SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 38 

A Revision of the Lichen Family 

Thelotremataceae in Panama 

Mason E. Hale, 3r. 

SMITHSONIAN INSTITUTION PRESS 

City of Washington 

1978 
ABSTRACT 

Hale, Mason E., Jr. A Revision of the Lichen Family Thelotremataceae in 

Panama. Smithsonian Contributions to Botany, number 38, 60 pages, 15 figures, 

1978.-The family Thelotremataceae comprises 99 species in Panama, distributed 

among four genera, Ocellularia, Phaeotrema, Thelotrema, and Leptotrema. It is 

most common in virgin rain forests at mid-elevations. Phytogeographically there 

are many species in common with the IVest Indies and South America, The following species are described as new: Ocellularia barroensis, 0. bullata, 0. congesta, 0. dactylifera, 0. dactyliza, 0. decolorata, 0. disszita, 0. esslingeri, 0. 

landronii, 0. minutula, 0. neei, 0. protocetrarica, 0. pseudoschizostorna, 0. 

puluerulenta, 0. referta, 0. subwrightii, 0. uniseptata, 0. viridis, Phaeotrema 

uulcani, Thelotrema hiatum, T. maximum, Leptotienza chiriquiense, L. hypoprotoceirariczim, L. pananzense, and L. stellatum. These new combinations are 

made: Ocellularia pachystoma (Nylander) Hale (basionjm: Thelotrema pachystomum Nylander) and 0. uerrucosa (Fee) Hale (basionym: Thelotrema uerrucoJuni 

Fee). 

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded 

in the Institution’s annual report, Smithsonian Year. SERIES COVER DESIGN: Leaf clearing from 

the katsura tree Cercidiphyllum japonicum Siebold and Zuccarini. 

Library of Congress Cataloging in Publication Data 

Hale, Mason E. 

A revision of the lichen family Thelotremataceae in Panama. 

(Smithsonian contributions to botany ; 38) 

Bibliography: p. 

Includes index. 

Supt. of Docs. no.: 1. Thelotremataceae. 2. Lichens-Panama. I. Title. 11. Series: Smithsonian Institution. 

QKl.SZ747 no. 38 [QK585.T5] 581’.08s 77-608159 [589’.1] 

SI 1.29:38 

Smithsonian contributions to botany ; 38. 
Contents 

Introduction ...................................................... Morphological Characters .......................................... Chemistry ........................................................ The Thelotreme Flora ............................................. Collecting Sites .................................................... 

Key to the Species of Ocellularia .................................. Phaeotrema Muller Argoviensis ..................................... Thelotrema Acharius ............................................... 

Key to the Species of Phaeotrema .................................. Leptotrema Muller Argoviensis ...................................... 

Key to the Species of Thelotrema .................................. 

Literature Cited ................................................... Index ............................................................ 

Ocellularia Meyer ................................................. 

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

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59 

... 

111 

A Revision of the Lichen Family 

Thelotremataceae in Panama 

Mason E. Hale, 3r. 

Introduction 

After completing a field program on the Thelotremataceae in Dominica (Hale, 1974a), I had an 

opportunity to visit Panama on three separate occasions in 1973, 1974, and 1975. There were only four 

specimens of the family from Panama on file in 

the herbarium at the Smithsonian Institution, and 

no species had ever been reported there (Imshaug, 

1956). The prospects for finding many specimens, 

therefore, seemed poor. Notwithstanding, I was able 

to collect nearly 600 specimens at 11 localities from 

the province of Chiriqui in the west to Panama in 

the east. Two excellent lichen collectors, Dr. Scott 

Mori (with Dr. A. Gentry) and Michael Nee, added 

small but valuable collections, some from the remote Darien region near Colombia, and Dr. T. L. 

Esslinger and Dr. M. Nakanishi, who accompanied 

me on different trips, added about 50 more specimens. All of these collections taken together represent 99 species among almost 700 numbers, more 

than twice as many as I had found in Dominica. 

It is very difficult and probably foolhardy to 

undertake a world monograph of the Thelotremataceae along the usual lines, namely, to locate and 

examine type collections, annotate available herbarium material, and prepare a monograph from 

this information. One of the more frustrating aspects of lichenology is that herbarium holdings of 

tropical groups are notoriously poor. While this is 

true for almost any plant group, it is especially 

Mason E. Hale, Jr., Department of Botany, Museum of 

Natural History, Smithsonian Institution, Washington, D.C. 

20560. 

true for crustose lichens that are rarely collected 

in abundance by phanerogamic botanists or naturalists, the main source of lichen specimens in the 

past. Even in the largest European herbaria, in the 

case of the Thelotremataceae, one finds mostly 

sheets of type collections from the 19th century and 

large holdings of Thelotrema lepadinum and some 

Leptotrema wight ii. 

I have in fact examined most of the type collections of the family Thelotremataceae, representing 

over 400 species. While about 100 of these have 

proved to be synonyms, I have already described 

or will soon add nearly 100 new species to the family and feel confident that another 100 will be discovered as time and resources permit further field 

work in the tropical rain forests. This particular 

forest biome, so rich in thelotremes, is being rapidly destroyed and many species will probably become extinct before they can even be collected. My 

approach has been and will continue to be a detailed examination of the thelotreme floras of specific areas where I have been able to conduct adequate field studies but still including data on 

synonymy at the world level. 

As usual, I am indebted to curators of museums 

and herbaria who sent collections on loan, as indicated by herbarium acronyms cited under “Specimens Examined.” I would like to mention in particular Dr. T. Ahti, Dr. Reino Alava, the late Dr. 

C. E. Bonner, Mr. M. S. Christiansen, Dr. H. Hertel, Mr. Peter James, Mme. Jovet-Ast, Dr. I. M. 

Lamb, Dr. H. Riedl, and Dr. R. Santesson. I am 

also especially thankful for transportation support 

provided by Dr. Robert Dressler, Smithsonian 

1 
2 SM’ITHSONIAN CONTRIBUTIONS TO BOTANY 

Tropical Research Institute, and by Dr. Michael 

Nee and Dr. Scott Mori, who were associated with 

the plant exploration program of the Missouri Botanical Garden at Summit Gardens. Dr. Ove Almborn has kindly read and corrected the Latin descriptions of the new species. Financial support was 

received from the Smithsonian Institution for the 

field studies. 

Morphological Characters 

I have already summarized the main morphological features of the family in my study of the 

Dominican species (Hale, 1974a) and this information will not be repeated here. The Panamanian 

species have the same broad characters and any 

unusual or interesting aspects are summarized below in discussions of the various genera. It is obvious, however, that the terminology for apothecial 

structure is rather confused at present and cannot 

be considered as final, As I use the term “lepadinoid,” for example, it is only intended to describe 

semiemergent to emergent apothecia with a free, 

persistent proper exciple. The “actinoid” columella 

would perhaps better be called “reticulate,” as 

Salisbury has suggested. “Chroodiscoid,” derived 

from the epiphyllous genus Chroodiscus, refers to 

species with an erect or recurved thalline margin, 

as in Ocellularia alborosella (Figure 4c), while 

“ascidioid” is used to describe strongly emergent 

apothecia. All of these terms could be dropped in 

favor of precise descriptions of the apothecial structures involved. 

Chemistry 

Chemistry is extremely helpful in routine identifications of specimens of the Thelotremataceae. 

While morphological characters may vary according 

to ecological or growth conditions, chemical content of each species is remarkably uniform and 

stable. The Panamanian species have a chemistry 

very similar to that of the family in Dominica. 

For example, the four most common acids, psoromic acid, stictic acid, hypoprotocetraric acid, and 

protocetraric acid, have almost identical frequency 

in both areas. Virensic acid, known from Leptotrema spondaicum (Nylander) Zahlbruckner in Dominica, does not occur in Panama, whereas lichexanthone, a component of five species in Panama, 

did not occur in Dominica. The acids are tabulated 

below by chemical groups and further notes will be 

found in the species discussions. All specimens were 

studied with precoated klerck silica-gel plates utilizing C. Culberson’s A and B solutions. Spots were 

visualized by spraying the plates with 10% sulfuric 

acid and heating them for 10 minutes at 110’C. 

When color tests are used, it is best to expose part 

of the amphithecial medulla for testing. The thallus is often too thin to evoke a clear test. 

DEPSIDEs.-This group of lichen substances is 

extremely rare in the family. For example, only 

one of 400 known species, Ocellularia americana 

Hale, contains gyrophoric acid. Of the species in 

Panama, 0. olivacea contains an unidentified acid 

(or acids) that I presume to be depsidic in nature 

because of the chromatographic behavior. 

DEPSIDONES.-This group is of frequent occurrence in the Thelotremataceae. The commonest 

acid is psoromic acid, which occurs with or without 

norpsoromic acid (Keogh, 1976) in the following 

23 species, nearly one-fourth of the family represented in Panama: Ocellularia antillensis, 0. berkeleyana, 0. calvescens, 0. comparabilis, 0. glaucophaena, 0. isidiifera, 0. latilabra, 0. minutula, 0. 

myriotremoides, 0. pachystoma, 0. pulverulenta, 

0. ripleyi, 0. subcavata, 0. subwrightii, 0. terebratula, and 0. uniseptata; Phaeotrema auberianum; and Thelotrema clandestinum, T. concretum, 

T. conforme, T. cryptotrema, T. maximum, and 

T. wrightii. 

The next most common acid is stictic acid, often 

accompanied by constictic acid in 16 species: Ocellularia bullata, 0. conglomeiata, 0. dactylifera, 0. 

dilatata, 0. exanthismocarpa, 0. neei, 0. pseudoschizostoma, and 0. sublilacina; Thelotrema erumpens, T. glaucopallens, T. myriocarpum, T. stylotlzecium, and T. subpraestans; and Leptotrema 

deceptum, L. reclusum, and L. trypaneoides. 

Closely following stictic acid, hypoprotocetraric 

acid, which is the only P- depsidone, is the main 

component in 15 species (frequently accompanied 

by 4-0-demethylnotatic acid (Culberson and Hale, 

1973), by unidentified spots, or in Phaeotrema 

glyphicum by lichexanthone): Ocellularia actinota, 

0. azirnlenta, 0. congesta, 0. domingensis, 0. fecunda, 0. glaucula, 0. microporella, 0. referta, 

and 0. rhodostroma; Phaeotrema glyphicum and 
NUMBER 98 3 

P. leiostomum; Thelotrema myrioporoides and T. 

occlusum; and Leptotrema chiriquiense and L. 

hypoprotocetraricum. 

Protocetraric acid occurs in the following eight 

species: Ocellularia dactyliza, 0. perforata, 0. protocetrarica, and 0. verrucosa; Thelotrema auratum; 

and Leptotrema bahianum, L. laeviusculum, and 

L. panamense. Two of these, Thelotrema auratum 

and Leptotrema panamense, also contain epihymenial pigments. 

Norstictic acid occurs in only one species, Thelotrema norsticticum. 

Unidentified (and presumed) depsidones were 

found in Thelotrema praestans (the “praestans” 

unknown) and in Ocellularia cavata, 0. crocea, 0. 

decolorata, 0. rhabdospora, and 0. subemersa (the 

“cinchonarum” or “cavata” unknown). Both of 

these react P+ red. 

XANTH0NES.-The only xanthone in the Thelotremataceae is lichexanthone, which reacts brilliant 

orange under longwave ultraviolet radiation. It is 

a component of the cortex in five species: Ocellularia glaucula, Phaeotrema glyphicum, Thelotrema 

myrioporoides, Leptotrema metaphoricum, and L. 

microsporum. 

ANTHRAQUINONES AND OTHER PrcMEws.-Various 

red or orange anthraquinone pigments, of which 

only endocrocin has been positively identified, are 

present in Ocellularia crocea, 0. mordenii, 0. rhodostroma, and 0. xanthostroma (all in the medulla) 

and in Thelotrema auratum and Leptotrerna panamense (epihymenial) and L. wightii (medullary). 

Unidentified yellow medullary or amphithecial 

pigments, perhaps related to entothein, are known 

in Ocellularia aurulenta, 0. cavata, and 0. fecunda. 

UNIDENTIFIED SUBSTANCES.-Three species, Ocellularia esslingeri, 0. recondita, and 0. viridis, show 

distinct P-, H,SO,+ gray spots, presumably phenolic in nature and probably new substances. 

NO SUBSTANCES PRESENT.-Twenty-one species, 

as listed below, reacted negative in color tests and 

produced no substances reacting H,SO, positive 

on the chromatographic plates: Ocellularia alba, 

0. alborosella, 0. barroensis, 0. chionostoma, 0. 

concolor, 0. dissuta, 0. landronii, 0. papillata, and 

0. turgidula; Phaeotrema meiospermum and P. 

vulcanum; Thelotrema adjectum, T. conveniens, 

T. hiatum, T. lepadinum, T. leucomelaenum, T. 

pycnoporellum, and T. tenue; and Leptotrema 

lepadodes, L. stellatum, and L. urceolare. 

The Thelotreme Flora 

One of my goals in tropical field work with the 

Thelotremataceae has been an attempt to estimate 

how many species one can reasonably expect to 

collect out of the total lichen flora. In Dominica 

I made deliberate efforts to recollect certain areas, 

make random transects, and otherwise satisfy myself that the number of species finally identified (48) 

came close to representing the total flora. Rough 

statistical extrapolations showed that there are 

probably about 55 species of thelotremes on Dominica and that the seven not yet found could be collected on additional excursions, but since time and 

financial resources are limiting factors, especially in 

the tropics, one must obviously be satisfied with a 

less than complete sampling. 

I found that similar estimates are not now possible in Panama, a much larger, more diverse, and 

less thoroughly collected region than Dominica. 

As in Dominica, new species comprise 30% of the 

flora, but I found that each new locality in Panama added at least one new species, not to mention 

new records of species for Panama. In other words, 

we are still on the slope of a theoretical sigmoid 

curve of species distribution, but an estimate of 

130-140 species in the flora may not be far off. 

The question cannot be answered until more 

forest sites have been investigated, and the problem 

of accessibility to virgin forests in Panama prevents 

us from reaching this goal quickly. I made samples 

at points where roads were being opened up and 

felled trees could be easily reached, but the minuteness of this sample is all too evident when one 

realizes that the 12 collecting sites were taken along 

a nearly continuous range of forest extending about 

600 km. 

There is little doubt, however, that a general 

lichen collector, one who tries to collect all lichen 

families in a limited amount of time, does justice 

to none of them, especially with crustose groups 

such as the Thelotremataceae, which can often 

only be recognized in the field after careful, timeconsuming searching with a hand lens. A general 

collector could probably be expected to collect no 

more than 10-3070 of the total number of species 

in the Thelotremataceae in Panama, especially if 

he did not have access to freshly felled trees. 

Details on the number of specimens collected 

and comparisons with Dominica are given below 
4 SMITHSONIAN CONTRIBUTIONS TO BOTASY 

under discussions of the genera. About 30 collections were sterile and could not be assigned to 

genus. An additional 20-30 specimens have not 

been identified to species level yet for various reasons and will be left for future study. 

Collecting Sites 

My main aim in Panama was to work only in 

the virgin rain forests, which occur in large unbroken tracts in the low mountains along the Caribbean side of the country. Unfortunately these sites 

are often remote from the main east-west Panamerican Highway and can only be reached where new 

roads are being cut across the isthmus, as at Santa 

Fe, El Valle, Cerro Jefe, etc., and at Volcin Chiriqui. The entire Pacific side was left unexplored; 

much of it is dry savanna used for cattle ranches, 

but good rain forest is reported to occur at higher 

elevations in the Ocu Peninsula. Nor did I enter 

the roadless Darien region but had access to collections by Xlori and Gentry. My first collecting 

trip was made in 1973 and included Chiriqui, the 

two road cuts at Cerro Jefe and El Llano-Carti, 

and Barro Colorado Island. The second trip in 

1974 included the Gamboa area, Cooh, Santa Rita 

Ridge, and El Valle. The third trip with Dr. T. L. 

Esslinger covered the sites near Alto Piedra above 

Santa Fe, the road cut to the Rio Calevobora, and 

second visits to Chiriqui and Barro Colorado Island. 

The localities are coded as follows in the list of 

specimens examined. 

1. Virgin oak cloud forest at 1900-2200 m elevation, t’olcan 

Chiriqui, along the trail from Cerro Punto to Boquete, 

Province of Chiriqui. The first trip is indicated as 1, the 

second as la. On the second trip the oak forest was already 

being logged and I was able to collect more extensively in 

the canopy. 

2. Recently logged rain forest at about 750 m elevation 

at La Mesa, north of El l.alle, Province of Cocle. This was 

a very rich locality for the family and much forest remains 

in spite of recent clearing for chicken farms. 

3. Mixed mature secondary and primary rain forest at 

100-150 m eletation, Barro Colorado Island, Canal Zone (3 

represents the first visit, 3a the second). Specimens were 

collected primarily along several main trails on the southern 

half of the island. This forest reserve has recovered from 

disturbances previous to the closing of the area when the 

canal xvas built about 80 years ago. 

4. Mature secondary rain forest along the Naval Pipeline 

Road at 150 m, 9 km northwest of Gamboa, Canal Zone. 

This forest is very similar to that on nearby Barro Colorado 

but few canopy species were collected because of the absence 

of windfalls. 

5. Open secondary groves at Summit Gardens, Canal Zone. 

As expected, this lvas a poor site for the family. 

6. Recently logged primary forest and remnant stands at 

about 350 m elevation, Santa Rita Ridge, about 6 km from 

Roosevelt Highway, Province of Colon. Rather extensive 

forests remain in this interesting area. 

7. Secondary rain forest near sea level, a few kilometers 

northwest of Colon, Province of Colon. This was the poorest site visited for no obvious reasons. 

8. Road cut through primary rain forest at about 300 m 

elevation, 1S20 km along the El Llano-Carti road, Province 

of Panama. This was a magnificent locality with easy access 

to understory as well as to freshly felled trees. 

9. Roadcut through primary rain forest at about 700 m 

elevation in the vicinity of Cerro Jefe, Province of Panamh. 

This was also a rich locality, similar to 8 but with fewer 

felled trees. 

10. Disturbed primary forest and pasture at base of 

mountain west of Alto Piedra at about 850 m elevation, 

Province of l’eraguas. There was no access to felled trees 

here and thelotremes were poorly represented. 

11. New road cut through primary rain forest at about 

800 m elevation, 4-8 km from Alto Piedra going toward Rio 

Calevobora, Province of Veraguas. This was an excellent 

area since several hectares of trees had just been felled. 

12. Primary rain forest in the Serrania del Darien region 

(near the Tacarcuna River) at about 1800 m elevation, 

Province of Darien. Specimens from this remote locality, 

reached only by boat and long hikes, were collected by Mori 

and Gentry. 

Specimens collected by Hale, Mori, and Nee are 

filed in the Smithsonian Institution (US); those by 

Esslinger are in his private herbarium. Standard 

herbarium acronyms indicate location of type specimens seen. Many of these types are illustrated in 

Figures 1 and 2 and others will be found in Hale 

(1 974a). 

Full citations for synonyms of species that also 

occur in Dominica (Hale, 1974a) are not usually 

repeated here. Statements on distribution of species 

outside of Dominica and Panama are based on my 

own determinations of specimens in BM, FH, G, 

H, L, LD, hf, PC, S, UPS, US, and W. Provisional 

determinations are indicated by “cf.” All species 

descriptions are taken from populations in Panama 

and deviations from the types are noted under 

“Remarks.” The chemical components, however, 

represent results of analyzing the types, and any 

differences with the Panamanian populations are 

noted under “Remarks.” All literature has been 

examined in the original to ascertain the exact 

designation of types. 

When identifying species in the Thelotremata- 
NUMBER 38 5 

FIGURE 1.-Type specimens of Thelotremataceae (listed alphabetically by species under the 

original basionym): a, Thelotrema actinotum Tuckerman (FH-Tuck); b, Thelotrema adjectum 

Nylander (H-Nyl); c, Myriotrema album Fee (G); d, Thelotrema annulatum Muller Argoviensis 

(G); e, Thelotwma auberianurn Montagne, (H-Syl); f, Thelotrema aui'atutn Tuckerman (FHTuck); g, Stegobolus berkeylanum Montagne (BM); h, Thelotrema calvescens Fee (G); i, Thelotrema chionostomum Nylander (FH-Tuck); j, Thelotrema citrinodiscum Redinger (S); k, Thelotrema clandestinum Fee (G); I, Thelotrema concretum Fee (G); m, Thelotrema connivens 

Nylander (S-Nyl); n, Ascidiunz croceum Krempelhuber (M); 0, Thelotrema cryptotrema Nylander (H-Syl); p, Ascidium domingense Fee (FH-Tuck); q, Thelotrema erumpens Magnusson 

(UPS); I, Thelotrema glaucophaenum Krempelhuber (M); Y, Thelotrema glauculum Nylander 

(H-Nyl); t, Thelotrema glyphicum Nylander (FH-Tuck). (1 mm scale for all species is given in It.) 
6 SMITHSONIAN CONTRIBUTIONS TO BOTASY 

FIGCRE 2.-Type specimens of Thelotremataceae (listed alphabetically by species under original 

basionym): a, Thelotrema iiitetvallatum Nylander (H-Syl); 6, Thelotrema leucotyliicm Sylander 

(EM); c, Thelotrema microporellum Nylander (FH-Tuck); e, Thelotiema my~iocn~pum Fee (PC); 

f, Thelotrema myioporoides Muller Agroviensis (G); h, Thelotremu occlusum h’ylander 

(H-Syl); i, Thelotrema oczilalum Vainio (TUR); j, Thelotrema opacuin Vainio (TUR); k, 

Thelotrema pachystomum Nylander (PC); I, Thelotrema porph,yrodiscum Zahlhruckner (tV); 

in, Thelotrema ppcnoporellum Nylander (H-X>l); 71, Thelotl-emu wclzcstirn Krempelhuber 

(H-Nyl); o, Thelotrema recotiditum Stirton (BSI); p, Thelotrema secoligella Muller Argoviensis 

(C); q, Thelotrema stylothecium Vainio (FH); r, Ocellularia subemersa Xuiller Argoviensis (G); 

s, Thelotrema trppaneoides Sylander (FH-Tuck); t, Thelotrema wriglitii Tuckerman (FH-Tuck). 

(1 mm scale for all species is given in Figure It.) 
NUMBER 38 

ceae, it is desirable to determine chemical contents 

with thin-layer chromatography, although thalline 

color tests are invoked as far as possible in the keys 

and will suffice up to a certain point. 

Ocellulmia Meyer 

Ocellularia, the most commonly collected genus 

in the family, includes about 175 species and the 

57 species in Panama represent almost 30% of those 

known. The 400 specimens comprise 60% of the 

total 700 collected in Panama. This compares with 

72% in Dominica, where the perhaps abnormally 

high frequency of 0. rhodostroma distorts the percentage somewhat. The 10 commonest species are 

as follows, compared with the ranking in Dominica 

(in parentheses): 1 (3), 0. perforata, 63 collections; 

2 (4), 0. papillata, 45; 3 (2), 0. fecunda, 24; 4 (not 

in Dominica), 0. ripleyi, 17; 5 (not in Dominica), 

0. sublilacina, 16; 6 (12), 0. concolor, 15; 7 (not 

in Dominica), 0. glaucophaena, 15; 8 (5), 0. subcavata, 12; 9 (not in Dominica), 0. latilabra, 10; 

10 (not in Dominica), 0. microporella, 10. All of 

the 10 commonest species in Dominica, excepting 

0. nigropuncta, also occurred in Panama. 

Of the 57 species reported here, 21 (37y0) are 

new, either described here or in Hale (1947b), compared with 8 of 22 (36y0) in Dominica, virtually 

identical percentages. 

Ocellularia was usually collected at each locality 

in abundance except for the secondary forest at 

Summit Gardens. Where sites were visited twice, 

there was not a high level of duplication, suggesting that while a single visit may turn up a majority 

of the species, certainly not all will be collected. 

For example, I collected nearly equal numbers 

of specimens (about 50) on two separate visits to 

Barro Colorado Island. On the first I discovered 

16 species. On the second trip, seven of these (0. 

comparabilis, 0. glaucophaena, 0. papillata, 0. 

perforata, 0. recondita, 0. ripleyi, and 0. sublilacina) were rediscovered. These were also the seven 

commonest species both times; however, a total of 

nine species (0. antillensis, 0. berkeleyana, 0. calvescens, 0. concolor, 0. dilatata, 0. glaucula, 0. 

olivacea, 0. unispetata, and 0. verrucosa) were not 

recollected on the second visit. Most of these species 

were represented by a single collection. At the same 

time 5 of the 12 species collected on the second visit 

(0. barroensis, 0. latilabra, 0. minutula, 0. subwrightii, and 0. xanthostroma) had not been collected on the first trip. 

The richest locality was, in fact, Barro Colorado 

Island with 21 species, followed by El Valle with 

18, Cerro Jefe with 17, El Llano-Carti road with 16, 

the Rio Calevobora road and Santa Rita both with 

15, and Gamboa with 12. Only eight species were 

collected in the cloud forest at Volcan Chiriqui, 

where Thelotrema and Leptotrema predominated. 

This reinforces my impression that Ocellularia is 

primarily a genus of low to mid elevation rain 

forest, overshadowed by Thelotrema at higher 

elevations. 

The morphology of the genus follows lines similar to those in Dominica. Isidia are clearly developed in 0. dactylifera and 0. isidiifera and sorediate structures in 0. pulverulenta. The thallus is 

distinct and epiphloeodal in all species except 0. 

chionostoma, 0. concolor, 0. exanthismocarpa, 

and 0. neei, which are hypophloeodal. The species 

may be grouped according to apothecial structure 

as discussed in Hale (1947a:Z). Chroodiscoid species, those with an open disc, erect to recurved 

thalline margin, and an often persistent proper 

exciple (Figures 4c, 5h, Sb), include 0. alborosella, 0. chionostoma, 0. conglomerata, 0. dilatata, 

0. dissuta, 0. esslingeri, 0. neei, 0. pseudoschirostoma, 0. referta, and 0. sublilacina. The lepadinoid type apothecium, where the walls are more 

incurved to form a more or less distinct pore, 

through which a free proper exciple is visible, is 

characteristic of 0. exanthismorcarpa and 0. turgidula. The myriotremoid species have immersed 

or flush apothecia: 0. alba, 0. glaucula, 0. microporella, 0. olivacea, 0. terebratula, and 0. uniseptata. Here the proper exciple may or may not 

persist, and there is no carbonized tissue. 

Columellate species include 0. antillarum, 0. 

cavata, 0. concolor, 0. Eandronii, 0. papillata, 0. 

perforata, 0. recondita, 0. ripleyi, 0. subcavata, 

and 0. subemersa. A weakly developed columella 

may be present in 0. isidiifera, 0. minutula, and 

0. mordenii. The reticulate or actinoid columella 

(Figure la) occurs in 0. actinota, 0. berkeleyana, 

and 0. latilabra and is imperfectly developed in 

0. burroensis, 0. cornparabilis, 0. protocetrarica, 

0. subwrightii, and 0. uiridis (Figure 3f). 

The species of Ocellularia often have strong 

habitat preferences, as I have found in Dominica, 

and species in common to these two regions have 
8 SMITHSONIAN CONTRIBUTIONS TO BOTASY 

FIGURE 3.--Cross section of apothecia of Ocelitilurin species: n, 0. bnyroensis (Hnie 46069) 

(x 125) (hymenium pulled away from hypothecium in left part); h, 0. dactylifera (Hale 43303) 

(x 125); c, 0. dactyliza (Hale 43559) (X 125): d, 0. minutzcln (Hale 43341) (X 125); e, 0. protocetraTicci (Hale 44786) (X 100); f, 0. viridis (Hale 38538) (X 125). 

similar behavior. Added to data on habitats of type 

collections, especially elevation, where this can be 

determined, it is possible to categorize the more 

common Panamanian species as follows: restricted 

to higher elevations (more than 2000 m) are 0. 

domingenyis, 0. isidiifera, 0. pachystoma, and 0. 

rhabdospora; restricted to low elevations (less than 

300 m) are 0. glaucophaena, 0. minutula, 0. recondita, 0. ripleyi, 0. sublilacina, 0. uniseptata, and 

0. verrucora. The remaining species occur in the 

rain forests at 300 to 1800 m elevation. 

