SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 16 

Morden-Smithsonian Expedition 

to Dominica: 

The Lichens (Thelo trema t aceae) 

Mason E. Hale, 3r. 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1974 
ABSTRACT 

Hale, Mason E., Jr. Morden-Smithsonian Expedition to Dominica: The Lichens 

(Thelotremataceae). Smithsonian Contributions to Botany, number 16, 46 

pages, 20 figures, 1974.-A revision is made of the lichen family Thelotremataceae 

in Dominica, based on previously published records by Elliott and on collections by the author. The family comprises 48 species in 4 genera, Leptotrema, 

Ocellularia, Phaeotrema, and Thelotrema. The following 15 species are described as new: Ocellularia antillensis, 0. conglomerata, 0. dominicana, 0. maculata, 0. mordenii, 0. nigropiinrta, 0. rimosa, 0. yorediata, Phaeotrema aggregatum, P. obscurum, Thelotrema confusum, T. dominicanum, T. papillosum, 

T. tenue, and Leptotrema deceptum. Three new combinations, Ocellularia fecunda (Vainio) Hale, Phaeotrema disciforme (Leighton) Hale, and Leptotrema 

occult um (Eschweiler) Hale, are also made. Morphological characters are 

reviewed and the chemistry of each species is presented in detail. The family 

is exceptionally well developed in mature rain forest. 

OFFICIAL PI~BLICATIOK DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution’s annual report, Srnithsonian Year. SI PRESS NUMBER 5043. SERIES COVFR DESIGN: 

Leaf clearing from the katsura tree Cercidiphyllum juponicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data ~~ 

Hale, Mason EMorden-Smithsonian Expedition to Dominica: the lichens (Thelotremataceae). 

(Smithsonian contributions to botany, no. 16) 

Bibliography: p. 

1. Thelotremataceae. 2. Lichens-Dominica, I. Title. 11. Series: Smithsonian Institution. 

QKl.S2747 no. 16 [QK585.T5] 581’.08s [589’.1] 73-22465 

Smithsonian contributions to botany, no. 16. 

For sale by the Superintendent of Documents, US. Government Printing Office 

Washington, D.C. 20402 Price $1.05 (paper cover) 
Contents 

Page 

Introduction ...................................................... 1 

Morphological Characters ................................ 

Thallus Structure ............................................... 2 

Apothecial Characters ...................... 

Spores ............................................ 

Generic Limits ........ ............... ................. 

Chemistry ............................................ 

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

The Thelotreme Flora .......................................... 13 

Collecting Localities ..................................... 

Ocellularia ............ ................................ 

15 

Phaeotrema .................................................... 28 

28 

Thelotrema ...................................................... 31 

31 

Leptotrema .......................... ................... 38 

38 

Literature Cited .............................. ....... 43 

Index ........................................................... 46 

Key to the Species of Ocellularia .................................. 

Key to the Species of Phaeotrema ................................. 

Key to the Species of Thelotrema 

Key to the Species of Leptotrema ................................ 

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

iii 

Morden- Smi t hsonian Expedition 

to Dominica: 

The Lichens (Thelotremataceae) 

Mason E. Hale, Jr. 

Introduction 

The Thelotremataceae comprise a very large 

family of crustose lichens that has attracted the 

attention of many lichenologists, beginning with 

Acharius (Hale, 1972). They have unique craterlike apothecia, but this characteristic feature can 

usually only be seen under magnification. As a 

matter oi fact, these inconspicuous lichens will 

oEten be overlooked in the field unless a hand 

lens is used. Furthermore, because of the small 

size of the apothecia, very few species can be recognized at sight. All must be collected and examined later for morphological and chemical 

characters. Thallus color, however, is rather constant, light greenish or ashy gray. Only a few species 

have the deep green color of the equally common 

pyrenocarpous lichens, which, together with the 

Graphidaceae and Thelotremataceae, account for 

the majority of crusts that are collected in lowland tropical regions. 

This paper is part of an on-going project on a 

modern treatment of the lichen flora of Dominica. 

As I mentioned in the first part on the Parmeliaceae (Hale, 1971a), where climate and topography 

are discussed, Dominica has the greatest remaining 

areas of virgin or relatively undisturbed forest of 

all the islands in the Lesser Antilles. Though 

Mason E. Hale, Jr., Department of Botany, A'ntional Museum 

of Natural Histoy, Smithsoninn Institution, Washington, D.C. 

20560. 

easily accessible by car and well-kept trails, these 

forests have as yet suffered little from human activity. Logging operations in the Dleau Gommier 

Forest Reserve, for example, have ceased, at least 

temporarily. Other islands in the Antilles are by 

contrast much more intensively cultivated and undisturbed forests are confined to higher, often 

inaccessible mountain slopes. 

On my first trip to Dominica in 1969 I was interested primarily in foliose lichens, which proved 

to be relatively poorly developed. Many of the 

numerous crusts that I collected more or less randomly and blindly were actually thelotremes. The 

obvious richness of the family inspired me to undertake a second trip to the island in December 

197 1, once again under the Morden-Smithsonian 

Expedition, to concentrate exclusively on this family. A final brief trip for the same purpose was 

made in May 1972 under National Geographic 

Society support. Altogether, nearly 550 specimens 

of thelotremes were collected, making this one of 

the most intensive collecting programs ever undertaken for a crustose lichen group. 

Special thanks are due Mrs. William J. Morden, 

who supported most of the field work for this 

project. Assistance is also gratefully acknowledged 

from the National Geographic Society for a broader 

study OC the evolution of the Thelotremataceae in 

the Lesser Antilles, which included Dominica. 

Curators at a number of museums and universities 

have sent loans of type-specimens, and their help 

1 
2 SiMITHSONIAN CONTRIBUTIONS TO BOTANY 

and patience are deeply appreciated: Dr. T. Ahti 

(H), Dr. Reino Alava (TUR), Dr. Ove Almborn 

(LD), Dr. C. E. Bonner (G), Mr. hl. Skytte Christiansen (C), Mr. Peter James (BM), Mme JovetAst (P), Dr. I. hl. Lamb (FH), Dr. R. A. Maas 

Geesteranus (L), Prof. H. Merxmuller (M), Dr. 

Harald Riedl (W), and Dr. R. Santesson (UPS). 

I wish also to thank Dr. Chicita Culberson for 

help in identitying or verifying most of the chemical substances of the various specimens. 

Morphological Characters 

After curating and labeling each specimen, I 

studied the various morphological characters that 

appear to be most important for taxonomic purposes. The descriptions, therefore, are not exhaustive but should be adequate for species 

identification. Redinger (1936 ) and Salisbury 

(1972a, 1972b) have drawn up useful summaries 

of the essential features of this family and a number of the species which may also be consulted. 

THALLUS STRUCTURE 

The anatomy of crustose lichens is not as complex as that found in typical foliose lichens. Cortical structure is less distinct, and, lacking a free 

lower surface, the medullary hyphae usually penetrate the outer layers of bark periderm (or rock 

substratum). The algal layer (Trentepohlia) may 

also be indistinct. Large colorless crystalline inclusions are commonly found in the medulla, as in 

Ocellularia nigropuncta, new species, and Thelotrema interpositzim. 

Sections examined under a scanningelectron 

microscope show some of these structures vividly. 

Ocellularia olivacea, 0. perforata, and Thelotrema 

confusum, new species (Figure la,b,c), all with 

well-developed epiphloeodal thalli, have a cortex 

consisting of several layers of heavily gelatinized 

cells appearing almost paraplectenchymatous; the 

loosely organized medulla penetrates the periderm. 

The surface is essentially smooth and featureless 

(Figure 2a-d). In some species the cortex is similar 

but peculiar aculeate hyphae protrude from the 

surface in greater or lesser abundance (Figure 

3a,b). They appear to be collapsed, perhaps because of vacuum treatment needed for specimen 

preparation. This aculeate orientation, reported 

here for the first time, is characteristic of Myriotrema species (Ocellularia terebratula, Thelotrema clandestinzim) as well as a few species with 

emergent apothecia (T. depressum, T. praestans). 

I have observed similar orientation in several 

species of Graphis and Graphina. Most thelotremes, 

however, have an irregularly organized surface, as 

in 0. mordenii, new species (Figure 3c,d), and a 

poorly defined cortex. 

Leptotrema wightii has an unusual cortex previously described by many lichenologists. The 

thallus is very thick (up to 1 mm) and has a grainy 

surface (Figure 54. The cortex is arranged in tall 

columns with algae scattered vertically (Salisbury, 

1971) (Figure Id). 

The so-called hypophloeodal species lack the 

structures described above. The external thallus 

is reduced to a microscopic layer of hyphae spread 

thinly over the bark surface (Figure 4), and these 

hyphae penetrate several layers of periderm with 

very sparce symbiotic algae apparently located just 

below the periderm surface. Representatives of this 

group from Dominica include Ocellularia concolor, 0. pyrenuloides, Phaeotrema disciforme, new 

combination, Thelotrema leucomelaenum, and T. 

tenue, new species. 

APOTHECIAL CHARACTERS 

Several morphologists have studied the internal 

structure and probable ontogeny of ascocarps in 

the Thelotremataceae (Redinger, 1936; Johnson 

and Brown, 1941; Letrouit-Galinou, 1966). The most 

commonly studied species is Thelotrema lepadinum, which has a distinct exciple breaking away 

free from the apothecial wall. The following DOminican species have more or less similar structure: Ocellularia conglomerata, new species, 0. 

dominicana, new species, 0. exanthisrnocarpa (Figure 5a), Leptotrema occultum, L. subcompunctum, 

and Phaeotrema aggregatunz, new species. I will 

call this arrangement “lepadinioid” in the descriptions; Salisbury (1972a) uses the term “Thelotrema 

lepadinum group.” 

What appears to be an extreme stage of this 

configuration of apothecial wall and exciple is seen 

in the Chroodiscus-type apothecium (here termed 

“chroodiscoid”). The outer wall or margin is erect 

and becomes recurved (Salisbury, 1972b), almost 

as in Geaster, and the exciple usually persists as a 
NUMBER 16 3 

FIGURE 1 .-Cross-sections of epiphloeodal lichens viewed with a scanning-electron microscope 

(a, b, and c, sectioned obliquely; d, Fertically): a, Ocellularia olivacea (Hale 35226) (X 1000); 

6, 0. perforata (Hale 38145) (X 1000); c, Thelotrema cotzfusum, new species (Hale 37697) 

(X 500); d, Leptotrema ulightii (Hioram 5850 from Cuba) (X 500). 

second inner rim. The disc is widely exposed and foliicolous species of the Thelotremataceae, innothing resembling the discrete pore characteristic cluding in Dominica Ocellularia alborosella and 

of the other thelotremes remains (Figure 5b). This 0. clilatata. It is callcd the “Tlzelotrema platy. 

type was first recognized among the foliicolous cui-pzim group” by Salisbury (1972b). In the ablichens, but it seems identical to that in the non- sence of more detailed ontogenetic research, 
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 2.-Surfaces of epiphloeodal lichens viewed with a scanningelectron microscope: a, b, 

Ocellutaria nigropuncta, new species (Hale 35365) (X 500 and X 2000); c, d, 0. dominicana, 

new species (Hale 37967) (X 500 and X 2000). 

however, we cannot yet say that Chroodiscus is a 

good genus, that the Chioodisczwlike nonfoliicolous thelotreines should be transferred to Chroodiscus, or that Clzioodiscus should be transferred to 

Ocellulaiiu. Chroodiscoid species, incidentally, are 

distributed among the four presently recognized 

genera, Ocellzih in, Leptotrema, Phaeotieinu, and 

Theloti emu, on the world level. 

The majority of species in the Thelotremataceae 

are neither lepadinioid or chroodiscoid. The exciple is fused with the receptacle wall and the 

main rim or margin closes over most of the disc 
NUMBER 16 5 

FIGURE 3.--Surfaces of epiphloeodal lichens viewed with a scanning-electron microscope: a, b, 

Ocellularia olivacea (Hale 38054) (X 500 and X 2000); c, d, 0. mordenii, new species (Hale 

35764) (X 500 and X 2000). 

leaving only a small constricted pore. The upper ized apothecia and a pore flush with the thallus 

wall is usually carbonized. There are, broadly surface (Figure 5c,d). The apothecia are always 

speaking, three groups that can be recognized for quite small, 0.1-0.3 mm in diameter. Typical 

practical purposes on the basis of degree of emer- examples in Dominica are Ocellularia olivacea and 

gence from the substratum. One, represented by Thelotrema clandestinum. 

the “Myriotrema” type, has immersed noncarbon- A second group, which includes the majority 
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 4.-Surface and cross-sections of hypophloeodal lichens viewed with a scanning-electron 

microscope: a, Thelotrema Zeucorneiaenum, surface view (Hale 39491) (X 2000); b, T. leucomelaenurn, cross-section (Hale 35491) (X 500); c, d, T. tenue, new species (Hale 35430) (X 500 

and X 2000). 

of the species, has emergent apothecia, neither X third group, typified by 0cellular.ia rhobasally constricted or strongly raised (Figure 5e,f). dostroma and 0. cavata, has strongly emergent apoThey are usually 0.4-1.0 mm in diameter, although thecia Tvitli a partially constricted base (Figure 

in a few species (for example, Thelotrema praestans 55). The pore is small and sometimes annulate or 

in Figure 17e) nearly 2 mm in diameter. open at maturity, as in 0. fecunda (Figure llh,i). 
NUMBER 16 7 

FIGURE 5.-Types of apothecia in the Thelotremataceae (all X 10): a, Ocellularia exanthisniocarpa (type of 0. multilocularis); b, 0. alborosella (Hale 353ila); c, 0. olivacea (Hale 35243); 

d, Leptotrema wightii, (type of L. prevostianum); e, 0. perforata (Hale 37i27); f, L. bahianum 

(type of Thelotrema rudius var. dominicanum); g, 0. rhodostroma (Hale 38122); h, Phaeotrema 

leiostomum (Hale 37956); i, 0. coinparabilis (Hale 37372). 

Diameter varies between 0.6 and 2.0 mm. This Although Kylander continued to use Ascidiunz 

group was called Ascidium by Fee (1824), not so for strongly emergent species, no modern lichenmuch because of emergence but because of a dupli- ologists have employed it. The type of the genus, 

cate membrane in the perithecium (the nature of A. cinchonarum Fee, is synonymous with Ocelluwhich is not clear to me) and a marginate pore. hia cavata. Naturally there are intergradations 
8 S.\fITHSOSIAN COXTRIBUTIOXS TO BOTANY 

walls and periderm, acting a5 a kind of ramrod. 

Not all species with periderm inclusions have a 

columella, but as Redinger found for the Brazilian 

species the correlation is high. 

The range in deLelopment of a coluinella is very 

great (Figure 6), from the Tveak thin columella in 

Ocellzilal-ia mordenii, new species, to the multiple 

act i n oi cl 0 ce 1/21 la ?.in co mpnra b i 1 is 

(Figure 5i) and Leptotrema fisszim (Figure 19a,c). 

The most highly developed species is Ocellzi Inria 

glnziovii (Muller-A4rgau known from South America 

Vainio (1921: 184) recognized these species under 

a separate genus, Rlinbdodzsczis. 

s t r uc t iires of 

between the Ascidium-type and the more common 

emergent type, but I believe the descriptive term 

“ascidioid” is still useful. 

A final group forms a transition to the Graphidaceae. The apothecia become irregularly elongate 

with a lirelliform dehiscence. In Dominica this 

includes Phaeotrema leiostomum, which can produce both round and elongate apothecia (Figure 

511). Other species from South America have this 

trait developed much more conspicuously and have 

been described in the Graphidaceae (as Phaeographis chionodisca Redinger (1936:69)). They 

should probably be transferred to the Thelotremataceae. 

Redinger (1936) further divides the Thelotremataceae into two groups that differ in position 

of the ascogonial initial. In one group the initial 

is located on or just below the periderm surface 

so that at maturity the apothecial wall develops 

on and apart from the unbroken periderm (for 

example, Ocellzilaiia rlzodostroma and Phaeotrema 

disciforme, new combination). The other group, 

including most nonchroodiscoid species in Dominica, has ascogonial initials deeply embedded in 

the periderm. The maturing apothecia must break 

through this layer to reach the surface. Several 

layers of periderm cells are thereby included in 

the apothecial receptacle and can be easily recognized in sections (Figure 6f). This character deserves much more detailed study. 

Development of a central cylindrical columella 

is particularly characteristic of the Thelotremataceae, and for the most part presence or absence 

of a columella is a good species character (Figure 

6). As a rule, columellate species do not have perfect counterparts that lack a columella. Noteworthy 

exceptions in Dominica are Oce11zilaria papillata 

and 0. perfomto, each of which has a large population where a columella is clearly developed and 

a smaller one where it is either absent or very 

difficult to find. I have not recognized such 

ecolumellate species as new without correlating 

morphological or chemical characters. Columellate 

species always have heavily carbonized apothecial 

walls. 

Redinger (1936:6) proposed an interesting hypothesis that the columella plays a role in breaking 

open the periderm layer in species that develop 

deeply embedded. As the apothecia mature, the 

columella pushes up against the upper carbonized 

SPORES 

Spore size is an importar,t and relatively constant character. The range of spore length, for 

example, is usually quite small except for very 

large spores (more than 10Op) which may vary 

between 100~ and 300u. ’IVe can distinguish four 

very broad categories among the Dominican species: 

(1) spores less than 20~ long with 4-6 transverse 

locules, eg., Lcplot~enza wightii, Ocelliilnria terebi.atiila, Tliclotwnza clandestinum; (2) spores 

20y-36;t long with 6-10 transverse locules, e.g., 

Ocellzilarin perfoycita (Figure 7a), Thelotrema 

leiiconielaeizi(ln; (3) spores 40p-9Op long with 

12-24 transverse locules, e.g., Leptotyema decepturn, new species (Figure 5d), Ocellzihria exanthismocarpa; and (4) spores 80p-300p long with 

numerous locules, eg., Ocelliilnria rhodostyoma 

(Figure fb), Thelotrema praestans, T. tiiberculif e I’ ii m. 

The number of longitudinal septations in 

Leploti~ema and Theloti.ema varies according to 

length, from 1 or 2 septae in small spores (up to 

about 30p long) to 4-6 or more in large spores. 

