: Rio de Janeiro, Brazil, Leyland (BM, lectotype; G, 

isolectotype)]. 

DEscRwTION.-Thallus adnate on bark, whitish 

mineral gray, 4-6 cm in diameter; lobes sublinearelongate, 0.7-1.5 mm wide; bulbate cilia dense, 

strongly inflated, becoming apically branched; 

upper surface plane, shiny, continuous; lower 

surface black, densely short rhizinate, the rhizines 

branched. Apothecia adnate, 1-4 mm in diameter, 

the exciple ecoronate; spores 8, bicornute, 3-4 X 

15-19 pm. 

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

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

HABITAT.-on trees in open forest. 

DIsmIBuTioN.-Brazil. 

REMARKS.-The most distinctive feature of this 

rare species-it is still known only from the typecollection-is the bicornute spores. The other two 

species in the genus with bicornute spores, B. 

schiffneri and B. semilunata, have different chemistry and much narrower lobes. It is probably 

unrelated to lecanoric acid-containing B. confoederata, a North American endemic which has 

ovoid spores and narrow lobes (0.5-1.0 mm wide). 

Muller measured the spores as listed above in 

the description, but my own examination gave 

smaller spores, about 8 pm long. 

Bulbothrix bulbochaeta 

FIGURE 3e 

Bulbothrix bulbochaeta (Hale) Hale, 1974:480. 

Parmelia bulbochaeta Hale in Hale and Kurokawa, 1964:138 

[tjpe collection: Shembaganur, Madurai District, India, 

Armsthi 4347 (Awasthi herbarium, holotype; US, isotype)]. 

DEscRIPTIo”.-Thallus adnate to loosely adnate, 

corticolous, coriaceous, mineral gray, 4-8 cm 

broad; lobes sublinear, crowded, 1.5-2.5 mm wide; 

bulbate cilia conspicuous, simple or apically 

branched; upper surface plane, shiny, continuous, 

usually heavily pycnidiate; lower surface black 

except for a narrow brown zone along the margins, densely rhizinate, the rhizines dichotomously 

branched. Apothecia adnate, 2-3 mm in diameter, 

the exciple coronate; spores 8, 4 X 5 pm. 

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

C-, KC-, P-, atranorin. 

HABITAT.-on trees in secondary forest at 2000 

m elevation. 

DIsTRIBuTI0N.-India. 

REMARKS.-This is the only member of Bulbothyix endemic to India. It is characterized by the 

rather coriaceous thallus and lack of medullary 

substances. I attempted to recollect the species in 

1974 in the Palni Hills of South India, but it is 

either very rare or has been (or soon will be) 

destroyed by continued deforestation in the 

region. 

Bulbothrix chowoensis 

FIGURE 3f 

Bulbothrix chowoensis (Hale) Hale, 1974:480. 

Parnzelia chowoensis Hale, 1972:343 [type collection: Nyika 

Plateau, Zambia, Jellicoe 53 (BM, holotype; US, isotype)]. 

DEsCRiPTIoN.-Thallus adnate, corticolous, coriaceous, light mineral gray, 4-6 cm broad; lobes 

subirregular, contiguous or subimbricate, 2-2.5 

mm wide; upper surface plane, shiny, emaculate; 

lower , surface brown, moderately rhizinate, the 
SMITHSONAN COXTRIBUTIONS TO BOTANY 12 

rhizines brown, simple. Apothecia numerous, adnate to subpedicellate, ecoronate, 2-4 mm in 

diameter; spores 8, 3-4 X 6-7 ym. 

C-, P+ red, atranorin and protocetraric acid. 

HABITAT.-oII branches of trees in open forest. 

DIsTRIsuTIoN.-Zambia and Mozambique. 

Rnf.mKs.-This species superficially resembles 

B. hypocraea but has smaller spores and different 

chemistry (B. hypocraea is K+ red with salazinic 

acid). One other species in the genus, isidiate B. 

subinflata, contains protocetraric acid but would 

not be considered related to B. cho?uoensis because 

of narrower sublinear lobes and occurrence in the 

rain forests of Southeast Asia. 

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

SPECIlfESS EX~\rrsrD.-Mozambique: SandrOne (F, us). 

Bulbothrix confoederata 

FIGURE 4a 

Btclbothrix confoederatn (Culberson) Hale, 1974:480. 

Parmelia confoederata Culberson, 1961:169 [type collection: 

Shallotte, Brunswick County, North Carolina, Culbersoti 

9010 (DUKE, holotype: FH, TNS, US, isotypes)]. 

DEscRiPTIoN.-Thallus closely adnate on twigs, 

light mineral gray, 2-5 cm broad; lobes sublinear, 

0.5-1 mm wide; bulbate cilia dense, conspicuously 

inflated, apically branched; upper surface plane, 

continuous; lower surface black, densely rhizinate, 

the rhizines dichotomously branched. Apothecia 

common, 1-3 mm in diameter, the exciple ecoronate; spores 8, 3-5 X 5-7 pm. 

CHE;MIsm\r.-Cortex K + yellow, medulla K - , 

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

HABITAT.-on tree branches and small shrubs 

near sea level in open areas. 

DlsTRIBuT1oN.-Southeastern United States. 

REhfARKS.-This is the only species in the genus 

endemic to North America. It had been completely overlooked prior to its description in 1961 

although sparsely represented in herbaria under 

such names as “Parmelia laevigata.” It occurs 

commonly on small exposed shrubs in a zone back 

from the shoreline along the Gulf and Atlantic 

coasts. Because of the similar Chemistry, one 

might be tempted to compare it with B. bicornitta, 

but the spores are ovoid, not bicornute, and the 

thallus significantly smaller. One might also consider it as the nonisidiate parent morph of lecanoric acid-containing B. laevigatula, a somewhat 

larger ecoronate species surely not far removed. 

SPECIMENS EXAMIXED.-united States: Virginia, Reed 59656 

(Reed herbarium); Xorth Carolina, Czdberson 5169 (DUKE, 

US); South Carolina, Culberson 10022 (DUKE, US); Georgia, 

Skorepa and Folkerts 2475 (US): Alabama, Evans 98 (US), 

Hale 7262, 34193; Florida, Calkins 32 (WIS), Hale 16710, 

17050, 17643, 21661, 21823, 21868, 21868, 21943, Rapp (FLAS, 

US); Texas, Fisher 50159 (US). See Moore (1968:219) for 

additional records from Florida. 

Bulbothrix coronata 

FIGURE 4b 

Bitlbothrix coronata (Fee) Hale, 1974:480. 

Parmelia coronata Fee, 1824:123 [type collection: South AmerParmelia glandzrlifera Fee, 18243123 [type collection: Tropical 

Pnrnzelin appressa Zenker in Goebel and Kunie, 1827:157 

Parmelia relicina var. coronata (FCe) Muller Argoviensis, 

ica (G, lectotype)]. 

.4merica, Humboldt and Bonpland (G, lectotype)]. 

[tjpe collection: Peru (L, lectotype)]. 

1887b:318. 

DEscRIPTIoN.-Thall~is closely adnate on bark, 

3-6 cm in diameter; lobes sublinear-elongate, 

crowded, 0.5-2 mm wide; bulbate cilia distinct; 

upper surface plane to rugulose, continuous to 

faintly maculate, cracked on older lobes; lower 

surface black, moderately rhizinate, the rhizines 

moderately to densely branched. Apothecia adnate, 

1-3 mm in diameter, the exciple coronate; spores 

8, 5 X 6-8 ym. 

CHEMIsTRY.-Medulla K - , C + rose, KC 4- red, 

P--, atranorin and gyrophoric acid. 

HARITAT.-on trunks and branches of trees in 

semi-arid woodlands up to 2300 m elevation. 

DIsTRInUTIoN.-United States, Mexico, Brazil, 

Paraguay, and South Africa. 

REMARKS.-This was the first species described 

in the genus and the marginal bulhate cilia were 

recognized by Fee when he used the epithet 

glandulifern. It occiirs in the New World with a 

disjunct locality in South Africa. No vegetative 

morphs have been discovered. The only confusable species, B. bicornitta, B. schiflneri, and B. 

seniilunata, are all distinguished by bicornute 

spores. 

