SMITHSONIAN CONTRIBUTIONS TO BOTANY NUMBER 16 Morden-Smithsonian Expedition to Dominica: The Lichens (Thelo trema t aceae) Mason E. Hale, 3r. SMITHSONIAN INSTITUTION PRESS City of Washington 1974 ABSTRACT Hale, Mason E., Jr. Morden-Smithsonian Expedition to Dominica: The Lichens (Thelotremataceae). Smithsonian Contributions to Botany, number 16, 46 pages, 20 figures, 1974.-A revision is made of the lichen family Thelotremataceae in Dominica, based on previously published records by Elliott and on collec- tions by the author. The family comprises 48 species in 4 genera, Leptotrema, Ocellularia, Phaeotrema, and Thelotrema. The following 15 species are de- scribed as new: Ocellularia antillensis, 0. conglomerata, 0. dominicana, 0. ma- culata, 0. mordenii, 0. nigropiinrta, 0. rimosa, 0. yorediata, Phaeotrema aggre- gatum, P. obscurum, Thelotrema confusum, T. dominicanum, T. papillosum, T. tenue, and Leptotrema deceptum. Three new combinations, Ocellularia fe- cunda (Vainio) Hale, Phaeotrema disciforme (Leighton) Hale, and Leptotrema occult um (Eschweiler) Hale, are also made. Morphological characters are reviewed and the chemistry of each species is presented in detail. The family is exceptionally well developed in mature rain forest. OFFICIAL PI~BLICATIOK DATE is handstamped in a limited number of initial copies and is recorded in the Institution?s annual report, Srnithsonian Year. SI PRESS NUMBER 5043. SERIES COVFR DESIGN: Leaf clearing from the katsura tree Cercidiphyllum juponicum Siebold and Zuccarini. Library of Congress Cataloging in Publication Data ~~ Hale, Mason E- Morden-Smithsonian Expedition to Dominica: the lichens (Thelotremataceae). (Smithsonian contributions to botany, no. 16) Bibliography: p. 1. Thelotremataceae. 2. Lichens-Dominica, I. Title. 11. Series: Smithsonian Institution. QKl.S2747 no. 16 [QK585.T5] 581?.08s [589?.1] 73-22465 Smithsonian contributions to botany, no. 16. For sale by the Superintendent of Documents, US. Government Printing Office Washington, D.C. 20402 Price $1.05 (paper cover) Contents Page Introduction ...................................................... 1 Morphological Characters ................................ Thallus Structure ............................................... 2 Apothecial Characters ...................... Spores ............................................ Generic Limits ........ ............... ................. Chemistry ............................................ Ecology and Habitats ........................................ The Thelotreme Flora .......................................... 13 Collecting Localities ..................................... Ocellularia ............ ................................ 15 Phaeotrema .................................................... 28 28 Thelotrema ...................................................... 31 31 Leptotrema .......................... ................... 38 38 Literature Cited .............................. ....... 43 Index ........................................................... 46 Key to the Species of Ocellularia .................................. Key to the Species of Phaeotrema ................................. Key to the Species of Thelotrema Key to the Species of Leptotrema ................................ ................................ iii Morden- Smi t hsonian Expedition to Dominica: The Lichens (Thelotremataceae) Mason E. Hale, Jr. Introduction The Thelotremataceae comprise a very large family of crustose lichens that has attracted the attention of many lichenologists, beginning with Acharius (Hale, 1972). They have unique crater- like apothecia, but this characteristic feature can usually only be seen under magnification. As a matter oi fact, these inconspicuous lichens will oEten be overlooked in the field unless a hand lens is used. Furthermore, because of the small size of the apothecia, very few species can be rec- ognized at sight. All must be collected and ex- amined later for morphological and chemical characters. Thallus color, however, is rather con- stant, light greenish or ashy gray. Only a few species have the deep green color of the equally common pyrenocarpous lichens, which, together with the Graphidaceae and Thelotremataceae, account for the majority of crusts that are collected in low- land tropical regions. This paper is part of an on-going project on a modern treatment of the lichen flora of Dominica. As I mentioned in the first part on the Parmelia- ceae (Hale, 1971a), where climate and topography are discussed, Dominica has the greatest remaining areas of virgin or relatively undisturbed forest of all the islands in the Lesser Antilles. Though Mason E. Hale, Jr., Department of Botany, A'ntional Museum of Natural Histoy, Smithsoninn Institution, Washington, D.C. 20560. easily accessible by car and well-kept trails, these forests have as yet suffered little from human ac- tivity. Logging operations in the Dleau Gommier Forest Reserve, for example, have ceased, at least temporarily. Other islands in the Antilles are by contrast much more intensively cultivated and un- disturbed forests are confined to higher, often inaccessible mountain slopes. On my first trip to Dominica in 1969 I was in- terested primarily in foliose lichens, which proved to be relatively poorly developed. Many of the numerous crusts that I collected more or less ran- domly and blindly were actually thelotremes. The obvious richness of the family inspired me to un- dertake a second trip to the island in December 197 1, once again under the Morden-Smithsonian Expedition, to concentrate exclusively on this fam- ily. A final brief trip for the same purpose was made in May 1972 under National Geographic Society support. Altogether, nearly 550 specimens of thelotremes were collected, making this one of the most intensive collecting programs ever under- taken for a crustose lichen group. Special thanks are due Mrs. William J. Morden, who supported most of the field work for this project. Assistance is also gratefully acknowledged from the National Geographic Society for a broader study OC the evolution of the Thelotremataceae in the Lesser Antilles, which included Dominica. Curators at a number of museums and universities have sent loans of type-specimens, and their help 1 2 SiMITHSONIAN CONTRIBUTIONS TO BOTANY and patience are deeply appreciated: Dr. T. Ahti (H), Dr. Reino Alava (TUR), Dr. Ove Almborn (LD), Dr. C. E. Bonner (G), Mr. hl. Skytte Chris- tiansen (C), Mr. Peter James (BM), Mme Jovet- Ast (P), Dr. I. hl. Lamb (FH), Dr. R. A. Maas Geesteranus (L), Prof. H. Merxmuller (M), Dr. Harald Riedl (W), and Dr. R. Santesson (UPS). I wish also to thank Dr. Chicita Culberson for help in identitying or verifying most of the chemi- cal substances of the various specimens. Morphological Characters After curating and labeling each specimen, I studied the various morphological characters that appear to be most important for taxonomic pur- poses. The descriptions, therefore, are not ex- haustive but should be adequate for species identification. Redinger (1936 ) and Salisbury (1972a, 1972b) have drawn up useful summaries of the essential features of this family and a num- ber of the species which may also be consulted. THALLUS STRUCTURE The anatomy of crustose lichens is not as com- plex as that found in typical foliose lichens. Corti- cal structure is less distinct, and, lacking a free lower surface, the medullary hyphae usually pene- trate the outer layers of bark periderm (or rock substratum). The algal layer (Trentepohlia) may also be indistinct. Large colorless crystalline inclu- sions are commonly found in the medulla, as in Ocellularia nigropuncta, new species, and Thelo- trema interpositzim. Sections examined under a scanningelectron microscope show some of these structures vividly. Ocellularia olivacea, 0. perforata, and Thelotrema confusum, new species (Figure la,b,c), all with well-developed epiphloeodal thalli, have a cortex consisting of several layers of heavily gelatinized cells appearing almost paraplectenchymatous; the loosely organized medulla penetrates the periderm. The surface is essentially smooth and