Spores are colorless or brown. It is sometimes 

a problem to determine color in large-spored 

species of Leptotrema and Thelotyema where, on 

the one hand, a Thelotwma species might have 

some senile spores turning brown and on the other 

a Leptotrema will haye some colorless spores (Figure 7d). Jn Phaeotwma disciforme, new combination, in fact, one often finds a mixture of colorless, mature brown, and shriveled brown spores 

(Figure 7c). Shriveling (Figures 7c,4 is characteristic of brown-spored species; I know of no cases 

where Thelotrema or Ocellularia spores shrivel. 
NUMBER 16 

FIGURE 6.-Cross-section of apothecia (all about X 130): a, Ocellularia nigropuncta, new 

species (Hale 35365); b, 0. inordenii, new species (Hale 37764); c, 0. cavata (Hale 35134); d, 

Phaeotrerp leiostoniun (Hale 35203); e, Thelotrema leuconlelaenuin (Hale 35491); j, T. tenrte, 

new species (Hale 35430). 

9 
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 7.--Spores of the Thelotremataceae: a, Ocellularia perfornta (Hale 37784) (X 400): 

b, 0. rhodostroma (Hale 35440) (X 130): c, Phaeotrema discifortne, new combination, with a 

mixtute of colorless spores, brown spores, and shii>eled brown spores (Hale 37939) (X 400); 

d, Leptotrenia deceptutn, new species, with colorless muriform spores on the right (Hale 38110) 

(X 400). 

In any event, transitional species should be judged nation, or 0. sitbcaunla on Dominica) should not 

intliviclually, and the presence of a few anomolous be totally unexpected; nor shoultl these isolated 

brown-spored specimens among many colorless cases prejudice the value of spore color as a generic 

specimens (as in Ocellzrlnria feczinda, new conibi- character. 
NUMBER 16 

Generic Limits 

The generic delimitation of the Thelotremataceae has undergone many changes since the 

classic work of Acharius. Fee (1824), for example, 

established two genera, in addition to recognizing 

Theloti enla, based on external features and apothecial characters: Ascidizim and My? iotrema, both 

presently considered to be synonyms of Ocellzlla, fa. lfullei--Argau (1887a) proposed the most farleaching changes, using spore septation and 

coloration to separate four genera (in addition 

to foliicolous Chq oodisctis): Ocellzilaria (transtrersel y septate colorless spores), Phaeotrema 

(transFersely septate brown spores), Thelotrema 

(muriform colorless spores), and L e p t o t i e m u (muriform brown spores). 

TVhile Vainio, one of the truly creative thinkers 

in lichen classification, rejected many “spore 

genera,” almost all authors since Muller-Argau 

ha1 e accepted these genera (and, indeed, sporebased genera in other families). Poelt (1973:613) 

refers to these as “artificial genera,” yet almost 

an) character we select is in a sense artificial. I 

don’t believe this term does justice in that it 

implies spore characters are artificial. 

TVe are, in reality, faced with at least thiee 

choices for generic delimitation in the family: (1) 

one genus (Tlielotremcr ) to accommodate all thelotreinataceous species; (2) four spore-based genera 

(Ocelliilnria, Phaeolrema, Thelotrema, and Leptot)en70); (3) five or more genera (Ascidi~m, 

C 11 rood isc 11 Y, Myio t I em n, R h 0 b dod i ~c us, antl T 11 e lotren10) based on apothecial emergence and excipular structures. Most lichenologists, as I said, have 

opted for spore-based genera. TVhile a majority 

opinion does not necessarily prove the naturalness 

or validity of such genera, they do repiesent one 

possible biological differentiation, broadly correlated with chemical, morphological, and ecological 

characters (as I will discuss briefly under each 

genus). 

I cannot agree with Salisbury’s (1952a) recent 

rejection of spore-based genera in favor of a single 

massi\e genus Tl7elotrema with 400 species subdivided into “groups” according to excipular structure, a character that is difficult to identify. His 

kejs and descriptions show how cumbersome this 

approach is, for in each instance one must desig11 

nate the spore type, e.g., TIzelotiemu wightii with 

Leptot)ema-type spoies, etc. If- one is forced to 

make this critical distinction every time a species 

is identified and described, then they should be 

tieatetl as distinct genera, a more coherent antl 

certainly a inore pragmatic, working solution. 

Spore characters are well proven, even critical, in 

the taxonomy of noiilichenized fungi. A retrogressive gene1 ic concept in lichens will hardly encourage a better understanding of their biology and 

evolution. Mj experience in the Thelotremataceae 

suggests that spore-based genera are reasonably 

natural, easily manageable units that are probably 

diverging in terms of habitat requirements and 

biogenetic characters as reflected in chemical differences. 

Chemistry 

in aliiiost eier) other gioup of lichens studied, 

chemistr) is an impoi tant and fundamental character in uncle1 standing the evolution and population biolog) ol the Thelotremataceae. The family 

has an unexpectedly iich chemistry with a pretloniinance of P + depsidones and related compounds (Culberson and Culberson, 1968; Culberson and Hale, 1973). I Trill discuss the possible 

significance of this chemistry below under the 

species and hope to exploi e the broader aspects 

oh biochemical evolution of the family in another 

paper. 

-411 of the specimens were initially tested with 

p-phen) lenetliaiiiine and chromatographed on 

Brinkinan thin-1a)er plates according to the P reaction. I ~roultl recommend, however, that lichenologists consider using this reagent as little as 

possible because of the well-known carcinogenic 

propel ties of rolatile amines. Identification of the 

lichen substances can be made just as easily with 

TLC rrithout recouise to a P test except in doubtful cases. 

Standard thin-layer solvent systems (hexaneether-formic acid, 130:80:20 v/v, and benzenedioxane-acetic acid, 180:45:5, I./.) xvere used to 

identify the chemical content of each specimen. 

NOTATIC Acr~.-Ocellzrlaria feciinda, new combination, (hypoprotocetraric acid predominating), 0. 

rhodost 1 om0 (4-O-demethylnotatic acid predominating), antl Pliaeotremu leiostomzim (4-O-demeHYPOPROTOCETRARIC ACID ASD 4-o-DEhIETHYL- 
12 SXITHSONIAN COSTRIBUTIONS TO BOTANY 

and Thelotrema dominicanuna, new species. Several species of Ocellularia and this single 

Thelotrema react Pt red, but neither protocetraric 

acid or fumarprotocetraric acid can be demonstrated with TLC. Instead there are at least t\vo 

gray spots above both of these acids on the plates 

(and often a high H,SO, yellow spot). These are 

apparently derivatives of protocetraric acid. 

“PRAESTASS” UsKsowN.-Thelotrema praestans. 

This unidentified depsidone reacts P+ red orange 

and forms a very distinct orange to reddish brmvn 

spot below psoromic acid in both solvents. 

“OLIVXCEA” USKNO~t.l\.S.-OCellllla~~a olivacea. 

This P- compound produces a distinct H2S04 

gray spot below lecanoric acid in both solvents. 

It may be a depside. TVhile the lectotype of Ocellzilnria olivacea contains only this one spot, all West 

Indian collections, otherwise morphologically indistinguishable, contain a lower gray spot in addition. All specimens have thick white acetone 

residues. 

OTHER P - UsK~o~t.n.s.-Ocellziluria rimosa, new 

species, and 0. sorediatn, new species. The unidentified spots in these species may or may not represent lichen substances. Some species were examined 

for fatty acids by dipping the plates in water and 

trying to detect white water-repellant spots as the 

water evaporated. Some spots resembling fatty acids 

Tvere seen but microchemical tests with G.E. and 

G.A.TV. xvere unsuccessful. Another major group 

of P- compounds is terpenes. All TLC plates 

shou.ed numerous terpenes, identifiable as purple, 

blue, or brovm spots falling generally above RF 

.5 and fading Tvithin a few hours. Probably all of 

these originate from the bark and are not lichen 

products, as can be proved by co-chroinatographing 

the lichens Jvith lichen-free bark samples from the 

same specimen, 

0. concolor, 0. dominicnnn, new species, 0. macuInto, new species, 0. mordenii, new species, 0. 

pnpillata, Phaeotiwnn aggregntum, new species, P. 

d iscif o im e, new com bi na t i on, T 17 e 1 o t re m a d ep resslim, T. leticomelnenum, T. papillosum, new 

species, T. tenue, new species, and T. tuberculiferum. 

PicsrENTs.-OcellziInria rnordenii, new species 

and Leptotrema wightii (anthraquinones) and 0. 

fecunda, new combination, 0. rhodostroma, and 

Phaeotremn leiostomum (K- yellow compounds). 

h’0 SUBSTASCES PRESENT.-OCel/Zilnria albOYoSelk7, 

thylnotatic acid sometimes lacking). These acids 

were recently examined in detail by Culberson 

(Culberson and Hale, 1973), who discusses the 

biogenesis. Ocelluiarza fecunda and 0. rhodostroma 

were not included in that study. 

NORSTICTIC AcIn.-Leptotyenaa occultum. This is 

a rare acid in the Thelotremataceae (especially in 

contrast to the Graphidaceae) and is largely confined to Leptotrema on the world level. 

PROTOCETRARIC AcIn.-Ocellularia nigropuncta, 

new species, 0. perforata, Leptotl-emu baliianum, 

and Theloti~ema confusum. This well-known P + 

red acid is easily identified on TLC plates, It is 

sometimes accompanied by a higher gray spot that 

I have termed the “amplior” unknown because of 

its constant occurrence in Ocellula~ia amplior (Nylander) Redinger. Culberson believes it to be protocetraric acid combined with a carboxylic acid, 

comparable to funiarprotocetraric acid or the recently described succinprotocetraric acid. 

PsoRoarIc AcID.-Ocellulaiia antillensis, new 

species, 0. conipaq.abilis, 0. subcavata, 0. terebratula, Leptotrema fissum, Thelotrema carassense, T. 

clandestinum, T. conforme, T. leucinum, and T. 

interpositurn. This is the commonest P+ acid in 

the Thelotremataceae. It is almost alivays accompanied by an unknown compound “conpsoromic 

acid,” appearing as a spot below psoromic acid. 

It, too, is P+ yellow. 

STICTIC AND CONSTICTIC Acins.-Ocellzilnria conglomerata, new species, 0. dilatata, 0. exanthismocarpa, 0. pyveniiloides, Phneotrema obscurzim, new 

species, Tlielotwma glaucopallens, Leptotrema dcceptum, new species, L. microglaenoides, and L. 

szibcomptinctiim. This widespread and well-known 

acid is actually rather rare in the Thelotremataceae 

except for the genus Leptotrema. Constictic acid, 

appearing as a H,SO,+ yellow spot just below 

salazinic acid, is a constant accessory compound. 

One or two other gray spots, as yet unidentified, 

may occur between stictic and constictic acids on 

TLC plates. Another rarer accessory is the highest 

H,SO, t orange-red spot in the “quintaria” series 

(Hale, 19Tlb), still unidentified but being discovered in a variety of genera. 

VIRENSIC Acm.-Leptotrema spondaicum. This 

acid is PS ‘red. It was previously unknown in the 

Thelotrcmataceae, having been first identified in 

Alectoria. 

“CINCHONARUM” UNK~own.s.-Ocellzilaria cavata 
NUMBER 16 13 

zone lvhere rain I‘orest is optimally developed, the 

trees being 30-40 m high (Figures 2 and 3 in 

Hale, 19Sla, illustrate this forest type). It would 

appear that this lichen family is totally dependent 

on undisturbed rain forest as a primary habitat 

and site for speciation. When this forest is destroyed, it seems inevitable that the lichens associated with it will become extinct. In Dominica, 

I have seen only two species able to invade disturbed or planted forests; namely, Leptoti.ema 

spondaicum and Ocellularia exanthismocarpa. 

They have a highly successful means of propagation and become established very quickly. 

The anthraquinones are generally produced in the 

thallus, while the yellow compounds occur in the 

medulla of the apothecia and not in the thallus. 

Ecology and Habitats 

I paid considerable attention to the ecology of 

the species and habitat requirements in Dominica 

in the hope that this information would help in 

understanding the distribution and evolution of 

the species on the island. Lichenologists have all 

too often prepared revisions of tropical lichens on 

the basis of preserved herbarium specimens alone 

with little if any opportunity to observe the species 

in the field. Labels on specimens collected by nonprofessionals rarely give pertinent information on 

ecology and habitat. For example, tropical rain 

forests have distinct layering and great vertical 

variation. in microclimate. Elevational differences 

are also important and once again such data are 

rarely given in full on herbarium labels. 

Base level species in Dominica form a distinctive group. They grow on roots, lianas, buttresses, 

prop roots, and saplings within 2-3 m of the 

forest floor. Characteristic species are Ocellularia 

concolor, 0. papillata, 0. perforata, Thelotrema 

glaucopallens, T. tenue, new species, and Leptotrema deceptum, new species. At the opposite extreme we find canopy species that occur high above 

the forest floor in tree tops, exposed to great 

changes in insolation and wetting-drying cycles. 

These include Ocellularia mordenii, new species, 

0. olivacea, 0. subcavata, 0. terebratula, Phaeotrema leiostomum, Thelotrema clandestinum, and 

T. praestans. Other species on Dominica have less 

stringent habitat requirements or have been collected too infrequently to categorize. 

On a broader scale altitudinal differences are 

also pronounced. The rain forest becomes stunted 

in Dominica above about 800 m and gradually 

merges into an elfin mossy forest best developed 

above 1000 m. Thelotremes become scarce in this 

wind-swept zone where rain falls almost every day. 

Two species, however, Thelotrema leucomelaenurn 

and T. tubewuliferzim, are restricted to the highest 

elevations. Ocellularia rhodostroma and Thelotrema tenue, new species, occur here as well but 

also at lower elevation. 

The vast bulk of thelotreme collections were 

made between 300 and 800 m on Dominica, the 

THE THELOTREME FLORA 

As I had found with Paimelia. (Hale, 1971a), 

almost all previous collections of the Thelotremataceae on Dominica were made by 1%‘. R. Elliott 

in the 1890s. H. A. Imshaug also made about ten 

collections in 1963 which are preserved at Michigan 

State University. I have not had access to these. 

Elliott collected five specimens, identified by Vainio 

(1896, 1915) as follows: 

Thelotiema excavatum Vainio var. impressulum 

Vainio (hlorne Anglais, the type-locality, Elliott 

160, specimens in Bill and TUR). This is synonymous with Ocellularia perforata. 

Thelotiema leucomelaenum var. elevatum Vainio 

(Morne Anglais, Elliott 1534, specimen in TUR). 

This is T. leucomelaenum. 

Theloti emu rhodostromzim (hlontagne) Vainio 

(Morne Anglais, Elliott 521, specimens in BM and 

TUR, and Prince Rupert, Elliott 1305, specimen 

in TUR). These are Ocellularia rhodostroma. 

Thelotiema rudius Vainio var. dominicanum 

Vainio (Prince Rupert, Elliott 1303, specimen in 

TUR). This is the same as Leptotrema bahianum. 

The thelotreme flora of Dominica now includes 

48 positively identifiable species (and three additional species represented by poor material). I collected all of these except for Leptotiema bahianum. 

No one locality, of course, had anywhere near this 

number of species (see p. 14, “Collecting Localities”). The richest area was one site (16b) in the 

Dleau Gommier Forest Reserve which was being 

logged in December 1951; I collected a total of 

19 species there over an area of about 3 hectares. 

An adjacent site had 14 species as did the trail 

to Morne Anglais. Dleau Gommier locality 15 had 
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

12 different species. A logged rain forest on the 

Layou Road (12) had 11 species. Five other localities (11, 16c, 18a, 21, and 24a), all between 350 

and 750 m, had 9 species each. 

The homogeneity of the flora is remarkably high. 

Locality 16, for example, contains three adjacent 

collecting sites. Out of a total flora of 21 species 

there, 8 were found in common at each site. Three 

occurred at two of the three sites and 10 were 

collected at only one of the three sites. The greatest variance in homogeneity obviously comes about 

when we collect at two very different elevations or 

sites where habitat conditions are significantly 

different. 

It should be evident that we have not yet inventoried the whole thelotreme flora of Dominica. 

I would estimate that there are at least 55 species 

(including the three unidentified collections as 

possible additions). It would require at least one 

more intensive collecting trip, possibly two, to 

reach this number. 

COLLECTING LOCALITIES 

Specimens were collected at 28 separate localities 

on the island (Figure 8), and multiple visits were 

made at different times to six of these, giving a 

total of 35 collecting sites. These are coded as 

follows for convenience in citing specimens, going 

from localities at low elevation in the scrub forest 

to those at highest elevations. 

1. Scrub forest, Rodneys Rock, sea level. 

2. Scrub forest near Mero, elevation 15 m. 

3. Scrub forest on slopes of Barbers Block, ele4. Scrub forest near Calibishie, elevation 15 m. 

5. Palm forest at Rosalie Bay, elevation sea level. 

6. Rain forest along trail to Madjini, elevation 

7. Virgin rain forest logging area, Newfound8. Rain forest above Newfoundland, elevation 

9. Rain forest logging area, Castle Bruce Road, 

10. Dleau Gommier rain forest near Emerald 

11. Felled rain forest along new road cut about 

vation 60 m. 

30 m. 

land, elevation 250 m. 

300 m. 

elevation 300 m. 

Pool, elevation 450 m. 

2 km northwest of Pont Casse., elevation 

430 m. 

12. Rain forest logging area on Layou Road, elevation 420 m. 

13. Mixed secondary forest logging area at Brantridge Estate, elevation 500 m. 

14. Rain forest logging area just east of Pont 

Casse., elevation about 550 m. 

15. Virgin rain forest area at Dleau Gommier Forest Reserve, elevation 500 m. 

16a, 16b, and 16c. Virgin rain forest logging at 

Dleau Gommier Forest Reserve, elevation 

350-370 m. 

17. Remnants of rain forest at Felicite, elevation 

370 m. 

18a and 18b. Remnants of rain forest along road 

to Jean Estate, elevation 600-700 m. 

19. Citrus grove at Laudat, elevation 400 m. 

c 172128 

L 

Scotts Head 4+-./0-5 

FIGURE 8.-Outline map of Dominica showing localities where 

specimens of Thelotremataceae were collected. 
NUMBER 16 

20a and 20b. Remnants of rain forest in pastures 

above Giraudel, elevation 550-600 m. 