Species restricted to the canopy include 0. glau- 
NUMBER 38 

cula, 0. microporella, 0. mordenii, 0. olivacea, 

0. subemersa, 0. subcavata, 0. terebratula, and 

0. uniseptata. A few species occur mostly at base 

level in dense rain forest: 0. dilatata, 0. concolor, 

0. papillata, 0. perforata, and 0. sublilacina. 

Much more field work will be needed to categorize 

the remaining species, although it is obvious that 

most grow out of normal reach on the mid to 

upper trunk areas. 

Any statements on the phytogeography of Ocellularia must be regarded as provisional and subject to modification as more collections of the 

group are made. One may generalize by saying that 

9 

the Ocellularia flora shows considerable affinity 

with those of both the Caribbean and South American area. For example, 0. domingensis, 0. pachystoma, 0. recondita, and 0. verrucosa were previously known only from South America. Ocellularia 

actinota, 0. fecunda, 0. glaucula, 0. landronii, 

0. microporella, 0. rhodostroma, 0. subemersa, 

and 0. xanthostroma are all typical Caribbean 

species. The pantropical element is represented by 

0. alba, 0. berkeleyana, 0. cavata, 0. concolor, 

0. crocea, 0. exanthismocarpa, 0. glaucophaena, 

0. olivacea, 0. papillata, 0. perforata, 0. subcavatu, and 0. terebratula. 

Key to the Species of Ocellularia 

1. Spores very large, 100-250 pm long. 

2. Medulla white, unpigmented. 

3. Apothecia emergent; thallus finely verrucose; P+ yellow (psoromic acid) .... 48 0. dpleyi 

3. Apothecia strongly emergent to ascidioid; thallus smooth; P- or P+ red. 

4. Distinct central columella present. ............................................................ 45. 0. rhabdospora 

4. Columella lacking. ............. 22. 0. domingensis 

2. Medulla pigmented yell 

5. Medulla yellow, K- or more intensively yellow. 

6. Apothecia strongly emergent; pore gaping, to 0.4 mm broad. .... 25. 0. fecunda 

6. Apothecia semi-emergent; pore less than 0.1 mm wide. ............................ 5. 0. aurulenta 

7. Pigments yellow orange. ................................................................................ 57. 0. xanthostromu 

7. Pigments red or deep pink. 

.46. 0. rhodostroma 

.... 33. 0. mordenii 

9. Apothecia chroodiscoid with an erect or recurved thalline margin and often a persistent 

proper exciple; columella never present (0. conglornerata. 0. exanthismocarpa, and 0. 

turgiduh have conspicuous lepadinoid apothecia that may appear chroodiscoid; 0. latilabra has an open disc but it is actinoid-columellate); thalline margin noncarbonized. 

50. 0. sublilacina 

11. Apothecia less than 0.3 mm in diameter; disc heavily pruinose. .... 11. 0. chionostoma 

11. Apothecia 0.5-1.0 mm wide; disc, if pruinose, remaining distinct. 

5. Medulla red or orange, K+ reddish or purple. 

8. Apothecia strongly emergent; pore annulate; medulla deep pin 

8. Apothecia semiemergent; pore not annulate; medulla blood 

1. Spores smaller, less than 100 pm long. 

10. Disc very broad, 24 mm wide; spores 60-80 pm long. 

10. Disc less than 2 mm broad; spores less than 30 pm long. 

12. Stictic acid present. 

13. Proper exciple persistent, free. 

14. Apothecia aggregated: disc partially filled with excipular material 

14. Apothecia separate; disc open 

15. Thalline rim very coarse, recurved. ............................................ 20. 0. dilatata 

15. Thalline rim crumbling apically, erect. 41. 0. psewloschizostoma 

.......................... 3. 0. alborosella 

...................................................................................................... 15. 0. conglo 

13. Proper exciple evanescent. 

12. Stictic acid lacking. 

16. Thalline margin delicate, recurved. . 

16. Thalline margin coarse, erect. 

17. Thalline margin entire, uniform1 

17. Thalline margin irregular, splitting. ............................................ 21. 0. dissuta 

9. Apothecia with an incurved thalline margin and more or less closed disc, flush, semiemergent, or ascidioid with a discrete pore; disc, if open, with an actinoid columella. 

18. Columella present. 
10 SMITHSONIAN CONTRIBUTIONS TO BOT.4NY 

Key to the Species of Ocellularia (cont’d) 

19. Central reticulate columella present; disc relatively open (0. burroensis and 0. protocetrarica have weakly carbonized, mostly superficial actinoid structures; 0. comparabilis and 0. subemersa have a main columella with some secondary actinoid 

formation). 

........................................ 28. 0. isidiifera 

21. Thallus and disc P - ............................................................................... 1. 0. actinota 

21. Thallus and disc P+ yellow (psoromic acid). 

22. Thalline rim irregularly split. ....................................................... 30. 0. latilabra 

22. Thalline rim entire. 

23.Apothecia 2-5 mm broad. .................................................... 51. 0. subwrightii 

23. Apothecia less than 2 mm broad. ........................................ 7. 0. berkeleyana 

19. Central columella simple (becoming partially actinoid in 0. comparabilis and 0. 

subemma); disc more or less closed with a distinct pore. 

20. Thallus isidiate. 

20. Thallus lacking is 

24. Thallus and apothecia P - . 

25. Apothecia strongly emergent, urceolate. ............................................. 56. 0. viridis 

25. Apothecia nearly flush to semiemergent. 

26. Thallus dull, largely hypophloeodal; pore to 0.4 mm wide, black inside. 

.............................................................................................................. 13. 0. concolor 

26. Thallus distinct, shiny, epiphloeodal; pore 0.1-0.2 mm wide. 

27. Apothecia semiemergent to emergent; pore white annulate. ............ 

........................................................................................................ 43. 0 rec 

27. Apothecia nearly flush to semiemergent; pore not annulate. 

28. Thallus dark greenish; apothecia sunken. .................... 29. 0. landronii 

28. Thallus whitish mineral gray; apothecia flush to slightly emergent. 

.......................................... 38. 0. papillata 

24. Thallus and apothecia P+ orange or yellow. 

29. Thallus and apothecia P+ yellow (psoromic acid). 

30. Pore annulate; base of columella heavily carbonized, the upper part only 

partially .......................................................................................... 48. 0. subcavata 

30. Pore not annulate; columella uniformly carbonized. 

31. Pore round, 0.1 mm or less in diameter; thallus thin, minutely verrucuose. .............................. 4. 0. antillensis 

31. Pore larger, irregular, t mm wide; thallus rather thick, smooth. 

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

................................................................................................ 12. 0. comparabilis 

29. Thallus and apothecia P+ red (psoromic acid lacking) 

32. Apothecia quite emergent, becoming urceolate; spores 4-loculate, 12-15 pm 

long. ................................................................................................ 49. 0. subemersa 

32. Apothecia barely emergent to strongly emergent; spores 6-18 loculate, more 

than 20 pn long. 

33. Apothecia small, 0.4-0.6 mm broad, flush to semiemergent; protocetraric 

............................ 10. 0. cmata 

on the surface of the disc). 

34. Medulla orange throughout ....................................... ................ 16. 0. crocea 

34. Medulla white. 

35. Spores 50-70 pm long, 12-28 loculate. 

36. Thallus fragile, densely bullate-pustulate ........................................ 8. 0. bullata 

36. Thallus smooth (coarsely isidiate in 0. dactyliza). 

37. Thallus coarsely isidiate. ................................................................ 18. 0. dactylixo 

37. Thallus smooth, nonisidiate. 

38. Stictic acid present ......................................... . 0. exanthismocarpa 

38. No lichen substances present .................... .......... 33. 0. turgidula 

35. Spores 6-30 bm long, 1-9 loculate. 

39. Apothecia more or less emergent, large, 0.5-2.0 mm in diameter. 

40. Pore relatively wide and open, filled with excipular material. 
NUMBER 38 11 

41. Apothecia aggregated, emergent with a distinct semierect thalline 

41.Apothecia mostly separate and only w gent; thalline margin 

42.Thallus P+ red 19. 0. decolorata 

42. Thallus P- ................................................................................... 44. 0. referta 

43. Apothecia becoming strongly emergent with an erect, apically crum43. Apothecia emergent, the apex not crumbling. 

margin. ........................................................ ... 15. 0. conglomerata 

not raised. 

40. Pore discrete, without intruding excipular material. 

bling thalline margin. ...................................................... 37. 0. pachystoma 

6. 0. barroensis 

45. Protocetraric acid present. 

46. Yore usually black rimmed: thallus roughened and verrucose. 

55. 0. verrucosa 46. Pore not black rimmed; thallus smooth 40. 0. protocetradca 

47. Thallus becoming coarsely and sparsely sorediate. ...................... 42. 0. pulverulenta 

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

45. Psoromic acid present. 

48. Pore about 0.1 mm wide, becoming annulate. ........ 

48. Pore 0.1-0.3 mm wide, not annulate. .... 26. 0. glaucophaena 

.............................................. 9. 0. calvescms 

39. Apothecia immersed to flush, usually not more than 0.4 mm wide. 

49. Thallus P - . 

50. No lichen substances present (using TLC). 

51.Apothecia about 0.2 mm wide, flush. ........................................ 2. 0. alba 

51. Apothecia 0.2-0.4 mm wide, becoming slightly emergent. .................... ...... 38. 0. papillata 

50. Lichen substances present (using TLC). 

52. “Olivacea” unknowns present. .............................................. 36. 0. olivocea 

52. Hypoprotocetraric acid present. 

53.Apothecia to 0.3 mm wide (or wider in 0. referta): pore rather 

54. Apothecia semiemergent, the pore open without intruding ex54.Apothecia nearly flush with a broad pore area filled wtih 

53.Apothecia 0.1-0.2 mm wide, flush; pore 0.05-0.1 mm in diameter. 

55. Lichexanthone present (UV+ orange); proper exciple often 

open. 

cipular material. ...................................................... 14. 0. congesta 

excipular material. .................................................... 44. 0. referta 

persistent. .................................................................. 27. 0. glaucula 

55. Lichexanthone absent; proper exciple evanescent. ...................... 31. 0. microporella 

56. Thallus subsidiate: stictic acid present (a trace of stictic acid may occur 

56. Thallus smooth, nonisidiate; psoromic acid present. 

49. Thallus P+ orange or yellow. 

with psoromic acid in 0. uniseptata). ...................... 

57.Spores 4-6 pm long, mostly 2 loculate. 

57. Spores 10-20 pm long, 4-6 loculate. 

54. 0. uniseptata 

58. Thallus surface dull, finely scabrid; pore more or less annulate. 

58. Thallus shiny, not scabrid; pore not annulate. 

...... 34. 0. myriotremoides 

59. Apothecia flush to semiemergent at maturity. ............................ 

...... 26. 0. glaucophaena 

long. .................................................................. 32. 0. minutula 

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

60. Apothecia sparsely developed; spores more than 20 pm 

60. Apothecia numerou pm long. ................ .................................. 52. 0. terebratula 
12 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

1. Ocellularia actinota 

FIGURE 4a 

Ocellularia actinota (Tuckerman) Muller Argoi iensis, 1887b: 

397. 

Thelotrema actinotum Tuckerman, 1862:411 [tbpe collection: 

Cuba, Wright 132 (FH, 1ectot)pe: isotype in L) (Figure 

141. 

DEsCRIPTIoN.-Thallus corticolous, epiphloeodal, 

shiny, smooth to irregularly warty and cracked, 

dull greenish mineral gray, 6-8 cm broad; apothecia 

solitary to clumped in 2-3s, emergent, up to 2 mm 

in diameter, the thalline margin semierect, the columella reticulate but not strongly carbonized, the 

exposed surface and apothecial rim whitish ivory, 

lightly pruinose; spores 8, 6 X 15 pm, 3-4 loculate, 

I+ blue. 

CHEMISTRY.-Hypoprotoce traric and 4-0-demethylnotatic acids. 

REMARKS.-This is the only reticulate columellate species with hypoprotocetraric acid. Superficially it resembles 0. berkeleyana (Montagne) Zahlbruckner, which has psoromic acid. Previously 

known only from Cuba, 0. actinota was collected 

by Scott Xfori in the remote Darien region of eastern Panama at about 1800 m. 

SPECIMEN EXASIISED.-~~: Mori 4462. 

2. Ocellularia alba 

FIGURE 4b 

Ocellularia alba (Fee) hfuller Argoviensis. 1887a:6. 

.Ifyriotreina album Fee, 1824:104 [type collection: “America 

meridionali, ad corticem Bonplandiae trifoliatae (ITild.)” 

G, lectoype; isolectotype in H) (Figure lc)]. 

Thelotrema myriotrema Sylander, 1897: 107. [Based 011 ltfyriotrema album Fee]. 

Thelotrema my>-ioporum Tuckerman, 1862:112 [type collection: Cuba. Wright 129 (FH-Tuck, lectotype; isolectotypes 

in BM, G, L, PC, UPS)]. 

Thelotrema album (Fee) Sylander, 1868:35. 

Thelotrevia subcrassulunz I’ainio, 1929:38 [type collection: 

El Yunque, Puerto Rico, Fink 729 (AIICH, lectotype; isolectotypes in F, FH, SY, US)]. 

DEsCRIPTIoN.-Thallus corticolous, epiphloeodal, 

thin, shiny, grenish mineral gray, 3-5 cm broad; 

apothecia numerous, immersed, dispersed, 0.2-0.4 

mm in diameter, noncarbonized, a columella lacking, the pore round, 0.05-0.1 mm in diameter; 

spores 8, uniseriate, 5-6 X 16-18 pm, 4-5 loculate, 

I+ blue. 

CHEMISTRY.-NO substances present, 

REMARKS.-The flush, noncarbonized apothecia 

are similar to those of 0. oliuacea (F6e) Muller 

Argoviensis and 0. terebratula (Nylander) hluller 

Argoviensis, both of which have a thicker thallus 

and different chemistry. I have identified the species 

from Brazil, the Philippines, Sarawak, the Solomon 

Islands, and Australia, occurring from sea level to 

1200 m elevation. In Panama I collected it only 

once, on the base of a large tree at 700 m. Redinger 

(1936:43) places Thelotrema uil-idialbum Krempelhuber in synonymy under 0. alba, but I found 

that Krempelhuber’s type in RI has muriform 

spores, coarse, almost isidiate outgrowths, and 

lichexanthone. He also regarded 0. costaiicensis 

hfuller Argoviensis as a form of 0. alba, but this 

species contains psoromic acid, and, as Redinger 

noted, is columellate. 

SPECIME\ EXA\II\ED -9: 38433. 

3. Ocellularia alborosella 

FIGURE 4c 

Ocellularia alborosella (Nylander) Santesson, 1952:308. 

Graphis alborosella Nylander, 1863b:372 [type collection: Colombia, Lindig 2694 (H, lectotype; isotypes in FH-Tuck 

and UPS)]. 

Thelotrema filatycarpellum I’ainio, 1923: 138. 

Ocellulaiia p1at)carpella (I’ainio) Zahlbruckner, 1923:998. 

[For full citations see Hale (1974a:16),] 

DESCRIPTIox.-Thallus corticolous, epiphloeodal 

but very thin, shiny and smooth, whitish gray, 5-8 

cm broad; apothecia chroodiscoid, up to 1.5 mm 

across, round to irregularly shaped, the recurved 

thalline margin conspicuous, the inner exciple 

free, entire, the disc flesh colored to white pruinose; 

spores 8, 5-8 X 10-20 /*m, 5-6 loculate, I& weakly 

blue. 

CHEMISTRY.--NO substances present. 

REMARKS.-spOre size is variable in this widespread species, averaging 10-20 pm long but rarely 

as short as 6-8 pm or as long as 36 pm. The significance of this degree of variation has yet to be 

determined. The delicate recurved rim and the 

rather broad pruinose disc are diagnostic. Ocellularia chionostomum (Nylander) Riddle (see page 

16) has smaller apothecia and a less conspicuous 
NUMBER 38 13 

FIGURE 4.--Specjes of Ocellzrln~ia: rr, 0. actinota (,!fori 4462); 6, 0. nlba {Hnle 38433): c, 0. 

nlborosella (Hale 46096); d, 0. antillensis (Hale 43961); e, 0. aurulenta (Hale 43567); f, 0. 

bai,roensis (Hale 46069); g, 0. berkeleyana (Hale 38461); h, 0. bullatn (Hale 34878); i, 0. 

calvescens (Hale 38757); j, 0. cauata (Hale 38466); k, 0. chionostomum (Hale 43455): 1, 0. 

cornparabilk (Hale 44905) (X 10). 
14 S!dITHSONIAN CONTRIBUTIOSS TO BOTANY 

recurved rim. Ocellularia dilatata Xliiller Argoviensis is much larger and coarser and contains stictic 

acid. The species occurs on lower trunks and saplings in primary and secondary forest at 700-1000 

m in Panama. I have also seen specimens from 

Louisiana, the West Indies, Venezuela, and Brazil, 

where it occurs up to 2500 m. 

SPECIME~S EXAMISED.--~: 43494; 10: 46042; 11: 46056 

ILLUSTR4TlOhSHale, 1974a: 18, figs. 5b, 96. 

4. Ocellularia antillensis 

FIGURE 4d 

Ocellularia antillensis Hale, 1974a: 17 [type collection: DOminica, Hale 35612 (US, holotype)]. 

DEscRIPTioh’.-Thallus corticolous, epiphloeodal, 

minutely verrucose, shiny, grayish white, 3-6 cni 

broad; apothecia numerous, separate and regularly 

dispersed, 0.3-0.6 mm in diameter, carbonized, the 

columella well developed, about 150 pm in diameter, the pore round, 0.1-0.12 mm in diameter, black 

inside; spores 8, 7 X 12-15 pm, 5-6 loculate, I& 

blue. 

CHEMISTRY.-PSOrOmiC and norpsoromic acids. 

REMARKS.-The rather open pore and the clearly 

visible black top of the columella separate this 

species from 0. subcauata (Nylander) Zahlbruckner, 

which has a distinct annulate pore, a partially carbonized columella, and a smooth thallus. Ocellzilaria antillensis was collected in Panama on the 

upper trunks of palm and evergreen hardwoods at 

low to middle elevations. 

SPECIMENS EXAMI&ED.-~: 43561; 3: 38659. 

ILLUSTFATIOSS.-Hak, 1974a: 18, fig. 9g. 

5. Ocellularia aurulenta 

FIGURE 4e 

Ocellularia auruhtn Hale, 1974b:490 [t) pe collection: El 

Llano-Carti Road, Province of Panama, Panama, Hale 

38948 (US, holotype)]. 

DESCRIPTIoN.-Thallus corticolous, epiphloeodal, 

thick and irregularly bullate, almost appearing 

coarsely isidiate, pale greenish or whitish mineral 

gray, up to 15 cm broad; apothecia not common, 

occurring inconspicuously among the bullae, semiemergent, about 0.5 mm in diameter, carbonized 

but without a columella, the pore round, 0.1 mm 

in diameter; spores 1-2, 25-35 X 150-200 pm, 15-19 

loculate, I + blue. 

CHE~~ISTRY.-HypOprOtOCe tararic and 4-0-demethynotatic acids with unidentified pale yellow 

pigments. 

KEMARKS.-This species has an unusual bullate 

thallus that is not fragile but can be easily abraded 

to reveal the yellow medulla. It is related to 0. 

feczinda (Vainio) Hale, at least in chemistry. The 

distribution area is quite restricted, the low mountains east of the Canal Zone, 

SPECI\iE\S EXA\II\ED.-6: 43467; 8. 38528, 38936, 38548, 

ILLusTRATIoxs.-Hale, 1974b:490, fig. 2. 

38587. 9: 38441, 38452, 38486, 38495. 

6. Ocellularia barroensis, new species 

FIGURE 4f 

DEsCRIP.rIoN.-Thallus corticola, epiphloeodes, 

nitidus, laevis vel verruculosus, continuus, pallide 

cinereo-viridis, 6-9 cm latus; apothecia (Figure 3a) 

numerous, separata vel aggregata, emergentia, 0.6- 

0.9 mm diametro, margin thallino suberecto, integro, decolorato, columella nulla, excipulo proprio 

pro parte evoluto sed vulgo evanescenti, disco carneo, superficie actinoideo-columellato, albo-pruinoso; ostiolum apertum, 0.2-0.5 mm diametro; 

hymenium ca. 110 pm altum; sporae 8:nae, 6 X 

17-20 pm, 5-6 loculatae, I+ caeruleae. 

CHEMISTRY.-NO substances present. 

HOLOTYPE.-TWO meters up trunk of tree in 

rain forest, Zetek Trail, Barro Colorado Island, 

Canal Zone, Panama, elevation 100 m, M. E. Hale 

46069,20 February 1975 (US). 

REMARKS.-This species is externally similar to 

0. glaucophaena, which contains psoromic acid, 

but the thalline margin is more erect, thin, and 

entire. The apothecia tend to be clumped and at 

maturity an actinoid pseudocolumella may develop 

on the surface of the disc. It occurred at the base 

of a tree at this single locality. 

7. Ocellularia berkeleyana 

FIGURE 4g 

Ocellularia Berkeleyanci (hfontagne) Zahlhruckner, 1905: 118. 

Stegoholtts heikele>an us Montagne, 184534 [type collection: 

Philippines, Berkeley 2185 (BM, lectotype; isolectotype in 

G) (Figure lg)]. 
NUMBER 38 15 

Thelotrema leucotylium Nylander, 1873: 166 [type collection: 

Andaman Islands, Kurr 101 (H, lectotype; isolectotypes in 

BM, UPS, W, ZT) (Figure 2b)l. 

Ocellularia leucotylia (Nylander) Muller Argokiensis, 1891b: 

51. 

Ocelhlmia radians Muller Argoviensis, 1887b:397 [type collection: Apiahy, Brazil, Puiggari 3049 (G, lectotype)]. 

DEscRIPTIoN.-Thallus corticolous, epiphloeodal, 

thin and shiny, light greenish mineral gray, 4-8 

cm broad; apothecia not common, irregularly dispersed, semiemergent to strongly emergent and 

basally constricted, 1-2 mm in diameter, the thalline rim entire, distinct, the open flat disc divided 

by the reticulate columella, becoming white pruinose, the columella carbonized; spores 8, 4-8 X 

8-10 pm, 4 loculate, I+ blue. 

CHEmsTRY.-Psoromic acid with or without 

norpsoromic acid. 

RExfARKS.-This was the first reticulate columellate Ocellularia to be described. As Salisbury 

(1975:59) correctly points out, the genus Stegobolus 

is older than and takes precedence over Rhabdodiscus Vainio, if one wished to recognize these 

species in a separate genus. Ocellularia berkeleyana 

is an extremely variable species and intergrades 

with several other populations. For example, 0. 

comparabilis (Krempelhuber) Muller Argoviensis 

may develop a rather strongly actinoid disc but 

would have less emergent apothecia. Ocellularia 

latilabra (Tuckerman) Muller Argoviensis has n 

more erect to recurved thalline margin and a less 

carbonized actinoid disc. I have identified the species definitely only from Cuba, Guadeloupe, Java, 

and South Africa. In Panama it occurs in the mid 

to upper bole area on larger trees at low to mid 

elevation. 

SPECIMENS EXAMINED.-3: 38693: 9: 38461 

8. Ocellularia bullata, new species 

FIGURE 4h 

DEscRIP'rIo;u.-Thallus corticola, epiphloeodes 

vel evanescens, tenuis, viridi-cinereus, 6 cm latus, 

isidiato-bullatus, isidiis solidis, integris, usque ad 

0.3 mm altis; apothecia rara, emergentia, 0.4-0.5 

mm diametro, ecolumellata; ostiolum rotundatum, 

0.1 mm diametro; excipulum apice fuligineum; 

hymenium ca 70 pm altum; sporae 4-8:nae, 10 X 

50-65 pm, 7-9 loculatae, I+ caerulescentes. 

CmMIsTRY.-stictic and constictic acids. 

HOLOTYPE.-&~atUre rain forest, Cerro Jefe, 

Province of PanamP, Panama, elevation 700 m, 

M. E. Hale 38478, 5 April 1973 (US). 

RE;MARKs.-Externally this species has some resemblance to 0. fragilis Hale, a Colombian species 

that has psoromic acid, small spores (12 pm long), 

and a more fragile thallus. It was collected on the 

lower trunk of a tree in mature rain forest. 

9. Ocellularia calvescens 

FIGURE 41 

Ocellularici calvrscens (Fee) Muller Argokiensis, 1857a:8. 

Thelotrenicr calvescem Fee, 1837:89 [type collection: "America 

ad cortices Cinchonarum" (G, lectotype; isolectotype in H) 

(Figure Ih)]. 

Ocellulnria sexlocularis Zahlbruckner in Magnusson and Zahlbruckner, 1943:48 [type collection: Haleakala, Hawaii, 

Faurie 678 (UPS, lectotype)]. 

DEscRIPTIos.-Thallus corticolous, epiphloeodal, 

shiny, smooth or becoming cracked with age, whitish mineral gray, to 10 cm broad; apothecia 

barely emergent to semiemergent, 0.5-0.8 mm in 

diameter, carbonized but without a columella, the 

pore round, about 0.1 mm in diameter, white annulate and vaguely to distinctly raised annulate; 

spores 8, 6-8 X 15-20 pm, 5-7 loculate, I+ blue. 

RE~IARKS.--AS with most of the Fee types, the 

original material of 0. calvescens in G is in very 

poor condition. It is distinguished by weakly 

emergent apothecia, a small more or less distinctly 

annulate pore, and psoromic acid. The spores, as 

in the Panamanian material, are small. It is a variable species and its limits are still not clearly defined. The specimens collected in Panama, which 

include some tentatively identified, occurred mostly 

in the canopy and on lianas from 100 to 2000 m 

elevation. 

CHEMISTRY.-PSOrOmiC acid. 

SPECIMESS EXANISED.-~: 38716, 38753, 38757; 3: 38622: 

8: 35613; 9: 38506. 

10. Ocellularia cauata 

FIGURE 41 

Ocellularia cavatn (Acharius) hliiller Argoviensis, 1852:499. 

Thelotrema cavatuna Acharius, 1812:92 [type collection: Sierra 

Leone, Afzelius (H, lectotype; S, UPS, isolectotypes)]. 
16 SMITHSONIAN CONTRIBUTIONS TO BOTASY 

Thelotremu obturatum Acharius, 1812:92. 

Ascidiurn cinchonarum Fee, 1824:96. 

Ocellularia cinchonarum (Fee) Sprengel, 1827:242. 

Ocellulnria obturata (Acharius) Sprengel, 1927:242. 

Ascidium cinchonarum f. ii7trrmediiim Njlander, 1867:319 

Ocellularia lindigiana hluller Argo\.iensis, 188ia:9. 

Ocellularia ci~zchonaruni f. i?itet media (Sylander) Zahlbruck- 

[For full citations see Hale (1974a:l7).] 

ner, 1923:586. 

DEscRIPTIos.-Thallus corticolous, epiphloeodal, 

more or less verruculose, greenish to whitish mineral gray, up to 10 cm broad; apothecia emergent 

to strongly emergent, 0.7-0.9 mm in diameter, the 

thalline margin apically carbonized, the exposed 

medulla either white or pale yellow, a well-developed columella present, about 200 pm in diameter, 

the pore round, about 0.1 mm in diameter; spores 

8, variable, 6-12 X 24-70 pm, 7-18 loculate, I+ 

blue. 