Muller (1 8871~318) was the first lichenologist 

to synonymize Pnrmelia glanditlifera under P. 

coronatn. He then reduced them to P. relicina var. 

coronata, emphasizing the coronate apothecia 
NUMBER 32 

FIGURE 4.-Species of Brtlbothrix: a, B. confoederata (Hale 33845); b, R. coronata (Hale 20505): 

c, B. decurtata (isotype in US); d, Zj. fungicola (isolectotype of Parmelia coronata f. isidiosa in 

BM) (X 10); e, B. goebelii (lectotype of Parmelia acariospora in IV): f, R. goebelii (Ravenel 

in US). (Scale in mm.) 

13 
14 SMITHSONIAZI CONTRIBUTIONS TO BOTANY 

common to these species but disregarding the yellow color (and different chemistry) of P. relicina 

(= Relicina relicinula (Muller Argoviensis) Hale). 

SPECIMENS EXAMINED.-United States: Texas, Anderson and 

Shushan 18742 (COLO, US), Wetmore 18231 (MINX, US). 

Mexico: Chiapas, Hale 20505; Michoacan, lltis 1893 (US, 

WIS); Oaxaca, Hale 20827, Iltis 2691 (US). Jamaica: lmshaug 

14299, 15007 (MSC). Brazil: Spruce 158 (BM); Mato Grosso, 

Malme (LD, S, US). For additional records from Brazil see 

Hale (1960:13). Paraguay: Balansa 4192 (P). Union of South 

Africa: Cape Province, Almborn 2879 (LD). 

Bulbothrix decurtata 

FIGURE 4c 

Bulbothrix decurtata (Kurokawa) Hale, 1974:480. 

Parmelia decurtata Kurokawa in Hale and Kurokawa, 1964: 

139 [type collection: 10 miles Fouthwest of Lydenburg, 

Transvaal, Union of South Africa, Alnzborn 7388 (LD, 

holotype; US, isotype)]. 

DEsCRIPTIoN.-Thallus adnate to closely adnate 

on rock, mineral gray, 3-7 cm broad; lobes subirregular to sublinear, 1-3 mm wide; bulbate cilia 

sparse to moderate, conspicuously inflated; upper 

surface plane, continuous or cracked on older 

lobes, moderately isidiate, the isidia cylindrical to 

granular, often black-tipped; lower surface dark 

brown to black in the center, densely rhizinate, 

the rhizines simple. Apothecia not seen. 

CHEMISTRY.-cOrteX K+ yellow, medulla K+ 

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

atranorin and salacinic acid, rarely with norlobaridone. 

HABITAT.-on rocks in semi-arid regions. 

D~smiBuTIoN.-Africa. 

REMARKS.-RUlbothriX decurtata is apparently 

the only obligately saxicolous species in the genus. 

The thallus is usually quite adnate and collected 

with the rock substratum. No parent morph i? 

known. One anomalous specimen from the Ivory 

Coast (Santesson 10485b) contains norlobaridone 

in addition to salazinic acid but is otherwise indistinguishable from the normal population. 

SPFCIMEhS EXAhf1SED.-Ivory Coast: Man, Santesson 10485b 

(UPS, US). Uganda: Kigezi District, Swinscow U 23/73 (BM, 

US). 

Rulbothrix fungicola 

FIGURE 4d 

Bulbothrix fungicola (Lynge) Hale, 1974:480. 

Parmelia fungicola Lynge, 1914:129 [type collection: Santa 

Anna da Chapada, Mato Grosso, Brazil, Malme 2438B 

(S, lectotype)]. 

Parmelia coronata f. isidiosa Muller Argoviensis, 1888:56 

[type collection: Paraguay, Balansa 4157 (G, lectotype; BM, 

M, W, isolectotypes)]. 

Parmelia pseudocoronata Gyelnik, 1931:289 [based on P. 

coronata f. isidiosa Miiller Argoviensis]. 

DEscRIPTIoN.-Thallus closely adnate on bark, 

olivaceous mineral gray in the herbarium, 2-3 cm 

broad; lobes sublinear, crowded, 0.5-1.5 mm wide; 

marginal bulbate cilia dense; upper surface plane, 

shiny, densely isidiate, the isidia initially erect 

and cylindrical but soon lobulate and dorsiventral, 

the procumbent lobules narrow, dichotomously 

branched; lower surface black, moderately branched, 

the rhizines simple to sparsely branched. Apothecia 

adnate, 1-3 mm in diameter, the rim coronate; 

spores 8, 4-6 X 8-10 pm. 

rose, KC+ red, P-, atranorin and gyrophoric 

acid. 

CHEMISTRY.-cOrteX K+ yellow, medulla c+ 

HABITAT.-on trees in open forest. 

DISTRIBUTION.-~~Test Indies and South America. 

REMARKS.-ThiS small, fragile lichen is easily 

confused with B. wffixn, a more widespread species differing in lobule formation without isidia 

(Figure If) and lack of pycnidia on the apothecia1 rim. Both are rather rare and more collections 

are needed to decide on their exact relationship. 

My report of bicornute spores in West Indian 

material (Hale, 1971b: 15) is probably incorrect, 

although I have not checked the specimen cited. 

The type of Parmelia fungicola definitely has 

ovoid spores; the type of P. coronata f. isidiosa 

has apothecia but no spores. 

SPFCIMENS ExAMINED.-See Hale (1971b: 15) for records from 

Dominica and the Dominican Republic. 

Bulbothrix goebelii, new combination 

FIGURE 4e,f 

Parmelia goebelii Zenker in Goebel and Kunze, 1827:134 

[type collection: “Auf officinellen Rinden,” Couth America, 

without collector, (M, lectotype)]. 

Parmelia papyrina Fee, 1837: 121 [type collection: tropical 

.4merica, (G, lectotype)]. 

Parmelia subdissecta Nylander in Nylander and Crombie, 

1883:51 [type collection: Tanjong, Malacca, Maingay (H, 

Nylander herbarium 35161, lectotype; BM, isolectotype)]. 

Parmelia granatensis Nylander, 1885:613 [type collection: 
NUMBER 32 15 

Socorro, Colombia, Li77dig (H, Sylander herbarium 35170; 

AM, PC, isolectotypes)]. 

Parinelia scortella Sylander, 1883:615 [type collection: Texas 

(FH-Tuck, lectotype; H, isolectotype)]. 

Parmelia addenda Vainio, 1907: I69 [type collection: Lem Dan, 

Koh Chang, Thailand, Schmidt X (C, lectotype; TUR, 

isolectotype)]. 

Partnelia acariospora Zahlbruckner, 1909: 169 [type collection: 

near Barra Mansa, Itapecirica, Sao Paulo, Brazil Schiffner 

(W, lectotype; US, isolectotype)]. 

Parinelia marginalis Lynge, 1914: 112 [type collection: Santa 

Anna de Chapada, Mato Grosso, Brazil, Malme 2393**** 

Parinelk njalensis Dodge, 195936.5 [type collection: Kori, 

Njala, Sierra Leone, Deightdn M.5642 (BM, lectotype)]. 

Bulbothrix papyrina (Fee) Hale, 1974:480. 

Bulbothrix scortella (Nylander) Hale, 1974:481. 

Aulbothrix stibdissecta (Nylander) Hale, 19743481. 

(S, lectotype)]. 

DEscRIPTIos.-Thallus closely adnate on bark, 

fragile, whitish mineral gray to olive-buff, 2-8 cm 

broad; lobes sublinear-elongate, 0.5-1.5 mm wide, 

the margins often filiform-dissected; bulbate cilia 

dense, becoming apically branched; upper surface 

plane, dull to clearly niaculate, moderately isidiate, the isidia simple or branched, erect, cylindrical, to 0.7 mm high, sometimes blackening apically; lower surface variable, dark brown to black, 

moderately to densely rhizinate, the rhizines pale 

brown, black with pale tips, or black and shiny, 

moderately to richly branched. Apothecia rare, 

adnate to sessile, 1-2 mm in diameter, the rim 

ecoronate (amphithecium rarely pycnidiate); spores 

8, 4-6 X 8-10 pm. 

C+ rose, KC+ red, P-, atranorin and gyrophoric 

acid. 

HABITAT.-& trees and more rarely on rocks 

in primary or secondary forests at lower elevations. 

DIsTRIBuTIos.-Pan tropical. 