21. Virgin rain forest at Syndicate Estate, elevation 650-700 b. 

22. Mixed rain forest along trail through Middleham Estate, elevation 650-700 m. 

23. Rain forest at the base of Trois Pitons, elevation 650-700 m. 

24a and 24b. Mixed secondary forest along trail 

to Boiling Lake, elevation 650-700 m. 

25, and 25b. Rain forest-mossy forest along trails 

at Fresh Water Lake, elevation 800-900 in. 

26a and 26b. Rain forest-mossy forest along trail 

up Morne Anglais, elevation 800-1000 m. 

27. Stunted rain forest-mossy forest on slopes of 

Trois Pitons, elevation 900-1200 m. 

28. Mossy forest area on slopes of Morne Diablotin, elevation 1000-1200 m. 

Sixteen localities were visited on the first trip 

in January 1969 (1, 2, 5, 6, 7, 13, 14, 15, 17, 18b, 

19,'20b, 24b, 25b, 26b, and 28). On the second trip 

seven additional localities were found (3, 11, 12, 

16a-c, 22, 23, and 27) and five sites adjacent to 

localities previously were examined (18a, 20a, 24a, 

25a, 26a), all during December 1971. Four new 

localities were visited in May 1972 (8, 9, 10, 21). 

All specimens are deposited in US unless otherwise indicated and duplicates will be deposited 

in the British Museum (BM). 

15 

I have provided illustrations of the Dominican 

species as well as of most type-specimens for comparison. These photographs also serve to show the 

range of variation in pore size and apothecial 

emergence. I have found these X 10 photographs 

to be extremely useful in identification, more so 

than line drawings of cross-sections. All specimens, 

including the types listed, were tested with thinlayer chromatography. 

Ocellularia 

This is the largest genus in the family, comprising about 165 species on the world level or about. 

40 percent of the total 400 known species. I had 

almost the same percentage on Dominica (22 of 

48 species) but it makes up a disproportionately 

large percentage of the specimens actually collected, 410 out of 530 (77y0). Ecologically the 

genus is best adapted to mature rain forest and 

cannot withstand extremes of very dry lowland 

and wet exposed high elevation. The ten commonest species (Tvith number of specimens collected in 

parentheses) are: 0. yhodostroma (93), 0. fecunda, 

new combination (64), 0. perforata (55), 0. pupillnta (48), 0. siibcavata (40), 0. exanthismocarpa 

(25), 0. nigropzincta, new species (16), 0. olivacea 

(12), 0. mordenii, new species (lo), and 0. terebmtziZa (10). Rarities, known from single collections each, are 0. antillensis and 0. cavata. 

Key to the Species of Ocellularia 

1. Medulla of thallus and/or ascocarps pigmented. 

2. 

2. Medulla of ascocarps yellow or pink but thallus medulla white. 

3. 

3. 

1. Medulla of thallus and ascocarps white. 

4. Apothecia lacking a central columella. 

5. Apothecia immersed, 0.2-0.3 mm in diameter. 

Medulla and ascocarps entirely deep red .......................... 12. 0. mordenii, new species 

Pigment yellow; pore large, 0.3 mm wide ................ 10. 0. fecunda, new combination 

Pigment deep pink; pore tiny, 0.1 mm wide, annulate. ......... 18. 0. rhodostroma 

6. Thallus Pf yellow ....................................... ........ 22. 0. terebratula 

6. Thallus P- .......................................................................... 14. 0. olivucea 

5. Apothecia emergent, 0.6-2.0 mm in diameter. 

7. Disc open with a strongly recurved exciple. 

8. 

8. 

7. Disc closed, lacking an outer recurved exciple. 

9. Inner lepadinoid exciple strongly developed. 

10. Apothecia aggregated in groups. 

Exciple thick, disc irregularly elongate; stictic acid present 

Exciple thin and delicate, disc round; stictic acid absent .......... 

...... 7. 0. dilatata 

1. 0. alboroselln 

11. Apothecial groups round; stictic acid lacking ...................................... 

.......................................................................... 8. 0. dominicam, new species 
16 SMITHSOXIAN COSTRIBUTIONS TO BOTANY 

11. Apothecial groups oblong: stictic acid present 

................................................................. 6. 0. conglomerata, new species 

12. Spores large, 55p-75p .................................... 9. 0. exanthismocarpa 

12. Spores small, 15p ................................ 6. 0. conglomerata, new species 

10. Apothecia solitary. 

9. Inner exciple not developed. 

13. Thallus P-. 

14. Thallus distinctly fissured: pore area black rimmed ............................ 

14. Thallus continuous; pores not black rimmed .............. 15. 0. papillata 

15. Pore area black rimmed .................. 13. 0. nigropuncta, new species 

15. Pore area not black rimmed ................................ 16. 0. perforata 

4. Apothecia with a well-developed black central columella. 

16. Thallus with large scattered verrucose outgrowths becoming sorediate .................... 

20. 0. sorediata, new species 

16. Thallus lacking soredia. 

............................................................................. 19. 0. rimosa, new species 

13. Thallus P+ red. 

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

15. Apothecia large, 0.5-1.1 mm in diameter, more or less emergent. 

18. Thallus dull tan, hypophloeodal, P- ........................... 5. 0. concolor 

18. Thallus distinctly epiphloeodal, P+. 

19. Columella becoming actinoid: thallus P+ yellow .... 4. 0. comparabilis 

19. Columella remaining simple: thallus P+ red ................. 3. 0. cavata 

17. Apothecia smaller, 0.2-0.5 mm in diameter, barely emergent to immersed. 

20. Pore annulate: spores small with crowded septae, 5p-6p X 9p-llp ................ 

21. 0. subcavata 

20. Pore not annulate: spores generally 20p-30p long (except smaller in 0. 

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

maculata). 

21. Thallus P-. 

22. Pore tiny, less than 0.1 mm in diameter; spores 4-loculate .............. 

22. Pore larger, 0.1-0.2 mm wide; spores 6-7-loculate . 15. 0. papillata 

21. Thallus P+ red or orange yellow. 

23. Thallus very thin to hypophloeodal; Pt orange .................. 

.......................................................... 11. 0. maculata, new species 

........................................................... 17. 0. pyrenuloides 

23. Thallus thick, epiphloeodal. 

24. Thallus P+ red: apothecia barely emergent ....................... 

24. 

.................................................................. 16. 0. perforata 

Thallus P+ yellow; apothecia strongly emergent ............... 

..... 2. 0. antillensis, nev species 

1. Ocellularia alborosella 

FIGURES 5b, 96 

Ocellu la ria alborosella (N ylander) San tesson, 1952: 308. 

Grapliis alborosella Sjlander, 18633352 [tj pe-collection: Colombia, Lhfdig 2ti94 (H, lectotypc: FH-Tuck, UPS, isotypes); 

Figure 9b]. 

Thelotre!tia platycnrpllwt! L'ainio, 1923: 138 [typc-collection: 

Arinia, Trinidad, TIiaxter 57 (TUR, lectotype); Figure gal. 

Ocellularia platjcarpella (Vaino) Zahlbruckner, 1923:598. 

Thallus whitish gray, thin but continuous, shiny, 

smooth, forming colonies 5-8 cm broad; apothecia 

chroocliscoid, up to 1.5 min across, round to irregular, the marginal flap conspicuous, the disc flesh 

colored to white pruinose; hymenium about 60p 

high; spores 8, 5-6 loculate, 4p-5~ X 8p-15p, I01 weakly blue (Figure 5b). 

C~~sr~si RY.--No substances present. 

HAnriAr.-Tree trunks at mid or higher elevation in rain forest. 

This was the first corticolous chroodiscoid 

species to be described. It is characterized by the 

small spores and lack of lichen substances. The 

Dominican collections have a wider disc than the 

type froin Colombia but they fall within a predictable range of variation. It is a rare and easily 

overlooked species. 

SPECIMENS EXARIINED.-~ (38143), 16b (37883), 

2613 (3537la). 
NUMBER 16 

2. Ocellularia antillensis Hale, new species 

FIGURE 9g 

Thallus corticola, albidus, epiphloeodes, planus 

vel minute verrucosus, nitidus, 3-5 cm latus; apothecia numerosa, emergentia, 0.3-0.5 mm diametro, 

apice obfusca, columellata, columella 0.15 mm lata; 

ostiolum 0.1-0.12 mm diametro, intus nigrum; 

hymenium 7Op-801.1 altum; sporae 8: nae, transversim 7 loculatae, 7p X 20p, I+ coerulescentes 

(Figure 9g). 

CHEMrsTRY.-Psoromic and conpsoromic acids. 

HoLOTwE.-Trail to Madjini, low elevation 

windward rain forest, Dominica, elevation about 

30 m, Hale 35612, January 1969 (US). 

HABITAT.-upper trunk of stunted rain forest 

trees (30m). 

The whitish minutely verrucose thallus and 

numerous emergent apothecia with the large contrasting black pore area distinguish this rare species. 

Ocellularia comparabilis var. microcarpa Redinger 

has larger spores (6p X 30p-34y), sparse apothecia, 

and a smooth thallus, while 0. terebrata (Acharius) 

Muller-Argau has barely emergent apothecia (see 

Hale, 1972). I later collected this species from 

Trinidad and Panama, both at low elevation. 

Grande, Trinidad, Hale 37444 (US); Barro Colorado Island, Panama, Hale 38659 (US). 

SPECIMENS EXAMINED.-3 miles NW Of Sangre 

17 

3. Ocellularia cavata 

FIGURES 6c, 9f 

Ocellularia cavata (Acharius) Muller-Argau, 1882133499. 

Thelotrema cabaturn Acharius, 1812:92 [type-collection: Sierra 

Leone, Afzelius (H, lectotype; S, UPS, isotypes)].-Hale, 

1971b, fig. la. 

Ascidium cinchonarum Fee, 1824:96 [type-collection: On 

Cinchona, South America (G, lectotype; H, L, M, P, isotypes); Figure gel. 

Ocellularia cinchonarum (Fee) Sprengel, 1827:242. 

Ascidium cinchonarum f. intermedium Nylander, I867:319 

[type-collection: Pie de Cuesta, Colombia, Lindig 5 (H, 

lectotype; BM, G, M, isotypes); Figure 9c]. 

Ocellularia lindigiana Muller-Argau, 1887a:9 [type-collection: 

Colombia, Lindig 2757 (G, lectotype; BM, FH, UPS, isotypes); Figure 94. 

Oeellularia cinchonarym (Fee) Sprengel f. intermedia (Nylander) Zahlbruckner, 1923:586. 

Thallus whitish mineral gray, epiphloeodal, continuous, more or less roughened, forming colonies 

up to 8 cm across; apothecia ascidioid, 0.7-0.9 mm 

in diameter, apically carbonized with a large 

columella (Figure 6c); pore round, 0.07-0.12 mm 

in diameter, the top of the columella clearly visible; 

spores 8, 6-8 loculate, 6p-8p X 22p-28p, IS blue. 

(Figure 9f). 

CmMIsTRY.-"Cinchonarum" unknowns A and 

B and a yellowish pigment. 

HABITAT.-canopy branch in mid-elevation rain 

forest (about 550 m). 

I had previously typified this distinctive but rare 

Acharian species. It appears to be rare in the 

Lesser Antilles. The chief diagnostic features are 

the unusual chemistry, small ascidioid apothecia, 

and the large columella which protrudes into the 

pore area. 

SPECIMEN EXAMINED.-14 (35134). 

4. Ocellularia comparabilis 

FIGURES 5i, 9i 

Ocellularia comparabilis (Krempelhuber) Muller-Argau, 1883: 

318. 

Thelotrema comparnbile Krempelhuber, 1876:220 [typecollection: Rio de Janeiro, Brazil, Glaziou 5463 (M, lectotype; BM, C, G, M, P, UPS, W, isotypes); Figure 9i]. 

Thallus pale greenish to whitish mineral gray, 

epiphloeodal, smooth, forming colonies 5-1 2 cm 

broad; apothecia emergent, 0.8-1.2 mm in diameter, 

the walls carbonized, columella simple 'to actinoid; 

pore open, 0.2-0.5 mm broad, the white pruinose 

disc clearly visible; hymenium 50y-60&i high; spores 

8, 4-5 loculate, 5p-6y X lOu--14~i, IS blue. (Figure 5i). 

CHEivIsTRY.-Psoromic and conpsoromic acids. 

HABITAT.-LOWer trunks in rain forest (300- 

700 m). 

The columella in this rare species is imperfectly 

actinoid. To judge from collections made in South 

America, the range of variation is very great with 

gradations into related psoromic acid-containing 

species. For example, Ocellulnria discoidea (Acharim) Muller-Argau (Hale, 1972) and 0. antillensis, 

new species, have a simple columella. Ocellzilaria 

efformata (Krempelhuber) Muller-Argau has a very 

similar actinoid columella but the outer margin 

is semi-erect, leaving an open rather lacerate pore: 

0. De?.keIeynna (Montagne) Zahlbruckner and 0. 

glnziovii Muller-Argau both have highly developed 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 9.-Specimens of Thelotremataceae (all about X 10): a, Ocellularia platycarpella (lectotype in TUR); b, 0. alborosella, (isotype in FH-Tuck); c, 0. cinchonarum, f. intermedia (Iectotype in H); d, 0. lindigiana (lectotype in G); e, 0. cinchonarum (lectotype in G); f, 0. cavata 

(Hale 35134); g, 0. antillensis, new species (Hale 35612); h, 0. conglomerata, new species (Hale 

37959); i, 0. comparabilis (isotype in BM); i, 0. concolor (lectotype in G); k, 0. concolor (Hale 

38167). 
NUMBER 16 

actinoid discs; and 0. latilabia (Tuckerman) 

Muller-Argau has an actinoid columella and a 

recurved margin. 

SPECIMENS EXAMINED.-~ (38128), 21 (37372, 

38144). 

19 

5. Ocellularia concolor 

FIGURE 9j,k 

Ocellularia concolor Meyen and Flotow, 1843:230. 

TYPE-CoLLEcTIoN.-;\fanila, Philippines, Meyen 

(G, lectotype) (Figure 9j). 

Thallus light grayish tan, dull, mostly hypophloeodal, forming colonies 3-4 cm broad; apothecia 

numerous, 0.8-1.1 mm in diameter, more or less 

emergent, apically carbonized, columella present, 

0.2-0.3 mm wide; pore open, round to somewhat 

irregular, 0.3-0.5 mm in diameter, the rim black 

and the flat black columnar disc easily seen; 

liymenium 14Op high; spores 8, 5-7 loculate, 5p8p X 13p-20p, I+ blue (Figure 9k). 

CHEI\IISTRY.--NO substances present. 

HABITAT.-Lianas, saplings, base and lower trunk 

of trees in dense rain forest (300-370 m). 

This easily recognizable species was first described from the Philippines and appears to be 

pantropical. The tannish hypophloeodal thallus 

and habitat are characteristic. There are no comparable species in this kind of habitat. 

16b (37801, 37809, 38075). 

SPFCILIEKS EXAMINED.-8 (38167 ), 16a (37665), 

6. Ocellularia conglomerata Hale, new species 

FIGURE 9h 

Tliallus corticola, cinereo-albidus, epiphloeodes, 

continuus, nitidus, 4-5 cm latus; apothecia primo 

solitaria, 0.6-0.9 mm diametro, semi-emergentia, 

mox 2-3: nae conglomerata, margine exteriore 

erecto, excipulo interiore evoluto, pulverulento, 

disco aperto, carneo; ostiolum irregulare, latum; 

11) menium 70p altum; sporae 8: nae, transversim 

6-7 loculatae, 5p-611 X 15p-l8p, I+ coerulescentes 

(Figure 912). 

CHEMISTRY.-stiCtiC and constictic acids. 

HOLoTwE.-Trail to hforne hglais, Dominica, 

elevation about 900 m, Hale 37959, 13 December 

1971 (US). 

Chemistry places this species near 0. exanthisrnocaypa but the exciple is not so clearly separate 

froin the thick, erect, pulverulent margin, leaving 

the disc open. The spores are much smaller. Another ilrest Indian stictic acid-containing species, 

Ocellularia albooliuacea (Vainio) Zahlbruckner, 

has a more typically recurved margin and a thin, 

entire exciple. 

7. Ocellularia dilatata 

FIGURE lla,b 

Ocellularia dilatata Miiller-Argau, 1895:452. 

TYPE-CoLLECTION.-Rio de Janeiro, Brazil, 

Glaziou 5531 pro parte (GI lectotype; BM, isotype). 

(Figure 1 la). 

Thallus whitish gray, thin, dull, in part hypophloeodal, forming colonies 4-6 cm broad; apothecia 

chroodiscoid, large, up to 2.5 mm broad, round 

to elongate, marginal flap rather coarse; disc white 

pruinose; hymenium about 60p high; spores 8, 

6-7 loculate, 5p-6p X 15p-l7p, I+ blue (Figure 

llb). 

CHEMIsTRY.-stictic and constictic acids. 

HABITAT.-LO\Wr trunk area in higher rain forest 

zone (850 m). 

This rare species is related to Ocellzila?.ia albo~osella, but it has a thicker recurved margin and a 

different chemistry. I have collected a number of 

specimens at low elevation in Trinidad, suggesting 

that the species is more at home on continental 

land masses. 

SPECIMENS EXAXIINED.--~~~ (37695, 37702). 

8. Ocellularia dominicana Hale, new species 

FIGURE 2c,d; 10 

Thallus corticola, brunneo- vel viridi-albidus, 

epiphloeodes, continuus, nitidus, 5-10 cm latus; 

apothecia semi-emergentia, 0.7-1.1 mm diametro, 

apice incolores, columella nulla, solitaria vel 2-3 

arcte aggregata, habitu similia apothecio singulari, 

excipulo interiore evoluto, ostiol~um complente; 

ostiolum rotundum, 0.1-0.5 mm diametro; hymenium 6Op-8Op altum; sporae 8: nae, transversim 

6-7 loculatae, 6p8p X 16p-25p, I + coerulescentes 

(Figure 10). 

CHEMISTRY.-NO substances present. 
20 S.\lII'HSONIAN CONTRIBUTIONS TO BOTANY 

Thelotrema homothecintm Vainio, 1921:190 [type-collection: 

Irosin, Mt. Bulusan, Pror. Sorsogon, Philippines, Elmer 

14852 (TUR, lectotype; FH, G, L, S, US, W, isotypes); Figure Ild]. 