featureless (Figure 2a-d). In some species the cortex is similar but peculiar aculeate hyphae protrude from the surface in greater or lesser abundance (Figure 3a,b). They appear to be collapsed, perhaps be- cause of vacuum treatment needed for specimen preparation. This aculeate orientation, reported here for the first time, is characteristic of Myrio- trema species (Ocellularia terebratula, Thelo- trema clandestinzim) as well as a few species with emergent apothecia (T. depressum, T. praestans). I have observed similar orientation in several species of Graphis and Graphina. Most thelotremes, however, have an irregularly organized surface, as in 0. mordenii, new species (Figure 3c,d), and a poorly defined cortex. Leptotrema wightii has an unusual cortex pre- viously described by many lichenologists. The thallus is very thick (up to 1 mm) and has a grainy surface (Figure 54. The cortex is arranged in tall columns with algae scattered vertically (Salisbury, 1971) (Figure Id). The so-called hypophloeodal species lack the structures described above. The external thallus is reduced to a microscopic layer of hyphae spread thinly over the bark surface (Figure 4), and these hyphae penetrate several layers of periderm with very sparce symbiotic algae apparently located just below the periderm surface. Representatives of this group from Dominica include Ocellularia con- color, 0. pyrenuloides, Phaeotrema disciforme, new combination, Thelotrema leucomelaenum, and T. tenue, new species. APOTHECIAL CHARACTERS Several morphologists have studied the internal structure and probable ontogeny of ascocarps in the Thelotremataceae (Redinger, 1936; Johnson and Brown, 1941; Letrouit-Galinou, 1966). The most commonly studied species is Thelotrema lepadi- num, which has a distinct exciple breaking away free from the apothecial wall. The following DO- minican species have more or less similar struc- ture: Ocellularia conglomerata, new species, 0. dominicana, new species, 0. exanthisrnocarpa (Fig- ure 5a), Leptotrema occultum, L. subcompunctum, and Phaeotrema aggregatunz, new species. I will call this arrangement ?lepadinioid? in the descrip- tions; Salisbury (1972a) uses the term ?Thelotrema lepadinum group.? What appears to be an extreme stage of this configuration of apothecial wall and exciple is seen in the Chroodiscus-type apothecium (here termed ?chroodiscoid?). The outer wall or margin is erect and becomes recurved (Salisbury, 1972b), almost as in Geaster, and the exciple usually persists as a NUMBER 16 3 FIGURE 1 .-Cross-sections of epiphloeodal lichens viewed with a scanning-electron microscope (a, b, and c, sectioned obliquely; d, Fertically): a, Ocellularia olivacea (Hale 35226) (X 1000); 6, 0. perforata (Hale 38145) (X 1000); c, Thelotrema cotzfusum, new species (Hale 37697) (X 500); d, Leptotrema ulightii (Hioram 5850 from Cuba) (X 500). second inner rim. The disc is widely exposed and foliicolous species of the Thelotremataceae, in- nothing resembling the discrete pore characteristic cluding in Dominica Ocellularia alborosella and of the other thelotremes remains (Figure 5b). This 0. clilatata. It is callcd the ?Tlzelotrema platy. type was first recognized among the foliicolous cui-pzim group? by Salisbury (1972b). In the ab- lichens, but it seems identical to that in the non- sence of more detailed ontogenetic research, 4 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 2.-Surfaces of epiphloeodal lichens viewed with a scanningelectron microscope: a, b, Ocellutaria nigropuncta, new species (Hale 35365) (X 500 and X 2000); c, d, 0. dominicana, new species (Hale 37967) (X 500 and X 2000). however, we cannot yet say that Chroodiscus is a good genus, that the Chioodisczwlike nonfoliico- lous thelotreines should be transferred to Chroodis- cus, or that Clzioodiscus should be transferred to Ocellulaiiu. Chroodiscoid species, incidentally, are distributed among the four presently recognized genera, Ocellzih in, Leptotrema, Phaeotieinu, and Theloti emu, on the world level. The majority of species in the Thelotremataceae are neither lepadinioid or chroodiscoid. The ex- ciple is fused with the receptacle wall and the main rim or margin closes over most of the disc NUMBER 16 5 FIGURE 3.--Surfaces of epiphloeodal lichens viewed with a scanning-electron microscope: a, b, Ocellularia olivacea (Hale 38054) (X 500 and X 2000); c, d, 0. mordenii, new species (Hale 35764) (X 500 and X 2000). leaving only a small constricted pore. The upper ized apothecia and a pore flush with the thallus wall is usually carbonized. There are, broadly surface (Figure 5c,d). The apothecia are always speaking, three groups that can be recognized for quite small, 0.1-0.3 mm in diameter. Typical practical purposes on the basis of degree of emer- examples in Dominica are Ocellularia olivacea and gence from the substratum. One, represented by Thelotrema clandestinum. the ?Myriotrema? type, has immersed noncarbon- A second group, which includes the majority 6 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 4.-Surface and cross-sections of hypophloeodal lichens viewed with a scanning-electron microscope: a, Thelotrema Zeucorneiaenum, surface view (Hale 39491) (X 2000); b, T. leuco- melaenurn, cross-section (Hale 35491) (X 500); c, d, T. tenue, new species (Hale 35430) (X 500 and X 2000). of the species, has emergent apothecia, neither X third group, typified by 0cellular.ia rho- basally constricted or strongly raised (Figure 5e,f). dostroma and 0. cavata, has strongly emergent apo- They are usually 0.4-1.0 mm in diameter, although thecia Tvitli a partially constricted base (Figure in a few species (for example, Thelotrema praestans 55). The pore is small and sometimes annulate or in Figure 17e) nearly 2 mm in diameter. open at maturity, as in 0. fecunda (Figure llh,i). NUMBER 16 7 FIGURE 5.-Types of apothecia in the Thelotremataceae (all X 10): a, Ocellularia exanthis- niocarpa (type of 0. multilocularis); b, 0. alborosella (Hale 353ila); c, 0. olivacea (Hale 35243); d, Leptotrema wightii, (type of L. prevostianum); e, 0. perforata (Hale 37i27); f, L. bahianum (type of Thelotrema rudius var. dominicanum); g, 0. rhodostroma (Hale 38122); h, Phaeotrema leiostomum (Hale 37956); i, 0. coinparabilis (Hale 37372). Diameter varies between 0.6 and 2.0 mm. This Although Kylander continued to use Ascidiunz group was called Ascidium by Fee (1824), not so for strongly emergent species, no modern lichen- much because of emergence but because of a dupli- ologists have employed it. The type of the genus, cate membrane in the perithecium (the nature of A. cinchonarum Fee, is synonymous with Ocellu- which is not clear to me) and a marginate pore. hia cavata. Naturally there are intergradations 8 S.\fITHSOSIAN COXTRIBUTIOXS TO BOTANY walls and periderm, acting a5 a kind of ramrod. Not all species with periderm inclusions have a columella, but as Redinger found for the Brazilian species the correlation is high. The range in deLelopment of a coluinella is very great (Figure 6), from the Tveak thin columella in Ocellzilal-ia mordenii, new species, to the multiple act i n oi cl 0 ce 1/21 la ?.in co mpnra b i 1 is (Figure 5i) and Leptotrema fisszim (Figure 19a,c). The most highly developed species is Ocellzi Inria glnziovii (Muller-A4rgau known from South America Vainio (1921: 184) recognized these species under a separate genus, Rlinbdodzsczis. s t r uc t iires of between the Ascidium-type and the more common emergent type, but I believe the descriptive term ?ascidioid? is still useful. A final group forms a transition to the Graphi- daceae. The apothecia become irregularly elongate with a lirelliform dehiscence. In Dominica this includes Phaeotrema leiostomum, which can pro- duce both round and elongate apothecia (Figure 511). Other species from South America have this trait developed much more conspicuously and have been described in the Graphidaceae (as Phaeo- graphis chionodisca Redinger (1936:69)). They should probably be transferred to the Thelotrema- taceae. Redinger (1936) further divides the Thelotre- mataceae into two groups that differ in position of the ascogonial initial. In one