REMARKs.-Bzdbothrix goebelii is a widespread, 

rather variable species characterized by isidia, 

branched rhizines, and gyrophoric acid. The New 

World and African populations tend to be very 

uniform with a brown to blackening lower surface 

and richly branched, often pale rhizines, as noted 

by Zenker in his description. The thallus is usually 

shiny pale olivaceous mineral gray. While some 

Asian specimens have these same characteristics, 

others, as in the types of Pal-melia addenda and 

P. szibdissecta and most specimens collected in the 

lowland dipterocarp forests of the Philippines, 

have a somewhat smaller, whitish thallus with a 

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

more consistently black lower surface and coarser, 

less branched rhizines. The amount of intergradation is so great, however, that 1 have decided to 

consider these extremes as part of a larger popiilation. Apothecia are not common in either the 

New or Old World. Of 80 specimens preserved in 

US only 9 were fertile. Seven of these were ecoronate and two had immersed pycnidin on the atnphithecium rather than strictly on the rim, much 

as in B. viridescens. 

SPECIMENS Ex.4xrIxrD.-~nited States: Tennessee, Sakanishi 

436 (Us); North Carolina, C2JlherSon .;051, 5276, 7701, 18016 

(DUKE), Ireland 4000 (US): South Carolina, ~illber,ioil 7801, 

10284 (DUKE), Hale 7614, 7625, 16,518, 16614, Ko.cJenel (ITS): 

Georgia, Hole 21982. Skorepa 2924 (US): Florida, Cnlkins 

(FLAS). Hale 21838, Ropp (FLAS, LTS); Alahama, Hole 7006, 

5197, 33919, 34025, 34102, Pursell 3916 (ITS); Mississippi, Hale 

7916: Louisiana, Hale 34010, Z.nngloit (US), Thieret 2461Gtl 

(US), Tucker 7247 (ITS): Texas, Hale 8169, 33803, 33885. 

Mexico: \'era Cruz, Hale 19401, 19979, 21230. Guatemala: 

Raja Vera Paz, Hale 38350. Panama: Chiriqui, Hale 38874, 

Maxon 5128 (US). Haiti: Ouest, Wetinore 2669 (MSC), 2685 

(MSC, US). Cuba: Pinar del Rio, I?mliaug 25291 (MSC): 

Oriente, Iinshaug 24952 (MSC, US). Jamaica: Iinshaug 14364 

(MSC). Dominican Republic: La Vega, Allard 16829, 16830, 

16847, 16857, 18032; Puerto Plata, Tl'efniore 3933 (US); Trujillo, Allard 17356.4, 1738i (US). Piierto Rico: Fink 1411 

(SY, US), Britton 750 (NY), Heller 293, 4338 (NY). Martinique: Imshaug 30496, 32597 (MSC). \'eneiuela: Fendler (FH); 

Distrito Federal, Sonfesson 6673 is. US): Merida, Hale 42752. 

Peru: San Martin, Allord 20701, 22581a (US). Brazil: Goias, 

Weddell (P); Mato Grosso, .lIoline 236,; is), 2435C (LD, S), 

2.509B (S, US), 2522F (S, LIPS), 2845 (S): S2o Paulo, Robert 

(BM), Schiffner (W). Paraguay: I3nlan.w 4176 (G). Chile: 

l'alparaiso, Santesson 2970 (S). Vnion of South Africa: Cape 

Province, Almborn 1361 (LD), 1381 (LD, US). Thailand: 

Schnzidt XXI (TUR, syntype of Porniclia addenda Vainio). 

Malaysia: Sabah, Hale 30359, 30365, 30.398, 90599, 30600. 

Indonesia: Java, Groenhnrt 3243 (L, LTS). Taiwan: Asahinn 

(TNS). Philippines: Agusan, Hale 24475, 24977, 25438; Basilan, Hale 24996; Cagayan, Hale 24411; Cavite, Hale 26839: 

Quezon, Hale 2691-5, 26918, 26983: Surigao tlel Sur, Hole 

24699; Zamhoanga del Sur, Hale 24314, 24776, 24824. New 

Guinea: Bailes (BM). Neiv Hebrides: Wilson (H). 

Bulbothrix hypocraea 

FIGURE 5a 

Bulbothrix hypocraea (Vainio) Hale, 1974:480. 

Parmelia hypocraea Vainio in TVelwitsch, 1901:400 [type 

collection: Monino River, Huila, .Angola, Tl'elwitsch 32 pro 

parte (TUR, lectotype: RM, isolectotype)]. 

Parrnelia continua Lynge, 1914:109 [type collection: Buriti, 

Serra de 'Chapada, Mato Grosso, Brazil, Malme (S, lectotype)]. 
16 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

Parmelia leptascea Steiner and Zahlbruckner in Zahlbruckner, 1926:514 [type collection: Lake Victoria, Bakoba, Africa, 

Schroder 319 (W, lectotype)]. 

Parmelia proboscidea var. saxicola Cengia Sambo, 1938:380 

[type collection: Mahulo, Kipengere, Tanzania, Eusebio 

13 bis (FI, lectotype)]. 

DEscRIPTroN.-Thallus adnate on bark, pale 

mineral gray, turning deep olive-buff in the herbarium, 3-5 cm in diameter; lobes dichotomously 

or irregularly branched, sublinear-elongate, of ten 

crowded, 1-3.5 mm wide; margins smooth to more 

or less crenate, with conspicuous, short bulbate 

cilia; upper surface plane and smooth, shiny, 

usually distinctly maculate; lower surface pale to 

dark brown, moderately rhizinate, the rhizines 

thick, brown, usually darker than the lower surface, simple, about 1 mm long. Apothecia nunierous, adnate, 1-5 mm in diameter, margins undulate, the amphithecium ecoronate, smooth; disc 

burnt umber; spores 8, 5-8 X 8-14 ym. 

CHEMIsmY.-Cortex K+ yellow, medulla K+ 

yellow turning red, C - , KC - , P + orange, atranorin and salazinic acid. 

HABITAT.-& trees in semi-arid woodlands at 

mid-elevations. 

DIsmInuTroN.-Brazil and Africa. 

REMARKS.-This species is characterized by the 

rather coriaceous, nonisidiate thallus and the pale 

brown lower surface with brown rhizines. It may 

be considered as the parent morph of B. tnbncinn. 

Although €3. semibizis is superficially very similar, 

it differs in having a black lower surface. 

SPECINENS EXAhIISED.-BraZi~: Mato Grosso, Mahe (US 

as a mixture with a specimen distributed as Parmelia corotiotn); Minas Gerais, Warming 263 (M); Sao Paulo, Eiteti 

29iGR, 2976C (US), Pzriggnri 1217 (G). Uganda: Toro District, 

Szhscori~ 21J 313 (Rbf, US). Zaire: Schmitz 8069 (US). 

Angola: nib, Degeliiis (Degelius herbarium); Huila, Degeliits 

(Degelius herbarium, US). Rhodesia: Hiieg (TRH), Schutle 

37 (LD, L'S), Wild 5806 (LD). Zambia: Jellicoe 50a (BM). 

Union of Soiith .4frica: Transvaal, Wager 1477 (LD). 

Bulbotlirix continua (Lynge) Hale, 1974:480. 

Bulbothrix imshaugii 

FIGURE 56 

Riclbotliriw itnslintigii (Hale) Hale, 1974:480. 

Pnrtnelia iuts/fai/gii Hale, 197la:31 [type collection: Mont%na Compana, Valparaiso, Chile, Itnsliaiig 36670 (MSC, 

holotype; US, isotype)]. 

DEscRIPTIoN.-ThallUs adnate to loosely adnate 

on bark, whitish ashy gray, 4-6 mm broad; lobes 

subirregular, 3-5 mm wide; upper surface plane 

to rugulose, shiny, maculate, becoming cracked 

with age, sorediate, the soralia laminal to subapical, diffuse, the soredia farinose (Figure Id); 

lower surface black except for a brown zone at 

the tips, moderately to densely rhizinate, the rhizines black, shiny, simple. Apothecia substipitate, 

ecoronate, 2-4 mm in diameter; spores 8, 4 X 8 

Pm. 

yellow turning red, C - , KC -, P + orange, atranorin and salazinic acid. 

CHEMISTRY .-Cortex K+ yellow, medulla K + 

HABITAT.-OII Nothofagii.f in open areas. 

D1sTRIBuTIoN.-Chile. 

REMARKS.-This species has still not been cob 

lected outside of Chile. It is unlike any other species in the genus because of the production of 

laminal soredia. 

SPECIhIENS ExAhrIxEo.-Chile: Curico, Mahii 3545 (us); 

I'alparaiso, Ritndel 7308 (US). 

Bulbothrix isidiza 

FIGURE 5c 

Zliilbothrix isidiza (Sylander) Hale, 19543480. 

Parttielin isidiza Kylander., 1884: 130 [type collection: Serra 

Chella, Angola, Nerc~fo~ (H, lectotype)]. 