Ocellularia homothecia (Vainio) Zahlbruckner, 1923:593. 

Ocellularia porinoides (hlontagne and von der Bosch) Zahlbruckner, 1923:599 [not 0. porinoides (Acharius) Sprengel, 

1827 : 2421. 

Thallus white to ashy, thin, smooth, in part 

hypophloeodal, forniing colonies 1-4 cm across; 

apothecia numerous, semi-emergent, the rim becoming erect, noncarbonized and without a 

columella, the inner exciple distinct, intact, forming an apical pore within the main pore; main pore 

open, 0.2-0.4 mm in diameter; hymenium 160p180p high; spores 4-8/ascus, 18-22 loculate, 12p18p X 6Op-lOOp, If deep blue (Figure llg). 

CmhiISTRY.-stictic and constictic acids and 

rarely the higher "quintaria" unknown B. 

HABITAT.-Trunks, canopy branches, fence posts, 

and planted trees (300-900 m). 

This common pantropical species is unique because of the pore-within-a-pore arrangement. The 

large spores are also distinctive but it is not unusual to find sterile specimens. It is also one of 

the few species in the family to invade secondary 

forests, fence posts, etc., in wet areas up to very 

high elevation. 

16b (37982), 17 (35605), 20a (37780, 37781, 37786, 

3i788, 38019, 38025), 25a (37708, 37709, 37713, 

37714, 37715, 37716, 37849, 37996, 38069), 25b 

(35473, 35516), 26a (37692, 37944), 27 (38042, 

38055). 

SPECIAIEXS EXAAIINED.-8 (381 14), 11 (38009), 

FIGURE lO.-OcelIularia dorninicana, new species (Hale 37841) 

(about X 10). 

HoLorYpE.-Submossy forest on upper slopes of 

Trois Pitons, Dominica, elevation about 1000 m, 

Hale 37967, 6 December 1971 (US). 

H.%nnxT.-Trunk and lower base of trees in submossy forest (700-1 100 m). 

The apothecia are so tightly aggregated that I 

first interpreted the wall between them as a 

columella. The combined outer margin and exciple 

are minutely pulverulent antl crowded, filling the 

broad disc but leaving a distinct pore area. The 

species isas rather corninon at Trois Pitons but not 

found elsewhere. 

SPECIMEKS EXAMIKED.--~~ (38053), 27 (37683, 

37688, 37829, 37832, 37841, 38041). 

9. Ocellularia exanthismocarpa 

FIGURES 50, llc,g 

Ocellulnria esa~ithisviocarpn (Leighton) Zahlbruckner, 1923: 

590. 

Thelotrema esn~~thisni~ca~.p~nz Leighton, 1869: 169 [typecollection: Central Province, Ceylon, Tlircnites C.L. 97 

(BM, lectotype; H, isotype); Figure llc]. 

Thelotrema porinoides hlontagne and I on der Bosch, 1855: 

484 [type-collection: Java, Junghuhn 151 (L, lectotype; 

FH, G, P, Ti', isotypes); Figure lle]. 

Ocellularia mziltiloczilnris Zahlbruckner, 1912:369 [typccollection: Lanai, Hawaii, Roqk li (IV. holotypc); Figure 

Ocellulnria isertii T'ainio, 1915:40 [type-collcction: Guadeloupe, Zsert 87 (C, lectotype; TUR, isotype); Figure Ilfl. 

5a]. 

10. Ocellularia fecunda (Vainio) Hale, 

new combination 

FIGURE 11 h,i 

Thelotrenu domingeiise var. fecu?iduni T'ainio, 1896:208 

[type-collection: Richmond Valley, St. Vincent, Elliotst 243 

(BM, lectotype; TUR, isotype); Figure llh]. 

Ocellularia donzingensis var. fecunda (Vainio) Zahlbruckner, 

1923:590. 

Thallus light greenish or whitish mineral gray, 

epiphlocotlal, smooth and shiny, forming large 

colonies up to 30 cin broad; apotliecia large, strongly emergent, 1.5-1.8 inin in diameter, the cortex 

soon breaking a~vay antl revealing a light yellowis11 orange medulla, walls heavily carbonized, 
KUMBER 16 21 

FIGURE 1 I .-Specimens of Thelotremataceae (all about X 10): a, Ocellularia dilatata (isotype in 

BM); b, 0. dilatata (Hale 37702); c, 0. exanthismocarpa (lectotype in BM); d, 0. homothecia 

(isotype in FH); e, 0. porinoides (lectotype in L); f, 0. isertii (lectotype in C); g, 0. exanthismocarpa (Hale 37996); h, 0. fecuxda, new combination (lectotype in BM); i, 0. fecuuda, new 

combination (Hale 35218); j, 0. maculata, new species (Hale 38015); k, 0. motdenii, new species 

(Hale 37887). 
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

columella absent or weakly and irregularly developed; pore gaping, round to irregular and jagged, 

0.4-0.8 mm in diameter, the white pruinose disc 

clearly visible; hymenium 2OOp-25Op high; spores 

1--4/ascus, 30-40 loculate, 12-20 X 80p-I50p, rarely 

turning brown, I+ blue (Figure lli). 

CHExmiRY.-Hypoprotocetraric acid and in 

lesser concentration 4-0-demethylnotatic acid 

along with unidentified pigments. 

HAuITAT.-Basal area of trees into the upper bole 

and canopy in the rain forest (300-850 m). 

This is one of the few species in the family that 

can be recognized at sight in the field. The apothecia are very large and the yellow-orange medulla 

is exposed by rubbing the cortex of€. It is the 

second commonest species in Dominica and forms 

very extensive colonies. As far as I have determined, 

the species is restricted to the West Indies and has 

no close relatives. 

37816, 37820, 38003, 38046, 38061), 12 (37902, 

37906, 37913, 37920, 37921, 37931), 15 (35155, 

35167, 35218), 16a (37658, 37942, 37970, 38050), 

16b (37796, 37802, 37810, 37874, 37881, 37954), 

16c (37765, 37766, 37774, 37943, 37980), 18a 

(37725, 37957, 37962, 37986, 38039), 21 (38080, 

38081, 38089, 38103, 38106, 38107, 38132, 38137), 

22 (37639, 37644, 37828, 37838, 37843, 37844, 

37844a, 37845, 37856, 37858, 37960, 38026), 23 

(37679), 24a (37740, 37743, 37063, 37064), 25a 

(37711), 26a (38056). 

SPECIMENS EXAMINED.-8 (38 120), 1 1 (378 1 1, 

11. Ocellularia maculata Hale, new species 

FIGURE llj 

Thallus corticola, cinereo-albidus, epiphloeodes, 

planus, continuus, nitidus, 9 cm latus; apothecia 

nunierosa, minuta, 0.2-0.3 nim diametro, apice 

ol)lusc;i, columellata vel columella parce evoluta; 

ostiolum rotundum, 0.05-0.1 mm diametro, anguste 

nigrocinctum, columella protrusa; hymenium 100p 

altum; sporae 8:nae, transversim 4 loculatae, 5y-7p 

X llp-l3p, I- (Figure llj). 

CHEXIISTRY .--No substances present. 

HoI.oTYI.r.-Remnants of rain forest in pastures 

above Giraudel, Dominica, elevation 650 m, Hale 

38015, 11 December 1971 (US). 

Occlliilnria mncirlata might seem at first glance 

to be 0. papillata but the apothecia and spores 

are much smaller. There are no related species. 

SPECIXE.N EXAMMINED.-lSb (35524). 

12. Ocellularia mordenii Hale, new species 

FIGURES 3c,d; 66; Ilk 

Thallus corticola, pallide cinereo- vel brunneoalbidus, epiphloeodes, continuus, aetate rimosus, 

nitidus, usque ad 12 cm latus; medulla omnino 

sanguinea; apothecia dispersa, vix emergentia, 

apice obfusca, extus crasse verrucosa, verrucis fragilibus, rumpentibus, columella nulla vel parce evoluta (Figure 6b); ostiolum rotundum, 0.08-0.12 mm 

cliametro; hymenium circa 2OOp altum; sporae 

1-2:nae, transversim 24-30 loculatae, 15p-20y X 

70p150p, I+ coerulescentes (Figure Ilk). 

CHExmTRY.-Unidentified anthraquinone pigments only. 

HoLoiyPE.-Canopy branches in primary rain 

forest, Dleau Gommier Forest Reserve, Dominica, 

elevation about 370 m, Hale 37764, 12 December 

HAwrAT.-Canopy branches in rain forest (370- 

800 m). 

‘The deep red pigment and large warts are distinctive. It is a conspicuous canopy species. A very 

similar species in Cuba, 0. xanthost~oma (Nylander) Zahlbruckner, differs mainly in containing 

n I<- pale yellow-orange pigment. This lichen is 

named in honor 01 Alrs. William J. Morden. 

SI’PCIMENS EXAhlINED.-l2 (37909), 15 (35198), 

16a (38051), 16b (37791), 16c (37762, 37886, 37887, 

37010), 26a (37863). 

1971 (US). 

13. Ocellularia ngropuncta Hale, new species 

FIGURES 2a,b; 6a; 12a 

Thallus corticola, pallide brunneo- vel viridicinei eus, epiphloeodes, continuus vel rarius I imosus, 5-8 cm latus; apothecia numerosa, emergentia, 

0.6-0.8 mni diametro, apice obfusca, columella 

nulla (Figure 6a); ostiolum rotundum, 0.1-0.2 mm 

tlianietro, nigrocinctum; hJmeniuin 100~1-110p 

altum; sporae 8:nae, transversim 6-8 loculatae, 6p10:~ X 2Oy-32p I+ coerulescentes (Figure 12a). 

CmwsTRY.-Protocetraric acid. 

HOLOTYPE.-T~ ail to summit of Rforne Anglais, 

Dominica, elevation about 900 m, Hale 35365, 

January 1969 (US). 
NUMBER 16 23 

FIGURE IZ.-Specimens of Thelotremataceae (all about X 10): a, Ocellularia nigropuncta, new 

species (Hale 35365); b, 0. rimosa, new species (Hale 38052); c, 0. oliuacea (lectotype in G); 

d, 0. papilla,ta (lectotype in BM); e, 0. papillata (Hale 38098); j, 0. perforata (lectotype in 

BM); g, 0. excavata (lectotype in BM); h, 0. rufocincta (lectotype in G); i, 0. terebrata var. 

abbreuiata (isotype in FH); j, 0. pyrenuloides (isotype in W); k, 0. pyrenuloides (Hale 35524). 
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

HABrT.AT.-Lianas, lower bole and base, symphonea roots, rarely canopy of trees in the rain 

forest (300-900 m). 

The black-rimmed pore area and smooth thallus 

are diagnostic features. Other ecolumellate protocetraric acid-containing species include Ocellulaiia 

bonplandiae (Fke) Muller-Argau, which has a 

grainy surface, and 0. ueirzIcosa (Fee) MiillerArgau, which has an irregular open pore and 

larger spores (over 30y long). Under the scanningelectron microscope 0. nigropuncta, new species, 

has a finely pitted but otherwise smooth surface 

(Figure 2a,b), whereas 0. uerizicosa appears to 

have an aculeate orientation and 0. bonplandiae 

is verrucose and fissured. 

SPECIMENS EXAMINED.-~ (381 15, 38158), 10 

(38169), 12 (37903), 16a (37660), 16b (37798), 16c 

(37755, 37770, 37963), 18a (37948), 21 (38092, 

38094, 38138), 22 (37852), 24a (37751). 

14. Ocellularia olivacea 

FIGURES la; 3a,b; 5c; 12c 

Ocellularia oliuacea (Fee) Miiller-Argau, 1887a:7. 

Myriot, emu olivacea Fee, 1824: 103 [type-collection: On Bonplandin trifoliata, South America (G, lectotype); Figure 

Ocellularia oliuacea (Fee) C. and D. Van Overeem-De Haas, 

1 2c] . 

1922: 118 [superfluous combination], 

Tliallus light greenish to ashy white, shiny, 

thick, up to 0.5 mm (Figure la), forming extensive colonies up to 50 cm broad, sometimes finely 

cracked and fissured; apothecia very numerous, immersed, 0.15-0.3 mm in diameter, noncarbonized 

and without a columella; pore small, flush, about 

0.1 mm in diameter; hymenium 65p-75p high; 

spores 8, more or less uniseriate, 4 loculate, 4y6p X 8y-12p, If blue (Figure 5c). 

CHE;\IIsTRY.-“Olivacea” unknown and a second 

lower spot. 

HABITAT.-canopy branches to mid-bole in rain 

forest (240-700 m). 

This is one of three “Myiiotiemn” species that 

occur in Dominica, the other two being 0. terebrat iila and Thelotrema clandestinttm, all typically 

found on canopy branches in dense rain forest. 

Ocellztlaria oliuacea is characterized by the rather 

small spores and unusual chemistry. The West 

Indian material all produces a second spot below 

the “olivacea” unknown, probably a related depside. The type-specimen of 0. oliuacea lacks this 

spot. The surlace under the scanning-electron 

microscope is strongly aculeate (Figure 3a,b). 

8 (38164), 9 (38154), 11 (38001), 12 (37965), 15 

(35205), 16a (37992, 38054), 18a (37965), 21 

(38113). 

SPECIMENS EXAMISED.-7 (35226, 35243, 35249), 

15. Ocellularia papillata 

FIGURE 12d,e 

Ocellularia papillata (Leighton) Zahlbruckner, 1923:597. 

Thelotrema pnpillatunz Leighton, 1869: 169 [type-collection: 

Central Province, Ceylon, Thwaites C.L. 129 (BM, lectotype; 

H, G, P, S, UPS, W, isotypes); Figure 124. 

Thallus whitish to pale greenish ashy, smooth 

to more or less warty, shiny, forming colonies 3-12 

cni broad; apothecia numerous, 0.6-0.8 mm in 

diameter, immersed to slightly emergent, the upper 

walls carbonized, columella present, thin to scarcely 

developed to absent; pore round, 0.1-0.2 mni in 

diameter; hymenium 50p-70p high; spores 8/ascus, 

7-9 loculate, 6p-lop X 2Oy-36p I+ blue. (Figure 12e). 

CHEJIISTRY.--NO substance present. 

HABITAT.-ktplingS, prop roots, lianas, and tree 

bases at mid-elevation in rain forest (300-900 m). 

This common species has few outstanding features. The columella is usually present, although 

often weakly developed and difficult to find. Some 

specimens, as indicated below, seem to lack it entirely. The apothecia are inconspicuous, almost 

immersed to slightly emergent, but the pore is 

always distinct. The thallus is well developed, 

usually quite whitish, and smooth. There are no 

close relatives i\rithout lichen substances, although 

0. perfomto, which has protocetraric acid, is very 

close in general aspect except perhaps that the 

apothecia are more often emergent. Both species 

occupy similar base level habitats in the rain forest 

and were collected together at 11 of the localities. 

At eight other localities only one of the species 

was found. 

SPECIMENS EXAMINED (columella present).-S 

(38123, 38130, 38165, 38168), 9 (38131), 11 (37825), 

16a (37795, 37991), 16b (37662, 37804, 37805, 

37808), 18a (37721, 37964); 21 (38084, 38085, 

38097, 38098, 38152), 22 (37645, 37837, 37842), 23 

(37671, 37681, 37682), 27 (37850, 38072), 28 

(35428). 
NUMBER 16 25 

SPECIMENS EXAMINED (columella absent).-7 

(35227), 10 (38127, 38150), 16a (37655, 37661), 

16b (37792, 38074), 16c (38045) 21 (38086, 38088), 

22 (37649, 37834, 37851, 37854, 37855), 23 (37677, 

38965), 24a (37746). 

16. Ocellularia perforata 

FIGURFS Ib, 5e, 7a, 12f 

Ocellularia perforata (Leighton) Muller-Argau, 1892284. 

Thelotrema perforatum Leighton, 1866:447 [type-collection: 

Casiquiari, Brazil, Spruce 254 (BM, lectotype); Figure 12fl. 

Thelotrema terebratum var. abbreuiatulum Vainio, 1890:83 

[type-collection: Caraqa, Minas Gerais, Brazil, Vainio 1551 

(TUR, lectotype; BM, FH, UPS, isotypes); Figure 12i]. 

Ocellularia rufocincta Muller-Argau, 1893: 146 [type-collection: 

Foret du Rio General, Tondoz s.n. (Pittier 6107) (G, lectotype); Figure 12hl. 

Thelotrema excavatum Vainio, 1896:208 [type-collection: Mt. 

St. Andrew, St. Vincent, Elliott 153 (TUR, lectotype; BM, 

FH, isotypes); Figure 12g]. 

Thelotrema excauatum var. impressulum Vainio, 1896:208 

[type-collection: Morne Anglais, Dominica, Elliott 169 (BM, 

lectotype)]. 

Ocellularia terebrata var. abbreviata (Vainio) Zahlbruckner, 

1923502. 

Thallus whitish to pale greenish ashy, smooth 

to more or less warty with age, shiny, forming 

colonies 3-10 cm broad; apothecia numerous, immersed to moderately emergent, 0.4-0.6 mm in 

diameter, the upper wall carbonized, columella 

usually developed, thin 50p-70p in diameter, but 

sometimes lacking or difficult to find; pore round, 

0.07-0.15 mm in diameter; hymenium 55p-70p 

high; spores 8/ascus, 6-8 loculate, 7p-lop X 20~- 

34p, I+ blue (Figure 5e). 

CHEMIsmY.-Protocetraric acid and the “amplior” unknown as an accessory substance. 

HABITAT.-Buttresses, saplings, and bole to 

rarely canopy branches in rain forest at midelevation (300-850 m). 

This is one of the commonest species in Dominica 

at base level. The smooth whitish thallus (see 

cross-section in Figure 2b) and the small semiemergent apothecia resemble 0. papillata, as discussed above. Some of the specimens as listed below 

lacked any sign of a columella. 

SPECIMENS EXAMINED (columella present).-8 

(38149), 11 (37812, 37817, 37997, 38030, 38033), 

12 (37899, 38078), 14 (35139), 16a (37651, 37666, 

38007, 38032), 16b (37884), 16c (37759, 37761, 

37775, 37952), 18a (37727, 37728, 37730, 37732, 

37535, 37961, 37969, 37983, 38020), 20a (37784, 

37785, 38016, 38017), 21 (38108, 38134, 38145), 22 

(37646, 37647, 37830, 35857, 38036), 23 (37675), 

24a (37752), 26a (37698, 37701), 27 (37686, 37827). 