group the initial is located on or just below the periderm surface so that at maturity the apothecial wall develops on and apart from the unbroken periderm (for example, Ocellzilaiia rlzodostroma and Phaeotrema disciforme, new combination). The other group, including most nonchroodiscoid species in Do- minica, has ascogonial initials deeply embedded in the periderm. The maturing apothecia must break through this layer to reach the surface. Several layers of periderm cells are thereby included in the apothecial receptacle and can be easily recog- nized in sections (Figure 6f). This character de- serves much more detailed study. Development of a central cylindrical columella is particularly characteristic of the Thelotrema- taceae, and for the most part presence or absence of a columella is a good species character (Figure 6). As a rule, columellate species do not have per- fect counterparts that lack a columella. Noteworthy exceptions in Dominica are Oce11zilaria papillata and 0. perfomto, each of which has a large popu- lation where a columella is clearly developed and a smaller one where it is either absent or very difficult to find. I have not recognized such ecolumellate species as new without correlating morphological or chemical characters. Columellate species always have heavily carbonized apothecial walls. Redinger (1936:6) proposed an interesting hy- pothesis that the columella plays a role in breaking open the periderm layer in species that develop deeply embedded. As the apothecia mature, the columella pushes up against the upper carbonized SPORES Spore size is an importar,t and relatively con- stant character. The range of spore length, for example, is usually quite small except for very large spores (more than 10Op) which may vary between 100~ and 300u. ?IVe can distinguish four very broad categories among the Dominican species: (1) spores less than 20~ long with 4-6 transverse locules, eg., Lcplot~enza wightii, Ocelliilnria tere- bi.atiila, Tliclotwnza clandestinum; (2) spores 20y-36;t long with 6-10 transverse locules, e.g., Ocellzilarin perfoycita (Figure 7a), Thelotrema leiiconielaeizi(ln; (3) spores 40p-9Op long with 12-24 transverse locules, e.g., Leptotyema decep- turn, new species (Figure 5d), Ocellzihria exanthis- mocarpa; and (4) spores 80p-300p long with numerous locules, eg., Ocelliilnria rhodostyoma (Figure fb), Thelotrema praestans, T. tiiberculi- f e I? ii m. The number of longitudinal septations in Leploti~ema and Theloti.ema varies according to length, from 1 or 2 septae in small spores (up to about 30p long) to 4-6 or more in large spores. Spores are colorless or brown. It is sometimes a problem to determine color in large-spored species of Leptotrema and Thelotyema where, on the one hand, a Thelotwma species might have some senile spores turning brown and on the other a Leptotrema will haye some colorless spores (Fig- ure 7d). Jn Phaeotwma disciforme, new combina- tion, in fact, one often finds a mixture of color- less, mature brown, and shriveled brown spores (Figure 7c). Shriveling (Figures 7c,4 is character- istic of brown-spored species; I know of no cases where Thelotrema or Ocellularia spores shrivel. NUMBER 16 FIGURE 6.-Cross-section of apothecia (all about X 130): a, Ocellularia nigropuncta, new species (Hale 35365); b, 0. inordenii, new species (Hale 37764); c, 0. cavata (Hale 35134); d, Phaeotrerp leiostoniun (Hale 35203); e, Thelotrema leuconlelaenuin (Hale 35491); j, T. tenrte, new species (Hale 35430). 9 10 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 7.--Spores of the Thelotremataceae: a, Ocellularia perfornta (Hale 37784) (X 400): b, 0. rhodostroma (Hale 35440) (X 130): c, Phaeotrema discifortne, new combination, with a mixtute of colorless spores, brown spores, and shii>eled brown spores (Hale 37939) (X 400); d, Leptotrenia deceptutn, new species, with colorless muriform spores on the right (Hale 38110) (X 400). In any event, transitional species should be judged nation, or 0. sitbcaunla on Dominica) should not intliviclually, and the presence of a few anomolous be totally unexpected; nor shoultl these isolated brown-spored specimens among many colorless cases prejudice the value of spore color as a generic specimens (as in Ocellzrlnria feczinda, new conibi- character. NUMBER 16 Generic Limits The generic delimitation of the Thelotrema- taceae has undergone many changes since the classic work of Acharius. Fee (1824), for example, established two genera, in addition to recognizing Theloti enla, based on external features and apo- thecial characters: Ascidizim and My? iotrema, both presently considered to be synonyms of Ocellzl- la, fa. lfullei--Argau (1887a) proposed the most far- leaching changes, using spore septation and coloration to separate four genera (in addition to foliicolous Chq oodisctis): Ocellzilaria (trans- trersel y septate colorless spores), Phaeotrema (transFersely septate brown spores), Thelotrema (muriform colorless spores), and Leptoti emu (mu- riform brown spores). TVhile Vainio, one of the truly creative thinkers in lichen classification, rejected many ?spore genera,? almost all authors since Muller-Argau ha1 e accepted these genera (and, indeed, spore- based genera in other families). Poelt (1973:613) refers to these as ?artificial genera,? yet almost an) character we select is in a sense artificial. I don?t believe this term does justice in that it implies spore characters are artificial. TVe are, in reality, faced with at least thiee choices for generic delimitation in the family: (1) one genus (Tlielotremcr ) to accommodate all the- lotreinataceous species; (2) four spore-based genera (Ocelliilnria, Phaeolrema, Thelotrema, and Lep- tot)en70); (3) five or more genera (Ascidi~m, C 11 rood isc 11 Y, Myio t I em n, R h 0 b dod i ~c us, antl T 11 e lo- tren10) based on apothecial emergence and excipu- lar structures. Most lichenologists, as I said, have opted for spore-based genera. TVhile a majority opinion does not necessarily prove the naturalness or validity of such genera, they do repiesent one possible biological differentiation, broadly corre- lated with chemical, morphological, and ecological characters (as I will discuss briefly under each genus). I cannot agree with Salisbury?s (1952a) recent rejection of spore-based genera in favor of a single massi\e genus Tl7elotrema with 400 species sub- divided into ?groups? according to excipular struc- ture, a character that is difficult to identify. His kejs and descriptions show how cumbersome this approach is, for in each instance one must desig- 11 nate the spore type, e.g., TIzelotiemu wightii with Leptot)ema-type spoies, etc. If- one is forced to make this critical distinction every time a species is identified and described, then they should be tieatetl as distinct genera, a more coherent antl certainly a inore pragmatic, working solution. Spore characters are well proven, even critical, in the taxonomy of noiilichenized fungi. A retrogres- sive gene1 ic concept in lichens will hardly encour- age a better understanding of their biology and evolution. Mj experience in the Thelotremataceae suggests that spore-based genera are reasonably natural, easily manageable units that are probably diverging in terms of habitat requirements and biogenetic characters as reflected in chemical dif- ferences. Chemistry in aliiiost eier) other gioup of lichens studied, chemistr) is an impoi tant and fundamental char- acter in uncle1 standing the evolution and popula- tion biolog) ol the Thelotremataceae. The family has an unexpectedly iich chemistry with a pre- tloniinance of P + depsidones and related com- pounds (Culberson and Culberson, 1968; Culber- son and Hale, 1973). I Trill discuss the possible significance of this chemistry below under the species and hope to exploi e the broader aspects oh biochemical evolution of the family in another paper. -411 of the specimens were initially tested with p-phen) lenetliaiiiine and chromatographed on Brinkinan thin-1a)er plates according to the P re- action. I ~roultl recommend, however, that lichen- ologists consider using this reagent as little as possible because of the well-known carcinogenic propel ties of rolatile amines. Identification of the lichen substances can be made just as easily with TLC rrithout recouise to a P test except in doubt- ful cases. Standard thin-layer solvent systems (hexane- ether-formic acid, 130:80:20 v/v, and benzene- dioxane-acetic acid, 180:45:5, I./.) xvere used to identify the chemical content of each specimen. NOTATIC Acr~.