Parmelin tiliacea var. liypoleitca Muller Argoviensis, 18943257 

[type collection: Usambara, Hdst 787 pro parte (G, lectoParinelia stibgla?idulifera Hue, 1899:144 [type collection: 

Madagascar, Graiididier (P, lectotype)]. 

Prtrmelin recztrwiscens Harmand, 1928:32G [type collection: 

Indochina, Deniange 36 (P, lectotype)]. 

Pnrnielin dettintigei Harmand, 1928:327 [type collection: Indochina, Detnange 37 (P. lectotype)]. 

Parttielin siibscortea Asahina, 19.57:99 [type collection: Keitau, 

Taiwan, Asnhiiin 3324 (TNS, lectotype)]. 

Pnrmelin gillettii Dodge, 1959:86 [type collection: Libah Mele 

Mountain, Somalia, Gillett 4699 pro parte (BM, lectotype)]. 

type)]. 

DEsCRrPTIoN.-Thal~iis adnate on bark or rock, 

yellowish glaucous to olive bluff, 5-10 cm in diameter; lobes subirregular to sublinear, 1-4 mm wide; 

bulbate cilia distinctly inflated; upper surface 

plane, continuous, faintly to distinctly maculate, 

densely isidiate, the isidia simple or branched; 

lower surface pale brown or darkening, moderately rhizinate, the rhizines simple, pale brown 

or darkening. Apothecia adnate, 1-3 mm in 

diameter, ecoronate; spores 8, 5-8 X 7-14 ym. 
NUMBER 32 

FIGURE 5.--Species of Bulbothrix: a, B. hypocraea (Degelius in US); b, B. imshaugii (isotype in 

US); c, B. isidiza (Hale 31756); d, B. klementii (holotype in US); e, B. laevigatula (Lunglois 

53 in US); j, B. rneizospora (Stainton 4045 in US). (Scale in mm.) 

17 
18 SMITHSONAN CONTRIBUTIONS TO BOTANY 

CHEwsmY.-Cortex K+ yellow, medulla K+ 

yellow-red, C-, KC-, P+ pale orange, atranorin 

and salazinic acid. 

HABITAT.-on trees (conifers and hardwoods) 

in pastures, cerrado woodland, and open areas at 

100-2000 m elevation. 

DISTRIBUTION .-Pantropical. 

REMARKs.-BzLlbothrix isidiza is especially conimon in Africa but also occurs widely in open 

secondary forests throughout the tropics. The 

lower surface is uniformly pale brown. Closely 

related B. tabacina has a black lower surface and 

a somewhat thinner, more fragile thallus. The 

parent morph is represented by B. hypocmea. 

Dodge (1959: 100) listed Parmelia tiliacea var. 

hypoleura as a synonym of his P. lythgoeana (= 

Hypotrachya lythgoeana (Dodge) Hale), evidently without seeing the type collection since H. 

lythgoeana is nonisidiate. Des Abbayes included a 

specimen identified as Parmelia isidiza in his 

Lichenes Madagascarienses Borbonici Selecti Exsiccati No. 10, but I have determined it to be 

Parrnelina ruallichiana (Taylor) Hale. 

SPECIhCEKS ExAJrIsED.-Mexico: Chiapas, Hale 20170, 20184. 

Guatemala: Baja Vera Paz, Hale 38373. Brazil: Rio de Janeiro, 

Ule 6 (G): Sao Paulo, Eiten 2976C (US). Paraguay: Balansa 

27 (G), 4176 (PC). Guinea: N’Zerekore Santesson 10562d 

(UPS). Ivory Coast: Man, Santesson 10616a (UPS). Uganda: 

East Mengo District, Lye L206 (BM, US); West Mengo District, Swinscow 2U 1/13 (BM, US). Kenya: Central Prov., 

hfaas Geesteranus 4434 (L), 4435 (L, LD, US): Nyanza Prov., 

Maas Geesteranus 10935 (L). Zaire: Hoeg (TRH). Angola: 

Bik, Degelius (Degelius herbarium, US); Huila, Degelius 

(Degelius herbarium): Moxico, Degelius (Degelius herbarium). Malawi: Jellicoe 32 (BM, US). Zambia: Jellicoe 44 

(BM). Tanzania: Hoeg (TRH). Rhodesia: Hoeg (LD, TRH, 

US) KoPer (LD). Union of South Africa: Natal, Almborn 

9642, 9767 (LD, US), Hoeg (TRH); Transvaal, Almborn 6442 

(LD), Hoeg (TRH): Cape Prov., Hoeg (TRH). Nepal: 

Awasthi 2179 (Awasthi herbarium), Norkett 7171 (BM), 

Stainton 3830 (BM). India: West Bengal, Thomson 49 (BM), 

Togashi (TNS, US); Maharashtra, Hale 43821, 43973; Tamil 

Nadu, Hale 40104, 43757. Malaya: Pahang, Hale 30098, 30113, 

30209, 30501. Indonesia: Java, Groenhart 5678 (L). Philippines: Benguet, Degelius (Degelius herbarium, US). Japan: 

Prov. Bungo, Kurokawa 62284 (TNS); Prov. Izu, Kurokawa 

58628 (TNS, US); Prov. Hyuga, Hale 29669, 29670. Tahiti: 

Bouffon (H). Hawaiian Islands: Kauai, Hale 31712, 31756: 

Maui, Hale 31247, 31273, 31532. 

Bulbothrix klementii, new species 

FIGURE 5d 

Thallus arcte adnatus, corticola, pallide 

stramineo-albidus, 1-2 cm latus, lobis sublinearibus, dichotome ramosis, separatis, margine conspicue bulbo-ciliatis, bulbis nitidis, orbicularibus, 

apice furcato-ciliatis, 0.6-1 .I mm latis; superne 

planus vel aetate transversim rimosus, nitidus, 

modice isidiatus, isidiis cylindricis, simplicibus; cortex superior 10-12 pm crassus, cellulis plus minusve 

verticalibus, stratum gonidiale 14 pm crassum, 

medulla alba, 80-95 pm crassa, cortex inferior 

paraplectenchymatus, 11-12 pin crassus; subtus 

pallide testaceus, dense rhizinosus, rhizinis pallidis, 

ca 24 pm crassis, dichotome furcatis. Apothecia 

ignota. 

Cm;MIsmY.-Cortex K+ yellow, medulla K--, 

C - , P-, atranorin and colensoinic acid. 

HoLorYPE.-Cerro Pavon, Rio Atabapo, Venezuela, elevation 110 m, K. Magdefmu 286, 13 

February 1958 (M; isotype in US). 

REMARKS.-This species has narrow lobes and 

densely branched bulbate cilia that form a mat 

around the edges of the lobes. The chemistry is 

unique in the genus: Colensoinic acid, identified 

by co-chromatography with Hypotrachyna livida 

(Taylor) Hale, is otherwise known only in Stereoraulon and Hypotrachyna. There are no close 

relatives in Bulbothrix. It is named in honor of 

Dr. Oscar Klement, who first saw the material at 

M and correctly placed it in the Parmeliaceae. 

Bulbothrix laevigatula 

FIGURE 5e 

Bulbothrix laevigatula (Sylander) Hale, 1974:480. 

Parmelin lnevigaiula Nylander, 1885:614 [type collection: 

French Guiana, Le Prieur 504 (H, Nylander herbarium 

35653, lectotype; PC, isolectotype)]. 

Parmelia hookeri Taylor, 1847: 169 [type collection: St. Vincent, Guilding (FH-Tayl, lectotype: BM, isolectotype): 

not Parmelin hookeri (Borrer) Sprengel, 1827:282]. 

DEscRIPTIoN.-Thallus adnate on bark, whitish 

mineral gray, turning cream-buff in the herbarium, 

4-9 cm in diameter; lobes irregularly branched, 

sublinear-elongate, 0.5-2 mm wide, the margins 

more or less crenate, with short marginal bulbate 

cilia less than 0.2 mm long, shiny, often apically 

branched, strongly inflated at the base; upper surface shiny, emaculate, isidiate, the isidia cylindrical, sometimes branched, less than 0.3 mm high; 

lower surface black, densely rhizinate, the rhizines 
NUMBER 32 19 

densely and finely branched, forming a woolly 

mat. Apothecia adnate, 1-5 mm wide, margins 

crenate and undulate, amphithecium rugose, 

ecoronate, isidiate; spores 8, 34 X 5-7 pm. 