CHEMISTRY.-The “cinchonarum” unknown, P + 

red unknowns, and a faint yellow pigment if pigmented. 

REMARKS.-AS recently typified (Hale, 1972: 188), 

0. cavata has a pale yellowish medulla in the apothecia (which I failed to emphasize) and spores in 

the range of 30-40 pm long. I have reexamined the 

type collections of Ascidium cinchonawm Fee 

(PC), 0 ce 1 lu la?-ia 1 indigian a M iiller Argoviensi s 

(FH, BM, UPS), and Thelotrema cauatum Acharius (S, UPS) and been able to confirm that all have 

a pale but distinct yellow medullary pigment in 

the amphithecium. Three specimens from Panama 

(38444, 38466, and 44921) compare well with the 

typical population. Two others from Darien (Mori 

4333 and 4459) have much larger spores (70-80 

pm long) and a white amphithecial medulla, although the chemistry is otherwise identical and 

the apothecia are externally indistinguishable. TYe 

obviously need to see more collections before the 

significance of these differences can be assessed. 

4459. 

SPECIMENS EXAXIISED.-1a: 44921; 9: 38444: 12: )\lOTi 4333, 

11. Ocellularia chionostoma 

FIGURE 4k 

Ocellularia chiorzostoma (Nylander) Riddle, 1917:325. 

Thelotwrna chionostornurn Sylander, 1863b:329 [type collection: Cuba, Wiight 3 (FH, lectotype) (Figure li)]. 

Ocellularia phlyctellacen Muller Argoviensis, 1893b: 146 [type 

collection: Buenos Aires, Costa Rica, Pittier 6106 (material 

in G Ic-ithout collector or number) (C, lectotype)]. 

DEscRIPTIos.-Thallus corticolous, thin and 

mostly hypophloeodal, whitish gray, 6-10 cm broad; 

apothecia adnate, chroodiscoid, the thalline and 

inner exciples short, becoming granular, not recurved, the disc pruinose, 0.2-0.3 mm in diameter; 

spores 8,4-6 X 10-18 pm, 4-3 loculate, I-. 

CHEMISTRY.--NO substances present. 

RExiARKS.-The main thallus rim is erect but 

short, sometimes virtually absent, in contrast, for 

example, to that of 0. alborosella (see above). The 

disc is always white pruinose. It is a lowland species 

growing near the base of trees in rain forest in 

Panama. 

SPECIMEYS EXAXIINED.4: 43455; 8: 38590. 

12. Ocellularia comparabilis 

FIGURE 41 

Ocellularia comparnbilis (Krempelhuber) Muller Argoviensis, 

1883:3 18. 

Thelotrema cornparabile Krempelhuber, 1875:220 [type collection: Rio de Janeiro, Brazil, Glariou 3463 (M, lectotype: 

isolectotypes in BM, C, G, M, PC, UPS, LV)]. 

DESCRIPTIoS.-Thallus corticolous, epiphloeodal, 

smooth, continuous or cracked with age, whitish 

mineral gray, 5-12 cm broad; apothecia numerous, 

emergent, 0.8-1.1 mm in diameter, carbonized with 

a simple or partially actinoid columella, the pore 

round and quite open, 0.1-0.2 mm in diameter, 

the pruinose tip of the columella visible; spores 

8, 6-8 X 15-20 pm, 4-6 loculate, I+ blue. 

CHE1LzISTRY.-PSOrOmiC acid with or without norpsoromic acid. 

REMARKs.-OCellUlaria coniparabilis is a variable 

species, the apothecia varying from nearly immersed 

to semiemergent and having a distinct thalline 

margin, open pruinose disc, and imperfect development of an actinoid columella. It intergrades 

most noticeably with 0. efformata (Krempelhuber) 

Muller Argoviensis, which has a jagged, erect thalline margin. It has a broad ecological range in 

Panama, from near sea level to 2000 m. 

SPTCI411:NS EXA\gISED.-la: 44905, 46025, Esdinger 4460; 3: 

35684; 3a: 46072, 44846, 44849. 
NUMBER 38 17 

13. Ocellularia concolor 

FIGURE 5a 

Ocellularia concolor Meyen and Flotow, 1843:230 [type collection: Manila, Philippines, Meyen (G, lectotype)]. 

DEscRIPTIoN.-Thahs corticolous, hypophloeodal, tannish gray and dull, 3-8 cm broad; apothecia 

numerous, dispersed, emergent, 0.8-1.1 mm in diameter, carbonized with a distinct columella about 

160 pm in diameter, the pore round to somewhat 

irregular, 0.3-0.5 mm broad, the rim blackening 

and the black tip of the columella easily visible; 

spores 8, 5-8 X 13-20 pm, 5-7 loculate, I+ blue. 

CHEMISTRY.-~'O substances present. 

REMARKs.-The distinguishing characters are the 

tannish hypophloeodal thallus and the rather open 

pore and black disc. As in Dominica, it grows at 

the base of large trees, on saplings, etc., in low to 

mid elevation rain forest. 

SPLCI\fENS EXAMIXED.--~: 43370, 43579; 3a: 44770; 4, 43431, 

43452; 6: 43474, 43478; 8: 38571, 38588; 9: 38460; 11: 44818, 

44988, 44995, 46018, 46033, Esslinger 4629. 

ILLusTRATIoNs.-Hale, 1974a: 18, fig. 9i. 

14. Ocellularia congesta, new species 

FIGURE 56 

DEscRIPTIox.-Thallus corticola, epiphloeodes, 

crassus, continuus, cinereo-albus, 6-10 cm latus; 

apothecia primus immersa, demum congesta et aggregata, semi-emergentia, 0.2-0.4 mm diametro, 

ecolumellata; ostiolum rotundatum, apertum, 0.1- 

0.2 mm latum; excipulum distinctum vel evanescens; hymenium 85-95 pm altum; sporae 8:nae, 

8 x 18 pm, 4-loculatae, I+ caerulescentes. 

CHEMISTRY.-HYPOprOtOCetrariC and 4-0-deme thylnotatic acids. 

HOLoTYPE.-Mature secondary rain forest, La 

iMesa, north of El Valle, Province of Cocle, elevation 750 m, M. E. Hale 43470, 13 February 1974 

RE3xARKS.-This species is very close to Thelotrema concretum Fee in external appearance, but 

T. concretum, also a canopy species, contains psoromic acid and has small longitudinally septate 

spores. Ocellularia microporella (Nylander) Zahlbruckner has hypoprotocetraric acid but the apo- 

(US). 

thecia are completely immersed and the spores 

smaller. 

SPECIMEN EXAMINED.-~ 38470 (cf.). 

15. Ocellularia conglomerata 

FIGURE 5c 

Ocellularia conglomerata Hale, 1974a: 18 [type collection: 

Dominica, Hale 37959 (US, holotype)]. 

DEscRrpTroN.-Thallus corticolous, epiphloeodal, 

thin and shiny, continuous, ashy white, 4-6 cm 

broad; apothecia not numerous, becoming aggregated in 2s and 3s, 0.5-0.8 mm in diameter, noncarbonized and without a columella, the thalline 

rim erect, the inner exciple distinct, pulverulent, 

more or less filling the disc; spores 8, 5 X 15 pm, 

6-loculate, It blue. 

CHEMISTRY.-stictic and constictic acids. 

REMARKs.-The conspicuous exciples and the 

aggregated apothecia distinguish this rare species. 

It is undoubtedly related to 0. exanthismocarpa 

(see below) but has much smaller spores and consistently aggregated apothecia with a semierect thalline margin. It occupies the same habitat as the 

type in Dominica, small branches of understory 

trees in rain forest at 800 m elevation. 

SPECIMEN EXAMINED.-~ 1 : 44952. 

ILLUSTRATIONS.-HP1f?, 1974a: 18, fig. 9h. 

16. Ocellularia crocea 

FIGURE 5d 

Ocellularia crocea (Krempelhuber) C. and D. van Overeem- 

'4scidiurn croceum Krempelhuber, 1875:25 [type collection: 

Phaeotrema croceum (Krempelhuber) Zahlbruckner, 1923:606. 

de Haas, 1922:118. 

Sarawak, Beccari 31c (M, lectotype) (Figure In)]. 

DEscRIPTio;v.-Thallus corticolous, epiphloeodal, 

rather thick, shiny, smooth to minutely verruculose, tannish gray, the exposed medulla pale reddish orange, 4-8 cm broad; apothecia numerous, 

solitary or crowded in 2s and 3s, more or less emergent, 0.4-0.7 mm in diameter, the thalline rim with 

a dull orange cast, a columella lacking, the inner 

exciple persisting and more or less filling the pore, 

the pore 0.1-0.2 mm in diameter; spores 8, 5-8 X 

18-20 pm, 5-7 loculate, I+ blue. 
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FICCKE 5.--Spwies of Orelldoria: a, 0. coucolor (Hale 46018); 6, 0. congesta (Hale 43470); C, 

0. conglomerata (Hale 449.52); d, 0. rrocea (Hale 43091); e, 0. doctilifrm (Hale 43503); f, 0. 

dactvliza (Hale 43559); g. 0. derolorata (Hale 43462); h, 0. dilatata (Hale 38640); i, 0. dissuta 

(Hale 43520): j, 0. dorningensis (Hale 46028): k, 0. esslingeri (Esslinger 4626): I, 0. exanthisrnocarpa (Hale 43542) (X 10). 
NUMBER 38 19 

CHEhiIsTRY.-"Cinchonarum" unknown and unidentified pigments. 

REMARKS.-The material from Panama is identical with the rather fragmentary type from Sarawak. 

The medulla and apothecial tissues are brilliant 

orange red and the pore nearly filled with excipular material. It is apparently confined to the upper 

trunk and canopy. 

SPECIMENS EXAMINED.-2: 43583, 43550, 43591. 

17. Ocellularia dactylifera, new species 

FIGURE 5e 

DESCRIPTION.-ThalIus corticola, epiphloeodes, 

opacus, minute granulosus, viridi-cinereus, 10-1 2 cm 

latus, sparse isidiatus, isidiis cylindricis, usque ad 

0.5 mm altis; apothecia (Figure 3b) numerosa, immersa vel leviter emergentia, dispersa vel paulo 

congregata, ecolumellata, 0.1-0.15 mm diametro, 

margine thallino albo-pruinoso; ostiolum rotundatum, 0.05 mm diametro; excipulum decoloratum, separatum vel pro parte connatum; hymenium 

90 pm altum; sporae 8:nae, 5 X 12 pm, 5-loculatae, 

I+ caerulescentes. 

CHEMISTRY.-Stictic and constictic acid. 

HoLoTYPE.-Rain forest, Santa Rita Ridge, 

Province of Cooh, Panama, elevation 350 m, M. E. 

Hale 43503, 12 February 1974 (US). 

REMARKS.-This is one of only several isidiate 

species in the genus. Ocellularia bullata Hale, also 

from Panama, has peculiar inflated, hollow isidia 

and stictic acid. From higher elevations we have 

0. isidiifera Hale, which has long, thin isidia and 

psoromic acid. Ocellularia dactyliza Hale (below) 

has large solid isidia and different chemistry. 

18. Ocellularia dactyliza, new species 

FIGURE 5j 

DESCRIPTIoN.-ThaIIus corticola, epiphloeodes, 

nitidus, pallide olivaceo-viridis, 8 cm latus, isidiatus, isidiis crassis, cylindricis vel irregulariter inflatis, facile disrumpentibus; apothecia (Figure 3c) 

emergentia, 0.7-1.1 mm diametro, ecolumellata, 

amphithecio laevi vel rugoso isidiatoque; ostiolum 

rotundatum vel irregulare, 0.1-0.3 mm latum, 

nigro-cinctum; excipulum apice fuligineum; hymenium 120 pm altum, paraphysibus granulatoencrustatis; sporae 6-8:nae, 12 X 50 pm, 12-15 

loculatae, I+ caerulescentes. 

CHEMIsTRY.-Protocetraric acid. 

HoLonPE.-Rain forest, Santa lii ta Ridge, Province of Colon, Panama, elevation 350 ni, M. E. 

Hale 43559, 12 February 1974 (US). 

REMARKS.-This species is characterized by the 

rather sparse, coarse isidia that break open apically 

and are easily dislodged, leaving white spots on the 

thallus. The ostiole is black-rimmed, much as in 

another protocetraric acid-containing species, 0. 

uerrucoSa (Fee) Hale, which has a roughened but 

never isidiate thallus and smaller spores (30-42 

pm long). It was collected 2 m up the trunk of a 

medium-sized tree in rain forest. 

19. Ocellularia decolorata, new species 

FIGURE 5g 

DESCRIPTIoN.-Thallus corticola, epiphloeodes, 

continuus, nitidus, albo-viridis, 6-9 cm latus; apothecia numerosa, irregulariter dispersa, immersa vel 

vix emergentia, ecolumellata, 0.4-0.6 mm diametro, 

margine thallino erecto, crasso, scabro; ostiolum 

irregulare, ca 0.1 mm latum; excipulum separatum, 

laceratum; hymenium 75-85 p,m altum; sporae 

8:nae, 6 X 18-22 pm, 5-6 loculatae, I+ caerulescentes. 

CHEMISTRY.-"CinChOna~Ulll" unknowns. 

HoLoTYPE.-secondary rain forest, La Mesa, 

north of El Valle, Province of Cocle, Panama, elevation 750 m, M. E. Hale 43462, 13 February 1974 

l<EMARKS.-ThiS species is virtually identical to 

Ocellularia crocea (Krempelhuber) Zahlbruckner, 

which has the brilliant pigment endocrocin. The 

disc is largely filled with the apically finely lacerate thallin margin and exciple. The chemistry relates it to the 0. cauata group. It occurs on the 

lower trunks of trees in secondary rain forest at 

low elevation. 

(US). 

20. Ocellularia dilatata 

FIGURE 5h 

OrcI1uliit.in rlilot,rtn Mullcr Algo\icu5is, 1895c:542 [type collection: Rio tle Janeiro, Brazil, Glaziou 5531 pro parte (G, 

lcctotype; isolQctot)pc in nbl)]. 

DEsCRrPTIoN.-Thallus corticolous, epiphloeodal, 

thin, dull, in part evanescent, pale greenish mineral 

gray, 4-8 cm broad; apothecia not numerous, dis- 
20 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

persed, chroodiscoid, up to 3.0 mm broad, round to 

irregularly elongate, the thalline rim thick, erect 

to recurved, the inner exciple not persisting, the 

disc flesh colored to heavily white pruinose; spores 

8, 5-8 X 15-25 pni, 6-9 loculate, I+ blue. 

CHESfISTRY.-stiCtiC and constictic acids. 

REMARKS.-The thick, split, recurved thalline 

margin and irregular shape of the apothecia are 

typical of the species. The chemical characters are 

shared by 0. neei Hale, which has smaller apothecia with a delicate margin and persistent inner 

exciple, 0. pseudoschizostoma Hale with a thick, 

erect thalline margin, and 0. sublilacina (Ellis and 

Everhart) Zahlbruckner, a very large species with 

long spores. Redinger (1936: 16) erroneously regarded the species as a Graphis. Ocellularia dilatatii 

occurs on the lower trunks and on lianas at 100- 

700 m elevation in Panama. 

SPECIME~S EXAMIKED.-~: 38640: 9: 38520. 

ILLUsrRATIoss.-Hak, 19i4a:21, figs. 1 la, b 

21. Ocellularia dissuta, new species 

FIGURE 5i 

DEsCRIPTIoN.-ThallUs corticola, epiphloeodes, 

nitidus, continuus, pallide brunneo-olivaceus, 6-10 

cm latus; apothecia chroodiscoida, solitaria vel 

pauce aggregata, 0.6-0.8 mm diametro, margine 

thallino crasso, irregulariter fisso, non reflexo; ostiolum irregulare, 0.4-0.6 mm latum; excipulum pro 

parte separatum fissum; disco pallide livido, leviter albo-pruinoso; hymenium 60-70 pm altum; 

sporae 8:nae, 4-6 X 10-12 pm, 3-4 loculatae, I-. 

CHEMISTRY.-NO substances present. 

HoLoTYPE.-Rain forest, Santa Rita Ridge, Province of Colon, Panama, elevation 350 m, M. E. Hale 

43520, 12 February 1974 (US). 

REMARKS.-The thalline margin of this species 

is coarse and split but not recurved, unlike any 

other species in the genus. It is confined to the 

lower trunks of small to medium-sized trees in 

mature rain forests at lower elevations. 

SPECIMENS EXAxlISED.-6: 43424, 43511, 43532; 8: 38575. 

22. Ocellularia domingensis 

FIGURE 5j 

Ocellulnrin doiningemis (Fee in Nylander) Miiller Argo\ienAscidiuni doniirigense Fee zn Nylaiider, 185731 18 [nomen 

sis, 1887b:398. 

nudum; Lalid publication in h’ylander 1863a:455: type collection: Colombia, Lindig 2683 (FH-Tuck, neot)pe; isoneotype in Bhi, UPS) (Figure Ip)]. 

DEsCRIPTIoN.-Thallus corticolous, epipliloeodal, 

rather thick, shiny, verruculose, pale tan, 4-8 cm 

broad; apothecia fairly numerous, strongly emergent, sometimes subglobose at maturity, about 1 

mm in diameter, carbonized but without a columella, pore more or less recessed, about 0.1 mm 

in diameter, subannulate, the pore area whitish; 

spores 1-2, 15-45 X 80-200 pm, It blue. 

CHEMISTRY.-Hypoprotocetraric acid. 

RE-\IARKS.-OcellUlaria domingensis was based 

on an unpublished Fee herbarium name, presumably from Santo Doming0 (Dominican Republic?), 

but no specimen so labeled could be found in 

Nylander’s collections at H or PC or in Fee’s at 

G or PC. The species was validated by Nylander 

in 1863 in his study of Lindig’s Colombian lichens, 

and I have selected one of these as the neotype of 

0. domingensis. The ascidioid apothecia are large 

and conspicuous and a columella is lacking; the 

spores are very large. The only similar species is 

0. henatomma (Acharius) Muller Argoviensis from 

South Africa, which has a distinct columella. The 

species occurs in cloud forest in Panama and is now 

known from similar habitats in Guatemala, Venezuela, and Colombia. 

SPECIME\S EXA\lI\ED.-1a: 44761, 44892, 44983, 46028 

23. Ocellularia esslingeri, new species 

FIGURE 5k 

DESCRIPTIox.-Thallus corticola, epiphloeodes, 

minute verruculosus, pallide cinereo-viridis, 4 cm 

latus; apothecia dispersa, chroodiscoida, emergentia, 0.7-1.1 mm diametro, margine thallino crasso, 

apice pulverulento vel subsorediato, erecto vel pro 

parte recurvato, excipulo proprio evanescenti, disco 

leviter albo-pruinoso, carneo; hymenium ca. 90 

pm altum; sporae 8:nae, 5-6 X 18-21 5-6 

loculatae, I+ caeruleae. 

CHEMISTRY.-Unidentified P- compound. 

HOLoTYPE.-on bark in rain forest at base of 

ridge west of Escuela de Agricola, elevation 1000 m, 

Province of Veraguas, Panama, T. L. Esslinger 4626, 

24 February 1975 (US). 

REMARKS.-NO other chroodiscoid species in the 

genus has quite the same round apothecia with a 
NUMBER 38 21 

thick, more or less entire (as opposed to split and 

recurved) thalline margin that is subsorediate 

around the rim. The proper exciple, apparently 

quite thick, may fuse with the thalline margin or 

be evanescent. Ocellularia pseudoschizostoma Hale, 

which is superficially similar, has a white, mostly 

hypophloeodal thallus, stictic acid, and less emergent apothecia. 

24. Ocellularia exanthismocarpa 

FIGURE 51 

Ocellularia exanthisnocarpa (Leighton) Zahlbruckner, 1923: 

590. 

Thelotrema exanthismocarpum Leighton, 1869: 169 [type collection: Ceylon, Thwaites C. L. 97 (BM, lectotype; isolectotype in H)]. 

Thelotrema porinoides Montagne and van den Bosch, 1855: 

484. 

Ocellularia multilocularis Zahlbruckner, 1912:369. 

Thelotrema isertii l’ainio, 1915: 40. 

Thelotrema hornothecium I’ainio, 1921: 190. 

Ocellularia homothecia (Vainio) Zahlbruckner, 19233593. 

Ocellularia porinoides (Montagne and van den Bosch) Zahl- 

[For full citations see Hale, 1974a:20.] 

bruckner, 1923:599. 

DESCRIPTIoN.-Thallus corticolous, thin and in 

part hypophloeodal, ashy white, about 3 cm broad; 

apothecia not numerous, semiemergent, about 0.7 

mm in diameter, noncarbonized and without a 

columella, the thalline rim becoming erect, pulverulent, the inner exciple persistent, free, forming a 

pore within the main pore, the main pore open, 

0.2-0.4 mm in diameter; spores 4-8, 10 >( 40-50 pm, 

about 12 loculate, I+ blue. 

CHEMISTRY.-stiCtiC and constictic acids. 

REMARKs.-This very widespread species, so often 

developed in secondary forests, was collected only 

once in Panama, in contrast to its abundance in 

Dominica (Hale, 1974a:ZO). The pore-within-a-pore 

configuration of the thalline and inner excipular 

margins, chemistry, and large spores make identification unmistakable. 

SPECIMEN EXAMINED.-~: 43542. 

ILLUSTRATIOh.S.-Hale, 1974a:21, figs. 5a, 1 lC, g; Salisbury, 

1972b3265, fig. 2 (as Thelotrema porinoides). 

25. Ocellularia fecunda 

FIGURE 6a 

Ocellularia fecunda (l’ainio) Hale, 1974a:21. 

Thelotrema domingense var. fecundum Vainio, 1896:208 [type 

collection: St. l’incent, Elliott 243 (BM, lectotype; isolectotype in TUR)]. 

Ocellularia domingensis var. fecunda (Vainio) Zahlbruckner, 

1923:590. 

DEScRIPTIoN.-Thallus corticolous, epiphloeodal, 

smooth and shiny, pale geenish gray, 10-20 cm 

broad; apothecia numerous, solitary and conspicuous, strongly emergent, 1.5-2.0 mm in diameter, 

heavily carbonized but without a central columella, 

the exposed medulla light yellowish orange, the 

pore irregular, 0.4-0.8 mm in diameter; spores 4, 

12-20 X 80-150 pm, 15-30 loculate, I+ blue. 

CHEMISTRY.--Hypoprotocetraric acid and in 

lesser concentration 4-O-demethylnotatic acid, with 

unidentified yellow K- pigments. 

REMARKS.--I had found this to be a very common 

species in Dominica and other islands in the Lesser 

Antilles with a very wide amplitude in ecological 

requirements. The large apothecia that abrade easily to reveal a pale yellow-orange medulla make 

field identification easy. In Panama it was very 

common in the Cerro Jefe region but surprisingly 

did not occur west of the Canal Zone in quite similar forests. 

SPECIMENS EXAhIISED.-8: 38530, 38540, 38542, 38556, 38569, 

38584, 38952, 38598, Mori 4089; 9: 38430, 38440, 38442, 38448, 

38453, 38469, 38472, 38490, 38498, 38503, 38508, 38509, 38513; 

12: Mori 4268. 

1LLUSTRATIONS.-Ha1e, 1974a:21, fig. 1 lh, i. 

26. Ocellularia glaucophaena 

FIGURE 6b 

Ocellularia glaucophaena (Krempelhuber) Zahlbruckner, 1923: 

591. 

Thelotrema glaircophaeiium Krempelhuber, 1875319 [type 

collection: Sarawak, Beccari 92 (M, lectotype; isolectotype 

in G) (Figure lr)]. 

DEscRIPTIox.-Thallus corticolous, epiphloeodal, 

often thin, shiny and smooth, continuous or cracked 

with age, pale greenish mineral gray, 5-15 cm 

broad; apothecia numerous, separate, nearly immersed to semiemergent, 0.3-1 .O mm in diameter, 

the thalline rim sometimes semierect and thickening, noncarbonized, a columella lacking, the inner 

exciple free, sometimes persisting but not conspicuous, the flesh-colored disc visible through the pore, 

rarely with pseudoactinoid surface markings, the 

pore 0.1-0.2 mm in diameter; spores 8, 4-6 X 10-24 

pm, 5-6 loculate, I+ blue. 

CHEMISTRY.-PSOrOmiC acid. 
22 SXIITHSOSIAN CONTRIBUTIONS TO BOTANY 

FICLRE &--Species of Ocellularia: a, 0. fecunda (Mori 4268); b, 0. glaucophaena (Hale 43486); 

c, 0. glaucula (Hale 43450); d, 0. isidiifera (Hale 46026); e, 0. landronii (Landrdn 358A); f, 

0. latilabra (.llori 4474); g, 0. microporella (Hale 44768) 11, 0. niinutula (Hale 43504); i, 0. mordenii (Hale 43446); 1, 0. myriotremoides (Hale 46020); k, 0. neei (Nee 3710); 1, 0. olivacea (Hale 

38434) (X 10). 
NUMBER 38 23 

REMARKS.-while 0. glaucophaena was first described from Sarawak, it seems to be pantropical 

with records in US from Florida, Cuba, Java, and 

the Solomon Islands. There are no strong distinguishing features; the noncarbonized apothecia are 

usually small but may reach 1.0 mm in Panama. 

It is a lowland rain forest species occurring on 

lower trunks. This complex will obviously need 

more study in the future as more specimens become 

available. 

SPECIMENS EXAMINED.-2: 43576; 3: 38625, 38626, 38647, 

38675, 38699; 3a: 44863, 44949, 44998, 46067, Esslinger 4347, 

4363, 4370; 4: 43437, 43438, 43459, 43486, 43496. 

27. Ocellularia glaucula 

FIGURE 6c 

Ocellularia glaucula (Xylander) Zahlbruckner, 1923:591. 

Thelotrema glauculum Nylander, 1863b3332 [type collection: 

Cuba, Wright (H, Nylander herbarium 22639, lectotype) 

(Figure Is)]. 

Thelotrema Eeptoporum Nylander in Leighton, 1869: 167 [nomen nudum in Nylander, 1857:118; type collection: Cuba, 

Wright 128 (BM, lectotype; isolectotypes in FH-Tuck, G, 

L, PC, UPS, us, W)]. 

Ocellularia leptopora (Nylander) Zahlbruckner, 1923:594. 

DESCRIPTIoN.-Thal~us corticolous, epiphloeodal, 

rather dull, smooth and continuous or cracked, 

whitish mineral gray, 5-10 cm broad; apothecia numerous, separate, immersed, 0.1-0.2 mm in diameter, noncarbonized and ecolumellate, the inner 

exciple free, forming a minute pore within the 

main pore, the main pore round, about 0.1 mm in 

diameter; spores 8, 5-6 X 10-12 pm, 3-4 loculate, 

I + blue. 

CHEMISTRY.-HYPOprOtOCetrariC acid and lichexan thone. 

REMARKs.-This canopy species is similar to 0. 

microfiorella (Nylander) Zahlbruckner superficially, 

but the thallus is dull white and the inner exciple 

distinctly free. I have also seen material from Costa 

Rica. No. 38491 has larger, more emergent apothecia than normal. 

SPECIMENS EXAMINED.-Ia: ESSlinger 4503; 3: 38689; 4: 

43450; 8: 38555, 38566; 9: 38491 (cf.). 

28. Ocellularia isidiifera 

FIGURE 6d 

Ocellularia isidiifera Hale, 1974b:492 [type collection: Province of Chiriqui, Panama, Hale 38709 (US, holotype)]. 

DEscRIPTIoN.-Thallus corticolous, epiphloeodal, 

rather thick and cracked, whitish to tannish ashy 

gray, up to 20 cm broad, isidiate, the isidia simple, 

cylindrical, about 1 mm high, initially erect but 

becoming decumbent; apothecia rather common, 

semiemergent, 0.5-1 .O mm in diameter, the thalline 

rim short but becoming erect, a columella absent 

or imperfectly developed, the pore round to irregularly elongate, 0.2-0.6 mm in diameter, the exposed 

disc white pruinose; spores 8, 6-9 X 18-22 pm, 5-6 

loculate, I+ blue. 