-Ocellzrlaria feciinda, new combina- tion, (hypoprotocetraric acid predominating), 0. rhodost 1 om0 (4-O-demethylnotatic acid predomi- nating), antl Pliaeotremu leiostomzim (4-O-deme- HYPOPROTOCETRARIC ACID ASD 4-o-DEhIETHYL- 12 SXITHSONIAN COSTRIBUTIONS TO BOTANY and Thelotrema dominicanuna, new species. Sev- eral species of Ocellularia and this single Thelotrema react Pt red, but neither protocetraric acid or fumarprotocetraric acid can be demon- strated with TLC. Instead there are at least t\vo gray spots above both of these acids on the plates (and often a high H,SO, yellow spot). These are apparently derivatives of protocetraric acid. ?PRAESTASS? UsKsowN.-Thelotrema praestans. This unidentified depsidone reacts P+ red orange and forms a very distinct orange to reddish brmvn spot below psoromic acid in both solvents. ?OLIVXCEA? USKNO~t.l\.S.-OCellllla~~a olivacea. This P- compound produces a distinct H2S04 gray spot below lecanoric acid in both solvents. It may be a depside. TVhile the lectotype of Ocel- lzilnria olivacea contains only this one spot, all West Indian collections, otherwise morphologically in- distinguishable, contain a lower gray spot in addi- tion. All specimens have thick white acetone residues. OTHER P - UsK~o~t.n.s.-Ocellziluria rimosa, new species, and 0. sorediatn, new species. The uniden- tified spots in these species may or may not repre- sent lichen substances. Some species were examined for fatty acids by dipping the plates in water and trying to detect white water-repellant spots as the water evaporated. Some spots resembling fatty acids Tvere seen but microchemical tests with G.E. and G.A.TV. xvere unsuccessful. Another major group of P- compounds is terpenes. All TLC plates shou.ed numerous terpenes, identifiable as purple, blue, or brovm spots falling generally above RF .5 and fading Tvithin a few hours. Probably all of these originate from the bark and are not lichen products, as can be proved by co-chroinatographing the lichens Jvith lichen-free bark samples from the same specimen, 0. concolor, 0. dominicnnn, new species, 0. macu- Into, new species, 0. mordenii, new species, 0. pnpillata, Phaeotiwnn aggregntum, new species, P. d iscif o im e, new com bi na t i on, T 17 e 1 o t re m a d ep res- slim, T. leticomelnenum, T. papillosum, new species, T. tenue, new species, and T. tuberculi- ferum. PicsrENTs.-OcellziInria rnordenii, new species and Leptotrema wightii (anthraquinones) and 0. fecunda, new combination, 0. rhodostroma, and Phaeotremn leiostomum (K- yellow compounds). h?0 SUBSTASCES PRESENT.-OCel/Zilnria albOYoSelk7, thylnotatic acid sometimes lacking). These acids were recently examined in detail by Culberson (Culberson and Hale, 1973), who discusses the biogenesis. Ocelluiarza fecunda and 0. rhodostroma were not included in that study. NORSTICTIC AcIn.-Leptotyenaa occultum. This is a rare acid in the Thelotremataceae (especially in contrast to the Graphidaceae) and is largely con- fined to Leptotrema on the world level. PROTOCETRARIC AcIn.-Ocellularia nigropuncta, new species, 0. perforata, Leptotl-emu baliianum, and Theloti~ema confusum. This well-known P + red acid is easily identified on TLC plates, It is sometimes accompanied by a higher gray spot that I have termed the ?amplior? unknown because of its constant occurrence in Ocellula~ia amplior (Ny- lander) Redinger. Culberson believes it to be pro- tocetraric acid combined with a carboxylic acid, comparable to funiarprotocetraric acid or the re- cently described succinprotocetraric acid. PsoRoarIc AcID.-Ocellulaiia antillensis, new species, 0. conipaq.abilis, 0. subcavata, 0. terebra- tula, Leptotrema fissum, Thelotrema carassense, T. clandestinum, T. conforme, T. leucinum, and T. interpositurn. This is the commonest P+ acid in the Thelotremataceae. It is almost alivays accom- panied by an unknown compound ?conpsoromic acid,? appearing as a spot below psoromic acid. It, too, is P+ yellow. STICTIC AND CONSTICTIC Acins.-Ocellzilnria con- glomerata, new species, 0. dilatata, 0. exanthismo- carpa, 0. pyveniiloides, Phneotrema obscurzim, new species, Tlielotwma glaucopallens, Leptotrema dc- ceptum, new species, L. microglaenoides, and L. szibcomptinctiim. This widespread and well-known acid is actually rather rare in the Thelotremataceae except for the genus Leptotrema. Constictic acid, appearing as a H,SO,+ yellow spot just below salazinic acid, is a constant accessory compound. One or two other gray spots, as yet unidentified, may occur between stictic and constictic acids on TLC plates. Another rarer accessory is the highest H,SO, t orange-red spot in the ?quintaria? series (Hale, 19Tlb), still unidentified but being discov- ered in a variety of genera. VIRENSIC Acm.-Leptotrema spondaicum. This acid is PS ?red. It was previously unknown in the Thelotrcmataceae, having been first identified in Alectoria. ?CINCHONARUM? UNK~own.s.-Ocellzilaria cavata NUMBER 16 13 zone lvhere rain I?orest is optimally developed, the trees being 30-40 m high (Figures 2 and 3 in Hale, 19Sla, illustrate this forest type). It would appear that this lichen family is totally dependent on undisturbed rain forest as a primary habitat and site for speciation. When this forest is de- stroyed, it seems inevitable that the lichens asso- ciated with it will become extinct. In Dominica, I have seen only two species able to invade dis- turbed or planted forests; namely, Leptoti.ema spondaicum and Ocellularia exanthismocarpa. They have a highly successful means of propaga- tion and become established very quickly. The anthraquinones are generally produced in the thallus, while the yellow compounds occur in the medulla of the apothecia and not in the thallus. Ecology and Habitats I paid considerable attention to the ecology of the species and habitat requirements in Dominica in the hope that this information would help in understanding the distribution and evolution of the species on the island. Lichenologists have all too often prepared revisions of tropical lichens on the basis of preserved herbarium specimens alone with little if any opportunity to observe the species in the field. Labels on specimens collected by non- professionals rarely give pertinent information on ecology and habitat. For example, tropical rain forests have distinct layering and great vertical variation. in microclimate. Elevational differences are also important and once again such data are rarely given in full on herbarium labels. Base level species in Dominica form a distinc- tive group. They grow on roots, lianas, buttresses, prop roots, and saplings within 2-3 m of the forest floor. Characteristic species are Ocellularia concolor, 0. papillata, 0. perforata, Thelotrema glaucopallens, T. tenue, new species, and Lepto- trema deceptum, new species. At the opposite ex- treme we find canopy species that occur high above the forest floor in tree tops, exposed to great changes in insolation and wetting-drying cycles. These include Ocellularia mordenii, new species, 0. olivacea, 0. subcavata, 0. terebratula, Phaeo- trema leiostomum, Thelotrema clandestinum, and T. praestans. Other species on Dominica have less stringent habitat requirements or have been col- lected too infrequently to categorize. On a broader scale altitudinal differences are also pronounced. The rain forest becomes stunted in Dominica above about 800 m and gradually merges into an elfin mossy forest best developed above 1000 m. Thelotremes become scarce in this wind-swept zone where rain falls almost every day. Two species, however, Thelotrema leucomelaenurn and T. tubewuliferzim, are restricted to the highest elevations. Ocellularia rhodostroma and Thelo- trema tenue, new species, occur here as well but also at lower elevation. The vast