HABITAT.-On trees (conifers and hardwoods, 

Pandanzi.f, bamboo, coconut) and more rarely on 

rocks in secondary forests from sea level to 3000 

m elevation. 

DIsTRIBuTIoN.-United States, Mexico, wrest 

Indies, and South America. 

REMARKS.-ThiS isidiate species is one of the 

most common members of the genus in the West 

Indies and adjacent United States but it is rare in 

other parts of tropical America. The thallus is 

distinctly ashy white in comparison with gyrophoric acid-containing B. goebelii which is more 

greenish or olivaceous whitish gray. It is usually 

collected sterile, only 7 of 36 specimens in US bearing apothecia. Except for tlie somewhat broader 

lobes, B. Inevigntula might be considered as the 

isidiate morph of B. confoederata, a small lichen 

endemic to the southeastern United States. The 

only other species in Bulbotkrix with lecanoric 

acid, B. bicornutn, has bicornute spores. 

SPECIMENS EXalfINED.-United States: South Carolina, Hale 

7624; Georgia, Hale 16767, 16769, Skorepa 3113 (US); Alabama, Hale 33796, 33977, 33980, McCulloicgh 1193 (US); 

Florida, Hale 16731, 17742, 17745, 21695, 34117, 34180, Thornson 4457 (US, W'IS) (for additional records from Florida see 

Moore, 1968:221); Mississippi, Langlois 53 (US). Mexico: 

Vera Cruz, Hale 19833. Cuba: Pinas Del Rio, Imshazcg 25233 

(MSC, US). Jamaica: Zrnslzaug 13877 (MSC, US), 14306, 14319, 

15791 (MSC), 15932 (MSC, US), 16040, 16250 (MSC). Dominican Republic: Hassler (M71S); Cordillera Central, Imshaug 

23759 (MSC, US): La Vega, Allard 15992a, 16548a (US); 

Santiago, Imshaug 23862 (MSC, US), Wetmore 3883 (MSC); 

Trujillo, AIlard 17356 (US). St. Lucia: Evans 71, 83 (FH, 

US), Imshaug 29724, 29991 (MSC). Dominica: Hale 35411, 

35687. St. Vincent: Imshaug 30468, 30729 (MSC). Grenada: 

hsknug 16234 IMSC). Tobago: lmshaug 31388, 31555 (MSC). 

Trinidad: Hale 38206, Imshatcg 32405, 32431 (MSC). Colombia: Santander, Killip and Smith 18075 (US). Venezuela: 

Mkrida, Hale 42754. Peru: Churubamba, Mexia 8245a is); 

San Martin, Allard 22256b (US). Ecuador: Galapagos, Weber 

191 (COLO). Brazil: Rio de Janeiro, Eiten 7079 (US), Schindlet 4577a (US). 

Bulbothrix meizospora 

FIGURE 5f 

Bulbothrix meizospora (Nylander) Hale, 1974:480. 

Partnelia tiliacea var. meizospora Nylander, 1860:383 [type 

collection: Nilgherries Mountains, India (H, Nylander 

herbarium 35107, lectotype)]. 

Pnrmelia meizospora (Sylander) Sylander, 1869:292. 

Parrnelia amplectens Stirton, 1877-78:201 [type collection: 

Nilgherries Mountains, India, Watt (BM, lectotype; GLAhf, 

isolectotype)]. 

DEscRIPTIo~.-Thallus adnate on bark, pale 

glaucous-green, turning olive-buff in the herbarium, 5-8 mm broad; lobes irregularly branched 

to sublinear-elongate, 1.5-5 mm wide, tlie margins 

crenate, with sparse short bulbate cilia; upper 

surface plane and smooth, more or less shiny, 

faintly maculate; lower surface black, dark brown 

and papillate in a broad or narrow rone near the 

tips, densely to moderately rhizinate, the rhizines 

brown to black, simple. Apothecia numerous, 

adnate, 2-8 mm in diameter, ecoronate, amphithecium more or less wrinkled; disc vandyke-brown; 

spores 8, 7-11 X 14-21 pm. 

yellow turning red, C-, KC -, P+ orange, atranorin and salazinic acid. 

HABITAT.-on trees (rarely on rocks) in open 

woodlands. 

DISTRIBuTION.-z4frica, Pakistan, Nepal, and 

India. 

REMARI<s.-This Asian species is rather variable 

in lobe width and spore size with a close relationship to B. setschwnnensis. which has a uniformly 

pale brown lower surface and occurs at higher 

elevations in the Himalayan region. Bulbothl-ix 

meizospora has a broader distribution although 

it too grows at elevations of 2000 m and higher. 

SPECIMENS EXAxrINED.-Cameroon: Rutherford 22 (BM). 

Pakistan: Iqbal 828, 835, 844 (US). Sepal: Ranerjee 1403 

(Aimsthi herbarium), Norkett 5306 IRM, US), 7890, 8231 

(BM), Stainton 4045 (BM, US). India: Assam, Blott (Awasthi 

herbarium); Sikkim; Harn (TSS), Togaslri (TNS); Uttar 

Pradesh, Awasthi 474, 897, 2683, 3443, 3798 (Awasthi herbarium), Degelitts Aslll (Degelius herbarium, US), Hiieg 3390, 

3403 (Awasthi herbarium), Steruart 12298 INY, US); Tamil 

Kadu, Hale 43877. 

CHEMISTRY.-cOrteX K + yellow, medulla K + 

Bulbothrix pigmentacea 

FIGURE 6a 

Ritlbothrix pigmentacea (Hale) Hale, 1974:480. 

Parrnelia pigrnentacea Hale, 1968:325 [type collection: Pagbilao, Sierra Madre, Luzon, Philippines, Hale 26895 (US, 

holotype)]. 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

DEscRIPTIoN.-Thallus closely adnate on bark, 

thin and fragile, whitish mineral gray, 1-2 cm 

broad; lobes linear, discrete, 0.5-1 .O mm wide, the 

margins densely bulbate ciliate, the bulbae apically branched; upper surface plane, shiny, moderately isidiate, the isidia simple, to 0.1 mm high; 

lower surface black, densely rhizinate, the rhizines 

dichotomously branched. Apothecia unknown. 

CHEMIsmY.-Cortex K+ yellow, medulla negative with color reagents, atranorin only present 

with an unknown red pigment in the rhizines and 

lower cortex. 

HABITAT.-on canopy branches of trees in lowland evergreen rain forest from sea level to 300 

m elevation. 

DISTRIBUTIoN.-Philippines and Malaya, 

REMARKS.-This is the only species of Btilbothrix present in the virgin lowland forests of 

Southeast Asia. The pigment in the lower cortex 

and rhizines is exposed when the thallus is cut 

with a razor blade for cross sections. This unknown pigment and the very narrow lobes would 

separate it from superficially similar B. goebelii. 

There are no closely related species nor a parent 

morph. 

Philippines and Malaya. 

SPECIMENS Exk-rrIxED.-~ee Hale (1968) for localities in the 

Bulbothrix pustulata 

FIGURE 6b 

Biclbothrix pustulata (Hale) Hale, 1974:480. 

Purmelia prrstuluta Hale in Hale and Kurokawa, 1964:140 

[type collection: Urundi, Africa, Deuse (17.5, holotype)]. 

DEsCRlPTIoN.-Thallus adnate to loosely adnate 

on bark, light mineral gray, 6-10 cm broad; lobes 

more or less subirregular, subrotund, 4-6 mm wide; 

bulbate cilia moderate, distinctly inflated; upper 

surface plane to rugose, continuous, moderately 

isidiate-pustulate (Figure le), soredia absent; lower 

surface black and moderately rhizinate except for 

a narrow papillate or naked brown zone along the 

margins, the rhizines simple. Apothecia rudimentary, sessile, to 1.5 mm in diameter; spores not 

found. 

CHEmsmY.-Cortex K 4- yellow, medulla K+ 

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

atranorin and salazinic acid. 

HABITAT.-on trees in open forest. 

DISTRIBUTION.-Africa. 

REMARKs.-This is the only species in Bulbothrix with pustules. These are typical pustules, 

erupting and fragile but without soredial formation. The thallus, although broad lobed, is quite 

membranous. There is no extant parent or other 

comparable species in the genus. It is still known 

only from the type collection. 

Bulbothrix schifneri 

FIGURE 6c 

Bulbothrix schiffneri (Zahlbruckner) Hale, 1974:481. 

Parinelia schiffneri Zahlbruckner, 1909:167 [type collection: 

Barra Mama, Itapecirica, SZo Paulo, Brazil, Schiflner (W, 

lectotype)]. 