SPECIMENS EXAMINED (columella absent).-8 

(38160, 38163), 16a (38018), 21 (38093, 38140, 

38157), 23 (37674), 24a (38066), 27 (37684, 37847), 

26a (37700), 26b (35370). 

17. Ocellularia pyrenuloides 

FIGURE 12j,k 

Ocellularia fiyrenuloides Zahlbruckner in Magnusson and 

Zahlbruckner, 1943:46. 

TYPE-CoLLECTIoN.-\vaiIUkU, Maui, Hawaii, 

Fauyie 656 (UPS, lectotype; S, IV, isotypes) (Figure 12j). 

Thallus ashy mineral gray, thin to evanescent, 

continuous, forming colonies about 5 cm broad; 

apothecia numerous, nearly immersed to moderately emergent, 0.5-0.7 mm in diameter, walls 

heavily carbonized, columella present, 130p in 

diameter; pore round, 0.1-0.15 mm wide, pore area 

surrounded by a blackish ring, in part white pruinose, the pruinose top of columella visible through 

the pore; hymenium about 60p high; spores 8/ascus, 

6-7 loculate, 611 X 12p-I5y, I+ blue (Figure 15%). 

CHEMISTRY .-stictic and constictic acids. 

HAnITAT.-Tree branch in rain forest (650 m). 

The darkened pore area is similar to that in 

Thelotrema teniie, new species. There are no related species with stictic acid. It is still surprising 

to find a Hawaiian species on Dominica, but to 

me this suggests how poorly the Thelotremataceae 

have been collected. 

SPECIMEN EXAMINED.-18b (35524). 

18. Ocellularia rhodostroma 

FIGURES 5g, 7b, 13a 

Ocellularia rhodostromn (Montagne) Zahlbruckner, 1923:600. 

Ascidium rhodostromum Montagne, 1851:75 [type-collection: 

Cayenne, French Guiana, collection 1334 (P, lectotype); 

Figure 13~1. 

Thallus light greenish tan to gray, smooth to 

finely warty, continuous and epiphloeodal, the 

medulla white or pink, forming colonies 5-10 cm 
26 ShIITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 13.--Specimrns of Thelotremataceae (all about X 10): a, Ocellularia rhodostroma (lectotype in P); b, 0. subcavata (lectotype in H); c, 0. vaga (isotype in FH); d, 0. subca7jata (Hale 

37769); e, 0. terebratula (lectotype in H); f, 0. remanens (isotype in G); g, 0. sor~diata, new 

species (Hale 37908); h, Plraeotrema aggregatum, new species (Hale 35229); i, 0. terebratula 

(Hale 37793); 1, P. disciforme, new combination (lectotype in Bhl); k, P. aqztiliiium (lectotype 

in TUR). 
NUMBER 16 27 

broad; apothecia large and easily visible with the 

naked eye, ascidioid, 1.0-1.8 mm in diameter, the 

walls heavily carbonized and the medulla deep 

pink, readily exposed as the fragile cortex breaks 

away, columella lacking; pore round and small, 

0.05-0.1 mm in diameter, surrounded by a raised 

ring; hymenium 2OOp-275p high; spores l-2/ascus, 

25-30 loculate, 25p-4Op X lOOp-25Op, I+ blue 

(Figures 5g and 70). 

CHEMISTRY.-Hypoprotocetraric acid and in 

greater concentration 4-0-demethylnotatic acid 

along with unidentified pigments. 

HABITAT.-CanOpy branches to lower bole at all 

elevations on the island. 

Ocellularia rhodostroma is the commonest thelotreme on Dominica and the easiest to identify as 

well. The large ascidioid apothecia have a deep 

pink medulla exposed when the fragile cortex 

breaks. The pore is strongly annulate. While occurring at all elevations, it seems best developed 

at higher elevation into the mossy forest, perhaps 

because it competes with so few other species 

there. 

38146, 38166), 10 (38119), 11 (37815, 37823, 37824, 

38004, 38023, 38049), 12 (37894, 37916, 37917, 

37927, 37930, 37934), 13 (35279 35448), 14 (35111, 

35133), 15 (35160, 35170, 35188, 35314, 16a (37653, 

37938), 16b (37800, 37806, 37875, 37994, 37995), 

16c (37756, 37870, 37979), 18a (37729, 37731, 

37987), 18b (35517, 35522, 35523, 35587), 20a 

(37777, 38065), 21 (38035, 38147), 22 (37641, 37643, 

35650, 37835, 37861), 23 (37672, 38975), 24a (37736, 

37742, 37745, 37749, 37754, 38068), 24b (35629), 

25a (37705, 37707, 37720, 37840, 37846, 37848, 

35862, 37867, 37971, 37975), 25b (35469), 26a 

(37689, 37690, 37693, 37694, 37699, 37703, 37864, 

37866, 37885, 37947 38000, 38014), 26b (35354, 

35366, 35371, 35372, 35376), 27 (37831), 28 (35435, 

35440 ). 

SPECIMENS EXAMINED.-7 (35246), 8 (38122, 

19. Ocellularia rimosa Hale, new species 

FIGURE 12b 

Thallus corticola, brunneo- vel viridi-cinereus, 

epiphloeodes, nitidus, continuus sed aetate rimosus, 

usque ad 15 cm latus; apothecia numerosa, emergentia, 0.7-09 mm diametro, apice obfusca, columella nulla; ostiolum rotundum vel irregulare, 

0.1-0.2 mm diametro, nigrocinctum; hymenium 

110p-150p altum; sporae 8:nae, transversim 6-8 

loculatae, 7p-lop X 24p-32p, I + coerulescentes 

(Figure 12b). 

CHEMISTRY.--NO substances present. 

HoLoTYPE.-Base of tree in primary rain forest, 

Dleau Gommier Forest Reserve, Dominica, elevation 370 m, Hale 37978, 1 December 1971 (U.S.). 

HABITAT.-Prop roots, tree bases, and lianas in 

rain forest (370-430 m). 

Except for chemistry and slightly larger apothecia, this species is very close to 0. nigropuncta, 

new species. Both are base level species, but 0. 

Yimosa has a more restricted elevational range. 

The two species were collected together only at 

Dleau Gommier. 

SPECIMENS EXAMINED.-~ 1 (38052), 16a (37654), 

16b (37392, 37803), 16c (37758). 

20. Ocellularia sorediata Hale, new species 

FIGURE 13g 

Thallus corticola, viridi-albidus, epiphloeodes, 

continuus, planus, nitidus, usque ad 15 cm latus, 

sorediatus, soraliis sparsis, verrucosis, circa 1 mm 

latis; apothecia immersa vel semi-emergentia, 0.4- 

0.6 mm diametro, apice fuliginea, columellata; ostiolum, rotundum, 0.08-0.1 1 mm diametro; hymenium 6Op-7Op altuni; sporae 8:nae, transversim 

3-4 loculatae, 6p-8p X 12p-16p, I+ coerulescentes 

(Figure 13g). 

CHEMISTRY.-NO substances present. 

HoL0TYPE.-Logging area about 3 km northwest 

of Pont Cassi. on the Layou Road, Dominica, elevation 420 m, Hale 37908, 10 December 1971 (US). 

HAnrTAT.-hfid-bole of tree in rain forest; 

This is, as far as I know, the only sorediate 

species in the family. Large corticate tubercules 

are produced first, and some of these become 

coarsely sorediate. Although these peculiar structures seem to originate from the main thallus, there 

is still the possibility that they have an extraneous 

origin. 

SPECIMEN EXAMINED.-16b (38029). 

21. Ocellularia subcaoata 

FIGURE 13b,d 

Ocellularia subcavata (Nylander) Zahlbruckner, 1923:601. 
28 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 

22. Ocellularia terebratula Thelotrema subcavatum Nylander, 1876:561 [type-collection: 

Cuba, Wright 509 (H, lectotype; FH-Tuck, isotype); Figure 

13bl. 

Thelotrema vagum Vainio, 1896:209 [type-collection: Richmond Peak, St. Vincent, Elliobt 260 (BM, lectotype, FH, 

isotype); Figure 13~1. 

Ocellularia uaga (Vainio) Zahlbruckner, 1923:603. 

Thallus ashy whitish to greenish gray, shiny, 

more or less warty and cracked, forming colonies 

up to 12 cm broad; apothecia very numerous, emergent to ascidioid, 0.7-1.0 mm in diameter, crowded, 

walls carbonized, columella present, basally carbonized but usually colorless in the upper half; 

pore round, small, 0.1-0.2 mm in diameter, surrounded by a whitish, more or less raised ring, 

the center filled with the often protruding pruinose 

columella; hymenium about 1001~ high; spores 

8/ascus, usually uniseriate, 5-6 loculate with narrow locules, 4p-7p X 8p-2Op1 very rarely turning 

brown with age, I+ blue (Figure 13d). 

CHEMIsTRY.-Psoromic and conpsoromic acids. 

HABITAT.-Canopy branches and upper bole in 

rain forest (240-800 m). 

This is one of the commonest thelotremes in the 

canopy and can be recognized in the field with 

a hand lens because of the numerous crowded 

apothecia with a white annulate pore and protruding columella. The columella is well developed 

in cross-section but unique in being more or less 

noncarbonized in the upper half. 

(38161), 10 (38129), 12 (37893, 37896, 37898, 

37900, 37905, 37912, 37924, 37925, 37926, 37928, 

37929, 37933), 13 (35294), 15 (35158, 35209), 16a 

(37652, 37795, 38012), 16b (37662, 37794, 37865), 

16c (37767, 37769, 37772, 37773, 37924, 37941, 

37972), 18a (37724, 37889, 37949, 38038), 23 

(37680), 24a (37737, 37744, 37750), 26a (37691, 

37704). 

SPECIMENS EXAMINED.-7 (35248, 35252), 9 

FIGURE 13e,i 

Ocellularm terebratula (Nylander) Muller-Argau, 1887: 12. 

Thelotrema terebratuluin Nylander, 1867:315 [type-collection: 

Rio Negro, Colombia, Lindig 129 (H, lectotype; FH, C, hf, 

P, isotypes); Figure 13el. 

Thelotrema clandestinum f. remanens Nylander, 1867:315 

[type-collection: Monte del Moiro, Colombia, Lindig 90 

(H, lectotype; BM, G, P, isotypes); Figure l3fl. 

Thallus light greenish to ashy white, thick and 

continuous, smooth, shiny, often forming extensive colonies up to 15 cm broad; apothecia numerous, 0.2-0.4 mm in diameter, immersed, noncarbonized and without a columella; pore flush, 

0.05-0.1 mm in diameter, usually surrounded by 

a faint whitish ring; hymenium about 65p high; 

spores 8/ascus, 3-5 loculate, 5p-9p X 15p-20p, I+ 

blue (Figure 132). 

Ocellularia remanens (Nylander) Muller-Argau, 1887:7. 

CHE;MISTRY.-PsOromic and conpsoromic acids. 

HABITAT.-~~~o~~ branches and upper bole in 

rain forest (300-400 m). 

Ocellularia terebratula, previously known only 

from Colombia, is a typical canopy level species. 

It is very similar to Thelotrema clandestinum, as 

explained below, except for spore characters. 

SPECIMENS ExAMINED.-~ (38125), 10 (38126), 12 

(37904, 37918), 16a (37668, 37878, 37937, 38011, 

38040), 16b (37793). 

Phaeotrema 

This genus is very similar to Ocellularia in spore 

septation but the spores are brown and sometimes 

shriveled at maturity, especially in large-spored 

species. The iodine reaction often seems to be 

negative, while almost all species of Ocellularia 

are strongly I+ blue. On the world level, PhaeoKey to the Species of Phaeotrema 

1. Apothecia aggregated ........................................................ 23. P. aggregatum, new species 

1. Apothecia solitary. 

2. Apothecia large, about 0.6 mm in diameter, strongly emergent; columella lacking ........... 

2. Apothecia smaller, less than 0.5 mm in diameter, immersed; columella more or less 

developed. 

3. Pore 0.1-0.2 mm in diameter, sometimes elongate, with a white annulus ....................... 

......................................................................................................... 23. P. leiostomum 

3. Pore 0.5-0.1 mm in diameter, round, annulus lacking . ._ 26. P. obscurum, new species 

.......................................................................................... 24. P. disciforme, new combination 
NUMBER 16 29 

trema is represented by about 40 species, a rather 

small group, moreso because few of the species 

are well represented by herbarium material. In 

Dominica only six collections (four species) were 

found, these all occurring below 350 m elevation. 

23. Phaeotrema aggregatum Hale, new species 

FIGURE 13h 

Thallus corticola, pallide viridi-cinereus, epiphloeodes, planus, aetate rimosus, 8 cm latus; 

apothecia irregulariter aggregata, 0.4-0.6 mm 

diametro, immersa, intus incolores, columella nulla, 

excipulo interiore bene evoluto; ostiolum 0.1-0.2 

mm latum, in centro excipulo interiore congesto, 

albido-pruinoso; hymenium 110p-12Op altum; 

sporae 8:nae, transversim 4-5 loculatae, 8p X 16p 

(Figure 13h). 

CHEMISTRY.-NO substances present. 

HOL0TYPE.-Logging area at Newfoundland, 

Dominica, elevation 250 m, Hale 35229, January 

HABITAT.-upper branches of tree in rain forest. 

This unusual species has densely aggregated, 

immersed apothecia. The pore is usually distinct 

but may be obscured by the pulverulent exciple 

and margin, giving the appearance of a Phlyctis. 

1969 (US). 

24. Phaeotrema discif orme (Leighton) Hale, 

new combination 

FIGURES ic, 131, 14a 

Thelotrema disciJornze Leighton, 1869: 170 [type-collection: 

Central Province, Ceylon, Thwaites (BM, lectotype; H, 

isotype); Figure 1311. 

Thelotrema exalbrduin Stirton, 1881 : 184 [type-collection: 

Assam, Il’att s.n. (BM, lectotype)]. 

Thelotretna aquilinum Vainio, 1915:137 [type-collection: St. 

Croix, Boergesen s.n. (TUR, lectotype; C, isotype)]. 

Leptotrema exalbidum (Stirton) Zahlbruckner, 1923:634. 

Thelotrema galactinum Vainio, 1926:24 [type-collection: 

Maloapan, Mexico, Liebmann 7712 (C, lectotype); Figure 

13k]. 

Phaeotrema galactinum (Vainio) Zahlbruckner, 1932:245. 

Thallus whitish, dull, very thin and evanescent 

to hypophloeodal, forming colonies 2-4 cm broad; 

apothecia large, emergent, 0.7-1.1 mm in diameter, 

upper walls becoming suberect, carbonized, columella lacking, the inner exciple weakly developed 

and pulling away from the main margin, the 

white-pruinose disc clearly visible; pore open, 

round, 0.2-0.5 mm in diameter, the rim usually 

darkened; hymenium 120p-140p high; spores apparently 4/ascus, 20-24 loculate, 7p-12p X 40p551.1, colorless when immature but mostly turning 

brown and shriveled at maturity, I- (Figure 14a). 

CHEMISTRY.-NO substances present. 

HABITAT.-Lower trunk area (sea level to 370 m). 

A thorough examination of the lectotype showed 

that the spores are transversely septate without any 

longitudinal septae. Old shriveled spores, however, 

give the false impression of being muriform. It is 

a widespread pantropical species characterized by 

the large emergent apothecia without periderm 

inclusions, wide pore, pruinose rim and disc, and 

lack of a columella. 

SPECIMENS EXAMINED.-4 (Wirth Sen.), 16b 

(37939). 

25. Phaeotrema leiostomum 

FIGURES 5h, 64 14b,c 

Phaeotrema leiostomum (Tuckerman) Zahlbruckner, 1923: 

Thelotrema leiostornum Tuckerman, 1862: 407 [type-collection: 

608. 

Cuba, Wright 149 (FH, lectotype); Figure 14bl. 

Thallus ashy to pale greenish white, epiphloeodal, continuous, smooth and shiny, forming colonies 

2-5 cm broad; apothecia immersed to slightly emergent, 0.3-0.6 mm in diameter, weakly carbonized, 

columella absent to well developed, noncarbonized (Figure 64; pore round to irregularly elongate, surrounded by a raised ring, 0.1-0.3 mm 

long; hymenium 130p-150p high; spores 8/ascus, 

brown, 4 loculate, 8p-10~ X 16p-l8p, I- (Figures’ 5h, 14c). 

CHEMISTRY.-HypOprOtOCetrariC acid with or 

without 4-0-demethylnotatic acid along with pale 

unidentified pigments. 

HABITAT.-canOpy and mid-bole branches in 

rain forest (350-500 m). 

A raised annulate pore is characteristic of this 

species. Elongation of the apothecia was best developed in collection 37956. The extreme form can 

be seen in Phaeotrema lirelliforme (Tuckerman) 

Zahlbruckner, which lacks any lichen substances. 

When apothecia are lacking, the thallus of Phaeotrerna leiostomum alone would not permit identification. 
30 SMMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 14.--Specimens of Thelotremataceae (all about X 10): a, Phaeotrema disciforme, new 

combination (Hale 37929); 6, P. leiostomzim (lectotype in FH); c, P. leiostomum (Hale 33203): 

d, P. obscurum, new species (Hale 38091); e, Thelotrema cara~sense (isotype in FH); f. T. 

carassense (Hale 37873); g, T. clandestinuni (lectotype in G); h, T. subcaesium (lectotype in H); 

i, T. clandestinum (Hale 33136): j, T. confornie (lectotype in G); k, 7’. consanguineum (lectotype in G); 1, T. conforme (Hale 38062). 
NUMBER 16 31 

SPECIMENS EXAMINED.-15 (35203), 16a (37956), 

16b (37993). 

26. Phaeotrema obscurum Hale, new species 

FIGURE 14d 

Thallus corticola, pallide brunneo-albidus, tenuis 

vel hypophloeodes, 5 cm latus; apothecia modice 

emergentia, 0.3-0.5 mm diametro, apice fuliginea, 

columellata; ostiolum rotundum, 0.1 mm diametro, intus albido-pruinosum; hymenium circa 

12Oy altum; sporae 8:nae, transversim 5-6 loculatae, 

lop-14p X 24p-28p (Figure 14d). 

CHEMrsTnY.-stictic and constictic acids. 