bulk of thelotreme collections were made between 300 and 800 m on Dominica, the THE THELOTREME FLORA As I had found with Paimelia. (Hale, 1971a), almost all previous collections of the Thelotrema- taceae on Dominica were made by 1%?. R. Elliott in the 1890s. H. A. Imshaug also made about ten collections in 1963 which are preserved at Michigan State University. I have not had access to these. Elliott collected five specimens, identified by Vainio (1896, 1915) as follows: Thelotiema excavatum Vainio var. impressulum Vainio (hlorne Anglais, the type-locality, Elliott 160, specimens in Bill and TUR). This is synony- mous with Ocellularia perforata. Thelotiema leucomelaenum var. elevatum Vainio (Morne Anglais, Elliott 1534, specimen in TUR). This is T. leucomelaenum. Theloti emu rhodostromzim (hlontagne) Vainio (Morne Anglais, Elliott 521, specimens in BM and TUR, and Prince Rupert, Elliott 1305, specimen in TUR). These are Ocellularia rhodostroma. Thelotiema rudius Vainio var. dominicanum Vainio (Prince Rupert, Elliott 1303, specimen in TUR). This is the same as Leptotrema bahianum. The thelotreme flora of Dominica now includes 48 positively identifiable species (and three addi- tional species represented by poor material). I col- lected all of these except for Leptotiema bahianum. No one locality, of course, had anywhere near this number of species (see p. 14, ?Collecting Locali- ties?). The richest area was one site (16b) in the Dleau Gommier Forest Reserve which was being logged in December 1951; I collected a total of 19 species there over an area of about 3 hectares. An adjacent site had 14 species as did the trail to Morne Anglais. Dleau Gommier locality 15 had 14 SMITHSONIAN CONTRIBUTIONS TO BOTANY 12 different species. A logged rain forest on the Layou Road (12) had 11 species. Five other locali- ties (11, 16c, 18a, 21, and 24a), all between 350 and 750 m, had 9 species each. The homogeneity of the flora is remarkably high. Locality 16, for example, contains three adjacent collecting sites. Out of a total flora of 21 species there, 8 were found in common at each site. Three occurred at two of the three sites and 10 were collected at only one of the three sites. The great- est variance in homogeneity obviously comes about when we collect at two very different elevations or sites where habitat conditions are significantly different. It should be evident that we have not yet in- ventoried the whole thelotreme flora of Dominica. I would estimate that there are at least 55 species (including the three unidentified collections as possible additions). It would require at least one more intensive collecting trip, possibly two, to reach this number. COLLECTING LOCALITIES Specimens were collected at 28 separate localities on the island (Figure 8), and multiple visits were made at different times to six of these, giving a total of 35 collecting sites. These are coded as follows for convenience in citing specimens, going from localities at low elevation in the scrub forest to those at highest elevations. 1. Scrub forest, Rodneys Rock, sea level. 2. Scrub forest near Mero, elevation 15 m. 3. Scrub forest on slopes of Barbers Block, ele- 4. Scrub forest near Calibishie, elevation 15 m. 5. Palm forest at Rosalie Bay, elevation sea level. 6. Rain forest along trail to Madjini, elevation 7. Virgin rain forest logging area, Newfound- 8. Rain forest above Newfoundland, elevation 9. Rain forest logging area, Castle Bruce Road, 10. Dleau Gommier rain forest near Emerald 11. Felled rain forest along new road cut about vation 60 m. 30 m. land, elevation 250 m. 300 m. elevation 300 m. Pool, elevation 450 m. 2 km northwest of Pont Casse., elevation 430 m. 12. Rain forest logging area on Layou Road, ele- vation 420 m. 13. Mixed secondary forest logging area at Brant- ridge Estate, elevation 500 m. 14. Rain forest logging area just east of Pont Casse., elevation about 550 m. 15. Virgin rain forest area at Dleau Gommier For- est Reserve, elevation 500 m. 16a, 16b, and 16c. Virgin rain forest logging at Dleau Gommier Forest Reserve, elevation 350-370 m. 17. Remnants of rain forest at Felicite, elevation 370 m. 18a and 18b. Remnants of rain forest along road to Jean Estate, elevation 600-700 m. 19. Citrus grove at Laudat, elevation 400 m. c 172128 L Scotts Head 4+-./0-5 FIGURE 8.-Outline map of Dominica showing localities where specimens of Thelotremataceae were collected. NUMBER 16 20a and 20b. Remnants of rain forest in pastures above Giraudel, elevation 550-600 m. 21. Virgin rain forest at Syndicate Estate, eleva- tion 650-700 b. 22. Mixed rain forest along trail through Middle- ham Estate, elevation 650-700 m. 23. Rain forest at the base of Trois Pitons, ele- vation 650-700 m. 24a and 24b. Mixed secondary forest along trail to Boiling Lake, elevation 650-700 m. 25, and 25b. Rain forest-mossy forest along trails at Fresh Water Lake, elevation 800-900 in. 26a and 26b. Rain forest-mossy forest along trail up Morne Anglais, elevation 800-1000 m. 27. Stunted rain forest-mossy forest on slopes of Trois Pitons, elevation 900-1200 m. 28. Mossy forest area on slopes of Morne Dia- blotin, elevation 1000-1200 m. Sixteen localities were visited on the first trip in January 1969 (1, 2, 5, 6, 7, 13, 14, 15, 17, 18b, 19,'20b, 24b, 25b, 26b, and 28). On the second trip seven additional localities were found (3, 11, 12, 16a-c, 22, 23, and 27) and five sites adjacent to localities previously were examined (18a, 20a, 24a, 25a, 26a), all during December 1971. Four new localities were visited in May 1972 (8, 9, 10, 21). All specimens are deposited in US unless other- wise indicated and duplicates will be deposited in the British Museum (BM). 15 I have provided illustrations of the Dominican species as well as of most type-specimens for com- parison. These photographs also serve to show the range of variation in pore size and apothecial emergence. I have found these X 10 photographs to be extremely useful in identification, more so than line drawings of cross-sections. All specimens, including the types listed, were tested with thin- layer chromatography. Ocellularia This is the largest genus in the family, compris- ing about 165 species on the world level or about. 40 percent of the total 400 known species. I had almost the same percentage on Dominica (22 of 48 species) but it makes up a disproportionately large percentage of the specimens actually col- lected, 410 out of 530 (77y0). Ecologically the genus is best adapted to mature rain forest and cannot withstand extremes of very dry lowland and wet exposed high elevation. The ten common- est species (Tvith number of specimens collected in parentheses) are: 0. yhodostroma (93), 0. fecunda, new combination (64), 0. perforata (55), 0. pupil- lnta (48), 0. siibcavata (40), 0. exanthismocarpa (25), 0. nigropzincta, new species (16), 0. olivacea (12), 0. mordenii, new species (lo), and 0. tere- bmtziZa (10). Rarities, known from single collec- tions each, are 0. antillensis and 0. cavata. Key to the Species of Ocellularia 1. Medulla of thallus and/or ascocarps pigmented. 2. 2. Medulla of ascocarps yellow or pink but thallus medulla white. 3. 3. 1. Medulla of thallus and ascocarps white. 4. Apothecia lacking a central columella. 5. Apothecia immersed, 0.2-0.3 mm in diameter. Medulla and ascocarps entirely deep red .......................... 12. 0. mordenii, new species Pigment yellow; pore large, 0.3 mm wide ................ 10. 0. fecunda, new combination Pigment deep pink; pore tiny, 0.1 mm wide, annulate. ......... 18. 0. rhodostroma 6. Thallus Pf yellow ....................................... ........ 22. 0. terebratula 6. Thallus P- .......................................................................... 14. 0. olivucea 5. Apothecia emergent, 0.6-2.0 mm in diameter. 7. Disc open with a strongly recurved exciple. 8. 8. 7. Disc closed, lacking an outer recurved exciple. 9. Inner lepadinoid exciple strongly developed. 10. Apothecia aggregated in groups. Exciple thick, disc irregularly elongate; stictic acid present Exciple thin and delicate, disc round; stictic acid absent .......... ...... 7. 0. dilatata 1. 