DEscRIPTIoN.-Thallus closely adnate on bark 

and mosses, light mineral gray, 1-2 cm broad; 

lobes dichotomously branched, sublinear, 0.3-0.5 

mm wide, the margins densely bulbate ciliate, the 

bulbae becoming apically branched; upper surface 

plane, shiny; lower surface black, densely rhizinate, the rhizines moderately to densely branched. 

Apothecia numerous, adnate, 1-1.5 mm in diameter, the amphithecium coronate; spores 8, 5 X 

I2 pm, bicornute with short upturned tips. 

CHEMIsTRY.-cortex K+ yellow, medulla K-, 

C+ rose, KC+ red, P-, atranorin and gyrophoric 

acid. 

HABITAT.-on trees in cerrado vegetation. 

DIsTRrBuTIoN.-Brazil. 

REMARKS.-This very small lichen is quite similar to B. semilzinata and is separated from it 

essentially by the presence of gyrophoric acid. It 

is still known only from the type collection. 

Bulbothrix semilunata 

FIGURE 6d 

RttlBothrix semilunntn (Lynge) Hale, 1974:479. 

Pnrtnelin serniluriatn Lynge, 1914:23 [type collection: Buriti, 

Serra da Chapada, Mato Grosso, Brazil, Malrne (S, lectotype)]. 

DEsCRIPTIoN.-Thallus closely appressed on bark, 

whitish mineral gray, 1.0-1.5 cm broad; lobes 

sublinear, 0.2-0.3 mm wide, the marginal bulbate 

cilia with a branched, ciliate tip; upper surface 

plane, smooth; lower surface black, moderately 
NUMBER 32 21 

FIGURE 6.-Species of Bulbothrix: a, B. pigmentacea (holotype in US); b, B. pustulata (holotype 

in US); c, B. schiffneri (lectotype in W); d, B. semilunata (Malme in US) (X 10); e, B. sensibilk 

ides Abbayec in US); f, B. setschwanensis (Awasthi 3316 in US). (Scale in mm.) 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

rhizinate, the rhizines branched. Apothecia numerous, adnate, about 1 mm in diameter; spores 

8, semi-lunate or cornute, 2-3 X 9-12 ym. 

CHEMIsmY.-cortex and medulla negative with 

color tests, only skyrin present. 

HABITAT.-on trees in cerrado vegetation. 

DIsmIsuTIoN.-Brazil. 

REMARKS.-This species is extremely close to 

B. schiflneri in thallus size and lobation. It would 

be differentiated by the C negative reaction in the 

medulla and the narrower, more semi-lunate spore 

outline. Neither species has been collected outside 

its type-locality. 

SPECIMENS EXAMINED.-Brazil: Mato Grosso, Malme (US) 

(discovered as a mixture in a specimen distributed as “Parmelia coronata” and probably part of the type collection). 

Bulbothrix sensibilk 

FIGURE 6e 

Bulbothrix sensibilk (Steiner and Zahlbruckner) Hale, 1974: 

481. 

Parmelia sensibilis Steiner and Zahlbruckner in Zahlbruckner, 

1926:522 [type collection: Bura, British East Africa, Schroder 285 (W, lectotype)]. 

DEsCRIPTIoN.-Thallus adnate on bark, pale 

olive-buff to pale olive-gray, turning deep olivebuff in the herbarium, rather coriaceous, 4-10 cm 

in diameter; lobes irregularly branched, more or 

less imbricate, 1.5-5 mm wide, the margins more 

or less crenate; bulbae rather sparse, shiny, simple; 

upper surface more or less shiny, very faintly to 

moderately maculate, irregularly rugose and 

cracked on older lobes; lower surface black, 

densely rhizinate, the rhizines black, simple. 

Apothecia adnate, 2-6 mm in diameter, the margins more or less undulate, the amphithecium 

ecoronate, rugose; disc warm-sepia; spores 8, 

5-12 X 7-18 ym. 

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

and salazinic acid. 

HABITAT.-on trees in open woodlands at 500- 

2000 m elevation. 

DISTRIBUTION.-venezuela and Africa. 

REMARKS.-Bulbothrix sensibilis resembles B. 

hypocraea closely in overall appearance but has a 

distinctly black lower surface. Both have a South 

CHEMISTRY.-cOrteX K+ yellow, medulla K+ 

America-Africa distribution pattern, B. sensibilis 

being the much rarer of the two species. It is or 

would fall very close to the parent morph of B. 

ta bacina. 

SPECIMENS EXAMINED.-VeneZUela: Tachira, Hale 45727. 

Guinea: des Abbayes (US). Zaire: Hoeg (TRH). Malawi: 

Brass 17788 (US). Zambia: Jellicoe 52 (BM). Angola: Benguela, Degelius (Degelius herbarium); Bie, Degelius (Degelius 

herbarium, US). 

Bulbothrix setschwanemis 

FIGURE 6f 

Bulbothrix setschmanensis (Zahlbruckner) Hale, 1974-481. 

Pnrtnelia setschivanensis Zahlbruckner, 1930: 184 [type collection: Otang, Kwapi, Setchwan, China, Handel-Mazzetti 

2739 (WU, lectotype)]. 

DEsCRIPTIoN.-Thallus adnate on bark, light 

mineral gray but turning dark olive-buff in the 

herbarium, 5-10 cm in diameter; lobes irregularly 

branched, sublinear-elongate, 2-5 mm wide, the 

margins crenate with sparse short black bulbate 

cilia; upper surface plane and smooth, more or 

less shiny, not maculate, irregularly wrinkled on 

older lobes; lower surface pale brown, densely rhizinate, the rhizines pale brown, coarse, simple, 

1-1.5 mm long. Apothecia adnate, 1-4 mm in 

diameter, margins and amphithecium smooth, 

ecoronate; disc vandyke brown; spores 8, 6-9 X 

12-19 pm. 

yellow turning red, C-, KC-, P+ orange, atranorin and salazinic acid. 

HABITAT.-on trunks and branches of trees in 

open forests at 2000 m elevation or higher. 

DIsTRIsuTIoN.-India, Nepal, and China. 

REMARKS.-This species is endemic to the Himalayan region and adjacent China. It has a large 

coriaceous thallus, very similar to that of B. meizospora (see above), which has a blackening lower 

surface. These two species may indeed prove to 

be synonymous when careful field studies are 

conducted. 

CHEMISTRY.-cOrteX K+ yellow, medulla K+ 

SPECIhfENS EXAMINED.-India: Uttar Pradesh, Awasthi 3316 

(Awasthi herbarium, US); West Bengal, Togashi (TNS). 

Nepal: Awasthi 2216 (Awasthi herbarium). China: Szechuan, 

Handel-Mazzetti 2017 (W, syntype of Parmelia setschwanensis Zahlbruckner). 
NUMBER 32 

Bulbothrix subcoronata 

FIGURE ia 

Bulbothrix subcoronata (Muller ..lrgoviensis) Hale, 1974:481. 

Parrnelia subcoronata Muller Argoviensis, 188ia: 135 [type 

collection: South America (G, lectotype)]. 

DESCRIPTION.-T~~~~US closely adnate on bark, 

buff mineral gray but turning pale to deep olivebuff in the herbarium, 3-5 cm in diameter; lobes 

irregularly branched, sublinear-elongate, 1-3 mm 

wide, the margins more or less crenate, with moderate short bulbate cilia; upper surface plane and 

smooth, shiny, slightly to moderately maculate; 

lower surface brown, moderately rhizinate, the 

rhizines brown to dark brown, simple. Apothecia 

adnate, 1-4 mm in diameter, the margins undulate, 

the amphithecium coronate; disc verona-brown; 

spores 8, 5-6 X 7-11 pm. 

yellow turning red, C - , KC - , P + orange, atranorin and norstictic acid. 

HABITAT.-on trunks and branches of trees in 

open woodland at lower elevations. 

DIsTRIBuTIoN.-~outh America. 

REMARKS.-This is a rare South American species, occurring apparently in cerrado forests. It is 

easily recognized by the presence of norstictic acid. 

Without a chemical test it might be mistaken for 

B. hypocraea, an ecoronate species with salazinic 

acid. Muller reported smaller spores (to 5 pm 

long) in the original description, but all tlie specimens I examined had larger spores as indicated. 