HoLoTYPE.-Forty centimeters up trunk of a 

small tree, Rosalie Road above Newfoundland, elevation 300 m, Hale 38091, 10 December 1972 (US). 

HAnITAT.-Tree in rain forest. 

This tiny species could easily be overlooked. 

It is unusual to find a well-developed columella 

in apothecia this small. There are no other species 

in the genus with similar morphology. Phaeotrema 

albidulum (Nylander ) Muller-Argau also has 

stictic and constictic acids, but the apothecia are 

immersed and clustered and the spores larger 

(40P). 

Thelotrema 

Thelotiema is a common genus in Dominica, 

represented by 14 species among 80 collections. 

On the world level there are about 130 species. 

Variation in spore size is as great as in Ocellularia 

but longitudinal septae are always clearly developed (except in T. clandestinum). The six commonest species were T. praestans (21 collections), 

T. glaucopallens (16), T. clandestinum (lo), T. 

interpositum (7), T. leucomelaenum (7), and T. 

depressum (4). 

As I noted for Ocellularia, Thelotrema avoids 

the dry lowlands but is widespread in the rain 

forest. Thelotrema pupillosum and T. tuberculiferurn are restricted to the mossy forest. 

Key to the Species of Thelotrema 

1. Central columella present. 

2. Thallus thin, hypophloeodal; apothecia not emergent, small, less than 0.6 mm in diameter. 

3. 

2. Thallus distinct, epiphloeodal; apothecia emergent, 0.5-1.5 mm in diameter. 

4. Spores 2Op-4Op long, 4-8/ascus. 

3. Columella narrow; spores less than 2Op long .......................... 39. T. tenue, new species 

Columella very broad; spores more than 30p long ...................... 36. T. leucomelaenum 

5. Spores about 20p long; pore not annulate but becoming black rimmed .................. 

35. T. leucinum 

5. 

4. Spores more than loop long, l/ascus. 

6. Thallus smooth, P- or P+ orange. 

7. 

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

Spores about 40p long; pore becoming annulate ................................ 29. T. confurme 

6. Thallus warty-granular, Pf yellow ........................................... 34. T. interpositum 

Thallus P- ........................ :, ........................................................ 31. T. depressurn 

7. Thallus P+ orange ................................................................... 38. T. praestans 

9. Thallus greenish, thin, and shiny ................................................ 33. T. glaucopallens 

10. Spores less than 2Op ......................................................... 30. T. confusum, new species 

1. Central columella lacking. 

8. Apothecia immersed, the pore flush with thallus surface. 

9. Thallus whitish, thick, dull ............ .................... 28. T. clandestinum 

8. Apothecia emergent, pores not flush wi 

10. Spores more than 70y long. 

11. Thallus P-; collected in mossy forest zone. 

12. Apothecia 1 mm tall, eroding around a large area of the pore ...................... 

................................................................................................... 40. T. tuberculiferum 

12. Apothecia barely emergent, pore area not eroding ........................................ 

................................................................................. 37. T. pupillosum, new species 

11. Thallus P+ orange or yellow; collected in rain forest zone. 

13. P+ yellow; apothecia up to 1.0 mm in diameter ................... 27. T. carassense 

13. P+ red; apothecia about 0.5 mm in diameter .................................................. 

......................................... ..................... 32. T. dominicanum, new species 
32 SMITHSOXIAS CONTRIBUTIONS TO BOTANY 

27. Thelotrema carassense 

FIGURE 14e,f 

Thelotrema carassense Vainio, 1890:79. 

TYPE-COLLECTION.-Carassa, Brazil, Vainio 1523 

(TUR, lectotype; BM, FH, isotypes). (Figure 14e). 

Thallus greenish ashy mineral gray, epiphloeodal, 

continuous and shiny, forming colonies up to 15 

cm broad; apothecia semi-emergent, 0.8-1.0 mm 

in diameter, upper wall weakly carbonized, 

columella lacking; pore about 0.1 mm in diameter, 

with a weak raised ring; hymenium IlOy-13Op; 

spores 8/ascus, muriform, 1-2 X 8-10 loculate, 

12p-15p X 30p-40p, I+ blue (Figure 14f). 

CmMIsTRY.-Psoromic and conpsoromic acids. 

HABITAT.-Mid and lower bole of trees in rain 

forest (370-800 m). 

The Dominican material is not a perfect fit for 

the South American type, but it agrees in essential 

features, lack of columella, intermediate spores, 

and presence of psoromic acid. The apothecia are 

larger and spores smaller than in Vainio’s specimen 

(about 70y long). The taxonomy of the ecolumellate psoromic acid-containing Thelotremnta is 

unfortunately still incomplete. Thelotrema postpositum Nylander has large spores (more than 

loop), but all other species, excluding T. carassense, have small spores (less than 20p long). 

26b (37873). 

SPECIMENS EXAMINED.-16b (37859), 24b (37738), 

28. Thelotrema clandestinum 

FIGURE 14g,i 

Thelotrema clandestinum Fee, 1837:90. 

Thelotrema subcaesium Nylander, 1869: 120 [type-collection: 

Brazil, Glariou 2193 (H, lectotype; BM, C, FH-Tuck, UPS, 

US, W, isotypes); Figure 14hl. 

Thelotrema concretum Fee var. subcaesiurn (Nylander) Redinger, 1936:96. 

TYPE-COLLECTION.-on CinChOnU hZ?ZCifOliU in 

America, without collector (G, lectotype). (Figure 

Thallus whitish to greenish ashy gray, shiny, 

thick, and continuous, forming colonies up to 20 

cm broad; apothecia numerous, immersed, 0.15- 

0.3 mm in diameter, noncarbonized and without 

a columella; pore flush, round, 0.07-0.1 mm in 

diameter, usually with a whitish rim; hymenium 

14g) * 

65p-75y high; spores 8/ascus, uniseriate, 4-5 loculate, muriform with 1-2 longitudinal septae in at 

least one of the locules, 6p-IOp X 15p-20y, I+ 

blue (Figure 14i). 

CHEMrsTRY.-Psoromic and conpsoromic acids. 

HABITAT.-canOpy branches in rain forest (500- 

670 m). 

Thelotrema clandestinum has small spores with 

few longitudinal septations. It would, in fact, be 

classified as an Ocellularia unless the spores are 

carefully examined. Chemically similar 0. terebratula has only transverse septae. Both species have 

a thick, continuous thallus and occupy the same 

kind of habitat. They were collected at a total 

of ten localities but never together in the same 

one. 

(35103, 35105, 35136), 15 (35207), 16c (37768, 

37872, 37981 ), 24b (35624a). 

SPECIMENS EXAMINED.-13 (35298, 35393), 14 

29. Thelotrema conforme 

FIGURE 14jJ 

Thelotrema conforrne Fee, 1837389. 

Thelotrema consanguineurn Miiller-Argau, 1885b3398 [typecollection: Apiahy, Brazil, Puiggari 477 (G, lectotype); 

Figure 14kl. 

TYPE-COLLECTION.-On Cinchona, America meridionalis, without collector (G, lectotype) (Figure 

141). 

Thallus whitish mineral gray, smooth and shiny, 

epiphloeodal, forming colonies up to 10 cm broad; 

apothecia emergent, 0.8-1.0 mm in diameter, apically carbonized, distinct columella present; pore 

round, 0.1-0.15 mm in diameter, surrounded by 

a whitish, sometimes raised ring; hymenium 170y200p high; spores 4/ascus, muriform, 8-10 loculate 

transversely, 2-4 loculate longitudinally, 18p-2Oy 

X 35y-50p, I+ blue (Figure 141). 

CHEMIsrRY.-Psoroniic and conpsoromic acids. 

HABrTAT.-Lower bole and trunk in open rain 

forest (670 m). 

The Dominican material is a satisfactory match 

for FCe’s type-specimen, which has slightly smaller 

apothecia and a less consistently developed annulate pore. The spores are definitely in the intermediate size range in contrast to related T. 

leucinum which has small spores. 

SPECIMENS ExAMINED.-24a (37739, 38062). 
NUMBER 16 33 

30. Thelotrema confusunt Hale, new species Lep~ieu~ ‘701 (Bhf, lectotype; H, isotype) (Figure 

FIGURES lc, 15 

Thallus corticola, cinereo- vel viridi-albidus, 

epiphloeodes, planus, continuus vel rimosus, 

nitidus, 4-8 cm latus; apothecia numerosa, emergentia, 0.8-1.0 mm diametro, apice obfusca, 

columella nulla; ostiolum rotundum, 0.15-0.2 mm 

diametro, margine nigrocinctum; hymenium 120~- 

130p altum; sporae 4:nae, murales, transversim 6-7 

loculatae, longitudinaliter 0-2 loculatae, 6p-10~ 

X 12pL-26y, I+ coerulescentes (Figure 15). 

CHEmsTRY.-Protocetraric acid. 

HoLoTYPE.-Trai1 to Boiling Lake, elevation 650 

m, Hale 37747, 9 December 1971 (US). 

HAnITAT.-Canopy branches or lower bole in 

open areas (650-800 m). 

The black-rimmed pore and fissured thallus resemble Ocellularia nigropuncta, but the spores are 

clearly muriform. This presents an interesting example ol close morphological and chemical convergence with spore septation being the crucial 

diagnostic character. There are no related species 

in Thelotrema. 

SPECIMENS ExAMINED.-~~~ (37807), 26a (37697). 

FIGURE 15.-Thelotrema confusum, new species (Hale 37747) 

(X 10). 

31. Thelotrema depressurn 

FIGURE 16b,c 

Thelotrema depressurn Montagne, 1851:73. 

TYPE-CoLLEcTIoN.-Cayenne, French Guiana, 

16b). 

Thallus and apothecia as in Thelotrema praestans (see below); spores l-2/ascus, muriform with 

numerous transverse and longitudinal locules, 351.1- 

50p. X 150p-300p, I+ blue (Figure 16c). 

CHEMISTRY.-NO substances present. 

HABITAT.-canopy branches of trees in the rain 

forest (300-500 m). 

The type-material of Thelotrema depresszim is 

very fragmentary but color and chromatographic 

tests established that no P+ compounds were 

present. The four specimens from Dominica are 

identical. They may represent only a chemical 

population of T. praestans, which is morphologically identical, as far as we can judge from 

available specimens, but contains a strong P+ 

orange-red unknown substance. Thelot?.ema pl-aestans is more common (20 collections) and has a 

broader altitudinal range (250-800 m). 

37922), 15 (35150). 

SPECIMENS EXAhIINED.-8 (38141), 12 (37901, 

32. Thelotrema dominicanurn Hale, new species 

FIGURE I& 

Thallus corticola, pallide brunneo-albidus, epiphloeodes, continuus, nitidus, 4-6 cm latus; apothecia numerosa, emergentia, 0.3-0.4 mm diametro, 

apice obfusca, columella nulla vel parce evoluta; 

ostiolum rotundum, 0.05 mm diarrietro; hymenium 

140p-160p altum; sporae 2-4:nae, murales, transversim 24-30 Ioculatae, Iongitudinaliter 2-3 loculatae, 15p-20p X 8Op-9Op, I+ coerulescentes (Figure 16a). 

stances. 

HoLoTwE.-Tree along trail to Morne Anglais, 

Dominica, elevation about 800 m, Haze 35355, 

January 1969 (US). 

HAsrTAT.-Tree bole in higher elevation rain 

forest. 

The most unusual feature of this rare lichen is 

the presence of the “cinchonarum” unknown, previously thought to be restricted to the genus 

Ocellularia. Without a chemical test one might 

identify it with T. carassense, which has larger 

apothecia and a wider pore. Otherwise there are 

no comparable species in the New World. 

CHEMISTRY.-“CinChOnarUm” P -k unknown sub- 
34 SMITI-ISONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 16.-Specimens of Thelotremataceae (all about X 10): a, Thelotrema dominicanurn, 

new species (Hale 35355); b, T. depressurn (isotype in H); c, T. depressurn (Hale 38141); d, T. 

glaucopallens (lectotype in FH-Tuck); e, T. pechueli (lectotype in G); f, T. butuanurn (lectotype 

in W); g, T. homopastoides (lectotype in TUR); h, T. glaucopallens (Hale 35147); i, T. interpositurn (isotype in L); j, T. interpositurn (Hale 35388); k, T. leucinum (lectotype in G); 1, T. 

leucinum (Hale 38100). 
NUMBER 16 35 

33. Thelotrema glaucopallens 

FIGURE 16d,h 

Thelotrema glaucopallens Nylander, 1863:327. 

Pyrenula clandestiiia Acharius, 1814: 10 [type-collection: on 

Cinchona flava, South America (H, lectotype; S, isotype)]. 

Thelotrema laevigans Kylander var. avertens Nylander, 1867: 

318 [type-collection: Tequendama, Colombia, Lindig 893 

(H, lectotype; BM, FH-Tuck, G, P, FV, isotypes)]. 

Thelotrema pechueli Miiller-Argau, 1880:34 [type-collection: 

Quillu River, Angola, Pechttel-Loesclie s.n. (C, lectotype); 

Figure 16el. 

Ocellularia clandestina (Acharius) Miiller-Argau, 1877a:$. 

Thelotrema homopastoides Vainio, 1896:207 [type-collection: 

Richmond Valley, St. Vincent, Elliott 327 (TUR, lectotype: 

BM, isotype); Figure 16gl. 

Thelotrema butuanum Vainio, 1921: 183 [type-collection: Luzon, Philippines, Fenix, BS-28347 (FV, lectotype); Figure 

16fl. 

TYPE-CoLLEcTIoN.-Cuba, Wright (FH-Tuck, 

lectotype, as 28; L, UPS, isotypes) (Figure 16d). 

Thallus light greenish gray, very thin and shiny, 

breaking away in thin sheets, forming extensive 

colonies up to 15 cm broad; apothecia numerous 

to rather rare, immersed to slightly emergent, 0.2- 

0.4 mm in diameter, uncarbonized or slightly carbonized apically, without a columella; pore flush 

to barely raised, round, rarely angular, 0.05-0.12 

nim in diameter; hymenium 6Op-8Op high; spores 

8/ascus, muriform, 6-8 loculate transversely, 2-3 

loculate longitudinally, 7p-lop X 15p-24~~ I - 

(Figure 16h). 

CHEMISTRY.-stictic and constictic acids. 

HARITAT.--Bu ttresses, base of large trees, exposed 

roots, lianas, rarely upper bole and canopy in the 

rain forest (450-800 m). 

The thallus of this pantropical species is unique. 

It is extremely thin and shiny and tends to break 

away in thin waxy sheets when bruised or cut. 

The color is a pyrenocarplike dark yellowish green. 

Apothecia are usually quite numerous with flush 

pores having considerable variation in development 

of a rim, from no distinct annulus to an obviously 

raised area. Some anomalous specimens may even 

have semi-emergent apothecia, but the small I - 

spores and the presence of stictic acid will positively identify them. 

SPECIMENS EXAMINED.--IO (38082), 11 (37999, 

38008, 38047), 15 (35147), 16a (37670, 37876, 

38043, 38044), 20a (37783), 21 (38083, 38095, 

38155, 38156), 23 (37678), 26b (35367), 27 (37891), 

28 (35431). 

34. Thelotrema interpositurn 

FIGURE 16i,j 

Thelotrema rnterposztum (Sylander) Muller-Argau, 1881:526. 

Asczdzum znterposztum Sylander, 1863 336 [type-collection: 

Cuba, Wrzght 28 (H, lectotype; BM, FH, L, M, P, UPS, 

isotvpes); Figure lGz]. 

Thallus light brownish ashy gray, epiphloeodal, 

smooth to minutely warty, forming colonies 5-8 

cm broad; apothecia strongly emergent, 0.8-1.1 mm 

in diameter, n alls heavily carbonized, columella 

present; pore round, 0.15-0.2 mm in diameter, 

opening into a tubelike area leading to the pruinose top of the columella; hymenium 2OOp high; 

spores 2/ascus, muriCorm with numerous transverse 

and longitudinal locules, 28p-40p X 85p-125~, 

I+ blue (Figure 16j). 

CHEMISTRY .-PSOrOmiC acid. 

HABITAT.-CanOpy branches and upper bole 

area (sea level to 670 m). 

The coarsely verrucose thallus and large emergent apothecia with large spores distinguish this 

species. Two other columnellate psoromic acidcontaining species, T. conforme and T. leiiciniim, 

hale a smooth thallus and much smaller spores. 

(35148, 35174), 16a (37656, 37657), 16c (37760), 22 

(37853, 37976). 

SPFCIXIFNS EXAMINED.--E) (35388), 7 (35231), 15 

35. Thelotrema leucinum 

FIGURE 16kJ 

Thelotrema leucinum Muller-Aigau, 188ia: 10. 

TYPE-COLLECTION.-on Cinchona, South America (G, lectotype) (Figure 16k). 

Thallus light ashy gray, epiphloeodal, continuous 

to cracked with age, shiny, forming a colony about 

10 cm wide; apothecia numerous, nearly immersed 

to semi-emergent, 0.4-0.6 mm in diameter, upper 

wall carbonized, columella present, 0.2 mm wide; 

pore round, 0.05-0.12 mm in diameter; hymenium 

1 IOp-120p high; spores 8/ascus, muriform with 4 

locules transversely, 2 locules longitudinally, 7pIlk X 12u-18p, I+ blue (Figure 162). 

CHEhiIsTRY.-Psoromic and conpsoromic acids. 

H.4nITAT.-canopy branches in rain forest (300 

The type of T. Zezicinum, one of Fee’s Cinchona 

in). 
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

bark specimens, is in very poor condition and it 

is unfortunate that Muller selected it. Identification with the Dominica material is at best provisional, depending largely on the wide columella, 

small spores, and presence of psoromic acid. A 

better understanding of the species will come when 

more specimens are available. 

SPECIXIEN EXARfINED.-g (38 100). 

36. Thelotrema leucomelaenurn 

FIGURES 4a,b; 6e; l'ia,d 

Thelotrema leucomelaenurn Nylander, 1863:3?9. 

Thelotrema pauperius Nylander, 1867:318 [type-collection: 

Rio Negro, Colombia, Lindig (H, lectotype; BM, M, isotypes); Figure l'ib]. 

Thelotrema leucomelaenum var. elevatum Vainio, 1915: 137 

[type-collection: Morne Anglais, Dominica, Elliott 1534 

(TUR, lectotype); Figure 17~1. 