0. alboroselln 11. Apothecial groups round; stictic acid lacking ...................................... .......................................................................... 8. 0. dominicam, new species 16 SMITHSOXIAN COSTRIBUTIONS TO BOTANY 11. Apothecial groups oblong: stictic acid present ................................................................. 6. 0. conglomerata, new species 12. Spores large, 55p-75p .................................... 9. 0. exanthismocarpa 12. Spores small, 15p ................................ 6. 0. conglomerata, new species 10. Apothecia solitary. 9. Inner exciple not developed. 13. Thallus P-. 14. Thallus distinctly fissured: pore area black rimmed ............................ 14. Thallus continuous; pores not black rimmed .............. 15. 0. papillata 15. Pore area black rimmed .................. 13. 0. nigropuncta, new species 15. Pore area not black rimmed ................................ 16. 0. perforata 4. Apothecia with a well-developed black central columella. 16. Thallus with large scattered verrucose outgrowths becoming sorediate .................... 20. 0. sorediata, new species 16. Thallus lacking soredia. ............................................................................. 19. 0. rimosa, new species 13. Thallus P+ red. ........................................................................................ 15. Apothecia large, 0.5-1.1 mm in diameter, more or less emergent. 18. Thallus dull tan, hypophloeodal, P- ........................... 5. 0. concolor 18. Thallus distinctly epiphloeodal, P+. 19. Columella becoming actinoid: thallus P+ yellow .... 4. 0. comparabilis 19. Columella remaining simple: thallus P+ red ................. 3. 0. cavata 17. Apothecia smaller, 0.2-0.5 mm in diameter, barely emergent to immersed. 20. Pore annulate: spores small with crowded septae, 5p-6p X 9p-llp ................ 21. 0. subcavata 20. Pore not annulate: spores generally 20p-30p long (except smaller in 0. ............................................................................................... maculata). 21. Thallus P-. 22. Pore tiny, less than 0.1 mm in diameter; spores 4-loculate .............. 22. Pore larger, 0.1-0.2 mm wide; spores 6-7-loculate . 15. 0. papillata 21. Thallus P+ red or orange yellow. 23. Thallus very thin to hypophloeodal; Pt orange .................. .......................................................... 11. 0. maculata, new species ........................................................... 17. 0. pyrenuloides 23. Thallus thick, epiphloeodal. 24. Thallus P+ red: apothecia barely emergent ....................... 24. .................................................................. 16. 0. perforata Thallus P+ yellow; apothecia strongly emergent ............... ..... 2. 0. antillensis, nev species 1. Ocellularia alborosella FIGURES 5b, 96 Ocellu la ria alborosella (N ylander) San tesson, 1952: 308. Grapliis alborosella Sjlander, 18633352 [tj pe-collection: Co- lombia, Lhfdig 2ti94 (H, lectotypc: FH-Tuck, UPS, isotypes); Figure 9b]. Thelotre!tia platycnrpllwt! L'ainio, 1923: 138 [typc-collection: Arinia, Trinidad, TIiaxter 57 (TUR, lectotype); Figure gal. Ocellularia platjcarpella (Vaino) Zahlbruckner, 1923:598. Thallus whitish gray, thin but continuous, shiny, smooth, forming colonies 5-8 cm broad; apothecia chroocliscoid, up to 1.5 min across, round to irregu- lar, the marginal flap conspicuous, the disc flesh colored to white pruinose; hymenium about 60p high; spores 8, 5-6 loculate, 4p-5~ X 8p-15p, I- 01 weakly blue (Figure 5b). C~~sr~si RY.--No substances present. HAnriAr.-Tree trunks at mid or higher eleva- tion in rain forest. This was the first corticolous chroodiscoid species to be described. It is characterized by the small spores and lack of lichen substances. The Dominican collections have a wider disc than the type froin Colombia but they fall within a pre- dictable range of variation. It is a rare and easily overlooked species. SPECIMENS EXARIINED.-~ (38143), 16b (37883), 2613 (3537la). NUMBER 16 2. Ocellularia antillensis Hale, new species FIGURE 9g Thallus corticola, albidus, epiphloeodes, planus vel minute verrucosus, nitidus, 3-5 cm latus; apo- thecia numerosa, emergentia, 0.3-0.5 mm diametro, apice obfusca, columellata, columella 0.15 mm lata; ostiolum 0.1-0.12 mm diametro, intus nigrum; hymenium 7Op-801.1 altum; sporae 8: nae, trans- versim 7 loculatae, 7p X 20p, I+ coerulescentes (Figure 9g). CHEMrsTRY.-Psoromic and conpsoromic acids. HoLOTwE.-Trail to Madjini, low elevation windward rain forest, Dominica, elevation about 30 m, Hale 35612, January 1969 (US). HABITAT.-upper trunk of stunted rain forest trees (30m). The whitish minutely verrucose thallus and numerous emergent apothecia with the large con- trasting black pore area distinguish this rare species. Ocellularia comparabilis var. microcarpa Redinger has larger spores (6p X 30p-34y), sparse apothecia, and a smooth thallus, while 0. terebrata (Acharius) Muller-Argau has barely emergent apothecia (see Hale, 1972). I later collected this species from Trinidad and Panama, both at low elevation. Grande, Trinidad, Hale 37444 (US); Barro Colo- rado Island, Panama, Hale 38659 (US). SPECIMENS EXAMINED.-3 miles NW Of Sangre 17 3. Ocellularia cavata FIGURES 6c, 9f Ocellularia cavata (Acharius) Muller-Argau, 1882133499. Thelotrema cabaturn Acharius, 1812:92 [type-collection: Sierra Leone, Afzelius (H, lectotype; S, UPS, isotypes)].-Hale, 1971b, fig. la. Ascidium cinchonarum Fee, 1824:96 [type-collection: On Cinchona, South America (G, lectotype; H, L, M, P, iso- types); Figure gel. Ocellularia cinchonarum (Fee) Sprengel, 1827:242. Ascidium cinchonarum f. intermedium Nylander, I867:319 [type-collection: Pie de Cuesta, Colombia, Lindig 5 (H, lectotype; BM, G, M, isotypes); Figure 9c]. Ocellularia lindigiana Muller-Argau, 1887a:9 [type-collection: Colombia, Lindig 2757 (G, lectotype; BM, FH, UPS, iso- types); Figure 94. Oeellularia cinchonarym (Fee) Sprengel f. intermedia (Ny- lander) Zahlbruckner, 1923:586. Thallus whitish mineral gray, epiphloeodal, con- tinuous, more or less roughened, forming colonies up to 8 cm across; apothecia ascidioid, 0.7-0.9 mm in diameter, apically carbonized with a large columella (Figure 6c); pore round, 0.07-0.12 mm in diameter, the top of the columella clearly visible; spores 8, 6-8 loculate, 6p-8p X 22p-28p, IS blue. (Figure 9f). CmMIsTRY.-"Cinchonarum" unknowns A and B and a yellowish pigment. HABITAT.-canopy branch in mid-elevation rain forest (about 550 m). I had previously typified this distinctive but rare Acharian species. It appears to be rare in the Lesser Antilles. The chief diagnostic features are the unusual chemistry, small ascidioid apothecia, and the large columella which protrudes into the pore area. SPECIMEN EXAMINED.-14 (35134). 4. Ocellularia comparabilis FIGURES 5i, 9i Ocellularia comparabilis (Krempelhuber) Muller-Argau, 1883: 318. Thelotrema comparnbile Krempelhuber, 1876:220 [type- collection: Rio de Janeiro, Brazil, Glaziou 5463 (M, lecto- type; BM, C, G, M, P, UPS, W, isotypes); Figure 9i]. Thallus pale greenish to whitish mineral gray, epiphloeodal, smooth, forming colonies 5-1 2 cm broad; apothecia emergent, 0.8-1.2 mm in diameter, the walls carbonized, columella simple 'to actinoid; pore open, 0.2-0.5 mm broad, the white pruinose disc clearly visible; hymenium 50y-60&i high; spores 8, 4-5 loculate, 5p-6y X lOu--14~i, IS blue. (Fig- ure 5i). CHEivIsTRY.-Psoromic and conpsoromic acids. HABITAT.-LOWer trunks in rain forest (300- 700 m). The columella in this rare species is imperfectly actinoid. To judge from collections made in South America, the range of variation is very great with gradations into related psoromic acid-containing species. For example, Ocellulnria discoidea (Acha- rim) Muller-Argau (Hale, 1972) and 0. antillensis, new species, have a simple columella. Ocellzilaria efformata (Krempelhuber) Muller-Argau has a very similar actinoid columella but the outer margin is semi-erect, leaving an open rather lacerate pore: 0. De?.keIeynna (Montagne) Zahlbruckner and 0. glnziovii Muller-Argau both have highly developed 18 SMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 9.