CHEMISTRY.-PSOrOmiC and norpsoromic asids. 

REMARKs.-Ocellularia isidiifera is amply distinguished by the tall thin isidia. I have collected it 

in cloud forests at 2500 m elevation in Venezuela 

as well. The type-locality in Panama was recently 

logged and the species may become extinct there. 

la: 46026, Esslinger 4489, 4509. 

SPECIMENS EXAMINED.-~: 38709, 38742, 38776, 38778, 38896; 

ILLUSTRATIONS-Hale, 1974bt492, fig. 7. 

29. Ocellularia landronii, new species 

FIGURE Ge 

DEscRIPTIoN.-Thal~us corticola, epiphloeodes, 

continuus, ni tidus, laevis vel minute verruculosus, 

cinereo-viridis, 6-8 cm latus; apothecia dispersa, 

immersa vel semi-emergentia, 0.8-1 .O mm diametro, 

apice fuliginea, columella evoluta, ca 120 pm diametro, apice nigra vel albo-pruinosa; hymenium 

80-90 pm altum; sporae 8:nae, 5-6 X 14-19 pm, 

5-6 loculatae, I+ caeruleae. 

CHEMISTRY.-NO substances present. 

HOLOTYPE.-on Dacryodes excelsa, El Verde, Luquill0 Mountains, Humacao District, Puerto Rico, 

I. Landron 358A (US). 

REMARKs.-This species is characterized by the 

smooth, shiny, dark greenish thallus and the peculiarly sunken apothecia. The distinct, white-tipped 

columella is typical of an emergent species, but 

here the apothecia are sunken in the thallus. There 

are no really comparable species, excepting perhaps 0. groenhartii Hale, where the apothecia are 

more clearly semiemergent (Hale, 1975: 173). Ocellularia landronii grows on the lower bole and base 

of trees in rain forests in both Puerto Rico and 

Panama. The species is named in honor of Dr. 

Ismael Landron. 

SPECIMEN EXAMINED.+: 43479. 
24 SMITHSONIAN CONTRIBUTIOSS TO BOTANY 

30. Ocellularia latilabra 

FIGURE 61 

Ocellularia latilabra (Tuckerman) Muller Argoviensis, 1895c: 

432. 

Thelotrema latilabrum Tuckerman, 1866:269 [type collection: 

Cuba, Wright 137 (FH-Tuck, lectotype; isolectotypes in 

BM, G, L, PC, UPS, It')]. 

DEscRIPTIos.-Thallus corticolous, epiphloeodal, 

shiny, continuous or cracked with age, pale greenish gray, 6-10 cm broad; apothecia numerous, separate, semiemergent, 1-2 mm in diameter, the thalline rim erect, firm, splitting with age and in part 

recurved, the disc densely actinoid-columellate, carbonized, the surface white pruinose, the pore open, 

irregular, 1-1.5 mm in diameter; spores 8, 6 X 12- 

24 pm, 5-6 loculate, I+ blue. 

CHEMISTRY.-PSOTOmiC acid. 

REhfARKs.-This actinoid species differs from 0. 

berkeleyana (Montagne) Zahlbruckner in having a 

tall, erect to recurved thalline margin. It is also related to poorly known 0. schizostoma (Tuckerman) 

Muller Argoviensis, described from Cuba, which 

has a similar irregularly erect, split thalline margin 

and an open disc. The disc, however, is not distinctly actinoid columellate; it lacks carbonization 

and is reticulately fissured. These three species are 

closely related and intergrade broadly. Only study 

of additional collections will decide the issue. The 

specimens from Panama occurred at lower to mid 

trunk in low to mid elevation rain forest. Specimens 44851, 44853, and 44860, all from the area 

north of Alto Piedra, have a distinct H,SO,+ yellow spot above psoromic acid on the TLC plates. 

SPECIMFNS EXAMINED.-~~: 44832, Esslinger 4350, 4377; 6: 

43475; 8: 38568; 11: 44838, 44851, 44853, 44860; 12: Mori 

4474. 

31. Ocellularia microporella 

FrGuRE 6g 

Ocellularia microporella (Nylander) Zahlbruckner, 1923:595. 

Thelotrema microporellurn Sylander, 1863b:327 [type collection: Cuba, Wright 126 (FH-Tuck, lectotype; isolectotypes 

in BM, G, L, PC) (Figure 2c)]. 

Thelotrema plurifnriuni Sylander, 1867:315 [type collection: 

Cuba, Wright (H, Nylander herbarium 22631, lectotype)]. 

Thclotrema suhterebratuin Sylander, 1886:174 [type collection: S. Thome, Moller 51 (H, Nylander herbarium 22772, 

lectotype; isolectotype in W)]. 

Ocellularia plurifaria (Kylander) Mu-ller Argoviensis, 1891b: 

Ocellularia subterebrata (Nylander) Zahlbruckner, 1923:601. 

75. 

DESCRIPTIoN.-Thallus corticolous, epiphloeodal, 

shiny, continuous and smooth, pale greenish mineral gray, 5-15 cm broad; apothecia numerous, immersed, 0.1-0.2 mm in diameter, noncarbonized 

and without a columella, the inner exciple usually 

evanescent but rarely well developed and filling the 

pore, the pore round, open, about 0.1 mm in diameter, the pale tan disc visible; spores 8, 5-8 X 12-24 

pm, 4-6 loculate, I+ blue. 

CHEMISTRY.-Hypopro tocetraric acid. 

REMARKS.-This species is obviously related to 

0. glauczila (see above), which contains lichexanthone in addition to hypoprotocetraric acid and has 

a persistent inner exciple. Both are canopy species, 

but 0. microporella generally occurs at higher elevations (380-800 m) than 0. glauczila. I have identified it from Jamaica also. 

4211; 11: 44768, 44807, 44811, 44827, 44984, 46032. 

SPECIMENS EXAVISED.-~: 43573, 43590; 8: 38611; 9: Molt 

32. Ocellularia minutula, new species 

FIGURE 6h 

DEscRrpTIos.-Thallus corticola, epiphloeodes, 

tenuis, continuus, pallide cinereo-viridis, 6-8 cm 

latus; apothecia (Figure 34 solitaria, sparsa, immersa vel semi-emergentia, 0.2-0.3 mm diametro, 

ecolumellata vel rudimentaliter columellata; excipulum non separatum, decoloratum; ostiolum discretum, rotundatum, 0.05 mm diametro, albo-cinctum; hymenium 120-130 pm altum; sporae 8:nae, 

7-10 X 22-24 pm, 6-8 loculatae, I+ caerulescentes. 

CHEMISTRY.-PSOrOmiC and norpsoromic acids. 

HoLoTYPE.-secondary rain forest, 9 km NW 

Gamboa along Naval Pipeline Road, Canal Zone, 

Panama, elevation 150 m, M. E. Hale 43341, 11 

February 1974 (US). 

REMARKS.-This inconspicuous species has much 

smaller apothecia than any other psoromic acidcontaining species with semiemergent fruiting bodies (nonmyriotremoid). The small but distinct pore, 

when strongly white annulate, resembles that of 0. 

umbilicata Muller Argoviensis, a Costa Rican species with a thicker, warty thallus and larger apothecia. The poorly known 0. albula (Nylander) Zahlbruckner from Brazil has a white pruinose thallus 
NUMBER 38 25 

and smaller spores. Ocellularia minutula occurs on 

saplings and lower trunks of trees in low to mid 

elevation rain forest. 

SPECIMENS EXA\fISED.-3a: 44885; 4: 43504; 7: 43603; 11: 

44789, 44801, 44897, 44941. 

33. Ocellularia mordenii 

FIGURE 6i 

Ocellulurza mordenii Hale, 1974:22 [type collection: Domitiica, Hale 37764 (US)]. 

DEscRIPTIoN.-Thallus corticolous, epiphloedal, 

continuous to cracked with age, rather warty and 

easily abraded, revealing the red medulla, greenish 

to tannish gray, about 8 cm broad; apothecia dispersed and inconspicuous, immersed to semiemergent in thallus warts, the surface very warty and 

easily abraded to reveal the red medulla, carbonized, ecolumellate or with a weakly developed columella, the pore round, about 0.1 mm in diameter; 

spores 1-2, 15-20 X 70-150 pm, 24-30 loculate, I+ 

blue. 

CHEMISTRY.-Unidentified anthraquinone pigments. 

REMARKS.-ThiS brilliantly pigmented species 

grows typically in the canopy in virgin rain forest. 

It is differentiated from 0. xanthostroma by color 

of the medulla (red rather than yellow orange) anti 

the somewhat less emergent apothecia. 

SPECIMEXS EXAMINED.-4: 43446, 43471; 6: 43501. 

34. Ocellularia myriotremoides 

FICLRE 67 

Ocellularia tnyriotremoides (h‘ylander) Zahlbruckner, 1923: 

596. 

Thelotrema mytiotremoides Nylander, 1859:221 [type collection: Weddell, Bolil ia (PC, lectotype) (Figure 2g)]. 

Thelotrema opncum l‘ainio, 1890:85 [type collection: Sitio, 

Minas Gerais, Brazil, T’nzno 808 (TUR, lectotype; isolectotypes in BM, FH, UPS) (Figure 2])]. 

Ocellula? in opaca (l’ainio) Zahlbruckner, 1923:597. 

Ocellulai !a albula (Sylander) Zahlbruckner f. opaca (\’ainio) 

Redinger, 1936:46. 

DEscRrPrIoN.-Thallus corticolous, epiphloeodal, 

continuous, dull, the surface becoming minutely 

scabrid, whitish mineral gray, 4-8 cm broad; apothecia numerous, separate to mostly irregularly aggregated, immersed to barely emergent, 0.3-0.4 mm 

in diameter, noncarbonized and without a columella, the pore round, surrounded by a broad white 

area, slightly raised and white pruinose, 0.1 mm in 

diameter; spores 8, 3-5 X 12 pm, 4-5 loculate, I+ 

blue. 

CHEMISTRY.-PSOrOmiC acid. 

RE&mRKs.-This species has a characteristic finely 

scabrid surface. The pore area is rather conspicuously annulate. Redinger (1936:46) reduced it to a 

forma of 0. albzila, but the type of 0. albula, a 

very poor fragment, has semiemergent apothecia 

with a small pore as well as a pruinose rather than 

scabride thallus surface. More closely related is 0. 

terebratula (Nylander) Muller Argoviensis, which 

has a smooth shiny thallus and at the fine structure 

level aculeate surface hyphae (Hale, 174a:5). The 

second collection by Tlieddell that Nylander identified as “Thelotrema myriotremoides” is in fact 0. 

terebratula, but I have selected the scabrid specimen as the lectotype since he describes the species 

as “thallo albiore magis opaco subfarinoso.” The 

surface of Thelotrema terebratulum, described in 

1867 from Colombia, is given as “laevigatus.” Ocellularia myriotremoides occurs at the mid trunk 

level in rain forest. 

SPECIMENS EXAMINED.-~: 38461; 11: 44788, 46020, Esdinger 

4651. 

35. Ocellularia neei, new species 

FIGURE 6k 

DEscRIPTIoN.-Thallus corticola, pro majore parte 

hypophloeodes, albo-cinereus, 5-7 cm latus; apothecia chroodiscoida, 0.4-0.7 mm diametro, margine 

thallino sat crasso, erect0 vel pauce reflexo; ostiolum latum, rotundatum, usque as 0.5 mm latum; 

excipulum separatum, distinctum, disco obscure 

ferrugineo vel albo-pruinoso, hymenium 60-70 pm 

altum; sporae 8:nae, 5 X 10-14 pm, 4 loculatae, 

I-. 

CHEMISTRY.-stictic and constictic acids. 

HOLOTYPE.-on Cochlospermum vitifolium in a 

thicket, Summit Gardens, Canal Zone, Panama, elevation about 30 m, M. Nee and S. Mori 3710, 18 

March 1971 (US). 

REzlARKS.-OCellZllaria neei differs from other 

chroodiscoid species such as 0. sublilacina (Everhart and Ellis) Zahlbruckner and 0. dilatata Miiller 

Argoviensis in being hypophloeodal and in having 

smaller spores. It is unusual in that it grows on 
26 SWTHSONIAN CONTRIBUTIONS TO BOTANY 

planted shrubs in an open, heavily used botanical 

garden near sea level, a lichenologically unpromising area. The species is named for Mr. Michael 

Wee, an observant lichen collector during his field 

studies in tropical America. 

36. Ocellularia olivacea 

FIGLRE 61 

Ocellularia olivacea (Fee) hfuller Argoviensis, 1887a:7. 

Mjriotrema olivaceum Fee, 1824: 103 [type collection: South 

America (C. lectotype)]. 

Ocellularia olivacea (Fee) C. and D. \'ail O\-ereern.De Haas, 

1922: 118. 

DEscRrPTroN.-Thallus corticolous, epiphloeodal, 

smooth and continuous or becoming cracked, up to 

0.5 mm thick, pale greenish or whitish ashy, 10-30 

cm broad; apothecia very numerous, immersed, 

0.13-0.3 mm in diameter, noncarbonized and lacking a columella, the inner exciple evanescent, the 

pore round, flush, about 0.1 mm in diameter; spores 

8, uniseriate, 4-6 X 6-12 pm, 4 loculate, I+ blue. 

CHEXIIsTRY.-"Olivacea" unknown and a second 

lower spot. 

RE;CIARKs.-Although this species is common in 

Dominica, in other islands of the Lesser Antilles, 

and in Cuba, I found it to be rather rare in Panama. It must be distinguished carefully with spore 

sections from Thelotrema clandestinzim F6e and 

with chemical tests from 0. terebratda (Nylander) 

Xluller Argoviensis and 0. uniseptata Hale (both 

psoromic acid present). They occupy the same habitat, mid bole to canopy of trees up to 750 m elevation in rain forest. 

SPECIIIENS EXAMINED.-Z: 43540; 3: 3864i; 9: 38434. 

ILLvsrRATross.-Hale, 1974a:23, fig. 12c; Redinger, 1936:48, 

fig. 24. 

37. Ocellularia pachystorna, new combination 

FIGURE 7a 

Thelotrema pachystomum Sylander, 1863b:325 [type collection: Pro\ ince Carabaya, Peru, Weddell (PC, lectotype) 

(Figure 2k)l. 

Phaeotrema pachystomum (Nylander) Zahlbruckner, 1923:609. 

DEscRImIox.-Thallus corticolous, epiphloeodal, 

continuous, shiny, light greenish mineral gray, 

about 8 cm broad; apothecia common, strongly 

emergent, separate to clumped in 2s and 3s, 0.7-1.4 

mm in diameter, the exciple carbonized, columella 

lacking, the thalline rim becoming erect, the pore 

round, the surrounding area broadly pulverulent 

or white pruinose, 0.1-0.2 mm in diameter, the pale 

disc surface visible; spores 8, 4 X 12 pm, 4-loculate, 

1 I blue. 

CHEMISTRY .-PsOrOmic acid. 

REMARKS.-The large emergent apothecia with a 

conspicuous, erect pulverulent thalline margin characterize this rare species. The t)pe has colorless, not 

brown, spores. I have collected it in Venezuela at 

2000 m elevation and it occurred in the Darien 

region at about 1800 m. 

SPECI\IE\ EXA\II\ED.-12: MOri 4439 

38. Ocellularia papillata 

FIGURE 7b 

Ocellularia pnpi/lata (Leighton) Zahlbruckner, 1923:597. 

Thelotvema popillotutn Leighton, 1869: 169 [type collection: 

Ceylon, Thwaites C. L. 129 (BM, lectotype; isolectotypes 

ill C, H. P, S, UPS, IC)]. 

DEscRIPTro,u.-Thallus corticolous, epiphloeodal, 

smooth and continuous, shiny, whitish to greenish 

gray, 3-12 cm broad; apothecia numerous, immersed 

to slightly emergent, carbonized, a columella usually present, 50-100 pm in diameter, the pore 

round, 0.1-0.2 mm in diameter; spores 8, 6-10 X 

20-32 pm, 6-8 loculate, I+ blue. 

CHEMISTRY.-NO substances present. 

REMARKs.-This was one of the commonest base 

level species in Panama as it was in Dominica 

(Hale, 1974a:24). It grows on prop roots, buttresses, 

lianas, and lower boles of trees in rain forest at low 

to mid elevation. All specimens had a columella. 

A chemical test should be used to separate it from 

0. pel-forata (see below). 

SPECIZIESS EXA\IISED.-3: 38627, 38631, 38657, 38679, 38701; 

3a: 41769, 44791, 44797, 44858, 44867, 44868, 44873, 44888, 

44889, 44896, 44934, 44937, 44975, 46071, 46075, 46084, Esslinger 4398; 4: 3436, 43481, 43499, 43500, 43502, 43587; 6: 43340, 

43429, 43463, 43505, 43529, 43531, 43558; 7: 43604, 43605, 

43606; 8: 38583; 10: Esslingel- 4630; 11: 44906, 44968, 45809, 

46043, 46031. 

ILLLSTR.4TIOS.-H3k, 1974a:23, figs. 12d, e. 

39. Ocellularia perforata 

FIGURE 'ir 

Ocellularia perforata (Leighton) Muller .4rgoviensis, 1892:284. 

Thelotrema perforntum Leighton, 18663447 [type collection: 

Casiquiari, Brazil, Spruce 254 (BM, lectotype)]. 
NUMBER 38 27 

FIGURE 7.-Species of Ocellularia: a, 0. pachystoma (Mori 4439); b, 0. papillata (Hale 43463); 

c, 0. perfnrata (Hale 46024); d, 0. protocetrarica (Hale 44786); e, 0. pseudoschizostoma (Hale 

38559); f, 0. puluerulenta (Hale 46073); g, 0. recotidita (Mori 4293); h, 0. referta (Hale 43566); 

i, 0. iliabdosf~ora (Mori 4448); j, 0. rhodostroma (Hale 44948); k, 0. ripleyi (Hale 44836); I, 

0. subcavata (Hale 38516) (X 10). 
28 SMITHSONIAN CONTRIBUTIOSS TO BOTANY 

Thelotrema cavatnm var. planius Nylander, 1867:316 [type 

collection: Colombia, Liridig 64 (BXf, lectotype; isolectotypes in FH-Tuck, PC)]. 

Theiotremn terebmtum var. abbreviatultim Vainio, 1890:83. 

Ocellztluria rufociucta Miiller ;\rgoviensis, 1893b: 146. 

The lot re ma excurm t uin l’ai no, 1 896 : 208. 

Th el 0 tre !?7 a excava t urn var. impressu lu rn l’aino, 1896: 208. 

Ocellularia cavata var. plaiiior (Nylander) Zahlbruckner, 

Oceliulaiia excnvata (Vainio) Zahlbruckner, 1923:590. 

Ocellularia terebrata var. abbreviatula (i’aino) Zahlbruckner, 

1923585. 

1923:602. [For full citations see Hale (1974a:25)]. 

DEscRIPTIox.-Thallus corticolous, epiphloeodal, 

shiny, smooth or becoming verrucose with age, 

whitish or greenish gray, 3-10 cm broad; apothecia 

numerous, immersed to somewhat emergent, 0.4- 

0.6 mm in diameter, carbonized, a columella usually 

developed but variable, 25-160 pm in diameter, or 

rarely lacking, the pore round, 0.07-0.13 mm in 

diameter; spores 8, 6-10 X 20-36 pm, 6-8 loculate, 

I + blue. 

CmwsTRY.-Protocetraric acid with or without 

the “amplior” unknown. 

REMARKs.-ThiS is the commonest species of 

Ocellzilaria and of the family in Panama, occurring 

on tree bases, buttresses, saplings, lianas, etc., from 

sea level to 2200 m in rain forest. About 10% of 

the specimens lacked a columella, much as in Dominica (Hale, 1974a:24). ’IVhere it occurs with externally similar 0. papillata-and the only major 

locality where they did not was the cloud forests 

of Volcin Chiriqui-0. perfornta can be distinguished by the P+ red test. I have identified this 

pantropical species from Cuba, the Lesser Antilles, 

Guatemala, Costa Rica, Brazil, and the Solomon 

Islands. 

SPECIMENS EX.AMINED.--I: 3871 1, 38725, 38737, 38544, 38764, 

38781; la: 44812, 44825, 44826, 44903, 44913, 44923, 44954, 

45842. 45877, Esslinger 4,502; 2: 3535, 43577; 3: 38616, 38623, 

38633, 38643, 38665, 38672, 38719; 3a: 44759, 44782, 44817, 

44829, 44843, 44872, 44893, 44908, 44953, 44971, 44989, 46076, 

46080, Esslinger 4339; 6: 43482, 43509, 43523, 43663; 8: 38609; 

9: 38437, 38485, 38488, 38496, 38507; 10: 46044, 46063; 11: 

44799, 44848, 44854, 44899, 44915, 44931, 44969, 44970, 44997, 

46022, 46024. 

ILLLSTRATIOSS.-Hale, 1974a:23, fig. 12f. 

40. Ocellularia protocetrarica, new species 

FIGURE id 

DEScRIPTIoN.-Thallus corticola, epiphloeodes, 

nitidus, laevis vel verruculosus, pallide viridi-albus, 

5-8 cm latus; apothecia (Figure 3e) numerosa, solitaria vel sparse aggregata, emergentia, 0.9-1.2 mm 

diametro, apice fuliginea, columella nulla vel rudimentariter actinoides, vix fuliginea, disco albopruinoso; ostiolum rotundatum, 0.2-0.5 mm diametro; hymenium ca 120 pm altum; sporae 8:nae, 

6-8 X 20-24 pm, 5-7 loculatae, I + eruleae. 

CHEwsmY.-Protocetraric acid. 

HoLoTYPE.-Primary rain forest on road to Rio 

Calovebora, 4-8 km from Alto Piedra, Province of 

Veraguas, Panama, elevation 800 In, M. E. Hale 

44786, 23 February 1975 (US). 

RExmKS.-The disc of the species at maturity 

has superficial pruinose actinoid divisions or fissures that may be slightly carbonized. The round 

pore is quite open and concolorous with the thallus. 

Two other protocetraric acid-containing species 

may cause confusion. Ocellularia vermcosa (see page 

33), which also occurs in Panama, has a black 

rimmed, rather irregular pore and a smooth, entire 

disc. Ocellularia nigropuncta Hale from Dominica 

has smaller apothecia and a small black-rimmed 

pore. 

SPECIMESS EXAMIISED.-~~: 46055; 12: 2bfOri 4270 (Cf.). 

4 1. Ocellularia pseudoschizostoma, new species 

FICLRE 7~ 

DEscRIPTIox.-Thallus corticola, epiphloeodes vel 

evanescens, albus, 3-5 cm latus; apothecia dispersa, 

singularia, chroodiscoida, 0.5-0.7 mm diametro, 

margine thallino crasso, erecto, non reflexo, irregulariter fisso; excipulum decoloratum, aetate non 

separatum; discus lividus, fissus, albo-pruinosis; 

hymenium 80-90 pm altum, paraphysibus granulato-encrustatis; sporae 8:nae, 12 X 24 ,,,m, 6-7 loculatae, I caerulescentes. 

CHEMISTRY.-stiCtiC and constictic acids. 

HoLoTYPE.-Primary rain forest, El Llano-Carti 

road at 16-20 km, Province of Panama, Panama, 

elevation 300 m, M. E. Hale 38559, 6 April 1973 

RExfARKS.-hother chroodiscoid species from 

Panama, 0. neei Hale, has similar chemistry but 

differs in having a distinct inner exciple and a dark 

disc, Ocellularia sublilacina (Everhart and Ellis) 

Zahlbruckner has much larger apothecia with a 

strongly recurved margin. Ocellularia pseudoschizostoma grows in the canopy in virgin rain forest. 

(US). 
NUMBER 38 29 

42. Ocellularia pulverulenta, new species 

FIGURE 7j 

DEscRIPTxolv.-Thallus corticola, epiphloeodes, 

verrucosus vel subgranulatus, rimosus, albido-viridis, ca 10 cm latus, pro parte isidiatus, isidiis minutis, sorediatis, soraliis irregularibus, sorediis granulosis; apotheca sparsa, emergentia, amphithecio 

verrucoso-pustulato, sorediascenti, 0.4-1.0 mm diametro, ecolumellata; ostiolum rotundatum, ca 0.1 

mm diametro; excipulum decoloratum; hymenium 

60-70 pm altum; sporae 8:nae, 6-10 X 18-24 pm, 

6-7 loculatae, I+ caerulescentes. 

CHEhfIsTRY.-Psoromic and norpsoromic acids. 

HoLoTYPE.-Secondary rain forest, 9 km NW of 

Gamboa along Naval Pipeline Road, Canal Zone, 

Panama, elevation 150 m, M. E. Hale 43468, 11 

February 1974 (US). 

REhfARKs.-The small irregular soredial patches 

and small crumbly masses of isidioid growths are 

unique in this species. There is, in fact, only one 

other sorediate species in the genus, 0. sorediata 

Hale from Dominica, which has large capitate soralia, immersed apothecia, and no chemical substances. The noncarbonized apothecia, at the same 

time, are similar to those of 0. glaucophaena 

(Krempelhuber) Zahlbruckner, although larger. 

Ocellularia pzilverulenta grew at about 1.5 m up 

on large trees in secondary low elevation rain 

forest. 

SPECIMEXS EX4MINED.-3a: 44821, 46073. 

43. Ocellularia recondita 

FIGURE 7g 

Ocellularia recondita (Stirton) Zahlbruckner, 1923:600. 

Thelotrema recondititm Stirton, 1876:162 [type collection: 

Amazons, Brazil, Trail (BM, lectotype) (Figure 20)]. 

DEscRxPTxo;v.-Thallus corticolous, epiphloeodal, 

shiny, minutely rugulose, greenish gray, 6-10 cm 

broad; apothecia numerous, emergent but not basally constricted, heavily carbonized, 0.6-1 .O mm in 

diameter, the columella distinct, thick, the pore 

round, 0.1-0.3 mm in diameter, usually white annulate, the white pruinose tip of the columella 

clearly visible; spores 8, 7-8 X 14-20 pm, 5-6 loculate, I+ blue. 

CHEMISTRY.-unknown P- substance. There is 

a single strong H,SO,+ gray spot near the “olivacea” unknown. 

REMARKS.-The emergent apothecia have a diStinctive white rim around the pore, but the most 

important character is the chemistry, a P- unknown. Redinger (1936: 37) regarded 0. recondita 

as a synonym of 0. cavata (Acharius) Miiller Argoviensis, noting the large apothecia. I do not believe 

the two species are related at all. Aside from the 

type, I have seen one specimen from a lowland 

rain forest in Colombia, the usual habitat in 

Panama. 

SPECIMENS EXA\IINED.-3: 38618, 38628, 38687; 3a: 46074, 

Esslinger 4366; 4: 43488; 6: 43517. 

44. Ocellularia referta, new species 

FIGURE 7h 

DEscRIPTros.-Thallus corticola, epiphloeodes, 

nitidus, rimosus, pallide olivaceo-viridis, 5-7 cm 

latus; apothecia semi-emergentia, subchroodiscoida, 

solitaria atque 2-3 confluentia, 0.5-0.7 mm diametro, margine thallino crasso, fisso, vix emergenti, 

non reflexo; ostiolum vix visibile, ca 0.1 mm diametro; excipulum pro parte separatum, fissum, 

apice fuligineum; hymenium ca 60 pm altum; 

sporae 8:nae, 8 X 24 pm, 5-7 loculatae, I+ caerulescentes. 

CHEMISTRY.-Hypoprotocetraric and 4-0-demethylnotatic acids. 

HOLOTYPE. Secondary rain forest, La Mesa, north 

of El Valle, Province of Cocle, Panama, elevation 

750 m, M. E. Hale 43566, 13 February 1974 (US). 