SPECIMENS ExAMlsED.-Brazil: Goias, Irwin 31977 (XY, us); 

Mato Grosso, Malrne 2271Ba (as a mixture with Parrnelia 

silvatica) (S), 2illBa (S); Rio de Janeiro, Glaziou 1825 

(UPS). Paraguay: Handel (G), Kuntze 42 (NY), Kzmlze (G), 

Balansa 4210 (C). Argentina: Missiones, Montes 10060 (MVM, 

CHEMISTRY .-cortex K + yellow, medulla K + 

US). 

Bulbothrix subinfluta 

FIGURE ib 

Biclbothrix srcbtnfinta (Hale) Hale, 1974:481. 

Parinelin stcbinflnta Hale, 1965b:ZOl [type collection: 50 miles 

south of Fabrica, Negros Occidentalis, Philippines, Hale 

26641 (V5, holotype; TNS, UPS, isot\pes)]. 

D~sCRIPT~oh..-Thallus closely adnate on bark, 

mineral gray, 4-7 cm broad, lobes short, sublinear, 

1-2 mm wide, marginally bulbate, the bulhae not 

23 

strongly inflated; upper surface shiny, emaculate 

or faintly maculate, becoming rugose with age, 

moderately isidiate, the isidia simple to sparsely 

branched, cylindrical, 0.1-0.2 mm high; lower surface pale brown, moderately rhizinate, the rhizines 

brown or darkening, simple. Apotliecia rare, 

adnate, to 1 mm in diameter, ecoronate; mature 

spores not seen. 

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

C-, KC+ rose or KC-, P+ red, atranorin and 

protocetraric acid. 

HABITAT.-on branches of evergreen hardwoods 

in mid-elevation rain forest at 300-1800 m elevation. 

DIsTRIBuTIoN.-Phi~ippineS and Malaysia. 

hMmKs.-Bztlbothrix szrbinpatn is the only species outside of B. pigmentacea which has evolved 

in the rain forests of Southeast Asia. It occurs at 

higher elevations than B. pigmentacea, for the most 

part in cloud forest. The marginal bulbae are not 

strongly inflated, as with B. cho70oensis, a broader 

lobed African species which also contains protocetraric acid. 

SPFCIhfF\S ExihrlNFD.-See Hale (196i:ZOl) for localities in 

the Philippines and Malaysia. 

Bulbothrix sufixa 

FIGURE 7c,d 

Rirlbotlzrix snfixa (Stirton) Hale, 1974:481. 

Parrnelia snffixa Stirton, 1877-78:299 [type collection: Knysna, 

Union of South .Africa, Knohel (Bhf, lcctotypc: GLAM, 

isolectotype)]. 

DEsCRIPTIoh..-ThalluS closely adnate on bark, 

fragile, whitish mineral gray, 2-6 cm broad; lobes 

dichotomously branched, sublinear, 0.5-1.3 mm 

wide, the margins more or less crenate; bulbate 

cilia dense, the bulbils shiny, inflated, often apically branched; upper surface plane and smooth, 

shiny, becoming densely lobulate, the lobules 

dorsiventral, oblong to spathuliform, marginally 

hulbate-ciliate (Figure If); lower surface black 

moderately rhizinate, the rliizines black, shiny, 

dichotomously branched. ,4pothecia adnate, the 

ampliitliecium becoming lobulate, tlie rim crenate, coronate; spores 8, 3-5 X 6-12 pm. 

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

C+ rose, KC+ red, P-, atranorin and gyrophoric 

acid (or if C- no substances present in medulla). 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

HABITAT.-On trees in open areas along trails 

and roads at up to 2400 m elevation. 

DIsTRIBuTIoN.-Tropical America and South 

Africa. 

REMARKS.-Bulbothrix sufixa is characterized 

by the production of dorsiventral lobules with 

dense marginal bulbate cilia (Figure If). There 

are no isidia. It has the same chemistry as B. 

fungicola, which is predominantly isidiate and 

isidiate-lobulate (Figure 34. As with other small 

species of Bulbothrix, it is not well represented 

in herbaria and may be subject to revision by 

future workers. Some specimens are C negative 

and lack gyrophoric acid but are morphologically identical with the C+ population. I am recognizing these for the present as an acid-free 

phase. 

SPECIMENS EXAMINED (gyrophoric acid).-Guatemala: Baja 

Vera Paz, Hale 38376. Honduras: Comayagua, Standley and 

Chacdn 5830 (US). Cuba: Wright 75 (UPS, US): Pinar del 

Rio, Zmshaug 25354 (MSC). Dominican Republic: Santiago, 

Zmshaug 23854, Wetmore 3881 (MSC). Jamaica: Zmshaug 

15111 (MSC). St. Lucia: Zmshaug 29820 (MSC). Dominica: 

Zmshaug 32758, 32760, 32762 (MSC). Venezuela: Merida, Hale 

42257a, 42565d. Union of South Africa: Cape Province, Aimborn 3444, 3971 (LD), 3961 (LD, US). Mauritius: Vaughan 

6 (BM). 

SPECIMENS EXAMINED (lacking gyrophoric acid).-Dominica: 

Zmshaug 33330 (MSC). Trinidad: Zmshaug 32078, 32088 

(MSC). Venezuela: Merida, Hale 42753. 

Bulbothrix tabacina 

FIGURE 7e 

Bulbothrix tabacina (Montagne and Bosch) Hale, 1974:481. 

Parmelia tabacina Montagne and Bosch in Montagne, 1856: 

327 [type collection: Java, Junghuhn (L, lectotype; P, isolectotype)]. 

Parmelia meizospora f. isidiosa Muller Argoviensis, 1884:620 

[nomen nudum]. 

Parmelia ochrovestita Zahlbruckner, 1928:200 [type collection: 

Botanical Garden, Bogor, Java, Ouereem 335 (W, lectoParmelia meizosporoides Dodge, 1959:83 [type collection: 

Imerina, Andrangolaoka, Madagascar, Hildebrandt (FH, 

lectotype)]. 

Parmelia sublaeuigatoides Dodge, 1959:88 [type collection: 

Mt. Eglon, Uganda (BM, lectotype)]. 

DEscRIPTIoN.-Thallus adnate to closely adnate 

on bark, rarely on rock, 4-7 cm in diameter; lobes 

sublinear to subirregular, 1.5-5 mm wide; bulbate 

cilia large, conspicuously inflated; upper surface 

type)]. 

plane, faintly to moderately maculate, sparsely 

isidiate, the isidia mostly simple, less than 0.5 mm 

high; lower surface black, shiny, moderately rhizinate except for a narrow papillate or naked 

zone along the margins, the rhizines mostly simple. 

Apothecia adnate, 1.54 mm in diameter, the 

amphithecium ecoronate, isidiate; spores 8, 5-8 X 

9-15 pm. 

CHEMIsmY.-Cortex K + yellow, medulla K -Iyellow turning red, C -, KC -, P + orange, atranorin, salacinic acid, and consalazinic acid. 

HABITAT.-oII trees (hardwoods and conifers) 

or rarely on rocks in open woodland and pastures 

at sea level to over 2000 m elevation. 

DIsTRIBuTI0N.-Pan tropical. 

REMARKs.-This isidiate species is one of the 

most commonly collected members of the genus. 

The lower cortex is black in contrast to that of 

B. isidiza, which is uniformly pale brown and has 

a somewhat more robust thallus. It may also be 

confused with B. ZaevigatuZa, which has dichotomously branched rhizines, a white cast to the 

thallus, and lecanoric acid (medulla C+ red). It 

is the only species of Bulbothrix so far discovered 

in Australia. 

Zahlbruckner (1928: 174) saw a “type specimen” 

in M and considered it to be close to Parmelia 

cetratn (= Parmotrema cetratum (Acharius) Hale, 

an amphigymnioid lichen with marginal cilia). 

Dodge (1959:83) cited Parmelia meizospora f. isidiosa Muller Argoviensis as a synonym of Parmelia 

meizosporoides Dodge, but Muller’s name is a 

nomen nudum without a description or designated 

tY Pe. 

SPECIMENS ExAMIh’En.-Mexico: Vera Cruz, Hale 19475; 

Chiapas, Hale 21067 (US), 20122 (S, US). Cuba: Oriente, 

Hiorant 5887 (US). Jamaica: Zmshaug 16014 (MSC). Haiti: 

Ouest, Zmshaug 22619 (MSC). Dominican Republic: La Vega, 

Allard 18169, 18172 (US). Tobago: Zmshatig 31332 (MSC). 