-Specimens of Thelotremataceae (all about X 10): a, Ocellularia platycarpella (lecto- type in TUR); b, 0. alborosella, (isotype in FH-Tuck); c, 0. cinchonarum, f. intermedia (Iecto- type in H); d, 0. lindigiana (lectotype in G); e, 0. cinchonarum (lectotype in G); f, 0. cavata (Hale 35134); g, 0. antillensis, new species (Hale 35612); h, 0. conglomerata, new species (Hale 37959); i, 0. comparabilis (isotype in BM); i, 0. concolor (lectotype in G); k, 0. concolor (Hale 38167). NUMBER 16 actinoid discs; and 0. latilabia (Tuckerman) Muller-Argau has an actinoid columella and a recurved margin. SPECIMENS EXAMINED.-~ (38128), 21 (37372, 38144). 19 5. Ocellularia concolor FIGURE 9j,k Ocellularia concolor Meyen and Flotow, 1843:230. TYPE-CoLLEcTIoN.-;\fanila, Philippines, Meyen (G, lectotype) (Figure 9j). Thallus light grayish tan, dull, mostly hypoph- loeodal, forming colonies 3-4 cm broad; apothecia numerous, 0.8-1.1 mm in diameter, more or less emergent, apically carbonized, columella present, 0.2-0.3 mm wide; pore open, round to somewhat irregular, 0.3-0.5 mm in diameter, the rim black and the flat black columnar disc easily seen; liymenium 14Op high; spores 8, 5-7 loculate, 5p- 8p X 13p-20p, I+ blue (Figure 9k). CHEI\IISTRY.--NO substances present. HABITAT.-Lianas, saplings, base and lower trunk of trees in dense rain forest (300-370 m). This easily recognizable species was first de- scribed from the Philippines and appears to be pantropical. The tannish hypophloeodal thallus and habitat are characteristic. There are no com- parable species in this kind of habitat. 16b (37801, 37809, 38075). SPFCILIEKS EXAMINED.-8 (38167 ), 16a (37665), 6. Ocellularia conglomerata Hale, new species FIGURE 9h Tliallus corticola, cinereo-albidus, epiphloeodes, continuus, nitidus, 4-5 cm latus; apothecia primo solitaria, 0.6-0.9 mm diametro, semi-emergentia, mox 2-3: nae conglomerata, margine exteriore erecto, excipulo interiore evoluto, pulverulento, disco aperto, carneo; ostiolum irregulare, latum; 11) menium 70p altum; sporae 8: nae, transversim 6-7 loculatae, 5p-611 X 15p-l8p, I+ coerulescentes (Figure 912). CHEMISTRY.-stiCtiC and constictic acids. HOLoTwE.-Trail to hforne hglais, Dominica, elevation about 900 m, Hale 37959, 13 December 1971 (US). Chemistry places this species near 0. exanthisrno- caypa but the exciple is not so clearly separate froin the thick, erect, pulverulent margin, leaving the disc open. The spores are much smaller. An- other ilrest Indian stictic acid-containing species, Ocellularia albooliuacea (Vainio) Zahlbruckner, has a more typically recurved margin and a thin, entire exciple. 7. Ocellularia dilatata FIGURE lla,b Ocellularia dilatata Miiller-Argau, 1895:452. TYPE-CoLLECTION.-Rio de Janeiro, Brazil, Glaziou 5531 pro parte (GI lectotype; BM, isotype). (Figure 1 la). Thallus whitish gray, thin, dull, in part hypoph- loeodal, forming colonies 4-6 cm broad; apothecia chroodiscoid, large, up to 2.5 mm broad, round to elongate, marginal flap rather coarse; disc white pruinose; hymenium about 60p high; spores 8, 6-7 loculate, 5p-6p X 15p-l7p, I+ blue (Figure llb). CHEMIsTRY.-stictic and constictic acids. HABITAT.-LO\Wr trunk area in higher rain forest zone (850 m). This rare species is related to Ocellzila?.ia albo~o- sella, but it has a thicker recurved margin and a different chemistry. I have collected a number of specimens at low elevation in Trinidad, suggesting that the species is more at home on continental land masses. SPECIMENS EXAXIINED.--~~~ (37695, 37702). 8. Ocellularia dominicana Hale, new species FIGURE 2c,d; 10 Thallus corticola, brunneo- vel viridi-albidus, epiphloeodes, continuus, nitidus, 5-10 cm latus; apothecia semi-emergentia, 0.7-1.1 mm diametro, apice incolores, columella nulla, solitaria vel 2-3 arcte aggregata, habitu similia apothecio singulari, excipulo interiore evoluto, ostiol~um complente; ostiolum rotundum, 0.1-0.5 mm diametro; hy- menium 6Op-8Op altum; sporae 8: nae, transversim 6-7 loculatae, 6p8p X 16p-25p, I + coerulescentes (Figure 10). CHEMISTRY.-NO substances present. 20 S.\lII'HSONIAN CONTRIBUTIONS TO BOTANY Thelotrema homothecintm Vainio, 1921:190 [type-collection: Irosin, Mt. Bulusan, Pror. Sorsogon, Philippines, Elmer 14852 (TUR, lectotype; FH, G, L, S, US, W, isotypes); Fig- ure Ild]. Ocellularia homothecia (Vainio) Zahlbruckner, 1923:593. Ocellularia porinoides (hlontagne and von der Bosch) Zahl- bruckner, 1923:599 [not 0. porinoides (Acharius) Sprengel, 1827 : 2421. Thallus white to ashy, thin, smooth, in part hypophloeodal, forniing colonies 1-4 cm across; apothecia numerous, semi-emergent, the rim be- coming erect, noncarbonized and without a columella, the inner exciple distinct, intact, form- ing an apical pore within the main pore; main pore open, 0.2-0.4 mm in diameter; hymenium 160p- 180p high; spores 4-8/ascus, 18-22 loculate, 12p- 18p X 6Op-lOOp, If deep blue (Figure llg). CmhiISTRY.-stictic and constictic acids and rarely the higher "quintaria" unknown B. HABITAT.-Trunks, canopy branches, fence posts, and planted trees (300-900 m). This common pantropical species is unique be- cause of the pore-within-a-pore arrangement. The large spores are also distinctive but it is not un- usual to find sterile specimens. It is also one of the few species in the family to invade secondary forests, fence posts, etc., in wet areas up to very high elevation. 16b (37982), 17 (35605), 20a (37780, 37781, 37786, 3i788, 38019, 38025), 25a (37708, 37709, 37713, 37714, 37715, 37716, 37849, 37996, 38069), 25b (35473, 35516), 26a (37692, 37944), 27 (38042, 38055). SPECIAIEXS EXAAIINED.-8 (381 14), 11 (38009), FIGURE lO.-OcelIularia dorninicana, new species (Hale 37841) (about X 10). HoLorYpE.-Submossy forest on upper slopes of Trois Pitons, Dominica, elevation about 1000 m, Hale 37967, 6 December 1971 (US). H.%nnxT.-Trunk and lower base of trees in sub- mossy forest (700-1 100 m). The apothecia are so tightly aggregated that I first interpreted the wall between them as a columella. The combined outer margin and exciple are minutely pulverulent antl crowded, filling the broad disc but leaving a distinct pore area. The species isas rather corninon at Trois Pitons but not found elsewhere. SPECIMEKS EXAMIKED.--~~ (38053), 27 (37683, 37688, 37829, 37832, 37841, 38041). 9. Ocellularia exanthismocarpa FIGURES 50, llc,g Ocellulnria esa~ithisviocarpn (Leighton) Zahlbruckner, 1923: 590. Thelotrema esn~~thisni~ca~.p~nz Leighton, 1869: 169 [type- collection: Central Province, Ceylon, Tlircnites C.L. 97 (BM, lectotype; H, isotype); Figure llc]. Thelotrema porinoides hlontagne and I on der Bosch, 1855: 484 [type-collection: Java, Junghuhn 151 (L, lectotype; FH, G, P, Ti', isotypes); Figure lle]. Ocellularia mziltiloczilnris Zahlbruckner, 1912:369 [typc- collection: Lanai, Hawaii, Roqk li (IV. holotypc); Figure Ocellulnria isertii T'ainio, 1915:40 [type-collcction: Guade- loupe, Zsert 87 (C, lectotype; TUR, isotype); Figure Ilfl. 5a]. 10. Ocellularia fecunda (Vainio) Hale, new combination FIGURE 11 h,i Thelotrenu domingeiise var. fecu?iduni T'ainio, 1896:208 [type-collection: Richmond Valley, St. Vincent, Elliotst 243 (BM, lectotype; TUR, isotype); Figure llh]. Ocellularia donzingensis var. fecunda (Vainio) Zahlbruckner, 1923:590. Thallus light greenish or whitish mineral gray, epiphlocotlal, smooth and shiny, forming large colonies up to 30 cin broad; apotliecia large, strong- ly emergent, 1.5-1.8 inin in diameter, the cortex soon breaking a~vay antl revealing a light yellow- is11 orange medulla, walls heavily carbonized, KUMBER 16 21 FIGURE 1 I .-Specimens of Thelotremataceae (all about X 10): a, Ocellularia dilatata (isotype in BM); b, 0. dilatata (Hale 37702); c, 0. exanthismocarpa (lectotype in BM); d, 0. homothecia (isotype in FH); e, 0. porinoides (lectotype in L); f, 0. isertii (lectotype in C); g, 0. exanthis- mocarpa (Hale 37996); h, 0. fecuxda, new combination (lectotype in BM); i, 0. fecuuda, new combination (Hale 35218); j, 0. maculata, new species (Hale 38015); k, 0. motdenii, new species (Hale 37887). 