REMARKS.-There are no other species with hypoprotocetraric acid similar to this one. Phaeotrema 

aggregatum Hale from Dominica has a similar filled 

disc but the apothecia are smaller and more aggregated and no lichen substances are produced. This 

species occurred at mid bole level in the rain 

forest. 

45. Ocellularia rhabdospora 

FIGURE 7i 

Ocellularia rhabdospora (Nylander) Redinger, 1936: 17. 

Ascidium rhabdosporum Nylander, 1867:320 [type collection: 

Pie de Cuesta, Colombia, Lindig (H. Nylantler herbarium 

22432, lectotype; isolectotype in BM)]. 
30 SMITHSOSIAN CONTRIBUTIOXS TO BOTANY 

Plzaeot? ema rliabdosporum (Nylander) Miiller Argoviensis, 

1895c:434. 

DEscRIPTIox.-Thallus corticolous, epiphloeodal, 

shiny, verruculose, rimose with age, brownish or 

greenish gray, 6-10 cm broad; apothecia numerous, separate and dispersed, strongly emergent and 

ascidioid, up to 2 mm in diameter, heavily carbonized with a distinct central columella, the pore 

round, 0.2-0.3 in diameter, slightly depressed and 

narrowly white annulate; spores 1-2, 12 X 120-160 

pm, 15-20 loculate, I+ blue. 

CHEMISTRY.-"Cinchonarum" P unknown substances. 

REMARKS.-In this Brazilian study Redinger 

(193626) described in detail the specimens on 

which Krempelhuber based Ascidium melanostomum (Phaeotrema melanostomum (Krempelhuber) 

3f tiller Argoviensis). I propose to typify this species 

with Glaziou 5573 (G). It contains hypoprotocetraric acid (as do the syntypes), even though the 

basic morphology is very close to that of 0. rhabdospora, which contains the Pt red "cinchonarum" 

series. With this hypoprotocetraric acid-containing 

population excluded and with proper typification 

of 0. rhabdospora, we must now refer to the large 

ascidioid species collected in Panama as 0. rkabdospoya. It is a rare species, occurring only in the 

type-locality in Colombia and in eastern Darien. 

SrEcI%ims Ex;\\riYrn.-lP: ,Ifo?-i 4448, 4457, 4460, 4463, 

4478, 4492. 

46. Ocellularia rhodostroma 

FIGURE 7j 

Ocellularia rhodostroma (Montagiie) Zahlbruckner, 1923:600. 

.4scidium rhodostromu~n Montagne, 1831 :75 [type collectioii: 

French Guiana, collection 1934 (PC, lectotype)). 

DEscRIPTIos.-Thallus corticolous, epiphloeodal, 

smooth to finely verruculose, continuous, pale 

greenish tan to gray, the exposed medulla deep 

pink, 5-10 cm broad; apothecia numerous, ascidioid, 1 .O-1.8 mm in diameter, heavily carbonized 

but without a columella, the thick deep pink niedulla easily exposed as the fragile cortex breaks 

away, the pore round, strongly annulate with a 

raised ring, 0.05-0.1 mm in diameter; spores 1-2, 

20-30 X 100-140 pm, 20-25 loculate, 1+ blue, 

CHE~IISTRY.-H)pOprOtOCetrariC acid and in 

greater concentration 4-0-demethylnotatic acid with 

unidentified K- pigments. 

RE;MARKS.-This was the commonest species of 

Ocellzilaria in Dominica (Hale, 1974a:27), but in 

Panama it was commonly collected only in the midelevation rain forests west of the Canal Zone and 

mostly in the canopy or mid bole area of the trees. 

Surprisingly I did not collect it at Chiriqui, an area 

of mossy forest not unlike that in Dominica where 

it had maximum abundance, and, as expected, it 

was absent from the low elevation rain forests. The 

pink medulla and large annulate-pored apothecia 

are so distinctive that the species can be identified 

positively in the field. 

SPI:CI>IF.\S EX \MISLD.--~: 43454, 43549, 43372, 43578. 43581; 

6: 43508, 43333, 43537; 9: 38438, 38443, 38438, 38462: 10: 44990; 

11: 44i8:,. ,44800, 44841, 44850, 44884, 44929. 44932, 44938, 

41948. 44963, 45814, 4GOj0, Esslinger 4648. 

Ir LL's-rR~~rross.-~a~e, 1974a:26, fig. 13a. 

47. Ocellularia ripleyi 

FICCKE 7k 

Oceiiukirici ripleyi Hale, 1974b:493 [type collection: Canal 

7oiie. Panama, Hale 38664 (US, holot\pe)J. 

DEscRIPTros.-Thallus corticolous, epiphloeodal, 

continuous, uniformly minutely yerrucose, dull, 

whitish gray, 8-14 cm broad; apothecia rather numerous, emergent, 1.0-1.5 mm in diameter, the 

amphithecium minutely verrucose, carbonized with 

a distinct columella, the pore round, 0.2 mm in 

diameter, white annulate; spores 1-2, 12-15 X 80- 

200 pm, 8-23 loculate, I+ blue. 

CHEalrsTRY.-Psoromic acid with or without norpsoromic acid. 

RE\fARKs.-This conspicuous understory species 

is the dominant Ocellularia on Barro Colorado 

Island. I collected it at only one other site, the adjacent mainland near Gamboa. This high localization suggests that it is a recently revolved population, very successful within a narrow environmental 

and geographic range, although one record has been 

made in Cuba. It is externally and chemically similar to Tlielotrema intei.positum Sylander, which 

does not occur at all in Panama. 

SPIXIMI-NS EXAMINED.-3: 38621, 38648, 38650, 38658, 38664, 

38688, 38700, Heire 14; %a: 44822. 44828, 44836, 44840, 44856, 

44991, 45813, 4GO81, EssliJiger 4353.1, 4359, 4374, 4406, 4409; 

4: 43434. 

ILLvsTRArroNs.-Hale, 1974b:493, fig. 8. 
NUMBER 38 31 

48. Ocellularia subcavata 

FIGURE 71

Ocellularia subcavata (Nylander) Zahlbruckner, 1923:601. 

Thelotrema subcavatum Nylander, 1876:561 [type collection: 

Cuba, Wright 509 (H. Nylander herbarium 22813, lectotype; isolectotype in FH-Tuck)]. 

Thelotrema vagum Vainio, 1896:209. 

Ocellularia vaga (Vainio) Zahlbruckner, 1923:603. 

DEScRIPTIoN.-Thallus corticolous, epiphloeodal, 

shiny, thick, warty and cracked, ashy white, up to 

12 cm broad; apothecia very numerous, emergent, 

0.6-1.0 mm in diameter, crowded, the walls carbonized but the central columella carbonized only at 

the base, the apex noncarbonized and white pruinose, the pore round, 0.1-0.2 mm in diameter, white 

annulate with a raised ring, the center filled with 

the white pruinose columella tip; spores 8, 4-8 X 

8-22 pm, usually uniseriate, 5-6 loculate with narrow locules, I+ blue. 

CHEMIsTRY.-Psoromic acid with or without norpsoromic acid. 

REMARKS.-ThiS species is easily recognized by 

the numerous, crowded annulate pored apothecia 

and the basally uncarbonized columella. It was very 

common in Dominica (Hale, 1974a:28) and has 

been identified from all the islands in the Lesser 

Antilles, Jamaica, and Venezuela, generally at mid 

elevation (300-800 m). I collected it in Panama 

only west of the Canal Zone, mostly at canopy 

level. 

SPECIMESS EXAMINED.-2: 43469; 8: 38529, 38543, 38551, 

38553, 38602, 38603; 9: 38479, 38487, 38500, 38516; 11: 46048. 

ILLUSTRATIOSS.-Hak, 1974a:26, fig. 13b. 

49. Ocellularia subemersa 

FIGURE 8a 

Ocellularia subemersa Muller Argoviensis, 1886:310 [type collection: Gordon Town, Jamaica, Hart 125 (G, lectotype) 

(Figure 2r)l. 

DEScRIPTIoN.-Thallus corticolous or partially 

muscicolous, epiphloeodal or very thin and in part 

hypophloeodal, shiny, fragile, tannish, or greenish 

gray, 5-8 cm broad; apothecia common, solitary or 

clumped in 2s and 3s, strongly emergent and becoming basally constricted, 0.4-0.7 mm in diameter, 

the thalline rim carbonized, erect, or becoming 

abruptly apically recurved, the entire rim white 

pruinose, the columella distinct, carbonized, the 

apex white pruinose, the pore open, irregular, 0.3- 

0.5 mm in diameter; spores 8, 5-6 X 12-15 pm, 

4-6 loculate, I+ blue. 

CHEMISTRY.-"Cinchonarum" unknowns. 

REMARKS.-This unusual species is characterized 

by the rather small but distinctly urceolate apothecia with an erect or slightly flared thalline margin surrounding the apically white pruinose columella. It is a canopy species at a 700-800 m 

elevation in Panama and also occurs in Jamaica, the 

type-locality, and Trinidad. 

SPECIMENS EXAMINED.-2: 43522, 43539, 43552; 9: 38474; 

11 : 44891, 46060, Esslinger 4646,4647. 

50. Ocellularia sublilacim 

FIGCRE 86 

Ocellularia sublilacina (Ellis and Everhart in Smith) Zahlbruckner, 1923:GOl. 

Karstenia sublilacina Ellis and Eberhart in Smith, 1896:49 

[type collection: Castillo, Nicaragua, Smith 49 (BPI, lectoThelotrema sublilacinum (Ellis and Everhart in Smith) 

type)]. 

l'ainio, 1923:137. 

DEscRIPTIoN.-Tha~~us corticolous, epiphloeodal, 

shiny, smooth, and continuous, dark olivaceous or 

greenish mineral gray, up to 15 cm broad; apothecia common, separate, conspicuous, chroodiscoid, 

2-5 mm in diameter, the thalline margin usually 

thick, erect or recurved, the inner exciple well developed, persistent, free, the disc pale tan to flesh 

colored; spores 2-4, 10-15 X 60-120 pm, 15-20 loculate, I + blue. 

CHEMISTRY.-stiCtiC and constictic acids with 

associated unknown substances. 

REMARKS.-oCellUhia sublilacina is one of the 

truly remarkable species in the genus. The chroodiscoid apothecia are so large as to appear as disc 

fungi, leading Ellis and Everhart to describe it as 

a fungus originally. The thalline rim and inner 

exciple remain free and form a conspicuous fringe 

around the disc. It occurs mainly on the lower 

trunk of trees in dense woods. I have also collected 

it in Trinidad. 

SPECISITNS EXAXIINED.-3: 38514, 38597, 38638, 38678; 3a: 

44760, 44771, 44794, 44803, 44874, 44901, 46077, Esslinger 

4352, 4357, 4387; 4: 43343, 43456; 8: 38596; 9: 38468; 11: 44795. 
32 SMITHSOSIAN CONTRIBUTIONS TO BOTANY 

FIGURE 8.--Species of Ocellularia: a, 0. subemersa (Hnle 37471); 0, 0. sublilocinn (Hole 38997); 

c, 0. srtbrcjTightii (Mori 4286); d, 0. terebrotitla (Hale 44iSi); c, 0. lii,gitiu/a (Hale 45832); f, 0. 

uniseptata (Hale 38671); g, 0. neri-ucosa (Hale 43453); h, 0. viridis (Hale 38338); i, 0. xanthostroma (Hale 44758) (X 10). 

51. Ocellularia subwrightii, new species 

FIC~RE 8c 

DEscRIPTioN.-Tha1lus corticola, epiphloeodes, 

sat tenuis et fragilis, laevis vel verruculosus, nitidus, 

continuus, pallide stramineo-viridis, 5-15 cm latus; 

apothecia rara, separata, rotundata vel irregulariter 

expansa, valde emergentia, aetate basi constricta, 

2-5 mm lata, margine tliallino crass0 (usque ad 1 

mm lato), scabroso, disco aperto, cameo, actinoidediviso, columellis subfuligineis, elevatis, pro parte 

superficialibus, livido-stramineo-pruinosis; liymenium ca. 120 Fm altum; sporae 8:nae, 5-7 X 14-22 

pm, 4-5 loculatae, I+ caeruleae. 

CmarmRv.-Psoromic acid and an unidentified 

H,SO,+ yellow spot near stictic acid in the benzene solvent. 

HOLOTYPE.-cerro Mali, Province of Darien, 

Panama, elevation 850 m, S. Mori and A. Gentry 

4286, 16 January 1975 (US). 

RExfARI(S.-The lai ge apothecia are pale yellowish white and stand out conspicuously from the 

more greenish thallus. The development of a 

coarse, partially carbonized, superficial columella is 

identical Tvith that in Thelotrema wl-ightiz (Tuckerman) Zahlbruckner. There are no close relatives in 

Ocellulal-ia; for example, 0. be? kelepnna (see page 

14) has smaller, round apothecia with a heavily 
NUMBER 38 33 

carbonized actinoid disc. It occurs rather widely 

in Panama from 100 m to 1100 m elevation on 

lower trunk and basal area of large trees. 

SPECIXIESS Ex.ilrIN~D.-3a: 44887; 6: 43519; 9: 38473; 11: 

46016, 44900 (cf.); 12: Afori 4254, 4255, 4279, 4286, 4305. 

52. Ocellularia terebratula 

FIGURE 8d 

Ocellularia terebratula (Sylander) Muller Argoriensis, 1887a: 

12. 

Thelotrema terebratdum Sylander, 1867:315 [type collection: 

Colombia, Lindig 129 (H, Sylander herbarium 22637, lectotype; isolectotypes in FH, G, M, PC)]. 

Thelotrema clandestinum f. remanens h’ylander, 1867:317. 

Ocellularia remanens (Sylander) Muller Argoviensis, 1887a:7. 

[For full citations see Hale (1974a:28)]. 

DEscRrPTroN.-Thallus corticolous, epiphloeodal, 

quite thick, shiny, smooth and continuous, pale 

greenish to whitish ashy, 4-8 cm broad; apothecia 

very numerous, immersed, 0.2-0.4 mm in diameter, 

noncarhonized and ecolumellate, the inner exciple 

evanescent or rarely persistent, the pore round, 

flush, 0.05-0.1 mm in diameter, usually faintly 

white annulate; spores 8, 5 X 12 pm, 3-5 loculate, 

I+ blue. 

CHE411STRY.-PSoromic and norpsoromic acids. 

RExfARKs.-This is one of the canopy species of 

the Ocellularia alba-Thelotrema clandestinum type 

that must be examined carefully for spores and 

chemistry for positive identification. It is distinguished by the flush apothecia, smooth shiny thallus, and presence of psoromic acid. The fine structure of the cortical surface, semierect aculeate 

hyphae (Hale, 1974a:2), is distinctive. Closely related 0. myriotremoides (#34) has a scabrid surface. I have identified specimens from Florida, 

Costa Rica, the Lesser Antilles, New Caledonia, 

Java, Sabah, and the Solomon Islands. 

9: 38464, 38476. 

SPECIMESS EXA>IISED.-2: 43556; 3a: 44787; 6: 43525; 

~I.I.rlsTRATIoss.-Hale, 1974a:26, fig. 13e. 

53. Ocellularia turgidula 

FIGURE 8e 

Ocellulm ia tttrgrrlrtla Muller Argoi iensis, 1893a;94. [type collection: i’vlont hfou, New Caledonia, Balansa (PC, lectotype; 

isolectotype in G)]. 

DEscRrPTrorv.-Thallus corticolous, epiphloeodal, 

thin, shiny, smooth to minutely verruculose, whitish ashy gray, 5 cm broad; apothecia numerous, 

separate to loosely clustered, semiemergent, 0.5- 

0.9 mm in diameter, the thalline exciple noncarbonized, somewhat raised, the inner exciple distinct, 

entire, free; the pore open 0.2-0.3 mm wide; spores 

4-8, 10 X 50-55 pm, 10-12 loculate, I+ blue. 

CHEMISTRY.-NO lichen substances present. 

REMARKs.-Externally 0. turgidula is quite similar to Thelotrema lepadinum (Acharius) Acharius 

but the thallus is more developed and the apothecia1 pore less open. It seems to occur in more open, 

secondary habitats. I have another record from 

Japan: Hale 29652, Inohae Valley, Hyuga Province. 

SPECIMEN EXAMINED.-~ 1: 45832. 

54. Ocellularia uniseptata, new species 

FIGURE 8f 

DEscRIPTIorv-Thallus corticola, epiphloeodes, 

continuus, minute verruculosus, albidus, 6-8 cm 

latus; apothecia numerosa, regulariter dispersa, immersa, 0.1-0.2 diametro, ecolumellata; ostiolum rotundatum, ca 0.07 mm diametro, leviter annulatum, 

albo-cinctum; excipulum decoloratum; hymenium 

45-50 pm altum; sporae 8:nae, 3-4 X 6 pm, 2-, 

Taro 3-loculatae, I+ caerulescentes. 

CHExfIsTRY.-Psoromic acid and a trace of stictic 

acid. 

HoLoTYPE.-Secondary rain forest, W. H. Wheeler 

trail, Barro Colorado Island, Canal Zone, Panama, 

elevation about 120 m, M. E. Hale 38671, 3 April 

1973 (US). 

REMARKS.-This species seems to be close to 0. 

terebratula (see above), a canopy species with a 

shiny, nongranular thallus and larger spores. Ocellularia uniseptata is unusual, however, in having 

small, mostly uniseptate spores and in occurring 

lower on the trunks in low elevation rain forest. 

SPECIMENS EXAMIINED.-.b: 44901, Esslinger 4392. 

55. Ocellularia verrucosa, new combination 

FIGURE 8g 

Thelotrema uerrucosum Fee, 1824:94 [type collection: “America, ad corticem Cinchonae lancifoliae (Mutis.)” (G, lectotype; isolectotype in L)]. 

Asridium cinchonarum yar. uemtcost~m (Fee) Fee, 1837:91. 

DEscRIPTIos.-Thallus corticolous, epiphloeotlal, 

thin, minutely verrucose, dull greenish or brown- 
34 SMITHSONIAN CONTRIBUTIONS TO BOTASY 

ish mineral gray, 5-7 cm broad; apothecia isolated, 

emergent, about 1.0 mm in diameter, carbonized 

but without a columella, the pore round, to 0.5 

mm in diameter, black rimmed, the lightly pruinose disc visible; spores 8, 8 X 22-32 pm, 6-8 

loculate, It blue. 

CHEmsmY.-Protocetraric acid. 

REMARKS.-The type of this species in G is in 

poor condition, but the apothecia can be adequately characterized as semiemergent, ecolumellate, and with a black rimmed pore. The granularverrucose thallus surface was noted by Fee in his 

original description. Fee himself later reduced it to 

a variety of Ascidiiim cinchonarum Fee, but that 

species contains the P + “cinchonarum” unknoTvns, 

not protocetraric acid, and is columellate. Muller 

(1887a:6) followed Fee’s intepretation, and as far 

as I can determine no one has investigated the problem since then. The material I collected in Panama 

matches Fee’s lectotype very closely, as does one 

other from Colombia, all collected in the canop) 

or upper bole of trees in lowland rain forest. The 

closest relative may be 0. lepadinioides (Leighton) 

Zahlbruckner, which has larger apothecia and 

spores. 

SPECIMEhS EXA\IINED.-3: 43492; 4: 43133, 43480; 8: 38544. 

56. Ocellularia viridis, new species 

FICGRE 8h 

DEScRIPTIoN.-Thallus corticola, epiphloeodes, 

nitidus, continuus, albo-viridis, 4-7 cm latus; apothecia (Figure 3f) numerosa, dispersa, emergentia, 

aetate basi constricta, globoso-urceolata, 0.6-0.9 mm 

diametro, ecolumellata vel demum rudimentaliter 

actinoide-columellata; ostiolum rotundatum, 0.1- 

0.2 mm diametro, plus minusve depressum, alboannulatum; excipulum decoloratum vel fuliginascens; hymenium 60-70 Fm altum; sporae 8:nae, 

4-6 X 7-14 pm, 4-5 loculatae, It caerulescentes. 

CHEMISTRY.-AII unknown substance. 

HOLoTYPE.-hfature rain forest, El Llano-Carti 

Road at 16-20 km, Province of Panamh, Panama, 

elevation 300 m, M. E. Hale 38538, 6 April 1973 

REhrARm.-The urn-shaped apothecia are somewhat similar to those of 0. subemersa (see above), 

a West Indian species with the “cinchonarum” unknown, but the pore is not open. The unknown 

(US). 

substance in 0. uiridis is P- and forms a weak 

gray spot in the usual solvent systems. The species 

grows at the mid bole and lower trunk levels in 

rain forest at low to mid elevation. 

SPECIMENS EXAMISELX-~: 43472; 8: 38554, 38601 

57. Ocellularia xanthostroma 

FICCRE 8i 

Oceilztlnrin Annthostroma (Nylander) Zahlbruckner, 1923:603. 

..lscidiicf?i sn?ithostrom~cm Sylander, 1891:ZO [type collection: 

Cuhn. TI’,-ight 33 (H, Sylander herbarium 22412, lectotype)]. 

DEscRIPTIox.-Tha1lus corticolous, epiphloeodal, 

shiny, verruculose, whitish mineral gray, the medulla bright reddish orange, 5-7 cm broad; apothecia numerous, separate, strongly emergent, 1 .I1.5 mni in diameter, the amphithecium smooth to 

warty, easily breaking open to reveal the pigmented 

medulla, the exciple carbonized, a coluiiiella lacking, the pore round, 0.1-0.2 mm in diameter, narrowly white annulate, somewhat depressed; spores 

2-4, 12 >< 100 pm, about 18 loculate, I- blue. 

CHEMISTRY.-Endocrocin and a second unidentified pigment. 

REwms-The cortical layer on the thallus and 

apothecia breaks away easily to reveal the brilliant 

orange-yellow medulla. The chief pigment is endocrocin. It differs from 0. mordenii (see page 25) 

in chemistry (different pigments that are red rather 

than orange) and apparently in having more discrete emergent apothecia, although we must see 

more collections to comprehend the full range of 

chemical and morphological variation in this complex. The one specimen from Panama was collected 

on a liana along an open trail on Barro Colorado 

Island. 

SPECIMEX EX.~MINED.-~~: 44755 

Phaeotrema Muller Argoviensis 

This genus is recognized by the brown, transversely septate spores. It is the smallest genus in 

the family, comprising about 50 species worldwide 

but only 5 in Panama. Of the 28 specimens collected, 21 are P. aziberianum, which occurred at 7 

of the 12 collecting localities. No other species is 

represented by more than tlvo collections scattered 

among nine localities, generally at mid elevation. 
NUMBER 38 35 

The genus is rare both at the highest localities in chroodiscoid or lepadinioitl. P1iaeoti.cnin njiberiChiriqui and in the lowland forests. nnuni and P. meios/iertn~~m are botli pantropical 

All of the species except P. vzilcani have a reticu- and the reinaining species neotropical in distribulate columella or a tendency to form one. None are tion. 

Key to the Species of Phaeotrema 

1. A columella lacking .................................................................................... P. vulcani 

1. .4 columella present. 

2. The disc open, 1-5 mm wide, often elongated, actinoid-columellate. 

3. Thallus P+ yellow (psoromic acid). .................................................................. 1. P. auberinnum 

3. Thallus P4. Columella broad, the top clearly visible through the open pore, the pore 0.2-0.4 nim 

wide 4. P. meiospermum 

4. Columella narrow or weakly developed, the pore 0. h a raised annulate 

rim. ............................................................ ......................... 3. P. leiostomum 

(hypoprotocetraric acid). .......... ......................................... 2. P. glyphicum 

2. Disc less than 0.4 mm wide, columella mostly simple. 

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

1. Phaeotrema auberianum 

FIGURE 9a, b 

Thelotrema glyphidellzcm Stirton, 1876:161 [type collection: 

Phaeographis clzioizodisca Redinger. 1935372 [type collection: 

Amazons, Brazil, Trail, (BM, lectotype)]. 

Santos, S5o Paulo, Maline 3559 (S, lectotype)]. 

Phaeotrenza auberianum (Montagne) Muller Argoviensis, DEScRIPTIoN.-Thallus corticolous, epiphloeodal, 

smooth to verruculose, shiny, fissured with age, 

common, irregularly dispersed, solitary, emergent 

1886:311. 

Thelotrema auberianuin Montagne zn Sagra, 1842:163 [type 

collection: Cuba, Auber (H, Sylander herbarium 22615, greenish mineral gray, 6-12 cm broad; apothecia 

lectotype; isolectotype in L) (Figure le)]. 

~ 

FIGURE 9.--Species of Phaeotrema: a, P. auberianum (Hale 43487); b, P. aubeizuiiiim with elongate apothecia (Hale 45581); c, P. glyphicum (Nakanishz 51); d, P. leiostomum (Hale 38600): 

e, P. meiospermum (Hale 46061); j, P. vulcani (Hale 43534) (X 10). 
SMITHSONIAN CONTRIBUTIONS TO ROTANY 

to strongly emergent at maturity, 1.0-2.5 mm wide, 

the tlialline margin thick, carbonized, tlie disc open, 

conspicuously actinoid-columellate, the columella 

carbonized; surface of the disc becoming heavy 

white pruinose; spores 8, 4-8 X 9-14 ,“m, 4 loculate. 

CHEMIsTRY.-PSOrOmiC acid. 

REsIARKS.-This is a conspicuous lichen, easily 

recognized and widespread in Panama and usually 

collected on lower trunks of trees in open forest. 

The actinoid disc is very similar to that of Ocellulaiin bel-heleynna (Alontagne) Muller Argoviensis. 

Range in size and shape of the disc is very great, 

Redinger’s type of Plrneogrnpliis cliionodiscn and 

my numbers 38564 and 38581 representing an estreme population Tvith long lirelliform apothecia 

up to 8 mm long (Figure 9b). It is a pantropical 

species. 

SPECI\IEXS Exarls~~.-3a: 44793, 44897, 44961; 4: 43487; 

6: 49525, $3543, 43,544; 8: 38,525, 38.530, 38579: 38.582. 38604: 

4257. 4413. 

9: 8473; 11: 44819, 44834, 44880. 44898. 44992, E?S/iJlgeJ2. Phaeotrema glyphicum 

FIGI.RE 9c 

Piioeotrenia g!sph;furn (Sylander) Zahlbruckner, 1923:607. 

Thrlorre~na gk.~ihicir?n Sylander. 1863a:453 [t)pe collection: 

\.illeta, Colombia. Lindig 826 (FH-Tuck, 1ectot)pe; isolectotypes in BM, PC, IV) (Figuic It)]. 

DEscRIPTIos.-Thallus corticolous, epiphloeodal, 

thick, smooth to verruculose, white-spotted, ivory 

to pale tan, 6-10 cm broad; apothecia numerous, 

more or less clustered, orbicular to lirelliform in 

outline, immersed to semi-emergent, up to 2 mm 

broad, the thalline margin becoming tvhite pruinose, forming a low rim, the disc open to somewhat 

closed, actinoid columellate, the columella carbonized; spores 8, 4-8 X 12-16 pm, 4 loculate. 

CHEa~IsTRY.-Lichexantlione witli or without 

hypoprotocetraric and 4-0-tlemetliylnotatic acids. 

I<EMARKs.-This rare species was previously 

known only from Brazil (Redinger, 1936:72) and 

the type collection in Colombia. The large irregularly shaped apothecia have a strongly developed 

actinoid disc. Hale 38365 contains both lichexanthone, the only coniponent in the type collection, 

and hypoprotocetraric and 4-0-demethylnotatic 

acids. The normal habitat seems to be the mid and 

upper trunk of mature trees in rain forest at higher 

elevations. 

SI’ECI\IENS EXA>fISED.-l: ~\’akanishi 51; la: 448ii; 9: 38465. 

3. Phaeotrema leiostomum 

FIGLRE 9d 

Phneolr PJIIO iciostornt~rri (Tuckerman) Zahlbruckner, 1923:608. 