Venezuela: Distrito Federal, Ernst (G): Merida, Hale 42514a; 

Miranda, Dennis 15244 (BM): Tachira, Hale 45724. Brazil: 

S%o Paulo, Osorio 4920 (MVM). Guinea: N’Zerekore, Santesson 10516a (UPS, US), 10507, 10573b, 10594b (UPS). 

Angola: Cuanza Norte, Degelius (Degelius herbarium). Tanzania: Santesson 23409, 23422 (UPS). Union of South Africa: 

Satal, Hoeg (TRH): Transvaal, Almborn 6546 (LD, US). 

India: LJttar Pradesh, Mehra 9 (US). Nepal: Poelt 117 (M). 

Malaya: Pahang, Hale 30134: Selangor, Hale 31182, 31183, 

Maingay (BM). Philippines: Leyte, Edano 16051, 16053 (US): 

Agusan, Hale 25052. Indonesia: Java, Groenhart 4356 (L, 

US), Neeruoert 926, 1414 (BOR), Ouereem 676 (BOR), van 

Steenis 5597 (BOR); Sumatra, Groenhart 9098 (BOR). Tai- 
SUMBER 32 25 

FIGURE 7.-Species of Bulbothrix: a, B. subcoronata (Montes 10060 pro parte in US); b, B. subinpata (Hale 33711); c, B. sufixa (lectotype in BM) (X 4); d, B. sufixa (Almborn 3961 in US); 

e, B. tabacina (Kurokawa 2730 in US); j, B. uentricosa (Hale 43080); g, B. uiridescens (Irnshaug 

42530 in US). (Scale in mm.) 
SMITHSONIAN CONTRIBUTIONS TO BOTANY 

wan: Kurokawa 1299, 1363, 2818, 2932 (TNS). Australia: 

Queensland, Bailey 32 (BM). 

Rulbothrix ventricosa 

FIGURE 7f 

Bulbothrix ventricosa (Hale and Kurokawa) Hale, 1974:481. 

Parmelia ventricosa Hale and Kurokawa, 1964:140 [based on 

P. isidiza var. domingensis Vainio]. 

Parmelia isidiza var. domingensis Vainio, 1915: 17 [type collection: La Cumbra, Dominican Republic, Raunkiaer 492 

(TUR, lectotype; C, isolectotype)]. 

DEScRIPTIoN.-Thallus adnate on bark, pale 

glaucous green, 3-5 cm broad; lobes subirregular, 

1-5 mm wide, the margins crenate; bulbate cilia 

distinctly inflated; upper surface plane, shiny, 

faintly maculate, moderately isidiate, the isidia 

fine, simple; lower surface black, moderately rhizinate, the rhizines black, mostly simple. Apothecia adnate, 2-3 mm in diameter, the exciple coronate; spores 8, 7-8 X 13-17 pm. 

CmMismY.-Cortex K+ yellow, medulla K f 

yellow turning red, C-, KC-, P+ orange, atranorin and norstictic acid. 

HABITAT.-on trunks and branches of trees in 

open primary cloud forest or secondary forest at 

900-2600 m elevation. 

DlsTRlBuTIoN.-MexiCo, Central America, Venezuela, and Union of South Africa. 

REMARKS.-B~~Z~O~~~.~X uentricosa occurs chiefly 

in higher elevation cloud forests in tropical America, although there is a disjunct record from South 

’4frica. While it would appear to be related to B. 

subrol’onata because of the presence of norstictic 

acid, it has much broader lobes and a black lower 

surface. No parent morph is extant. 

SPFCIMESS ExAarlNED.-Mexico: Chiapas, Hale 20166. Costa 

Rica: Cartago, Fletiniketz (US). Panama: Chiriqui, Hale 

38763, Snkatiishi (US). Venezuela: Mkrida, Hale 43080: 

Tdchira, Hale 4572ia. Iinion of South Africa: Transvaal, 

Alr~ihorn 6813 (LD). 

Btclbothrix viridescens 

FIGURE 7g 

Rulbothrix niridescens (Lynge) Hale, 19743481. 

Parmelza vzridescens Lynge, 1914:117 [type collection: Santa 

Anna da Chapada, Mato Grosso, Brazil, Malme 2453 (S, 

lectotype; W, isolectotype)]. 

Parmelia marginalis var. laeviuscula Rasanen, 1947:45 [type 

collection: Canelones, Carrasco National Park, Uruguay, 

Lamb 3081 (H, lectotype)]. 

DEscRIPTIoN.-Thallus closely adnate, corticalous, pale buff or olivaceous ashy, 3-5 cm broad; 

lobes sublinear, contiguous to imbricate, 1-2 mm 

wide, the marginal bulbate cilia distinct, round, 

becoming apically ciliate; upper surface shiny, 

plane to distinctly rugulose, becoming heavily 

pycnidiate; lower surface dark brown or blackening, moderately rhizinate, the rhizines pale brown 

or darkening, simple or branched. Apothecia numerous, sessile to substipitate, the amphithecium 

and rim pycnidiate (Figure Ig); spores 8, 4 X 

6 pm. 

CmMIsmY.-Cortex K + yellow, medulla negative with color tests, atranorin only present. 

REMARKS.-This is one of the few species of 

Bulbothrix lacking any medullary substances. It 

is a rare lichen with a limited range in Brazil and 

Uruguay but with more careful collecting should 

eventually be found in adjacent countries. 

SPFCIXfEhs ExAwhED.-Uruguav: Floresta, Herter 61467 

(S); Montevideo, Zmshaug 42530 (MSC, US). 

Nomen Inquirendum 

Parmelia stenophylla Muller Argoviensis 1893:128 [type collecParmelia ste?iophylli:ans Zahlbruckner, 1929375 [based on 

Parmelia stenophylla hluller Argoviensis, not “Parmelia 

stenophylla (Acharius) Heugel”]. 

The material in G is fragmentary and sterile 

but has distinct apically branched bulbate cilia. 

The thallus lacks isidia, the lobes are 0.17-0.25 

mm wide, the medulla reacts C-, KC-, and the 

lower surface is black with branched rhizines. It 

is very close to or the same as either B. schiflnel-i 

or B. semilunata, but a positive identification cannot be made without apothecia. 

tion: Boruca, Costa Rica, Pittier 5434 (G, lectotype)]. 
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Index 

1 continua, 15 

coronata, 12 

coronata var. denudata, 9 

coronata f. isidiosa, 14 

decurtata, 14 

demangei, 16 

fungicola, 14 

gillettii, 16 

glandulifera, 12 

goebelii, 14 

1 granatensis, 14 

hookeri, 18 

kypocraea, 15 

imshaugii, 16 

isidiza, 16 

isidiza var. domingensis, 26 

laevigatula, 18 

leptascea, 16 

lythgoeana, 18 

marginalis, 15 

(Synonyms in italics) 

Bulbothrix affixa, 9 

apophysata, 9 

atrichella, 10 

bicornuta, 11 

bulbochaeta, 11 

chowoensis, 11 

confoederata, 12 

continua, 16 

coronata, 12 

decurtata, 14 

fungicola, 14 

goebelii, 14 

hypocraea, 15 

imshaugii, 16 

isidiza, 16 

klementii, 18 

laevigatula, 18 

meizospora, 19 

papyrina, 15 

pigmentacea, 19 

pustulata, 20 

schiffneri, 20 

scortella, 15 

semilunata, 20 

sensibilis, 22 

setschwanensis, 22 

subcoronata, 23 

subdissecta, 15 

subinflata, 23 

suffixa, 23 

tabacina, 24 

ventricosa, 26 

marginalis var. laeviuscicla, 26 

tneirospora f. isidiosa, 24 

meizosporoides, 24 

njalensis, 15 

ochrovestita, 24 

papyrina, 14 

pigmentacea, 19 

Proboscidea var. saxicola, 16 

psettdocoronata, 14 

pustulata, 20 

recuruiscens, 16 

relicina var. coronata, 12 

schiffneri, 20 

scortella, 15 

semilimnto, 20 

sensibilis, 22 

setschwanensis, 22 

stenopkylla, 26 

stenopkyllizans, 26 

subcoronata, 23 

subdissecta, 14 

subglandulifera, 16 

suhinflata, 23 

sublaevigatoides, 24 

sztbscortea, 16 

sufiixa, 23 

tabacina, 24 

tilineen var. hypoleuca, 16 

tiliacea var. meizospora, 19 

?~entricosa, 26 

riiridescens, 26 

Parmelina wallichiana, 18 

U.S. GOVERNMENT PRINTING OFFICE: lS76-2ll-S5S/l9 

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