TypE-CoLLEcTIoN.-Fusagasuga, Colombia, Lindig 2777 (H, lectotype; BM, FH, G, M, P, UPS, 

isotypes) (Figure 17~). 

Thallus white, thin and in part hypophloeodal 

(Figure 4a,b), forming colonies up to 8 cm broad; 

apothecia immersed to semi-emergent, 0.6-0.9 mm 

in diameter, apically carbonized with a broad 

columella 0.2-0.3 mm wide; pore round to subirregular, 0.2-0.3 mm across, opening to the 

pruinose columella top (Figure 6e) ; hymenium 

130p high; spores 8/ascus, muriform, 4-6 loculate 

transversely, 2-3 loculate longitudinally, 12p-15y 

X 30~-3Gp, I- (Figure 17d). 

CHEMISTRY.-NO substances present. 

HABITAT.-Sma~kr branches of trees in mossy 

forest (800-900 m). 

A pantropical species, T. leucomelaenurn has 

been correctly identified by most lichenologists. The 

apothecia, while at most semi-emergent, stand out 

because of the black pore rim. The broad columella 

occupies most of the interior of the apothecia. 

37718, 37871), 25b (35491), 26a (38076). 

SPECIMENS ExAxfINE~.-25a (37706, 37710, 37717, 

37. Thelotrema papillosum Hale, new species 

FIGURE 17j 

Thallus corticola, cinereo-viridis, epiphloeodes, 

minute papillosus, nitidus, 6 cm latus; apothecia 

vix emergentia, 0.7-0.9 mm diametro, intus incolores, columella nulla; ostiolum rotundum vel 

irregulariter laceratum, 0.1-0.3 mm diametro; 

hymenium circa 2001~ altum; sporae l-2:nae, murales, loculis numerosis, 3Gp-40p X 14Oy-l50y, 

I- (Figure 17j). 

CHEMISTRY.--NO substances present. 

HoLoruPE.-Rain forest above Newfoundland, 

Dominica, elevation about 300 m, Hale 38124, 7 

May 1972 (US). 

HARITAT.-CanOpy branch in rain forest. 

This species would be easy to miss in the field. 

The small greenish warty thallus is not like any 

Thelotrema. The large noncarbonized apothecia 

are not numerous and seem lost among the thallus 

irregularities. The closest relative in terms of apothecial structure is Thelotrema tuberculiferum, 

which has large spores but strongly emergent eroding apothecia. 

38. Thelotrema praestans 

FIGURE 17e,g 

Thelotrema praestans Miiller-Argau, 1895:433. 

Thelotrema elliottii Vainio, 1896:207 [type-collection: Richmond Valley, St. Vincent, Elliott 246 pro parte (BM, lectotype); Figure 17fl. 

Tk.PE-CoLLECTIoN.-Rio de Janeiro, Brazil, Portelln s.n. (G, lectotype; BM isotype) (Figure 17e). 

Thallus light greenish to whitish gray, smooth 

and continuous, epiphloeodal, forming colonies 

5-10 cm broad; apothecia semi-emergent to strongly 

emergent, up to 2 mm in diameter, heavily carbonized with a thick columella; pore 0.15-0.3 mm 

in diameter, opening through a deep tube into the 

ascocarp; hymenium about 2OOp high; spores 

I-Z/ascus, muriform with many transverse and 

longitudinal locules, 3Op-45p X lOOy-275p, I+ 

blue (Figure 17g). 

CHEMIsTRY.-"Praestans" unknown. 

HABITAT.-CanOpy branches, less commonly bole, 

liana, or base in rain forest (250-800 m). 

This conspicuous thelotreme is related to T. dep7.esszim but differs in producing a P+ orange-red 

unknown substance just below psoromic acid in 

both liexane and benzene solvents. The apothecia 

are large and heavily carbonized. The fine structure of the cortex is aculeate, as in T. depressum. 

35273), 8 (38087, 38121, 38159), 9 (38135), I1 

SPECIMTXS EXAMINED.-7 (35237, 35244, 35254, 
NUMBER 16 37 

FIGURE lj.--Specimens of Thelotremataceae (all about X 10): a, Thelotrema leucornelaenuifz 

(lectotype in H); b, T. pauperins (lectotype in H); c, T. leucomelaenurn var. elemturn (lectotype 

in TUR); d, T. leucomelaenuni (Hale 35491); e, T. pmestuns (lectotlpe in G); f, T. elliottii 

(lectotype in Bhl); g, T. praestans (Hale 35273); h, T. ticberculiferum (lectotype in TUR); 

i, T. ,tuberculiferum (Hale 35519); 1, T. pupillosum, new species (Hale 38124); k, T. tenue, new 

species (Hale 35430). 
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

(37821, 37998, 38002, 38048), 12 (37907, 37911), 

15 (35171), 18a (37734, 37968), 18b (35521), 20a 

(37776), 26a (38070). 

39. Thelotrema tenue Hale, new species 

FIGURES 44; 6f; l'ik 

Thallus corticola, cinereo-albus, hypophloeodes 

(Figure 4c,d), nitidus, 7 cm latus; apothecia numerosa, immersa vel parce emergentia, 0.3-0.4 mm 

diametro, apice obfusca, columellata (Figure 6f) ; 

ostiolum rotundum, 0.05-0.1 mm diametro; hymenium 90p-1OOp altum; sporae 8:nae, murales, 

transversim 4-5 loculatae, longitudinaliter 1-2 

loculatae, 6p-7p X 17p-l9p, I- (Figure 17k). 

CHEMISTRY.--NO substances present. 

HOLOTYPE.--R/IOSS~ forest zons on trail to Morne 

Diablotin, Dominica, elevation about 1100 m, 

Hule 35430, January 1969 (US). 

HABITAT.-LOM'er trunk in rain forest to mossy 

forest. 

Superficially this species resembles Ocellularia 

pyrenuloides in having small blackish apothecia 

and a thin columella. Thelotrema leucomelaenurn, 

another hypophloeodal mossy forest species, has 

larger spores and a very broad columella. Other 

than these there are no related species. 

40. Thelotrema tuberculiferum 

FIGURE 17h,i 

Thelotrema tuberculiferum Vainio, 1915:136. 

TYPE-CoLLzCTI0N.-Savane-aux-Ananas, Guadeloupe, Duss 1497 (TUR, lectotype) (Figure 17h). 

Thallus ashy whitish, thin and in part evanescent, scattered, forming colonies 4-8 cm broad; 

apothecia strongly ascidioid, about 1.0 mm in 

diameter and 1.0 mm high, noncarbonized and 

lacking a columella, cortex eroding away toward 

the apex leaving a pulverulent area; pore round, 

0.1-0.2 mm in diameter, surrounded by a gray, 

sometimes raised ring and the broader pulverulent 

area; hymenium 140p-160p high; spores l-Z/ascus, 

muriform with numerous transverse and longitudinal locules, 30p-50p X 60p-120c(, I+ pale blue 

(Figure 17i). 

CHEMISTRY.-NO substances present. 

HABITAT.-Exposed moss-covered branches in 

mossy forest (800-900 m). 

This Thelotrema might be mistaken at first for 

a Pertusaria. It was first described from Guadeloupe and is probably endemic to the Lesser Antilles, occurring at high elevations in the upper 

limit of the mossy forest. 

SPECIMENS EXAMINED.-25a (377 19), 26a (37868). 

Lept otrema 

This genus differs from Thelotrema in having 

brown spores which tend to shrivel at maturity 

regardless of size and appear to be I - . Eight species 

occur on Dominica among my 36 collections (inKey to the Species of Leptotrema 

1. 

1. Columella lacking. 

Columella present, simple to actinoid .......................................................... 43. L. fissum 

Pore wide, 0.5-0.8 mm in diameter, the exposed disc dark brown to black ......................... 

Thallus thick with a columnar cortex; red crystalline inclusions often present ............. 

............................................................................................................................ 48. L. wightii 

Spores about 2Oy long ............................................................ 45. L. occultum 

Spores about 45p long ....................................................... 47. L. subcornpunctum 

2. 

2. Apothecia immersed to emergent with pores 0.1-0.2 mm in diameter. 

3. 

3. Thallus thin with a normal cortex lacking pigments. 

4. Apothecia with an inner lepadinoid exciple. 

5. 

5. 

4. Apothecia lacking a lepadinoid exciple. 

6. Pore distinct, 0.1-0.2 mm in diameter; spores 20~-30~ long 

6. Pore very tiny, less than 0.1 mm in diameter; spores large, more than 40y long. 

7. 

7. Pore not depressed, surrounding area the same color as the thallus. 

8. Apothecia immersed with flush pores ................ 47. L. subcompunctiim 

8. .4pothecia more or less emergent ................. 44. L. microgluenoides 

................................................................................................................ 46. L. spondaicum 

..41. L. bahianum 

Pore depressed in a darker area .......................... 42. L. deceptum, new species 
NUMBER 16 39 

cluding the collection of L. bahianum by Elliott). 

It is rarer than would guess from this number 

of collections since one species, L. deceptum, is 

represented by 2 1 specimens. Leptotrema spondaiczim was found 8 times, L. wightii 3 times, and 

tlie remainder once each. On the world level the 

genus has about 50 species. 

Leptotrema behaves in a peculiar way on 

Dominica. It is virtually the only thelotreme (5 

of 7 collections) in the dry scrub forest. It then 

essentially skips the rain forest zone and reappears 

in the submossy forest. Only 5 collections were 

made in tlie broad rain forest zone in contrast to 

23 above 900 m. 

41. Leptotrema bahianum 

FIGURE 5f 

Leptotrema bahiatiurn (Acharius) Muller-Argau, 188‘ia:12. 

Thelotrema lepadinurn var. bahianutrt Acharius, 1803: 132 

[type-collection: Bahia, Brazil, without collector (H, lectotype; L, UPS, isotypes)].-Hale, 19723193, fig. 2c. 

Thelotrema ritditis Vainio var. dominicanttm Vainio, 1915: 

136 [type-collection: Prince Rupert, Dominica, Ellioft 1303 

(TUR, lectotype); Figure 5fl. 

Tliallus corticolous, epiphloeodal, light tannish 

white, about 4 cm broad; apothecia numerous, 

emergent, 0.4-0.6 mm in diame:er, noncarbonized 

and without a columella; pore round, about 0.1 

mm in diameter; hymenium 150p high; spores 

8/ascus, muriform with 5-6 transverse locules and 

2-3 longitudinal locules, 12p-14p X 16y-24y, I- 

(Figure 5f). 

CHniIsmY.-Protocetraric acid. 

HABITAT.-Tree branch in dry scrub forest near 

sea level. 

This well-known Acharian species (Hale, 1972) 

seems to be very rare on Dominica since it has 

not been found since Elliott’s time. It is generally 

a loivland species in drier areas of the tropics. 

42. Leptotrema deceptum Hale, new species 

FIGURES ’id, 18 

Thallus corticola, brunneo-cinereus, epiphloeodes, planus, nitidus, continuus vel rimosus, usque 

ad 12 ciii latus; apothecia immersa, 0.3-0.4 mm 

diametro, intus incolores, columella nulla; ostiolum 

minutum, 0.02-0.05 mm diametro, depressum, 

ambitu obfuscans; hymenium 100p-120y altum; 

sporae obfuscae, 4:nae, murales, loculis numerosis, 

14p-28p X 4Op-7Op, I- vel I+ pallide coerulescentes (Figure 18). 

CHEXrIsmY.-Stictic and constictic acids. 

Hor,oTYPE.--Sloanea buttress, disturbed primary 

foiest at Middleham Estate, elevation about 670 m, 

Hole 37860, December 1971 (US). 

HABIrAT.-Buttresses and exposed roots, saplings, 

liarias, lower bole in higher elevation 1 ain forest 

(about 670 m). 

This species belongs to a large group of difficult 

species characterized by a very tiny pore and presence of stictic and constictic acids, including in 

Dominica L. micioglnenoides and L. subcompzinctzim. It is separated from related species by the 

smooth, usually fissured thallus and a distinct darker depressed area surrounding the pore, this area 

in turn surrounded by a whitish ring. Spent apothecia remain as large, open corticate pits. 

At first I had considered this (and similar 

5pecies) as having closer relationships to the Pyienulaceae. Mr. Richard Harris, a student of tlie 

Pyrenulaceae, gave the folio\+ ing opinion in support of placing L. deceptzim in the Thelotremataceae: 

hlost pFienocaipous lichens ha\e 5ome sort of carbonized 

“aall” surrounding the h\menium [I. deceptzctn is noncarbonized]. The spoies react nith IKI, a few pFrenocaips hale 

this reaction also but hate entirel\ different paraphFses. The 

paraph>ses [of L. deceptitrn] are unlike thme in the pyreno- 
40 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

carpous lichens in the relative lack of branching and especially in the very thick, coherent gelatinous wall or sheath 

around them (also found in the Graphidaceae). The asci are 

also unlike any pyrenocarpous asci. They are closest to 

Porina but lack any “chitinoid” apical ring. Thelotrema 

lepadinum has asci very like those in your specimen. Finally 

I have never found depsides or depsidones in any pyrenocarpous lichens. 

SPECIMENS EXAMINED.-21 (37373, 38104, 38105, 

38109, 38110, 38112, 38117, 38133, 38136, 38139, 

38142, 38148, 38151, 38153), 22 (37640, 37648, 

37836, 3i839, 38037), 23 (37673). 

43. Leptotrema fissum 

FIGURE 19a,c 

Leptotrema fissurn (Nylander) Muller-Argau, 1882a.333. 

Thelotrema fssuin NJlander, 18593258 [type-collection: Bouibon, Richard s.n. (H, lectotype; G, P, isotypes); Figure 

Leptotrema zntegtunz M~ller-~4rgau, 1887b:399 [type-collection: Australia, Sajer (G, lectotjpe, S, isotjpe); Figure 19b]. 

Leptotreina fisszcrn (Nylander) Zahlbiuckner, 1923.634 [supeiflous combination]. 

Thallus greenish ashy gray, epiphloeodal, smooth, 

continuous or cracked with age, forming colonies 

4-6 cm broad; apothecia semi-emergent, round to 

irregular, 0.7-1.2 mm wide, with thick erect to 

coarsely recurved carbonized upper walls, columella 

variable, barely developed to actinoid, the disc 

white pruinose; pore initially round but opening 

at maturity, up to 1 mm broad; hymenium about 

100p high; spores brown, muriform with 4-6 

transverse locules and 0-2 longitudinal locules, 

6p-7p X lOp--13p, I- (Figure 19c). 

CHEhlISTRY.-PSOrOmiC and conpsoromic acids, 

HABITAT.-canopy branches in rain forest (430 

The diagnostic features of this pantropical 

species are the open, often imperfectly actinoid 

disc, semi-erect lacerated margin, and presence of 

psoromic acid. It is close in these respects to Ocel121 la ria compara bilis. 

19aI. 

m). 

SPECIMEN EXAMINED.--~ 1 (38031). 

44. Leptotrema microgluenoides 

FIGURE 19d,e 

Leptotrerna microgluenoides (Vainio) Zahlbruckner, 1923:637. 

Thelotrema microgluenoides Vainio, 1896:206 [type-collection: 

St. Vincent, Elliott 266 (BM, lectotype); Figure 19dI. 

Thallus light whitish tan, epiphloeodal, continuous and shiny, forming a colony about 6 cm 

broad; apothecia semi-emergent, 0.8-1 .O mm in 

diameter, walls not carbonized, columella lacking; 

pore very tiny, 0.02-0.05 mm in diameter, depressed 

but with a tiny darker ring at the periphery; 

hymenium 15Op-17Op; spores brown, 4/ascus, muriform with numerous transverse and longitudinal 

locules, 18p-22p X 36p-48p, I- (Figure 19e). 

CHEwsTRY.-Stictic and constictic acids. 

HAnITAT.-Branches of tree in rain forest (430 

The lectotype of this species is rather fragmentary with very few well-developed apothecia. 

The thallus is continuous, but with a faint grainy 

appearance, thin and shiny. The Dominican material is’close to the type-specimen, although Vainio 

reported spores in the range of 120p long. I found 

spores about 50p long in the lectotype. The large 

complex of stictic acid-containing species (e.g. 

Leptotrenza monosporum (Nylander) hlullerArgau, L. phaeosporum (Nylander) Muller-Argau, 

and L. wclzisum (Krempelhuber) Zahlbruckner) is 

still far from resolved. 

m). 

SPEcmrrs EXAMINED.-~ 1 (37984). 

45. Leptotrema occultum (Eschweiler) Hale, 

new combination 

FIGURE 19f,g 

Thelotremu occulfum Eschweiler in Martius, 1833: 174 [typecollection: South America (M, lectotjpe; G, isotype); Figure 

19fl. 

Thallus tannish white, epiphloeodal, continuous, 

smooth to minutely pitted, forming colonies 3-5 

cm broad; apothecia numerous, immersed, 0.2-0.3 

mm in diameter, noncarbonized and without 

columella, inner lepadinioid exciple well developed and pulling away from the wall, lightly 

pruinose surface of hymenium visible; hymenium 

65p-7Op high; spores 4-8/ascus, brown, muriform 

with 5-7 transverse locules, 2-3 longitudinal 

locules, 9p-lop X 18p-21p, I- (Figure 19g). 

CHEMISTRY.-Norstictic acid. 

HABITAT.-Trees in dry scrub forest near sea 

level. 

Leptotrema occulturn can be typified with the 

Martius specimen in Munich and the isotype fragment in Geneva. It is virtually indistinguishable 
NUMBER 16 41 

FIGURE 19.--Specimens of Thelotremataceae (all about X 10): a, Leptotrema fissum (lectotype 

in H); b, L. integrum (isotype in S); c, L. fissuiit (Hale 38031); d, L. microgluenoides (lectotype 

in BM); e, L. microgluenoides (Hale 37984); f, L. occulturn (isotype in G); g, L. occz~ltz~rn (Hale 

35718); h, L. spondaicum (lectotype in FH-Tuck); i, L. spondaicum (Hale 35503). 

from pantropical L. compunctum (Acharius) 

Muller-Argau, which contains stictic acid in addition to norstictic acid. I am keeping L. occultum 

46. Leptotrema spondaicum 

FIGURE 19h,i 

Leptotrema spondaicum (Nylander) Zahlbruckner, 1923:640. separate until study of more material shows 

being generally smaller in norstictic acid-containing 

specimens) are valid or not. The L. compunctum 

group is very characteristic of dry scrub forest 

throughout the tropics. 

whether the and 'pore (spores Thelotrema spondaicum Nylander, 1863:330 [type-collection: 

Cuba, 

Thallus ashy white, epiphloeodal to evanescent, 

waxy, smooth, forming colonies 3-15 cm wide; 

apothecia numerous, round to irregular, 0.8-1.4 

35 (FH.Tuck, lectotype); Figure 19hl, 

SPECIMEN EXAMINED.-2 (35718). 
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

mm across, emergent, walls weakly carbonized, 

columella lacking; pore initially distinct, about 

0.2 mm wide, but soon opening, wide and irregular, 

to 0.5 mm wide, exposing the dark brown disc, 

outer rim becoming pulverulent; hymenium 150~- 

l7Op high; spores 1-2/ascus, brown, muriform with 

numerous transverse and longitudinal locules, 30p40y X 80~-130p, I- (Figure 19i). 