Tiieloti.eJrin leirJstoiJi IIJII Tuckerman, 1862:407 [type collectiou: Cuba. W~igizt 149 (FH-Tuck, lectotype)]. 

DEscRIPTIoN.-Thallus corticolous, epiphloeodal, 

continuous, smooth and shiny, about 5 cm broad; 

apotliecia slightly emergent, 0.3-0.6 mm in diameter, weakly carbonized and uith a weakly developed 

columella, tlie pore round, 0.1-0.3 mm in diameter, 

white annulate; spores 8, 8 X 15 pm, 4 loculate. 

CHEal1sTRY.-Hyproprotocetraric acid. 

REarARKs.-The Panamanian material, which 

lacks 4-0-demethylnotatic acid, is identical to the 

population in Dominica (Hale, 1974a:29) but with 

less variation in apothecial outline. It also has a 

similar habitat: mid bole on trees in rain forest at 

mid elevation. 

SPECIMEKS EX,k\IINED.-8: 38600. 

ILLCsTRATIOSS.-Ha1e, 1974a:30, figs. 14b, C 

4. Phaeotrema meiospermum 

FIGURE 

Plineotrerrin rri( io~fierrnunz (Sylaiider) Jiiiller Argoviensis, 

1886:311. 

lection: Cuba, Wright 136 (FH-Tuck, lectotype; isolectotypes in BM, (;, H. L, PC, CPS)]. 

Thelotreinn ~iiiaeospe~moides \.ainio, 1921: 185 [type collectioii: Pro!. Rizal. Philippines, Ramos and EdnF~o 29456 

(TUR. lectotvpe: isolectotypes iii BM, FH, US, I+’)]. 

Phneotrenin phaeospermoides (\.ainio) Zahlbruckner, 1923: 

609. 

T/ie/Ott’?,l?/~ Jl~?~~~~~f’l~lJli~l~f Xylander, 1863b3333 [type COlDEscRIPTIoN.-Thallus corticolous, epiphloeodal, 

smooth with a grainy surface, shiny, ivory colored, 

6-12 cm broad; apothecia not common, immersed 

to semiemergent, round to elongate in outline, 

0.3-0.5 mm in diameter, the thalline margin carbonized, a columella present, very broad, to 300 

pi in diameter, the pore open ivith the black columella surface clearly visible, round to irregular, 

0.2-0.4 mm ivide; spores 8, 10 X 16-18 pm, 4 

loculate. 
NUMBER 38 37 

CmmsTRY.-No substances present. 

RE3iARI;S.-The apothecia are quite variable in 

size and may have a single broad columella or a 

main columella ivi th secondary actinoid growths. 

I have identified the species from Florida, Cuba, 

Sri Lanka, Java, and Sarawak. In Panama it occurred on the lower trunk of trees in the rain forest 

at mid elevation. 

SPECIMESS EXA~II~EII.-~ 1 : 46033, 46061 

5. Phaeotrema vulcani, new species 

E’IGLRE 9f 

DEscRIPTIos.-Thallus corticola, epiphloeodes, 

continuus vel demum rimosus, glauco-viridis, 10 

cm latus; apothecia (Figure 10) numerosa, immersa 

vel semi-emergentia, solitaria vel 2-3 confluentia, 

0.5-0.7 mm in diametro, ecolumellata; ostiolum 

rotundatum vel elongatum, 0.2-0.3 mm latum, annu10 elevato, albocincto; excipulum decoloratum; 

hymenium 60-90 pm altum; sporae 8:nae, 4-5 X 

10-12 pm, 2-3 loculatae, I-. 

CHEIIISTRY.--NO substances present. 

HoLoTYPE.-secondary rain forest, La Mesa, 

north of El Valle, Province of Cocle, Panama, elevation 750 m, M. E. Hale 43534, 13 February 1974 

REJiARKS.-The raised, white, round to elongated ostiole is similar to that of Pheotrema leiostomum (Tuckerman) Zahlbruckner but much 

larger. There are no comparable species in the 

(US). 

FIGURE 10-Section of apothecium of Phneotrema uzclcani 

(Hale 43534) (X 125). 

genus. It is confined to canopy branches in the rain 

forest at mid elevations. 

SPECIhIEX EXAhlINED.-ll: 44763 

Thelotrema Acharius 

This is the second largest genus in the family, 

characterized by colorless muriform spores. On the 

world level there are about 140 species. The 23 

species in Panama were collected frequently at all 

the localities, excepting the Summit Botanical Garden and the pasture above Alto Piedra. About 120 

collections were made, 20y0 of the total for the 

family compared with 15y0 in Dominica. 

The floristic composition of the genus is quite 

different from that in Dominica. For example, 7 

of the 10 commonest species, T. erumpens (4 collections), T. occlusum (4), T. maximum (12), T. 

myrioporoides (6), T. stylothecium (6), T. subpraestans (17), and T. wrightii (5) have not yet 

been collected in Dominica. The three remaining 

species of the 10 most common, T. conforme (8), 

T. glaucopallens (31), and T. praestans (3) are 

among the 10 most common in Dominica. Four 

of the 23 Thelotremata are described as new from 

Panama, as compared with 4 of 14 in Dominica. 

The morphological variation of the apothecia 

is similar to that in Dominica. One species, T. 

hiatum, is clearly chroodiscoid, while T. conveniens, 

T. lepadinuni, T. myriocarpzim, and T. occlusum 

are lepadinoid. Thelotiema clandestinum, T. glaucopallens, T. novsticticum, and T. pycnoporellum 

have flush myriotremoid apothecia. Only T. leucomelaenum, T. praestans, T. stylothecium, and T. 

subpraestans are strongly columellate, and T. 

wrightii.has a weakly actinoid disc. 

The lowland species, as represented by the collections made on Barro Colorado Island and nearby 

Gamboa, include T. adjectum, T. cryptotrema, T. 

glaucopallens, T. hiatum, T. myrioporoides, and 

T. subpraestans. The strictly high elevation species include T. concretum, T. conveniens, T. erumpens, T. Eepadinum, T. leucomelaenum, T. occlusum, T. stylothecium, and T. tenue. Thelotrema 

is best developed at higher elevations, there being 

a total of eight species among 22 collections made 

at Chiriqui. Only five species make up the 39 collections made at Barro Colorado Island. 
38 SMITHSOXIAX CONTRIBUTIONS TO BOTANY 

FIGURE 11.-Cross sections of apothecia of Thelotrema and Ixptotrenrn: a, T. hiatnnr (Hnlr 

43458) (X 125); 0, T. niaxijniiui (Hale 43580) (X 100); c, L. chiriqiiie?ise (Hale 413035) (X 100); 

d, L. panamense (Hale 43584) (X 125). 

The following species are restricted to the canopy: T. clandestinum, T. cryptotrema, T. myrioporoides, T. praestans, and T. subpraestans. The remaining species grow lower on the trunk but only 

T. glaucopallens is strongly restricted to the base 

level. 

The phytogeography of Thelotrema is less well 

known than for Ocellularia, because feTver collections have been made. A rather large group of 

species were previously known only from South 

.4inerica, illustrating perhaps the more continental 

nature of the Panamanian flora as compared with 

that of Dominica: T. concretum, T. conveniens, 

T. cryptotrema, T. occlirsum, and T. stylothecizim. 

Caribbean species include T. adjectum, T. myriopol-oides, T. norsticticum, T. pycnoporellum, T. 

tentie, and T. wl-ightiz, most of which were previously known only from Cuba. Thelotrema auraturn, 

T. clandestinitm, T. el-urnpens, T. glaucopallens, 

and T. lepadinum form a pantropical element. 

Key to the Species of Thelotrema 

1. Apothecial disc open, the thalline rim erect or recurved. 

2. Apothecia large, 1-3 mm in diameter, the rim coarse. .......................................... 23. T. wriphtii 

2. Apothecia less than 1 mm wide, the rim thin. 10. T. hiaturn 

3. A distinct black central column present (weakly developed in T. confo~-me and T. tenue). 

5. Stictic acid present .......................................................................................... 21. T. subpraestnns 

1. .4pothecia with a distinct pore, the thalline rim not erect. 

4.Apothecia large and emergent, 2-3 mm in diameter; spores 200-300 ,urn long. 

5. “Praestans” unknown present. .......................................................................... 18. T. praestans 
NUMBER 38 39 

4. Apothecia 1 mm or less in diameter; spores less than 5Opm. 

6. Apothecia to 1 mni in diameter, emergent; psoromic acid present (P+ yellow). ........... 

5. T. conforme 

6. Apothecia 0.2-0.8 mm in diameter, nearly flush; stictic acid present (P+ orange) or 

7. Stictic acid present. .................................................................................... 20, T. stylothecium 

7. No lichen substances present. 

8. Pore small, 0.1 mn: wide; columella thin and tall; spores less than 20 pi. ........... 

..................................................................................................................................... 22. T. tenue 

8. Pore gaping 0.2-0.4 min wide; columella thick and broad; spores more than 30 

pm. ...................................................................................................... 12. T. Zeucotnelaenuni 

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

no substances present. 

3. Central columella lacking. 

9. Apothecia large and emergent, 1-4 mi11 in diameter. 

10. Disc covered with a yellow-orange pruina ..................................................... 2. T. autatum 

10. Disc unpigmented. 

11. Pore small, about 0.1 mm wide. ................................................................ 13. T. maximion 

11. Pore open, 0.2-0.4 mm wide, the proper exciple visible. .............. 11. T. lepadintcm 

9. Apothecia smaller, 0.2-0.8 mm in diameter, flush to semiemergent. 

12. Spores large and densely muriform, more than 50 pn: long. 

13. Thallus P+ yellow (psoromic acid). .................................................. 5. T. cryptotremn 

13. Thalluus P-. 

14. Spores less than 100 pm long. ................................................................ 1. T. adjectum 

14. Spores more than 100pm long. 

15. Pore distinct, the proper exciple forming a small inner pore. .......................... 

6. T. conveniens 

15. Pore broader, 0.2-0.4 mm wide, more or less filled with excipular material. 

.............................................................................................................. 17. T. occlusum 

16. Apothecia to 0.5 mm in diameter, slightly emergent and often apically carbon17. Apothecia with an open pore, to 0.2 mm wide; uncarbonized. .... 4. T. concretum 

17. Apothecia with a small pore, less than 0.1 mm wide; apically carbotiized 

18. Stictic acid present. ............................................................................ 8. T. erumpens 

18. No lichen substances present. ................ 

16. Apothecia 0.1-0.3 mm in diameter, flush, 11 

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

12. Spores smaller, less than 40 pm long. 

ized. 

in T pycnoporellum). 

19. Stictic acid present. 

19. Stictic acid lacking. 

21. Thallus P+ yellow or orange. 

22. Psoromic acid present. .......................................................... 3. T. clandestinicm 

..................... 16. T. norsticticum 

,.,15. T. myrioporoides 

.19. T. lrycnoporellum 

22. Norstictic acid present. ....... 

21. Thallus P-. 

23. Hypoprotocetraric acid present. 

23. No lichen substances present. ... 

1. Thelotrema adjectum 

FICLRE 120 

Thelotrer?~~ adjecticin Sylander, 1866:290 [type collection: 

Cuba, Wright ser. 2, 82 (H, Sylantler herbarium 22572, 

lectotype; isolectotype ii: UPS) (Figure 1 b)]. 

DESCRIPTION.-T~la~~L~S corticolous, epiphloeodal, 

the surface grainy to minutely verruculose, continuous, pale tannish or whitish gray, 6-12 cm broad; 

apothecia numerous, solitary, nearly immersed to 

semiemergent, 0.6-1.0 mm in diameter, the thalline 

rim coarse, becoming semierect, jagged, the inner 

exciple well developed, thickened, filling the pore 

area and forming a double pore, flesh colored, the 

pore irregular, 0.2-0.3 mm wide; spores 8, 12-15 X 

36-50 pin, 1-2 X 9-12 loculate, I-. 

CHEMISTRY.-XO substances present. 

REMARKS.-This inconspicuous species is char- 
40 SMITHSOSIAN CONTRIBUTIOKS TO BOTASY 

FIGURE 12.--Spccics of The/o/~c,rnn: (1, T. ndpfur71 (Hnle 44882); 0, T. anrcztu~ti (Hole 43544); 

c, T. clatrdestinuin (Hnic 44942); tl, 7'. miirr-etuiri (Hole 43549); e, T. co)ifor?rie (Hale 38463); 

f, T. conniTvtis (Hale 38741); g, 7'. rryptotreiria (Halr 43483); h, T. C~11~Il~~ll5 (Hale 38752j; i, 

T. ghcopo!/rns (Hnlc 44873); j, T. hiaturn (Hale 43458); k, T. /epndi?ltcr~~ (Hale 44902); /, 

T. leucornelaeiiuin (Hale 38765) (X 10). 
NUMBER 38 41 

acterized by the semiemergent apothecia and the 

filled disc. The type collection from Cuba has 

slightly larger spores, 57-78 pm long. In Panama 

T. adjectum occurs at the base of trees in the low 

elevation rain forest of Barro Colorado Island but 

at no other sites. It is also known in subtropical 

United States (Florida and Louisiana). 

SPECIMENS EXAlIISED.-3a: 44759, 44864, 44882. 

2. Thelotrema auratum 

FIGURE 12b 

Thelotrema auratum Tuckerman, 1862:408 [type collection: 

Cuba, Wright 133 (FH-Tuck, lectotype; isolectotypes in 

BM, G, L, PC, UPS) (Figure 101. 

Thelotrema intei.r~allatum Sylnnder, 1868:36 [type collection: 

Sew Caledonia, Maj ie (H, Sylander herbarium 22458, 

lectotype; isolectotypes in G, PC) (Figure 2a)l. 

Thelotrema oczilntztnz l’ainio, 1921:179 [type collection: POlillo, Philippines, Robinson 9082 (TUR, l.ainio herbarium 

26828, lectotype) (Figure 2i)I. 

Thelotrema porphyrodisczim Zahlbruckner, 1908:245 [type 

collection: Sawai, Samoa, Rechinger 2926 (W, lectotype) 

(Figure 21)]. 

Thelotrema citrinodiscum Redinger, 1936:78 [type collection: 

Corcovado, Rio de Janeiro, Brazil, hfalme 88 (S, lectotype) 

(Figure lj)]. 

DEscRIPTIoh’.-Thallus corticolous, epiphloeodal, 

shiny, continuous, pale brownish mineral gray, 

8-1 1 cm wide; apothecia solitary, dispersed, emergent, 0.9-1.2 mm in diameter, the exciple carbonized, columella lacking, the disc yellow-pruinose, 

the pore irregular, black rimmed, to 0.5 mm wide; 

spores 8, 12 X 24-26 pm, 0-2 X 6-7 loculate, I+ 

bluish. 

CHEMIsTRY.-Protocetraric acid and an unidentified epihymenial pigment. 

REMARKS.-This species is easily recognized by 

the large, emergent ascidioid apothecia with a yellowish or orange-red pruinose disc. The chromatograms of T. aziratum and all the types of the synonyms listed were run together and proved to be 

identical. Vainio (1921: 180) had actually indicated 

that his T. oculatzim was very closely related to 

T. intervallaturn Nylander. Redinger (1936:SO) 

compares T. citrinodiscum with T. porphyrodiscum Zahlbruckner, which he differentiates by a 

deep purplish disc. The disc pigmentation is in 

fact quite variable but the same pigments seem 

to be involved. The species is rare in Panama and 

occurred on upper trunks of trees in rain forest. 

SPECIMEXS EXAMINED.-~: 43544,43574 

3. Thelotrema clandestinum 

FICCRE 12c 

Thelotrema clandestinum Fee, 1837:90 [type collection: “In 

America ad corticem Cinchonae lancifoliae Mutis.” (G, 

lectotype) (Figure lk)]. 

Thelotrema subcnesium Nylander, 1869:120. 

Thelotiema concretum Fke var. subcaesium (Nylander) Red- 

[For full citations see Hale (1974a:32).] 

inger, 1936:96. 

DESCRIPTIoN.-ThallUS corticolous, epiphloeodal, 

thick and continuous, whitish to pale greenish gray, 

up to 20 cm broad; apothecia numerous, separate, 

immersed, 0.15-0.3 mm in diameter, noncarbonized and without a free inner exciple and columella, the pore round, 0.07-0.1 mm in diameter, 

usually more or less distinctly white annulate; 

spores 8, 6-7 X 14-22 lm, 0-1 X 4-5 loculate, I+ 

blue. 

CHEMrsmY.-Psoromic acid with or without norpsoromic acid. 

RmfARKs.-This canopy species has a thick continuous thallus similar to that of Ocellularia terebratzila (Nylander) Muller Argoviensis, and as I 

had found in Dominica, these species must be carefully separated by examining spores. I have identified T. clandestinum from Florida, Guatemala, the 

Caribbean region, Venezuela, Brazil, South India, 

the Philippines, and the Solomon Islands. 

SPECIlIENS EXAZZINED.-2: 43597; 1: 44942, 44959. 

ILLUSTRATIONS.-Half?, 1974a:32, figs. 14g, i. 

4. Thelotrema concretum 

FIGURE 12d 

The/otrema concreturrz Fee, 1837:90 [type collection: “In 

America meridionali ad cortices Cinchonae lancifoliae 

Mutis.” (G, lectotype; isolectotvpe in H-Nyl) (Figure ll)]. 

Thelotrema microporoides Nylander, 1863:326 [type collection: Colombia, Lindig 2622 (H, lectotype; isolectotypes in 

BM, FH, G, PC, S, UPS)]. 

DEscRIPTIoN.-Thalhs corticolous, epiphloeodal, 

smooth and continuous, shiny, pale greenish to 

tannish mineral gray, 10-12 cm broad; apothecia 

numerous, solitary or becoming clumped, immersed 

to semiemergent, 0.2-0.4 mm in diameter, noncar- 
42 SYITHSONIAN CONTRIBUTIONS TO BOT.INY 

bonized and lacking a columella, the inner exciple 

sometimes persistent, the low thalline rim raised 

and erect at maturity, the pore open, to 0.2 mm 

wide, the pale flesh-colored disc visible; spores 8, 

7 X 12-14 pm, 1 X 4-5 loculate, I+ blue. 

cHEMISTRY.-PSOrOmiC acid with or without norpsoromic acid. 

REMARKS.-Thelotrema concretzim has small, 

crowded, confluent semiemergent apothecia as so 

clearly illustrated by Fee (1837, pi. 37). It is part 

of a complex of similar species with psoromic acid: 

T. clandestinum Fee, which has entirely immersed 

apothecia, T. insigne Zahlbruckner (Redinger, 

1936:96), which has long thin isidia, and T. harfii 

%fuller Argoviensis, which has sparse, thick isidia. 

It also resembles the rare T. halei Tuckerman, 

which has a distinct inner exciple and numerous 

crystal inclusions in the thallus. Redinger (1936:96) 

also compares it with T. Zeucohymenilim Zahlbruckner, a hypoprotocetraric acid-containing species with entirely immersed apothecia, which I 

place in synonymy under T. myrioporoides Aliiller 

Argoviensis (see page 45). In addition, while Redinger regards T. szibcaesium Nylander as a variety 

of T. concretum, I feel it is closer to T. clandestinum (see above). Further studies of this species 

group and other psoromic acid-containing species 

such as T. quitoensis Fee from Ecuador should be 

made as more specimens become available for study. 

Incidentally, my spore measurements (taken in part 

from the lectotype in G) as well as those by Redinger (1936:95) are smaller than hfiiller’s (1887a: 

Thelotrema concretum is not a commonly collected species but is apparently pantropical. In 

Panama it occurs at mid elevations on mature trees, 

generally on the lower trunks. I have seen specimens from the cloud forests of Venezuela (about 

2200 m elevation) and from Brazil and Sabah. 

11). 

SPECIMEXS EXAMI~ED.-~: 38774: 2: 43949 

5. Thelotrema conforme 

FIGLRE 12e 

Thelotrema confof me Fee, 1837:89 [type collection: “In 

America meridionali ad cortices Cinchona~um” (G, lectoThelotrema consanguineum Muller Argoviensis, 1887b:398 

type)]. 

[type collection: Brazil, Puzggari 477 (G, lectotype)]. 

DEScRIPTIoN.-Thall~is corticolous, epiphloeodal, 

smooth and shiny, whitish mineral gray, up to 10 

cm broad; apothecia fairly numerous, solitary, 

emergent, apically carbonized, 0.8-0.1 min in diameter, the columella distinct to weakly developed, 

the pore round, 0.1-0.5 mni in diameter, often 

white annulate; spores 4, 10-16 X 24-50 pm, 2-4 X 

7-10 loculate, I+ blue. 

CHEMISTRY.-PSOrOmiC and norpsoromic acid. 

REhfARI;s.-This is another classic Fee species. 

It has rather large, more or less emergent apothecia 

with an annulate pore and intermediate sized 

spores. As Redinger notes (1936: 88), FCe describes 

the spores a5 “tetrasporis,” apparently referring to 

a 4-loculate spore. The lectotype material in G, 

however, examined by Muller (1887a: 11) and me, 

clearly has larger spores and more than four locules. 4 columella was weakly developed to absent 

in the specimens from Rio Calevobora in Panama, 

but variability in columella formation is not unusual in Thelotrema. The species is usually collected at mid bole or in the canopy and also occurs 

in Dominica, Trinidad, and Brazil (Redinger, 

1936:87). 

~PFCXMI \\ EX.AMINED.-~: 38.557. 38.399, 38608; 9: 38463: 11: 

44792, 44824, 44830, 46017. 

6. Thelotrema conueniens 

I.’IG~,IW 121 

7‘/2dolJC)f7n NJJlUC?lie?fS Sy-laiider, 1873:168 [type collectioll: 

Tequentlama, Colombia, Lifidig in 1863 (H, Nylander herbarium 22496, lectotype) (Figure 1 m)]. 

DEsCRIPTrox.-Thallus corticolous, epiphloeodal, 

verrucose to warty, shiny, pale tan, 5-7 cm broad; 

apothecia not numerous, solitary, emergent at maturity, the thalline margin warty, noncarbonized, 

0.6-1.0 mm in diameter, a columella lacking, the 

inner exciple well developed, entire, leaving a small 

inner pore, the main pore round, 0.2-0.3 mm in 

diameter; spores 1, 30 X 150 ILm, densely muriform, 

I + blue. 

CHEMISTRY.-NO substances present. 

REMARKs.-This species was first described in 

1873 froni the cloud forests near Bogoti and has 

not, as far as I know, been collected elsewhere. It 

is distinguished by the small lepadinoicl apothecia 

and the large spores. Thelotrema rugatulum Nylander, described at the same time by Nylander 
NUMBER 38 43 

from the Andaman Islands, has smaller apothecia, 

a less distinct, open pore, and smaller spores (75-85 

pn long). It could otherwise be confused with T. 

adjectum Nylander (see above), a lowland species 

with smaller spores and a filled pore. Thelotrema 

conueniens was collected only at Chiriqui on the 

mid bole of large oak trees at about 2000 m elevation. 

SPECIMENS EXAMINED.-~: 38741; la: Esslinger 4483 

7. Thelotrema cryptotrema 

FIGURE 12g 

Thelotrema cryptotrema Nylander, 1867:318 [type collection: 

French Guiana, Melinou (H, Nylander herbarium 22490, 

lectotype; isolectotype in PC) (Figure lo)]. 

Thelotrema secoligella Muller Argoi iensis, 1895a:31 [type collection: French Guiana, Leprieur 722 (G, lectotype) (Figure 

Thelotrema annulaturn Muller Argoviensis, 1895c:453 [type 

collection: Rio de Janeiro, Brazil, Glaziou 5474 (C, lectotype; isolectotype in BM) (Figure Id)]. 

DESCRIPTIoN.-Thallus corticolous, epiphloeodal, 

smooth, continuous to broadly cracked, whitish 

mineral gray, 8-12 cm broad; apothecia not common, immersed to semiemergent, 0.4-0.6 mm in 

diameter, noncarbonized and without a columella, 

the inner exciple more or less free, apically pruinose, filling much of the pore area, the pore round, 

0.05-0.1 mm in diameter; spores 1-4, 15-30 X 55-80 

pm, densely muriform, I-. 

2P)l. 

CHEMISTRY.-PSOrOmic acid. 

REMARKS.-The pore area is white and more or 

or less closed as in a Phlyctis. No other psoromic 

acid-containing species have this character. The 

descriptions of Muller’s two synonymous species are 

essentially identical; he compares T. annulaturn 

with “Th. disciforme Leighton,” a quite different 

species of Leptotrema (see p. 53). The species had 

not previously been reported outside of the typelocalities in French Guyana and Brazil. In Panama 

it occurred in tree tops at lower elevations (100- 

350 m). 

SPECIMENS EXAMINED.-3: 38692; 6: 43483. 

8. Thelotrema erumpem 

FIGURE 12h 

Thelotrema erumpens Magnusson, 1955:279 [type collection: 

Mauna Kea, Hawaii, Faurie 1909 (VPS, lectotype) (Figure 

Thelotrema leucornelaenuin Far. cathoinalizans Nylander, 

1863a:452 [type collection: Tequendama, Colombia, Lindig 

(H, Nylander herbarium 22581, lectotype; isolectotypes in 

BM, FH (as no. 857)]. 

DEScRIPTIoN.-Tha~~us corticolous, epiphloeodal, 

dull, smooth to verruculose, finely rimose, whitish 

mineral gray, 8-10 cm broad; apothecia numerous, 

dispersed to confluent, nearly flush to semiemergent, 0.5-0.7 mm in diameter, the exciple carbonized, columella lacking, the pore round, 0.05-0.1 

mm in diameter, becoming blackened; spores 4-8, 

12-15 X 36-50 pm, 1-3 X 6-8 loculate, I-. 

CHEMIsmY.--Stictic acid, constictic acid, and the 

high “quintaria” unknown. 

REMARKS.-The pore of this species is usually 

heavily carbonized and the surrounding area blackened. Magnusson (1955:279) pointed out its superficial resemblance to Ocellularia pyrenuloides Magnusson, which also contains stictic acid and is 

known from Hawaii and Dominica (Hale 1974a:25). 

I collected it in Panama on palm trees and evergreen hardwoods at 2200 m elevation in cloud 

forest. 

lS)J 

SPECIMESS EXASIISED.-~ : 38752; la: 44814, 44955, 45839. 

9. Thelotrema glaucopallens 

FIGURE 12i 

Thelotrema glaucopallens Nylander, 1863b:327 [type collection: Cuba, lliright 28 (FH-Tuck, lectotype; isolectotypes in 

L, M, UPS)]. 

Pyrenula clandestina Acharius, 1814:lO. 

Thelotrema laevigans Nylander, 1863a:451 [type collection: 

Bogota, Colombia, Lindig 893 (BM, lectotype; isolectotypes 

in FH, G, PC, U’)]. 

Thelotrema laevigans Nylander var, avertens Nylander, 1867: 

318. 

Thelotrema pechueli Muller Argoviensis, 1880:34. 

Ocellularia clandestina (Acharius) Muller Argoviensis 1887a: 

Thelotrema homopastoides l’ainio, 1896:207. 

Thelotrema buttianurn Vainio, 1921:183. 

[For full synonymy see Hale (1974a:35).] 

7. 

DEscRIPTIolu.-Thallus corticolous, epiphloeodal, 

very thin and shiny, greenish gray, up to 20 cm 

broad; apothecia usually numerous, separate, immersed to slightly emergent, 0.2-0.4 mm in diameter, noncarbonized and without a columella, the 
44 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

inner exciple evanescent, the pore round, 0.05-0.1 

mm in diameter; spores 8, 7-10 X 15-25 pm, 1-3 X 

6-8 loculate, I-. 

CHEZIISTRY.-stiCtiC and constictic acids. 

REMARKS.--AS I had discovered in Dominica, 

T. glaucopallens is one of the commonest base 

level species in the rain forests at lower elevations, 

Ivliere it competes with pyrenocarpous lichens. The 

species was very common in Dominica and I have 

identified collections from Guatemala, Guyana, 

Venezuela, Trinidad, Tanzania, Ivory Coast, and 

the Union of South Africa in addition to localities 

in the Philippines. It was the most common lichen 

in the rain forests of Barro Colorado Island in 

Panama, occurring on prop roots, buttresses, and 

lower trunks. 

SPLCI>IENS ExaMIsED.-3: 38522. 38624, 38646, 38656, 38662, 

38667, 38669, 38670. 38680, 38681, 38691: 3a: 447.58. 44766, 

44774, 4481 1, 44847, 44873, 41895, 44943, 460i9, Esslitzger 

4338, 4340, 4404: 4: 43342, 43447, 43465; 6: 43491, 43527, 

43555; 7: 43607; 8: 38606. 

10. Thelotrema hiatum, new species 

FICLRE 121 

D~sc~~~~~os.-Thallus corticola, epiphloeodes, 

tenuis, albidus, 8-12 cm latus; apothecia chroodiscoida (Figure 1 la), 0.4-0.7 mm diametro, margine 

thallino crasso, erecto; excipulum interius decoloratum, discretum vel evanescens, disco nigro vel 

levi ter albo-pruinoso; hymenium 180-200 p,m altum; sporae 4-8:nae, 6 X 12 pin, longitudine 0-1 

loculatae, transversim 2-3 loculatae, I+caerulescentes. 

CHEMISTRY.-NO substances present. 

HOLoTYPE.-on a felled Miconia, along h'aval 

Pipeline Road about 9 km NTf of Gamboa, Canal 

Zone, Panama, elevation 150 m, M. E. Hale 43458, 

11 February 1974 (US). 

RE>iARKS.-The only apparent relative of this 

chroodiscoid species is T. lepl-oca1-pirm Tuckerman, 

known from southern United States, which has 

large spores (30-80 long). The thallus is opaque 

and crumbly, similar to the whitish bark excrescences characteristic of Aliconia, a small melanostomaceous tree. It was surprising to find a Thelotrema on this weedy roadside tree at low elevation. 

1 1. Thelotrema lepadinum 

FIGURE 12k 

Thelotreinn [epadinum (Acharius) Acharius, 1803:132. 

Lichen lepndinzts Acharius, 1798:30 [type collection: Sweden, 

Arhnrizis (H-Ach., lectotvpe: isolectotypes in BXI, UPS)]. 

DEscRIPTIoh'.-Thallus corticolous, epiphloeodal 

but thin and in part evanescent, pale tannish, 2-5 

cm broad; apothecia dispersed, emergent, 0.5-1.0 

mm in diameter, noncarbonized, lacking a columella, the proper exciple well developed, free, splitting, the pore open, round, 0.2-0.4 mm in diameter; spores 4-8, 20-26 X 80 pm, with numerous 

locules, I- or bluish. 

CHEMISTRY.--NO substances present. 

RExfmKs.-This pantemperate lichen, by far the 

most commonly collected member of Thelotl-emu, 

occurs as a montane species in the tropics, usually 

in cloud forest. '4s expected, it was found only at 

Volcdn Chiriqui in Panama. 

SPECIMEX EXAMINED.-~: 38777; la: 44902. 45968 

12. Thelotrema leucomelaenurn 

FIGURE 121 

Thelotrema leucomelaenurn Nylander, 1863b:329 [type collection: Colombia, Lindig 2577 (H, Sylander herbarium 

22576, lectotype; isolectotypes in BJI, FH, G, PC, UPS)]. 

Theiotwma paupei-ius Sylander, 1863:329. 

Thelotreinn lezicomelneniain \ar. elerxtum Vainio, 1915:137. 

[For full citations see Hale (1974a:36).] 

DEscRIPTIox.-Thallus corticolous, mostly hypophloeodal, whitish, up to 8 cm broad; apothecia 

immersed to semiemergent, 0.6-0.9 mm in diameter, 

heavily carbonized with a broad columella about 

0.3 mm wide, the pore round, 0.2-0.3 mm in diameter, partially filled by the white-pruinose tip 

of the columella; spores 8, 12-15 X24-35 

2-3 X 4-6 loculate, I-. 

CHEMISTRY.--;~O substances present. 

REMARKS.-The columella is broad and flat, 

leaving a narrow ringshaped hymenial area (Redinger, 1936:92). Ifhile this was a rather common 

species at high elevations in Dominica, it is largely 

replaced by externally similar but chemically different T. stylothecium in Panama (see page 48). I 

have seen additional material of the species from 

Jamaica, Colombia, Venezuela, and Brazil. 

SPECIMEKS EXAMINED.-I: 38765; la: 44950. 
NUMBER 38 45 

13. Thelotrema maximum, new species 

FIGURE 13a 

DESCRIPTION.-ThallUS corticola, epiphloeodes, 

crassus, nitidus, continuus, viridi-cinereus, 6-10 

cm latus; apothecia (Figure 11 b) emergentia, basi 

non constricta, 1.5-2.0 mm diametro, ecolumellata 

vel rudimentaliter columellata; ostiolum rotundatum, 0.1 mm diametro, plus minusve annulatum, 

albo-cinctum; excipulum apice fuligineum; hymenium 200-220 pm altum; sporae I-2:nae, 30-40 X 

100-160 pm, dense murales, I+ caerulescentes. 

CHEMIsTRY.-Psoroniic and norpsoromic acids. 

HOLOTYPE.-TO~ of palm tree in secondary rain 

forest, La Mesa, north of El Valle, Province of 

Cocle, Panama, elevation 750 m, M. E. Hale 43580, 

13 February 1974 (US). 

REMARKS.-This is clearly a member of the T. 

conforme Fee complex. It is distinctive in having 

very large apothecia not at all basally constricted. 

Of the related species with psoromic acid, T. conform Fee usually has a distinct columella and 

spores only up to 36 pm long; T. intel-positurn 

Nylander has comparably large spores (I 15-160 

pm) but the thallus is conspicuously warty; and 

T. postposit urn Nylander has smaller (0.6-0.9 mm 

wide) apothecia with a constricted base (see Hale, 

1974a:32j. This species was very common at the 

one site in the Province of Cocle, mostly on canopy 

branches. 

SPECIXfENS EXAMINED.-~: 43427, 43495, 43516, 43521, 43541, 

43565, 43569, 43575, 43582, 43589,43595. 

14. Thelotrema myriocarpum 

FIGURE 13b 

Thelotrenza rnytiocarpurn FPe, 1824:94 [type collection: “In 

America meridionali ad corticem Cinchoilue rubrue” (PC, 

lectotype; isolectotypes in H, L) (Figure Ze)]. 

DEscRrPTror\i.-Thallus corticolous, epiphloeodal, 

dull, smooth to minutely verruculose, becoming 

rimose, whitish mineral gray, 4-8 cm broad; apothecia numerous, separate, immersed, 0.2 mm in 

diameter, noncarbonized, the inner exciple entire, 

free, forming an inner ring, the pore round, 0.05- 

0.1 mm in diameter; spores 8, 12 X 24-25 pm, 

1-2 X 5-7 loculate, I- or I+ weakly reddish. 

CHEMISTRY.-stiCtiC and constictic acid. 

REhfARKs.-Thelotrerna inyriocarpzirn was described by Fee from South America (probably 

Venezuela) and has not been reported since. As 

with most of Fee’s types now housed in G, the 

original material is in poor condition although 

the PC material is adequate to characterize the 

species. A single collection from Panama compares 

well with it; small, more or less immersed apothecia 

with a persistent inner excipular structure, although the spores in the lectotype are somewhat 

larger (about 36 pm long). The species is chemically 

identical with T. glaucopallens, which has similar 

apothecia but no remnants of an inner exciple. 

They are obviously clearly related but more specimens of the T. myriocarpum type must be seen 

to resolve the problem. 

SPECIMEN EXAMINED.-6: 43428. 

15. Thelotrema rnyrioporoides 

FIGURE 13c 

Thelotrema rnyrioporoides Muller Argoviensis, 1893b: 147 

[type collection: Boruca, Costa Rica, Tonduz in 1893 (C, 

lectotype) (Figure Zf)]. 

DESCRIPTIoN.-Tha~lus corticolous, epiphloeodal, 

thick and smooth, deeply fissured, whitish to pale 

greenish mineral gray, 6-10 cm broad; apothecia 

numerous, immersed, about 0.2 mm in diameter, 

noncarbonized, the inner exciple free, partially filling the pore, the pore round, white-annulate, 0.1 

mm in diameter; spores 8, 6-10 X 12-14 pm, 0.1 X 

3.5 loculate, I+ blue. 

CHEMISTRY.-Hypoprotocetraric acid and 4-0- 

demethylnotatic acid, with or without lichexanthone. 

REMARKS.--First described from Costa Rica, this 

species inhabits mid elevation rain forests in Panama. Superficially it resembles other canopy species with immersed apothecia such as Thelotrema 

clnndestinum Fee and Ocellularia terebratula (Nylander) Muller Argoviensis, as noted by Miiller 

(1893:147), who had probably seen the type of T. 

subcaesium Nylander, a synonym of T. clandestiniini. Thelotrema subconfol-me Nylander, also 

mentioned by Muller, is an Asian species lacking 

any lichen substances and having somewhat smaller, 

more crowded apothecia. One specimen from Panama (45806) is anomalous in producing lichexan- 
46 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

FIGURE 13.--Species of Thelotrema: a, T. maximum (Hale 43580); b, T. myriocarpum (Hale 

43428); c, T. myrioporoides (Hale 38420); d, T. norstictictcm (Hale 43592); e, T. occlusum (Hale 

38736); f, T. praestans (Hale 44784); g, 7'. pvcnoporellu?n (Hale 38505); 12, T. stylotliecium 

(Hale 38724); i, T. subpraestans (Hale 44878); j, T. tenue (Hale 44810); k, T. wrightii (Hale 

38449) (X 10). 
NUjMBER 38 

thone in addition to the normal components, a 

not unexpected combination. 

SPECIMENS EXA.\IINED.-4: 43435; 7: 43602; 9~38429, 38494; 

11: 45806; Hunter and Allen (Province of Panamii). 

16. Thelotrema norsticticum 

FIGURE 13d 

Thelotrema norsticticum Hale, 1973:417 [type collection: 

Trinidad, Hale 37369 (US, holotype)]. 

DEscRIPTIoN.-Thallus corticolous, epiphloeodal, 

about 0.6 mm thick, continuous, whitish mineral 

gray, about 6 cm broad; apothecia numerous, uniformly dispersed, immersed, 0.1-0.2 mm in diameter, noncarbonized, a columella lacking, the pore 

round, 0.06 mm in diameter, white annulate; spores 

8, 6-8 X 12-14 E*m, 0-1 X 4 loculate, Ii- blue. 

CHEMISTRY.-Norstictic acid. 

REMARKs.-This is the second record for the species. It is differentiated from T. clandestinum Fee 

by having a white annulate pore and different 

chemistry (T. clandestinum has psoromic acid). 

SPECIMEN Ex.w1Nm.--2: 43592. 

17. Thelotrema occlusum 

FIGURE 13r 

Tlielotrema occlusum Nylander, 1867:318 [type collection: 

Chucuri, Colombia, Lindig in 1863 (H, Nylander herbal - 

ium 22522, lectotype) (Figure 2h)l. 

DESCRIPTIoN.-Thallus corticolous, epiphloeodal, 

thin, verruculose, shiny, whitish to ivory colored, 

3-8 cm broad: apothecia solitary, dispersed, nearly 

immersed to semiemergent, 0.6-1 .O mm in diameter, 

the thalline margin sometimes suberect, the inner 

exciple well developed, crumbly, filling the pore, 

columella lacking, the pore round, open, 0.2-0.4 

mm in diameter; spores 1, 15-40 X 100-220 pm, 

densely muriform, I+ blue. 

CHEMISTRY.-Hypoprotocetraric acid and 4-0- 

demethylnotatic acid. 

REMARKS.-ThiS is still another of the Colombian species described by Nylander which occurs 

in the highlands of Panama. The disc is filled with 

the irregular, lacerate inner exciple, as in Phlyctis, 

and the spores are large. No other species in Thelotrema with hypoprotocetraric acid have this struc47 

ture. It occurs from base to mid bole on large evergreen hardwoods in both Panama and Venezuela. 

SPECIMENS EXAAIINED.-~: 38736; la: 44993, 45841, 45876. 

18. Thelotremu praestans 

FIGURE 13f 

Thelot).enza praestans Miiller .4rgoviensis, 1895c:453 [type 

collection: Rio de Janeiro, Brazil, Portella (C, lectotype: 

isolectotype in BM)]. 

Thelotrema elliottii Vainio, 1896:207 [type collection: St. 

['incent, Elliott 246 pro parte (BhI, lectotype)]. 

DEscRIPTIoN.-Thallus corticolous, epiphloeodal, 

smooth and continuous, greenish mineral gray, 6-12 

cm broad; apothecia widely dispersed, solitary, becoming strongly emergent, up to 2 mm in diameter, 

heavily carbonized with a thick columella, the pore 

round, 0.15-0.3 mm in diameter; spores 1-2, 

30-45 X 100-275 pm, densely muriform, I+ blue. 

CHEmsTKY.-"Praestans" unknown. 

REMARKs.-Thelostrema praestans is a large, 

easily recognized species. It has large spores, a 

heavily carbonized exciple and a thick columella, 

and a tiny depressed pore. As Redinger mentions 

(1936:91), Muller's type material has numerous 

cyphellalike white spots, apparently pycnidia, but 

I have not seen these in any other specimens. A 

very closely related species, T. subpraestans Hale, 

which occurs mainly in the lowlands, is even larger 

and contains stictic acid. Thelotrema praestans 

occurs generally at higher elevations in rain forest 

and is known from Guatemala, the West Indies, 

and Brazil. 

SPECIMENS EXAMINED.-~: 43524; 11: 44784; 12: Mori 4464; 

ILLUSTRATlOZIS.-Hak!, 1974a:36, figs. 17e, f, g; Redinger, 

Province of PanamA, Gentry 4210. 

1936:90, fig. 54. 

19. Thelotrema pycnoporellum 

FIGURE 13g 

Thetotremu pycnoporellurn Nylander, 1876:562 ([type collection: Cuba, Wright 69 (H, Nylander herbarium 22682, 

lectotype; isolectotypes in FH, G, UPS) (Figure 2m)l. 

DEscRIPTIoN.-Tliallus corticolous, epiphloeodal, 

thin, shiny, greenish mineral gray, 6-8 cm broad; 

apothecia numerous, separate, immersed, 0.1-0.5 

mm in diameter, the exciple barely carbonized, the 
48 SMITHSONIAN CONTRIBUTIONS TO ROTASY 

inner exciple evanescent, columella lacking, the 

pore round, 0.05 mm in diameter, slightly darkened; spores 8, 8-10 X 20-24 /Lm, 1-2 >: 5-6 loculate, II blue. 

CHEMISTRY.-NO substances present. 

I<E3fARKs.-This species was first described from 

Culia and is the only New TVorld Thelotrema with 

immersed apotliecia and no lichen substances. It 

is similar externally to T. myi$oporoides Muller 

Argoviensis. Zahlbruckner (1 923: 628) misspelled the 

specific epithet as “pycnocarpellum.” In Panama 

I collected it at the base and on fallen tops of 

evergreen hardwoods at 700-1000 m elevation. 

SPECIMENS EXAMIXED.-~: 38505: 11: 46019. 

20. Thelotrema stylothecium 

FIGLRI. 1312 

Theiotieiii~~ stylothecitcin Vaino, 1890:50 [type collection: 

Carassa, hlinas Gerais, Brazil, Vai7ro 1370 (TUR, lectotype; 

isolectotypes iii BN, FH, UPS) (Figure 2q)]. 

Thelotvema leuconielaenziiu var. stylotheciuiii (Vainio) Red. 

inger, 1936:92. 

DEscRIPTIoS.-Thallus corticolous, epiphloeodal 

but in part evanescent or hypophloeodal, whitish 

mineral gray, 4-8 cm broad; apothecia numerous, 

solitary, immersed to semiemergent, 0.4-0.i mm 

in diameter, the thalline margin coarse, sometimes 

becoming semierect, carbonized, the columella 

broad, about 250 pm in diameter with a whitepruinose apex partially filling the pore, the pore 

round, about 0.3 mm in diameter; spores 8, 

10-12 X 20-35 pm, 1-2 X 6-8 loculate, I-. 

CHEMISTRY.-stiCtiC acid with or without constictic acid. 

REMARKS.-h his original description Vainio 

(1990:81) did not discuss the differences between 

this species and the very close T. leucomelaenum. 

His Latin diagnoses are almost identical except 

for spore size, T. stylothecizim having smaller 

spores (15-17 pm vs. 28-32 pm long). However, I 

found that, aside from the basic chemical distinction, the spore sizes overlap to a much greater extent than either Vainio or Redinger (1936:92) 

showed and I do not believe that this is a valid 

difference. Both species occur in similar habitats 

in cloud forests at higher elevations (above about 

1500 m). I have identified T. stylothecium from 

St. Lucia and Guadeloupe in the West Indies-but 

not from Dominica-and from I’enezuela. 

SPECIMI 15 EXAXII\EU.-~: 38710, 35724, 38707, 35775, 38779; 

ILLLsTRArIos.-Kedinger, 1936:93, fig, i6. 

la: 45840. 

2 1. Thelotrema su bpraestans 

FICLKI 131 

Thelotwino aubprnestnns Hale. 19741):497 [t!pe collection: 

ProLince of Panama, Panama, Hale 38525 (US, holotype)]. 

DEscRImIox.-ThaIIus corticolous, epiphloeodal, 

rather thick, rimose with age, pale greenish or 

whitish gray, 10-20 cm broad; apothecia numerous, 

solitary, strongly emergent, up to 2.5 mm in diameter, carbonized, a thick central columella present, 

the pore round, inconspicuous and slightly depressed, 0.1 mm in diameter, white annulate; 

spores 1, 30-50 X 150-300 pm, densely muriform, 

I- blue. 

cHEMISTRY.-StiCtiC and constictic acids with or 

without the high “quintaria” unknown. 

RExtARKs.-Thelotrema szibpraestans is the predominant canopy species on Barro Colorado Island 

and also occurs at higher elevation to the east in 

the nearby Cerro Jefe region. It appears to represent a recent derivative, perhaps a mutant, of the 

more widespread T. praestans hIiiller Argoviensis, 

which contains an unknowm P+ compound and 

occurs generally at higher elevations. 

SPICIMENC ExaMls~D.-3: 35637, 38665, 38660, 38682, 38696; 

3a: 44737, 44705, 44809, 44878, Esslinger 4330, 4346, 4381; 

8: 38525, 38534, 38349; 9: 38432,38515. 

22. Thelotrema tenue 

FICCRE 131 

Thelotremn tenue Hale, 1974a:38 [type collection: Dominica, 

Hale 35439 (US. holotype)]. 

DEscRIPTIox.-Thallus corticolous, hypophloeodal, shiny, whitish gray, 6-8 cm broad; apothecia 

numerous, immersed to semiemergent, 0.3-0.4 mm 

in diameter, apically carbonized, the columella 

weakly developed to absent, the pore round, 0.05- 

0.1 mm in diameter; spores 8, 10-12 X 20-30 pm, 

1-2 X 5-6 loculate, I-. 

CHEMISTRY.-NO substances present. 
SUhIBER 38 

RExARlts.-Previously known only from Dominica, T. tenue is represented in Panama by a single 

collection in the cloud forest at Chiriqui. I have also 

collected it in Venezuela. The columella is .i\.eakly 

developed to lacking in the Panamanian material. 

51~1 CIUI.~ Ex.ahilrf.D.--la: 14810. 

It ~.us~~i~lohs.-Hale, 1974a:37, hg. 17k. 

23. Thelotrema wrightii 

FIGURE 13k 

Ti~elotrema wrightii Tuckerman, 1862:409 [type collection: 

Cuba, Li’right 143 (FH-Tuck, lectotype: isolectotypes in 

BM, G, L, PC, UPS) (Figure 2t)]. 

DEscRIPTIoN.-Thallus corticolous, epiphloeodal, 

thin and fragile, shiny, smooth to verruculose, pale 

tannish gray, 10-15 cm broad; apothecia conspicuous, chroodiscoid, becoming basally constricted, 1-3 

mm in diameter, the thalline margin coarse, erect 

to slightly recurved, the inner exciple evanescent, 

the disc white pruinose, becoming superficially 

actinoid with light carbonization; spores 8, 5-6 X 

10-18 pm, 0-1 X 4-5 loculate, IS blue. 

CHESfISTRY.-PSOrOmiC acid. 

REMARKS.-ThiS species forms large conspicuous 

colonies on the lower trunk to mid bole of large 

trees. The pale flesh-colored apothecia stand out in 

particular because of the broad chroodiscoid disc. 

The disc becomes irregularly ridged with age, giving the superficial actinoid appearance mentioned 

by Tuckerman (1862:410). It has been known before only from Cuba. Externally it is extremely 

close to Ocellularia subwrightii Hale (see page 32). 

SPECIMENS EXAMINED.-~: 38537, 38389; 9: 38449; 11: 44870, 

46052. 

Leptotrema Miiller Argoviensis 

Leptotrema is characterized by brown muriform 

Fpores with very close relationships to Thelotrema. 

I recognize approximately 65 species at the world 

level. In Panama I collected 14 species and 64 

specimens, about 9yo of the total 700 for the family, 

comparable to the 7y0 in Dominica. Four of these 

are new to science. 

The most commonly collected species was Leptotrema yeclusum, all 13 collections, however, being 

made at Barro Colorado Island. In the same locality L. laeuzuscztlzirn was collected eight times. While 

the genus is generally scattered at different localities, most collections were concentrated at just two 

localities, Chiriqui and Barro Colorado. Three 

species, L. chiriquzense, L. metaphoricurn, and L. 

urceolare, were confined to the highest elevations 

in Chiriqui, and L. laeviusculum, L. reclusum, and 

L. wightii can be considered as typical lowland 

species. 

The apothecial development in Leptotrema is 

varied. Leptotrema stellatum is the only strongly 

chroodiscoid species, while L. hypoprotocetraricurn 

is generally lepadinoid. Leptotrema metaphoricum 

has an actinoid columella and columellate L. micyosporum tends toward this condition. Flush 

myriotremoid apothecia are present in L. laeviusculum, L. tyypaneoides, and L. wightii. 

As with the other thelotremoid genera, Leptotrema has phytogeographical affinities with the 

Caribbean and South America. Leptotrema decepturn, L. laeviusculum, and L. trypaneoides form a 

Caribbean group, L. metaphoricum and L. microsporum are reported here for the first time outside 

of South America, and L. reclusum, I. urceolare, 

and L. wightii are pantropical. 

Key to the Species of Leptotrema 

1. Apothecia open and chroodiscoid, ecolumellate. ........................................................ 11. L. stellatum 

1. Apothecia not chroodiscoid: if open an actinoid columella present. 

2. Columella present. 

3. Disc open, large, actinoid, becoming irregularly elongate. ...................... 7. L. metaphoricurn 

3. Disc round, not actinoid or open. .................................................................... 8. L. microsporum 

4. Spores large, more than 150 pm long, densely muriform. 

2. Columella lacking. 

5. Pore 0.1 mm or less in diameter. ................................................................ 2. L. chiriquiense 

5. Pore open, 0.2-0.3 mm wide, the proper exciple filling the disc partially. ........................ 

...................................... 4. L. hypoprotocetraricum 

4. Spores less than 130 pm long. 

6. Apothecia generally large, emergent, 0.5-1.0 mm in diameter, the pore to 0.5 mm 

wide. 
50 SMITHSONIAN CONTRIBUTIONS TO BOTASY 

7. Thallus P+ red (protocetraric acid). 

8. Disc covered with an orange-yellow pruina. ..................... ...... 9. L. panamense 

8. Disc unpigmented. ........................................................................................ 1. L. bahianum 

9. Spores 80-100 pm large. ............ 

7. Thallus P--. 

9. Spores smaller, 30-30 Fm long. .... 13. L. urceolare 

6. Apothecia immersed to flush, 0.2-0.5 mm in diameter, the pore 0.05-0.1 mm wide. 

10 Red crystal inclusions in the cortex. ............................................................. 14, L. wightii 

10. Cortex lacking pigments. 

11. Spores about 100 prn long. .................................................................. 10. L. reclusum 

11. Spores ahout 25 long. 

12. Cortex shiny, thin; stictic acid present. ............................... 12. L. trypaneoides 

12. Cortex dull. thick and grainy; protocetraric acid present. .... 5. L. Zaeviusculum 

1. Leptotrema bahianum 

FIGURE 14a 

Leptotreina bahianum (Acharius) Muller r\rgoi iensis, 

Thelotrema lepadinunz tar. bahianum Acharius, 1803:132. 

Thelotrema deuelatum A'ylander, 1863a:454 [type collection: 

Colombia, Lindig 774 (BSI, lectotype; isolectotypes in FH, 

1887a: 12. 

PC, W)]. 

Thelotrema rudius Vainio var. dominicanum Vainio, 19153136. 

Leptotrema develatum (Nylander) Zahlhruckner, 1923:633. 

[For full citations see Hale (1974a:39).] 

DEScRIPTIoN.-Thallus corticolous, epiphloeodal, 

thin, light tannish or greenish gray, 4-10 cm broad; 

apothecia numerous, emergent, 0.4-0.8 mm in diameter, noncarbonized and ecolumellate, the pore 

round, about 0.2 mm in diameter; spores 8, 

12-14 X 16-24 pm, 1-3 X 5-6 loculate. 

CHEmsmy.-Protocetraric acid. 

REMARKS.-This is a rather uniform species. It 

is extremely close to L. urceolare (Acharius) Muller 

Argoviensis, which lacks any lichen substances. 

Leptotrema bahianum is most common in the Caribbean area but also occurs in Venezuela, the Philippines, Sarawak, and other tropical countries. It 

was collected in Panama over a broad range of 

elevation (near sea level to 2000 m) on trees in 

primary and secondary forest. 

SPECIMENS EXAMIXED.-~: 38726; 3a: 44869, Esslinger 4339; 

5: 43599; 11: 44845. 

Redinger, 1936:106, fig. 66. 

ILLCSTRATIONS.-Hak, 1972: 193, fig. 2C; 1974a:7, fig. 5f; 

2. Leptotrema chiriquiense, new species 

FIGURE 14b 

DEscRIPTIoN.-Thallus corticola, epiphloeodes, 

nitidus, verrucosus rimosusque, fragilis, stramineoalbidus, 4-6 cm latus; apothecia (Figure llc) numerosa, congesta, emergentia vel ascidioida, rotundata veI fere subglobosa, 0.7-1.0 mm diametro, 

apice fuliginea, columella nulla; ostiolum rotundatum, vix depressum, ca. 0.1 mm diametro; hymenium ca. 300 pm, altum; sporae, l-2:nae, 35-40 

X 160-190 pm, dense murales, I-. 

CHEmsTRY.-Hypoprotocetraric acid and apparently 4-0-demethylnotatic acid. 

HoLoTwE.-Base of exposed tree, remnants of 

oak forest in pasture at Alto Bespingo, Volcin 

Chiriqui, Province of Chiriqui, Panama, elevation 

2400 m, M. E. Hale 46035, 23 February 1975 (US). 

REMARKS.-T~~S species is externally somewhat 

similar to Ocelliilaria domingensis (Nylander) 

Muller Argoviensis, which also has hypoprotocetraric acid and large spores but with larger, less 

fragile apothecia. Both species were collected at 

Chiriqui. This is only the second Leptotrema reported to contain hypoprotocetraric acid, the other 

being L. hypoprotocetrariczim Hale from the 

Darien. 

SPECIMEN EXAMINED.-~: 38888. 