CmmsmY.-Virensic acid. 

HABITAT.-Trunks of primary and secondary 

trees in open pastures and citrus groves at higher 

elevations (600-700 m). 

Leptotrema spondaicum stands apart from all 

other thelotremes in having an open blackish 

disc surrounded by a thick, often pulverulent margin. One might be tempted to treat it as a species 

of Diploschistes. The chemistry is unique in the 

family. It is also different ecologically in that the 

primary habitat seems to be planted trees in disturbed areas. 

SPECIMENS EXAMINED.-lSb (35590), 19 (38077), 

20a (37779, 37789, 37790), 20b (35492, 35503), 26a 

(37696). 

47. Leptotrema subcompunctum 

FIGURE 20a,b 

Leptotrema su bcompunctum (Nylander) Zahlbruckner, 1923: 

640. 

Thetotrema subcompunctum Sylander, 1868:76 [typecollection: Lifu, Loyalty Islands, Thie'baut s.n. (H, lectotype; G, P, i$otypes); Figure 20al. 

Thallus light tannish mineral gray, epiphloeodal, 

continuous, smooth, dull, forming colonies 4-5 cm 

broad; apothecia numerous, 0.3-0.4 mm in diameter, immersed to slightly emergent, noncarbonized 

and without a medulla, the inner lepadinioid 

exciple clearly developed; pore round, 0.05-0.1 mm 

in diameter; hymenium 140p-160p high; spores 

FIGURE ZO.--Specimens of Thelotremataceae (all about X 10): a, Leptotrema subcompunctum 

(isotype in P); b, L. subcompunctum (Hale 37888); c. L. wightii (lectotype in FH); d, L. pavicans 

(lectotype in G); e, L. subconcretum (lectotype in BM); f, L. wightii (Hale 35882). 
NUMBER 16 43 

brown, I-Z/ascus, muriform with 8-1 1 transverse 

locules and 2-4 longitudinal locules, 10y-13y X 

45c(-55y, I- (Figure 20b). 

CHEmsTRY.-Stictic and constictic acids. 

HABITAT.-Tree bole in open rain forest (600 m). 

Except for the inner partially free exciple this 

species would probably be identified as L. microglaenoides. The thallus, however, is dull and the 

ascocarps barely emergent to immersed. This is 

the only collection that I have seen from the New 

World. 

SPECIMEN EXAMINED.-~~~ (37888). 

48. Leptotrema wightii 

FIGURES Id, 20c,f 

Leptotrema wightii (Taylor) Miiller-Argau, 1882:499. 

Endocarpon wightii Taylor, 1847: 155 [type-collection: 

Madras, India, Wight (FH, lectotype; BM, G, isotypes); 

Figure ~OC]. 

Thelotrema prevostianurn Montagne, 1849:292 [type-collection: 

Antilles, Prkvost s.n. (P, lectotype); Figure 5d]. 

Leptotrema prevostianurn (Montagne) Montagne, 1856:364. 

Theloftrema subconcretum Leighton, 1869: 169 [type-collection: 

Central Province, Ceylon, Thwaites 89 (BM, lectotype; H, 

G, P, S, UPS, W, isotypes); Figure 20e]. 

Phaeotrema subconcretum (Leighton) Miiller-Argau, 1887a: 

10. 

Leptotrema subconcretum (Leighton) Muller-Argau, 1891:277. 

Leptotrema fiauicans Miiller-Argau, 1888: 114 [type-collection: 

Guarapi, Paraguay, Balansa 4170 (G, lectotype; BM, M, 

W, isotypes): Figure 20d]. 

Thallus pale greenish mineral gray, thick and 

epiphloeodal, bulging up and flaking away with 

age, surface smooth but grainy in appearance, the 

columnar cortex with or without red crystal masses 

(Figure Id), forming colonies up to 10 cm broad; 

apothecia immersed, 0.2-0.3 mm in diameter, noncarbonized, without a columella; pore round, 0.08- 

0.11 mm in diameter, flush; hymenium about lOOy 

high; spores brown, 8/ascus, muriform with 4-6 

transverse locules and 1-2 longitudinal locules, 

9y-lly X 2Oy-28yL, I- (Figure 20f). 

CHEMISTRY.-NO substances present except for 

the unidentified anthraquinone pigment. 

HABITAT.-Tree trunks in dry scrub forest near 

sea level. 

This pantropical species has been discussed in 

detail by Salisbury (1971). It seems to occur at 

low elevations in dry scrub forest and extends 

northward into temperate forests of North America and southward into Argentina. 

SPECIMENS EXAMINED.--~ (35582, 35584), 3 

(3 29 1 8). 

Literature Cited 

Acharius, E. 

1803. Methodus qua onznes detectos Lichenes. 394 pages. 

Stockholm. 

1812. Anmarkningar vid Lafslagtet Thelotrema med 

Nogare Bestammande af Dess Arter. Kongliga Vetenskapliga Academiska Nya Handlingar, 1812:79- 

95. 

Synopsis Methodica Lichenurn. 392 pages. Lund. 

The Constituents of Some Species of the Thelotre. 

mataceae. The Journal of Japanese Botany, 43:316- 

323. 

1814. 

1968. 

Culberson, C. F., and W. L. Culberson. 

Culberson, C. F., and M. E. Hale, Jr. 

1973. 4-0-Demethylnotatic Acid: A New Depsidone in 

Some Lichens Producing Hypoprotocetraric Acid. 

The Bryologist, 76:77-84. 

Fee, A. L. 

1824. Essai sur les cryptogames des ecorces exotiques 

oficinales. 167 pages. Paris. 

1837. Essai sur les cryptogames des ecorces exotiques 

oficinales, II: Supplement et Revision. 178 pages. 

Paris. 

Hale, M. E., Jr. 

1971a. Morden-Smithsonian Expedition to Dominica: The 

Lichens (Parmeliaceae). Smithsonian Contributions 

to Botany, 4:l-25. 

1971 b. Studies on Parmelia Subgenus Xanthoparrnelia 

(Lichenes) in South Africa. Botaniska Notiser, 124: 

343-354. 

Typification of Species in the Lichen Family Thelotremataceae Described by Acharius. Botaniska 

Notiser, 125:186-198. 

1972. 

Johnson, G. T., and W. L. Brown 

1941. Stages in the Development of Thelotrema interpositurn. Mycologia, 33: 60 1-608. 

Krempelhuber, A. 

1876. Lichenes Brasiliensis Collecti a D. .4. Glaziou in 

Provincia Brasiliensi Rio Janeiro. Flora, 59:217-224. 

1866. Lichenes Amazonici et Andini. Transactions of the 

Linnean Society of London, 25:433-460. 

1869. The Lichens of Ceylon Collected by G. H. K. 

Thwaites. Transactions of the Linnean Society of 

Leighton, W. A. 

London, 27: 161-185. 
44 SMITHSONIAX CONTRIBUTIONS TO BOTANY 

Letrouit-Galinou, M.-A. 

1966. Recherches sur l’ontogknie et l’anatomie comparkes 

des apothkcies de quelques discolichens. Revue 

Bryologique et Lichdnologiques, 34:413-588. 

1943. Hawaiian Lichens. Arkiv for Botanik, 31A(1): 1-96. 

1833. Flora Brasiliensis. Stuttgart. [Lichens, by F. 

Magnusson, A. H., and A. Zahlbruckner 

Martius, C. F. P. von 

Eschweiler, I(part 1): 1-390.1 

Observationes in Itinere Circum Terram Institutae: 

Lichenes. Novorum Actorum Academiae Caesareae 

Leopoldino-carolinae h’aturae Curiosorum, Supplement 1, 19:209-232. 

Meyen, F., and J. Flotow 

1843. 

Montagne, J. F. 

1849. Sixieme centurie de plantes cellulaires exotiques 

nouvelles, cryptogamae taitensis. Annales Sciences 

Naturelles Bontanique, series 3, 12:285-320. 

Cryptogamia Guyanensis seu Plantarum Cellularium 

in Guyana Gallica Annis 1835-1849 a C1. Leprieur 

Collectarum Enumeratio Universalis. Annales Sciences Naturelles Botanique, series 3, 16:47-81. 

1856. Sylloge Generum Specierumque Cryptogamarum. 

498 pages. Paris. 

1851. 

Montagne, J. F., and R. von der Bosch 

1855. Lichenes. Pages 427-494 in F. Junghuhn, Enumeratio Plantarum Ques in Insulis Java et Sumatra 

Detexit. 

Muller-Argau, J. 

1880. Lichenes Africa Occidentalis a C11. Dr. PechuelUvesche et Soyou e Regione Fluminis Quilla et ex 

Angola Missi, in Mus. Bot. Reg. Berolinensi Servati. 

Linnaea, 43:31-48. 

1881. Lichenologische Beitrage XIV. Flora, 64:513-527. 

1882a. Lichenologische Beitrage XV. Flora, 65:327-336. 

188213. Lichenologische Beitrage XVI. Flora, 65:492-509. 

1883. Revisio Lichenum Meyeniorum. Jahrbuch der 

Konlige Botanische Gartens zu Berlin, 2:308-319. 

1887a. Revision Lichenorum Fkeanorum. Revue Mycologique, 35: 1-16 

1887b. Lichenologische Beitrage XXVI. Flora, 70:396-402. 

1888. Lichenes Paraguayensis. Revue Mycologique, 10: 

1891. Lichenes Victoryenses. h’uovo Giornale Botanico 

1892. Lichenes Exotici. Hedwigia, 31:276-288. 

1893. Lichenes. In J. Durand and H. Pittier, editors, 

Primitiae Florae Costaricensis. Bulletin Socidte‘ 

Botanique Bergique, 32: 122-173. 

Thelotremeae et Graphideae Novae Quae Praesertim 

ex Herb. Reg. Kewensi Exponit. Journal of the 

Linnean Society of London, Bo’tany, 303451-463. 

113-120. 

Italiano, 23276-279. 

1895. 

Nylander, W. 

1859. Lichenes Exotici. Annales Sciences hTaturelles Botunique, series 4, 11:203-264. 

1863. Lichenes. In J. Triana and J. E. Planchon, editors. 

Prodromus Florae Novo-Granatensis ou Enumbration des Plantes de la Nouvelle-Grenade. Annales 

Sciences Naturelles Botanique, series 4, 19:28&382. 

Lichenes. In J. Triana and J. E. Planchon, editors, 

Prodromus Florae Novo-Granatensis ou Enumkration 

des Plantes de la Nouvelle-Grenade. Additamentum. 

Annales Sciences Naturelles Botanique, series 5, 7: 

1868. Synopsis Lichenum Novae Caledoniae. Bulletin Societe Linnee Normandie, series 2, 2:39-140. 

1869. Lichenes in Brasilia a C1. Glaziou Lecti. Flora, 52: 

1876. Collemacei, Caliciei, Cladoniei et Thelotremei 

1867. 

301-354. 

117-126. 

Cubani novi. Flora, 59:558-562. 

Classification. Pages 599-632 in V. Ahmadjian and 

M. E. Hale, editors, The Lichens. New York: Academic Press. 

Poelt, J. 

1973. 

Redinger, K. M. 

1936. Thelotremataceae Brasiliensis. Arkiv for Botanik, 

28A(8): 1-122. 

Salisbury, G. 

1971. Thelotrema wightii (T. Tayl.) Nyl. Portugaliae 

Acta Biologica, series B, 11:35-37. 

1972a. Thelotrema Ach. sect. Thelotrema, 1: The T. 

lepadinum group. Lichenologist, 5:262-274. 

1972b. Thelotrema sect. Thelotrema, 2: The T. platycarpum group. Revue Bryologique et Lichdnologique, 38:281-290. 

Foliicolous Lichens, I: A Revision of the Taxonomy 

of the Obligately Foliicolous, Lichenized Fungi. 

Symbolae Botanicae Upsalienses, 12 (1):l-590. 

1827. Systema Vegetabilium 4. Edition 16. Part 2, 410 

Santesson, R. 

1952. 

Sprengel, K. 

pages. Gottingae. 

Stirton, J. 

1881. On Vegetable Parasites on the Tea Plant, More 

Especially That of Assam. Proceedings of the Philosophical Society of Glasgow, 13: 181-193. 

New Lichens, Principally from the Herbarium of 

W. J. Hooker. London Journal of Botany, 6:148-197. 

Observations on North American and Other Lichens. 

Proceedings of the American Academy of Arts and 

Sciences, 5:383-422. 

Etude sur la classification naturelle et la morphologie des lichens du Bresil. Acta Societas Fauna et 

Flora Fennica, 7(7): 1-247. 

1896. Lichenes .4ntillarum a \V. R. Elliott Collecti. 

Journal of Botany British and Foreign, 34:204-210. 

1915. Additamenta ad Lichenographia Antillarum 111~1~- 

trandam. Annales Academiae Scientiarum Fennicae, 

1921. Lichenes Insularum Philippinarum 111. Annaled 

Academiae Scientiarum Fennicae, 15A: 1-368. 

1923. Lichenes in Insula Trinidad a Professor R. Thaxter 

Taylor, T. 

1847. 

Tuckerman, E. 

1862. 

Vainio, E. A. 

1890. 

6.4 (7):1-226. 
NUMBER 16 45 

Collecti. Proceedings of the American Academy of und Lichenes. Bulletin du Jardin Botanique, series 

Arts and Sciences, 58: 129-147. 3, 4:l-146. 

1926. Lichenes Mexicani. Dansk Botanisk Arkiv, 4: 1-25. Zahlbruckner, A. 

Neue Flechten VI. Annales Mycologici, 10:359-384. 

1922. Verzeichnis der in Niederlandisch Ost-Indien bis 1923. Catalogus lichenurn universalis. 2:580-643. Leipzig. 

Catalogus lichenurn universalis. 8:244-247. Leipzig. 

Van 0vereem.De Haas, C. and D. 1912. 

dem Jahre 1920 Gefundenen Myxomycetes, Fungi 1932. 
Index 

(Index to main entries only: synonyms in italics) 

Ascidium cinchonarum, 17 

interpositum, 35 

rhodostromum, 25 

Endocarpon wightii, 43 

Graphis alborosella, 16 

Leptotrema bahianum, 39 

deceptum, 39 

exalbidum, 29 

fissum, 40 

pavicans, 43 

integrum, 40 

microglaenoides, 40 

monosporum, 40 

occultum, 40 

phaeosporum, 40 

prevostianum, 43 

reclusum, 40 

spondaicum, 41 

subcompunctum, 42 

subconcretum, 43 

wightii, 43 

Myriotrema oliuacea, 24 

Ocellularia alboolivacea, 19 

alborosella, 16 

antillensis, 17 

berkeleyana, 17 

bonplandiae, 24 

cavata, 17 

cinchonarum, 17 

cinchonarum f. intermedia, 17 

clandestina, 35 

comparabilis, 17 

comparabilis var. microcarpa, 17 

concolor, 19 

conglomerata, 19 

discoidea, 17 

doming~nsis var. fecunda, 20 

dominicana, 19 

efformata, 17 

exanthismocarpa, 20 

fecunda, 20 

glaziocii, 17 

homothecia, 20 

isertii, 20 

latilabra, 19 

lindigiana, 17 

maculata, 22 

mordenii, 22 

multilocularis, 20 

nigropuncta, 22 

olivacea, 24 

papillata, 24 

perforata, 25 

platycarpella, 16 

porinoides, 20 

pyremloides, 25 

remanens, 28 

rhodostroma, 25 

rimosa, 27 

rufocincta, 25 

sorediata, 27 

subcavata, 27 

terebrata, 17 

terebrata var. abbreviatula, 25 

terebratula, 28 

riaga, 28 

verrucosa, 24 

xanthostroma, 22 

albidulum, 31 

disciforme, 29 

galactinum, 29 

leiostomum, 29 

lirelliforme, 29 

obscurum, 31 

siibconcretum, 43 

Phaeotrema aggregatum, 29 

Pyrenula clandestina, 35 

Thelotrenia aquilinum, 29 

butuanum, 35 

carassense, 32 

cavatum, 17 

clandestinum, 32 

clandestinum f. remanens, 28 

cornparabile, 17 

concretum car. subcaesium, 32 

conforme, 32 

confusum, 33 

consanguineum, 32 

depressum, 33 

disciforme, 29 

domingense car. fecundum, 20 

dominicanum, 33 

elliottii, 36 

exalbidum, 29 

exanthismocarpum, 21 

excavatum, 25 

excavaticm var. impressulum. 13, 25 

fissum, 40 

galactinurn, 29 

glaucopallens, 35 

homopastoides, 35 

homotkecium, 20 

interpositum, 35 

laevigans var. avertens, 35 

leiostomum, 29 

lepadinum var. bahianum, 39 

leucinum, 35 

leucomelaenurn, 36 

leucomelaenum var. elevatum, 13, 36 

microglaenoides, 40 

occultiim, 40 

papillatun, 24 

papillosum, 36 

pnuperius, 36 

pechueli, 35 

perforatum, 25 

platycarpellum, 16 

porinoides, 20 

postpositum, 32 

praestans, 36 

prevostianum, 43 

rhodostromum, 13 

rzidius Lar. dominicaizum, 13, 39 

spondaicum, 40 

subcaesium, 32 

subcavatum, 28 

su bcompunctum, 42 

subconcretum, 43 

tenue, 38 

terebratuliam, 28 

terebratuin var. abbreviatulum, 25 

tuberculiferum, 38 

vagum, 28 

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