3. Leptotrema deceptum 

FIGURE 14c 

Leptotrema deceptum Hale, 1974a:39 [type collection: DOminica, Hale 37860 (US, holotype)] 

DEscRIPTIox.-Thallus corticolous, epiphloeodal, 

continuous to cracked with age, smooth and shiny, 

light tannish gray, 8-10 cm broad; apothecia numerous, immersed to barely emergent, about 0.4 

mm in diameter, noncarbonized and ecolumellate, 

the pore minute, about 0.05 mm in diameter, depressed, surrounded by a darkened area; spores 4, 
NUMBER 38 51 

FIGURE 14.--Species of Leptotrema: a, L. bahianum (Hale 44845); b, L. chiriquiense (Hale 46035); 

c, L. deceptum (Hale 43551); d, L. hypoprotocetraricum (Mori 4513); e, L. laeuiusculum (Hare 

44973); f, L. lepadodes (Hale 46045); g, L. nietaphoricum (Hale 44911); h, L. microsporum 

(Hale 44936); i, L. panainense (Hale 43585); j, L. reclusum (Hale 44773); k, L. stellaturn (Hale 

44999); I, L. trypaneoides (Hale 46038) (X 10). 
52 SMITHSONIAN CONTRIBUTIONS TO BOTANY 

25-30 X 60-80 pm, densely muriform, I+ pale blue. 

CHE\IISTRY.-StiCtiC and constictic acids. 

REm%RI(s.-This species was previously known 

only from Dominica and it occurs in Panama in 

the same type of habitat, base of large trees in rain 

forest at mid elevations. The pore area is slightly 

darker and broader than in the typical population. 

Two other species in Panama have stictic acid: L. 

recliiszim (Krempelhuber) Zahlbruckner with larger 

spores and a grainy appearing cortex and L. trypaneoides (Nylander) Riddle with flush apothecia 

and smaller spores. 

SPLCIMES Es~NINF~.-~: 43551. 

Ir~usm~~IoNs.-Hale, 1974a:39, fig. 18. 

4. Leptotrema hypoprotocetraricum, new species 

FICLRE 14d 

DEScRIPTIoS.-Tha~lus corticola, epiphloeodes, 

crassus, laevis vel aetate rimosus, pallide cremeoalbidus, 2-4 cm latus; apothecia dispersa, solitaria 

vel 2-3 aggregata, semi-emergentia, 0-8-1.5 mm diametro, margine thallino crasso, raro semi-erecto, 

apice pulverulento, albo, excipulo proprio plus 

minusve distincto, decolorato, separato, columella 

nulla; hymenium 310-320 pm altum; sporae 1-2: 

nae, 33-36 X 150-210 ILm, dense murales, I-. 

CHEMISTRY.-Hypoprotocetraric and 4-0-demethylnotatic acids. 

HoLoTYPE.-~~~~s~ ridge of Cerro Tacarcuna massif, Province of Darien, Panama, elevation 1700 m, 

S. Mori and A. Gentry 4513, 3 February 1975 (US). 

REifARKS.-The solitary to clumped apothecia, 

somewhat reminiscent of those in Ocellulnria conglornerata Hale, are whitish pulverulent apically 

and stand out conspicuously from the thallus. No 

other species of Leptotrema have this type of disc, 

and as pointed out above, only L. chiriqziiense also 

contains hypoprotocetraric acid. 

5. Leptotrema laeviusculum 

FIGURE 14.5 

Leptotremn laeviusculum (Nylander) Zahlbruckner, 19233635. 

Thelotrema laeviusculum Nylander, 19631):335 [type collection: Cuba, TTright 155 (FH-Tuck, lectotype: isolectotypes 

in BAI, G, L, PC, UPS)]. 

DEscRIPTIos.-Thallus corticolous, epiphloeodal, 

smooth and rather thick with a grainy appearance, 

sometimes breaking free of the substratum, brittle, 

pale greenish gray, 6-12 cm broad; apothecia numerous, immersed, separate, 0.2-0.3 mm in diameter, noncarbonized and without a columella, the 

pore round, 0.8 mm in diameter, white annulate; 

spores 8, 6-12 X 18-24 pm, 0-1 X 4-5 loculate. 

CmziIsTRY.-Protocetraric acid. 

REMARKS.-This species is characterized by the 

dull grainy thallus with crystal inclusions and the 

small immersed apothecia. T’ery superficially it 

might appear to be a member of the Ocellzilaria 

oliuacea group but the habitat is quite different, 

lianas, exposed roots, and base of trees in low elevation rain forest. I have identified specimens from 

Cuba, the type-locality, and Trinidad. 

SPECIhIFhS EXAhlINED.-3a: 44762, 44764, 44783, 44861, 44865, 

44960, 44973, Esslinger 4357. 

6. Leptotrema lepadodes 

FICVRE 14f 

1,eptotrenza iepadodes (Tuckerman) Zahlhruckner, 1923:636. 

Thelotrema lepnrlodes Tuckerman, 1862:405 [type collection: 

Cuba, Wright (FH-Tuck, lectotype)]. 

Thelotrema ~nonosporu~~~ Xylander f. album Sylander, 1861: 

46 [type collection: Bonin Islands, Wright (FH-Tuck, lectoThelotrema monosporum Tar. patulum Nylander, 1963a:452 

[type collection: Cune, Colombia, Lindig in 1860 (H, Nylander herbarium 22715, lectotvpe)]. 

Thelotrema disciforme Leighton, 1869:170 [type collection: 

Central Province, Ceylon, Thulnites (BM, lectotype)]. 

Leptotrema pntuluni (Nylander) Sfiiller Argoviensis, 1895b: 

315. 

Thelotretnn nquilinum T’ainio, 19153137 [type collection: St. 

Croix, Boergesen (TUR, lectotype)]. 

Leptotrerna monosporutn f. album (Sylander) Zahlbruckner, 

1923:638. 

Thelotreina rarotongae Rasanen ex Sbarbaro, 1939:lOl [type 

collection: Cook Island, Rarotonga, Parks 2 (H, lectotype)]. 

Lefltotrema pinnrocarpurn Zahlbruckner in Magnusson and 

Zahlhruckner, 1944:54 [type collection: Il’ailuku, IIaui, 

Hawaii, Faurie 679 (C‘PS, lectotype)]. 

type)]. 

Phneotrenia disciforme (Leighton) Hale, 1974a:29. 

DEscRIPTIoN.-Thallus corticolous, very thin and 

mostly hypophloeodal, grayish white, 4-6 cm broad; 

apothecia common, solitary, emergent, to 1.3 mm 

in diameter, the thalline margin thick, darker gray 

and decorticate apically, semierect, carbonized, the 

inner exciple distinct and persistent, surrounding 

the open, white pruinose disc that pulls away from 
NUMBER 38 53 

the wall, a columella lacking, the pore gaping, to 

0.7 mm in diameter, the disc easily visible; spores 

8, 2-4, 15-25 X 80-100 pm, densely muriform. 

CHEMISTRY.-NO substances present. 

REMARKS.-The apothecia of this species have 

an open, gaping pore and a disc that pulls away 

from the main wall at the periphery. In Panama 

it occurs in disturbed forest and at canopy level in 

rain forest. I have seen specimens from the United 

States, England, West Indies, Hawaii, India, Java, 

the Philippines, and the Solomon Islands in addition to localities represented by the type collections. 

When I first examined the type collection of 

Thelotrema disciforme Leighton, I apparently sectioned the only specimen with transversely septate 

spores and therefore placed the species in Phaeotrema. The remaining material (3 of the 4 pieces 

in the original collection in BM), however, has muriform spores, as later discovered and annotated by 

Dr. P. G. Patwardhan. Correctly typified as a species of Leptotrema, it is antedated by L. lepadodes. 

I have not yet determined the disposition of the 

Phaeotrema species. In this connection it should be 

pointed out that the type of Thelotrema Zepadodes 

Tuckerman in FH-Tuck definitely has muriform 

spores, not transversely septate as Salisbury (1972: 

272) reports. This is a difficult group that seems to 

cast doubt on the validity of the spore differences 

as a generic character. Much more material needs 

to be examined. 

SPECIXiENS EXAMINED.-3: 38673: 10: 46045. 

7. Leptotrema metaphoricum 

FIGURE 14s 

Leptotrema metaphoricum (Nylander) Zahlbruckner, 1923: 

637. 

Thelotrema rnetaphoricum Nylander, 1963a:455 [type collection: \’illeta, Colombia, Lindig 2814 (BM, lectotype: isolectotypes in FH-Tuck, G, PC, S, UPS, W)]. 

DEscRIPTIos..-Thallus corticolous, epiphloeodal, 

rather thick, smooth, fissured with age, pale tannish 

white, 4-10 cm broad; apothecia common, solitary 

or in part clumped, semiemergent, chroodiscoid, 

0.5-1.5 mm in diameter, round to irregularly elongate in outline, the thalline margin thick, semierect, 

the inner exciple evanescent, the disc broad, partially actinoid columellate; spores 8, 5-8 X 10-18 

pm, 0-1 X 3-4 loculate. 

CHEMISTRY.-LiCheXanthOne. 

REMARKS.-The disc is usually large and actinoid 

columellate with a short thalline margin, much as 

in Phaeotrema glyphicum (Nylander) Zahlbruckner. 

Older apothecia become irregularly lirelliform. 

This species was first described from Colombia but 

has been collected so far in Guatemala and Brazil 

and in Panama in cloud forest at 2000 m elevation. 

Only one other comparable species contains lichexanthone, L. microsporum Zahlbruckner; it has 

small round apothecia and a simple columella, but 

there are obvious and sometimes troublesome intermediates between the two species. 

SPECIMENS EXAXrINED.-la: 4491 1, 44916, 44928. 

8. Leptotrema microsporum 

FIGURE 14h 

Leptotrema microsporum Zahlbruckner, 1902:392 [type collection: Rio de Janeiro, Brazil, Hohnel 148 (W, lectotype) 

(Figure 241. 

Thelotrema rnicrosporoides Zahlbruckner, 1923:625 [based on 

“Thelotrema microsporum Zahlbruckner” (not T. microsporum Montagne).] 

DEscRIPTIoN.-ThalIus corticolous, epiphloeodal, 

rather thick and smooth to grainy, dull greenish 

mineral gray, 4-7 cm broad; apothecia mostly solitary, at first immersed but becoming slightly emergent at maturity, to 1 mm in diameter, round in 

outline, the thalline margin entire, thick, semierect 

with age, the disc broad, the columella simple to 

partially actinoid, carbonized, the surface white 

pruinose, the pore wide, 0.5-1.0 mm wide, white 

annulate, the tip of the columella often filling the 

pore; spores 8, 5-8 X 8-11 pm, 0-1 X 2-4 loculate. 

CHEMIsTRY.-Lichexanthone. 

REMARKS.-Typical apothecia, as determined 

from the type specimen, are flush and 0.5 mm or 

less in diameter. Some specimens, however, have 

larger apothecia, which, if strongly emergent, intergrade with those of L. metaphoricum (Nylander) 

Zahlbruckner (see above). These two species may 

ultimately prove to be part of a large variable 

population but too few specimens are on hand to 

make an intelligent judgment. The normal habitat 

seems to be the lower trunk of trees in open forest. 

I have identified the species from Mexico, Costa 

Rica, Cuba, and Colombia. 
54 SMITHSONIAN CONTRIBUTIOSS TO BOTANY 

SPECIMENS EXA\IIXED.-~: 38735, 38755 (both tentatively 

identified); la: Esslinger 4487, 4485; 3a: 44790, 44936: Mariaganti: M’illiarns 1103. 

9. Leptotrema panamense, new species 

FIGURE 14i 

DEscRrPrrou.-Thallus corticola, epiphloeodes, 

continuus sed demum rimosus, leviter verruculosus, 

olivaceo-albidus, 6-9 cm latus; apothecia (Figure 

1 Id) numerosa, dispersa, emergentia, 0.7-1.0 mm 

diametro, basi non valde constricta; ostiolum rotundatum, 0.1-0.2 mm diametro, plus minusve annulato, margin nigro-cinctum vel aurantiaco-cinctum; excipulum apice fuligineum, disco superficie 

aurantiaco; Iiymenium 130-140 pm altum; sporae 

6-8:nae, 10-12 X 24-28 pm, longtitudine l-loculatae, transversim 5-6 loculatae, I A caerulescentes. 

CHEairsTRu.-Protocetraric acid and an unidentified pigment. 

HoLoTYPE.-Secondary rain forest, La Mesa, 

north of El Valle, Province of Cocle, Panama, elevation 750 m, M. E. Hale 43584, 13 February 1974 

REMARKs.-The rimose thallus and black-rimmed 

ostiole are similar to those of 0. nigropzrncta Hale 

and 0. rimosa Hale, both from Dominica. The 

reddish orange disc contains a pigment identical 

to that of Thelotrema aztratunz. It is a canopy species at mid elevations. 

(US). 

SPECIVENS EXA\II\ED.-2: 43506, 43585; 8: 38567; 11: 46039, 

46040, Esslinger 4642. 

10. Leptotrema reclusum 

FIGURE 14j 

Leptotrerna xclusurn (Krempelhuber) Zahlbruckner, 1923: 

639. 

Thelotrema reclusurn Krempelhuber in Nylander, 1973:168 

[type collection: Andaman Islands, h’ur: 21 (M, lectotype; 

isolectotypes in EM, FH-Tuck, H-Syl, UPS, TV, 27) (Figure 2n)l. 

DEsCRIPTIor.-Thallus corticolous, epiphloeodal, 

well developed and appearing rather thick, broadly 

warty or bullate, fragile, deeply fissured, the surface 

shiny but appearing grainy, tannish to greenish 

mineral gray, 4-8 cm broad; apotliecia inconspicuous, immersed to semiemergent, 0.4-0.6 in diameter, 

noncarbonized and without a columella, the pore 

round, 0.05 mm in diameter; spores 1-2, 20-30 X 

70-130 pm, densely muriform. 

CHEXfISTRY.-stiCtiC acid, constictic acid, and infrequently the highest “quintaria” unknown. 

REMARKs.-Leptotrenza Yeclusum is one of a 

number of species in the genus with stictic acid, a 

very difficult group that has yet to be well comprehended (see Hale, 1974a:40). It is characterized by 

the large spores, the weakly bullate thallus with a 

grainyappearing cortex, and immersed apothecia. 

The Panamanian material, all collected on Barro 

Colorado Island at the base of trees, compares very 

well with the type from the Andaman Islands and 

specimens from Florida. 

SPECI\IE\S EX4\IINtD.-3: 38426, 38629, 38644, 38668, 38695: 

3a: 44767, 44772, 44773, 44775, 44776, 44780, Esslinger 4333, 

4354. 

11. Leptotrema stellatum, new species 

FIGURE 14k 

DEscRIPTIos.-Thallus corticola, epiphloeodes, 

continuus, nitidus, pallide stramineo-glaucescens, 5 

cm latus; apothecia rara, dispersa, chroodiscoida, 

rotundata vel oblonga, usque ad 2 mm lata, margine thallino crasso, stellato-fisso, recurvato, excipulo proprio evanescenti, disco obscuro vel leviter 

albo-pruinoso; hymenium 220-230 pm altum; sporae 2:nae, 22-26 X 105-120 pm, dense murales, I-. 

CHEMISTRY.-NO substances present. 

HOLOTYPE.-TOP of tree, primary rain forest 

along road from Alto Piedra to Rio Calovebora, 

Province of Veraguas, Panama, elevation 800 m, 

M. E. Hale 44999, 25 February 1975 (US). 

REMARKS.-The chroodiscoid disc is lightly pruinose and the thick thalline margin split and recurved as in a Geaster. There is some resemblance 

to L. mimbilis Zahlbruckner, which has smaller 

spores (42-46 pm long) and contains stictic acid. 

12. Leptotrema trypaneoides 

FIGURE 141 

Leptotiema trypaneoides (Nvlander) Riddle, 1916: 151. 

Tlielotrerna trypaneoides Nvlander, 186311:335 [type collection: Cuba, Tt’r-ight I56 (FH-Tuck, lectotype; isolectotypes 

in EM, FH, G, L, PC, UPS, US, TV) (Figure 2s)J. 
NUMBER 38 55 

Thelotrema subterebrans Nylander, 1876:561 [type collection: 

Cuba, Wright 520 (H, Nylander herbarium 22514, lecto- 

‘Y Pe) 1. 

Leptotrema subterebrans (Nylander) Zahlbruckner, 19!23:640. 

DEscRIPTroN.-Thallus corticolous, epiphloeodal, 

smooth, shiny and continuous, pale greenish gray, 

8-12 cm broad; apothecia numerous, separate, immersed, 0.15-0.2 mm in diameter, sometimes slightly 

carbonized apically, a columella lacking, the inner 

exciple evanescent, the pore round, sometimes black 

rimmed, about 0.1 mm in diameter; spores 8, 10-12 

X 24-30 pm, 1-2 X 5-6 loculate. 

CHEMIsTRY.-stictic and constictic acids. 

REMARKS.-The smooth shiny thallus and the immersed dark-pored apothecia distinguish this species. Leptotrema phaeosporum (Nylander) Miiller 

Argoviensis from New Caledonia, a close species, 

has small apothecia without a distinct bordered 

pore. It is a rare species in Panama, collected at 

the base of trees and on saplings in rain forest. 

SPECIMENS EXAMINED.-%: 44815; 11: 44958, 46038. 

FIGURE 15.4pecies of Leptotrema: a, L. urceolatum (Hale 

4493); b, L. wightii (Hale 38663) (X 10). 

often basally constricted apothecia and negative 

chemistry. I also have specimens from Hawaii and 

Sarawak. In Panama it occurred on buttresses and 

bases of large trees, most commonly in the cloud 

forest at Chiriqui but also near sea level at Barro 

Colorado. 

SPECIMENS EXAMINED.-~: 38712, 38740, 38766; la: 44939, 

44974, 45815, Esslinger 4466, 4500: 3: 38630. 

ILLUSTRATIONS.-Hak 1972: 195, fig. 3A. 

13. Leptotrema urceolare 14. Leptotrema wightii 

FIGURE 15a FIGURE 15b 

Leptotrema urceolare (Acharius) Miiller Argoviensis, 1887a: 12. 

Thelotrema urceolare Acharius, 1812:90 [type collection: 

“America” on Cinchona rubra (H-Ach, lectotype; isolectotypes in C, UPS)]. 

DEScRIPTIoN.-Thallus corticolous, epiphloeodal, 

thin, smooth and continuous, dull, light tannish 

gray, 6-12 cm broad; apothecia numerous, solitary 

to irregularly clumped, emergent and sometimes 

basally constricted to give an urceolate appearance, 

0.7-1.1 mm in diameter, carbonized, ecolumellate, 

the whitish pruinose disc usually visible through 

the pore, the pore wide, round, 0.2-0.5 mm in diameter; spores 8, 10-12 X 20-30 pm, 0-1 X 4-5 

locula te. 

CHEMISTRY.-NO substances present. 

REMARKS.-h my study of the Acharian species 

(Hale, 1972) I expressed reservation on the identity 

of this species. Since examining more collections, 

however, I feel that the Panamanian specimens are 

typical and help establish the characters of L. UYceolure. The main features are the semiemergent, 

Leptotrema wightii (Taylor) Miiller Argoviensis, 1882:499. 

Endocarpon wightii Taylor, 1847:155 [type collection: India, 

Wight (FH-Tayl, lectotype; isolectotypes in BM, G)]. 

Thelotrema prevostianum Montagne, 1849:292. 

Leptotrema prevostianum (Montagne) Montagne, 1956:364. 

Thelotrema subconcretum Leighton, 1869: 169. 

Thelotrema foraminulosum Krempelhuber, 1875:20 [type collection: Sarawak, Beccari 112 (M, lectotype; isolectotype in 

Phaeotrema subconcretum (Leighton) Miiller Argoviensis, 

Lefitotrema flavicans Miiller Argoviensis, 1888:114. 

Leptotrema subconcretum (Leighton) Miiller Argoviensis, 

Phaeotrema foraminulosum (Krempelliuber) Zahlhruckner, 

[For full citations see Hale (1974a:43).] 

G)l. 

1887a:lO. 

1891c:277. 

1923:607. 

DEscRIPTIoN.-Thal~us corticolous, epiphloeodal, 

thick, the surface smooth but with a grainy appearance, the cortex columnar, the medulla with conspicuous red crystal masses, greenish mineral gray, 

up to 10 cm broad: apothecia numerous, separate, 

immersed, 0.2-0.3 mm in diameter, noncarbonized 
56 ShlITHSOKIAN CONTRIBUTIONS TO BOTANY 

and without a columella, the pore round, about 0.1 

mm in diameter; spores 8, 9-11 X 20-28 pn, 0-2 X 

4-6 loculate. 

CHE;\IISTRY.-XO substances present except for 

REMARKS.-This pantropical species is rare in 

Panama. As a lichen most commonly collected in 

dry scrub forests, one would not expect to find it 

well del’eloPed in virgin rain forests. 

the unidentified red anthraquinone. SPECIMENS EXAMIXED.-.?: 38663; 3a: 44813. 
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1923. 
Index 

(Synonyms in italics) 

Ascidium cinchonarum, 16 

cinchonarum f. intermedium, 16 

cinchonarum var. verrucosum, 33 

croceum, 17 

domingense, 20 

melanostomum, 30 

rhabdosporum, 29 

rhodostromum, 30 

xanthostromum, 34 

Endocarpon wightii, 55 

Graphis alborosella, 12 

Karstenia sublilacina, 31 

Leptotrema bahianum, 50 

deceptum, 50 

develatum, 50 

flavicans, 55 

hypoprotocetraricum, 52 

laeviusculum, 52 

lepadodes, 52 

metaphoricum, 53 

microsporum, 53 

mirabilis, 54 

monosporum f. album, 52 

panamense, 54 

patulum, 52 

phaeosporum, 55 

pinarocarpum, 52 

prevostianum, 55 

reclusum, 54 

stellatum, 54 

subconcretum, 55 

subterebrans, 55 

trypaneoides, 52, 54 

urceolare, 55 

wightii, 55 

Lichen Eepadinus, 44 

Myriotrema album, 12 

Ocellularia actinota, 12 

olivaceum, 26 

alba, 12 

alborosella, 12 

albula, 24 

a1 bula f. opaca, 25 

antillensis, 14 

aurulenta, 14 

barroensis, 14 

berkeleyana, 14 

bullata, 15 

calvescens, 15 

cavata, 15 

cavata var. planior, 28 

chionostoma, 16 

cinchonarum, 16 

cinchonarum f. intermedia, 16 

clandestina, 43 

comparabilis, 16 

concolor, 17 

congesta, 17 

conglomerata, 17 

costaricensis, 12 

crocea, 17 

dactylifera, 19 

dactyliza, 19 

decolorata, 19 

dilatata, 19 

dissuta, 20 

domingensis, 20 

domingensis var. fecunda, 21 

efformata, 16 

esslingeri, 20 

exanthismocarpa, 21 

excavata, 28 

fecunda, 21 

fragilis, 15 

glaucophaena, 21 

glaucula, 23 

groenhartii, 23 

henatomma, 20 

homothecia, 21 

isidiifera, 23 

landronii, 23 

latilabra, 15, 24 

lepadinioides, 34 

leptopora, 23 

leucotylia, 15 

lindigiana, 16 

microporella, 24 

minutula, 24 

mordenii, 25 

multilocularis, 21 

myriotremoides, 25 

neei, 25 

nigropuncta, 28, 54 

obturata, 16 

olivacea, 26 

opaca, 25 

pachystoma, 26 

papillata, 26 

perforata, 26 

phlyctellacea, 16 

platycarpella, 12 

plurifaria, 24 

porinoides, 21 

protocetrarica, 28 

pseudoschizostoma, 28 

pulverulenta, 29 

pyrenuloides, 43 

radians, 15 

recondita, 29 

referta, 29 

remanens, 33 

rhabdospora, 29 

rhodostroma, 30 

rimosa, 54 

ripleyi, 30 

rufocincta, 28 

schizostoma, 24 

sexlocularis, I5 

sorediata, 29 

subcavata, 31 

subemersa, 31 

sublilacina, 31 

subterebratula, 24 

subwrightii, 32 

terebrata var. abbreviatula, 28 

terabratula, 33 

turgidula, 33 

umbilicata, 24 

uniseptata, 33 

vaga, 31 

verrucosa, 33 

viridis, 34 

xanthostroma, 34 

Phaeograp h is c hionodisca, 35 

Phaeotrema aggregatum, 29 

auberianum, 35 

croceum, 17 

disciforme, 52 

f orn m in ulosum, 55 

glyphicum, 36 

leiostomum, 36 

meiospermum, 36 

melanostomum, 30 

pachystomum, 26 

phaeospermoides, 36 

rhabdosporum, 30 

subconcretzrm, 55 

vulcani, 37 

Pyrenula clandestina, 43 

Stegobolus berkeleyanus, 14 

59 
SXIITHSONIAN COSTRIBUTIOSS TO ROTASY 

Thelotrema actinotum, 12 

adjectum, 39 

album, 12 

awn ula tum, 43 

aquilinum, 52 

auberianum, 35 

auratum, 41, 54 

butuanum, 43 

calvescens, 15 

cavatu m, 15 

cavatum bar. planius, 28 

chionost om it m, 16 

citrinodiscum, 41 

clandestinum, 41 

clandestinum f. remanens, 33 

comparabile, 16 

concretum, 41 

concretuin var. subcaesium, 41 

conforme, 42, 45 

consanguineum, 42 

conveniens, 42 

cryptotrema, 43 

develatum, 50 

disciforme, 52 

dorningense Tar. fecundum, 21 

elliottii, 47 

erumpens, 43 

exanthismocarpum, 21 

excnvatum, 28 

excavatum var. impressuliim, 28 

foraminulositm, 55 

glaucopallens, 43 

glaucophaenum, 21 

glaucitlum, 23 

glyphicitm, 36 

glyphidelluni, 35 

hiatum, 44 

homopastoides, 43 

homothecium, 21 

interpositurn, 30, 45 

in terva lla t um, 41 

isertii, 21 

laevigans, 43 

laevigans var. avertens, 43 

laeviusculum, 52 

latilabrum, 24 

leiostomum, 36 

lepadinum, 44 

lepadinum var. bahianum, 50 

lepadodes, 52 

leprocarpum, 44 

leptopo?um, 23 

leucomelaenum, 44 

leucomelaenum var. cathomalizans, 43 

leuconzelaeuitin var. elevatum, 44 

leucomelaenum var. sf!lotheciiirn, 18 

leucotylium, 15 

maximum, 45 

meiospermum, 36 

metaphoricuni, 53 

microporellum, 24 

microporoides, 41 

rnicrosporoides, 53 

monosporum f. album, 52 

monosporum var. patztlum, 52 

myriocarpum, 45 

myrioporoides, 45 

myrioporuni, 12 

myriotrema, 12 

norsticticum, 47 

obtiiratum, 16 

occlusum, 47 

oculatum, 41 

opacum, 25 

pachystomum, 26 

papillatum, 26 

pauperius, 44 

pechueli, 43 

peTfo ra t u m, 26 

phaeospermoides, 36 

platycarpellurn, 12 

plurifarium, 24 

porinoides, 21 

porphyrodiscum, 41 

postpositum, 45 

praestans, 47 

prevostianum, 55 

pycnoporellum, 47 

rarotongae, 52 

reclusum, 54 

reconditum, 29 

rudiits var. dominicanum, 50 

rugatulum, 42 

secoligella, 43 

stylothecium, 48 

subcaesium, 41 

subcavatum, 31 

subconcretum, 55 

subconforme, 45 

subcrassulum, 12 

sitblilacinum, 31 

suhpraestans, 48 

siibterebrans, 55 

sit btere bratztm, 24 

tenue, 48 

terebratitm var. abbreviatitlum, 28 

terebratzclum, 33 

trypaneoides, 54 

urceolare, 55 

vagum, 31 

verrucosum, 33 

viridialbum, 12 

wrightii, 49