22 SMITHSONIAN CONTRIBUTIONS TO BOTANY columella absent or weakly and irregularly devel- oped; pore gaping, round to irregular and jagged, 0.4-0.8 mm in diameter, the white pruinose disc clearly visible; hymenium 2OOp-25Op high; spores 1--4/ascus, 30-40 loculate, 12-20 X 80p-I50p, rarely turning brown, I+ blue (Figure lli). CHExmiRY.-Hypoprotocetraric acid and in lesser concentration 4-0-demethylnotatic acid along with unidentified pigments. HAuITAT.-Basal area of trees into the upper bole and canopy in the rain forest (300-850 m). This is one of the few species in the family that can be recognized at sight in the field. The apothe- cia are very large and the yellow-orange medulla is exposed by rubbing the cortex of?. It is the second commonest species in Dominica and forms very extensive colonies. As far as I have determined, the species is restricted to the West Indies and has no close relatives. 37816, 37820, 38003, 38046, 38061), 12 (37902, 37906, 37913, 37920, 37921, 37931), 15 (35155, 35167, 35218), 16a (37658, 37942, 37970, 38050), 16b (37796, 37802, 37810, 37874, 37881, 37954), 16c (37765, 37766, 37774, 37943, 37980), 18a (37725, 37957, 37962, 37986, 38039), 21 (38080, 38081, 38089, 38103, 38106, 38107, 38132, 38137), 22 (37639, 37644, 37828, 37838, 37843, 37844, 37844a, 37845, 37856, 37858, 37960, 38026), 23 (37679), 24a (37740, 37743, 37063, 37064), 25a (37711), 26a (38056). SPECIMENS EXAMINED.-8 (38 120), 1 1 (378 1 1, 11. Ocellularia maculata Hale, new species FIGURE llj Thallus corticola, cinereo-albidus, epiphloeodes, planus, continuus, nitidus, 9 cm latus; apothecia nunierosa, minuta, 0.2-0.3 nim diametro, apice ol)lusc;i, columellata vel columella parce evoluta; ostiolum rotundum, 0.05-0.1 mm diametro, anguste nigrocinctum, columella protrusa; hymenium 100p altum; sporae 8:nae, transversim 4 loculatae, 5y-7p X llp-l3p, I- (Figure llj). CHEXIISTRY .--No substances present. HoI.oTYI.r.-Remnants of rain forest in pastures above Giraudel, Dominica, elevation 650 m, Hale 38015, 11 December 1971 (US). Occlliilnria mncirlata might seem at first glance to be 0. papillata but the apothecia and spores are much smaller. There are no related species. SPECIXE.N EXAMMINED.-lSb (35524). 12. Ocellularia mordenii Hale, new species FIGURES 3c,d; 66; Ilk Thallus corticola, pallide cinereo- vel brunneo- albidus, epiphloeodes, continuus, aetate rimosus, nitidus, usque ad 12 cm latus; medulla omnino sanguinea; apothecia dispersa, vix emergentia, apice obfusca, extus crasse verrucosa, verrucis fragil- ibus, rumpentibus, columella nulla vel parce evo- luta (Figure 6b); ostiolum rotundum, 0.08-0.12 mm cliametro; hymenium circa 2OOp altum; sporae 1-2:nae, transversim 24-30 loculatae, 15p-20y X 70p150p, I+ coerulescentes (Figure Ilk). CHExmTRY.-Unidentified anthraquinone pig- ments only. HoLoiyPE.-Canopy branches in primary rain forest, Dleau Gommier Forest Reserve, Dominica, elevation about 370 m, Hale 37764, 12 December HAwrAT.-Canopy branches in rain forest (370- 800 m). ?The deep red pigment and large warts are dis- tinctive. It is a conspicuous canopy species. A very similar species in Cuba, 0. xanthost~oma (Ny- lander) Zahlbruckner, differs mainly in containing n I<- pale yellow-orange pigment. This lichen is named in honor 01 Alrs. William J. Morden. SI?PCIMENS EXAhlINED.-l2 (37909), 15 (35198), 16a (38051), 16b (37791), 16c (37762, 37886, 37887, 37010), 26a (37863). 1971 (US). 13. Ocellularia ngropuncta Hale, new species FIGURES 2a,b; 6a; 12a Thallus corticola, pallide brunneo- vel viridi- cinei eus, epiphloeodes, continuus vel rarius I imo- sus, 5-8 cm latus; apothecia numerosa, emergentia, 0.6-0.8 mni diametro, apice obfusca, columella nulla (Figure 6a); ostiolum rotundum, 0.1-0.2 mm tlianietro, nigrocinctum; hJmeniuin 100~1-110p altum; sporae 8:nae, transversim 6-8 loculatae, 6p- 10:~ X 2Oy-32p I+ coerulescentes (Figure 12a). CmwsTRY.-Protocetraric acid. HOLOTYPE.-T~ ail to summit of Rforne Anglais, Dominica, elevation about 900 m, Hale 35365, January 1969 (US). NUMBER 16 23 FIGURE IZ.-Specimens of Thelotremataceae (all about X 10): a, Ocellularia nigropuncta, new species (Hale 35365); b, 0. rimosa, new species (Hale 38052); c, 0. oliuacea (lectotype in G); d, 0. papilla,ta (lectotype in BM); e, 0. papillata (Hale 38098); j, 0. perforata (lectotype in BM); g, 0. excavata (lectotype in BM); h, 0. rufocincta (lectotype in G); i, 0. terebrata var. abbreuiata (isotype in FH); j, 0. pyrenuloides (isotype in W); k, 0. pyrenuloides (Hale 35524). 24 SMITHSONIAN CONTRIBUTIONS TO BOTANY HABrT.AT.-Lianas, lower bole and base, sym- phonea roots, rarely canopy of trees in the rain forest (300-900 m). The black-rimmed pore area and smooth thallus are diagnostic features. Other ecolumellate proto- cetraric acid-containing species include Ocellulaiia bonplandiae (Fke) Muller-Argau, which has a grainy surface, and 0. ueirzIcosa (Fee) Miiller- Argau, which has an irregular open pore and larger spores (over 30y long). Under the scanning- electron microscope 0. nigropuncta, new species, has a finely pitted but otherwise smooth surface (Figure 2a,b), whereas 0. uerizicosa appears to have an aculeate orientation and 0. bonplandiae is verrucose and fissured. SPECIMENS EXAMINED.-~ (381 15, 38158), 10 (38169), 12 (37903), 16a (37660), 16b (37798), 16c (37755, 37770, 37963), 18a (37948), 21 (38092, 38094, 38138), 22 (37852), 24a (37751). 14. Ocellularia olivacea FIGURES la; 3a,b; 5c; 12c Ocellularia oliuacea (Fee) Miiller-Argau, 1887a:7. Myriot, emu olivacea Fee, 1824: 103 [type-collection: On Bon- plandin trifoliata, South America (G, lectotype); Figure Ocellularia oliuacea (Fee) C. and D. Van Overeem-De Haas, 1 2c] . 1922: 118 [superfluous combination], Tliallus light greenish to ashy white, shiny, thick, up to 0.5 mm (Figure la), forming exten- sive colonies up to 50 cm broad, sometimes finely cracked and fissured; apothecia very numerous, im- mersed, 0.15-0.3 mm in diameter, noncarbonized and without a columella; pore small, flush, about 0.1 mm in diameter; hymenium 65p-75p high; spores 8, more or less uniseriate, 4 loculate, 4y- 6p X 8y-12p, If blue (Figure 5c). CHE;\IIsTRY.-?Olivacea? unknown and a second lower spot. HABITAT.-canopy branches to mid-bole in rain forest (240-700 m). This is one of three ?Myiiotiemn? species that occur in Dominica, the other two being 0. tere- brat iila and Thelotrema clandestinttm, all typically found on canopy branches in dense rain forest. Ocellztlaria oliuacea is characterized by the rather small spores and unusual chemistry. The West Indian material all produces a second spot below the ?olivacea? unknown, probably a related dep- side. The type-specimen of 0. oliuacea lacks this spot. The surlace under the scanning-electron microscope is strongly aculeate (Figure 3a,b). 8 (38164), 9 (38154), 11 (38001), 12 (37965), 15 (35205), 16a (37992, 38054), 18a (37965), 21 (38113). SPECIMENS EXAMISED.-7 (35226, 35243, 35249), 15. Ocellularia papillata FIGURE 12d,e Ocellularia papillata (Leighton) Zahlbruckner, 1923:597. Thelotrema pnpillatunz Leighton, 1869: 169 [type-collection: Central Province, Ceylon, Thwaites C.L. 129 (BM, lectotype; H, G, P, S, UPS, W, isotypes); Figure 124. Thallus whitish to pale greenish ashy, smooth to more or less warty, shiny, forming colonies 3-12 cni broad; apothecia numerous, 0.6-0.8 mm in diameter, immersed to slightly emergent, the upper walls carbonized, columella present, thin to scarcely developed to absent; pore round, 0.1-0.2 mni in diameter; hymenium 50p-70p high; spores 8/ascus, 7-9 loculate, 6p-lop X 2Oy-36p I+ blue. (Fig- ure 12e). CHEJIISTRY.--NO substance present. HABITAT.-ktplingS, prop roots, lianas, and tree bases at mid-elevation in rain forest (300-900 m). This common species has few outstanding fea- tures. The columella is usually present, although often weakly developed and difficult to find. Some specimens, as indicated below, seem to lack it en- tirely. The apothecia are inconspicuous, almost immersed to slightly emergent, but the pore is always distinct. The thallus is well developed, usually quite whitish, and smooth. There are no close relatives i\rithout lichen substances, although 0. perfomto, which has protocetraric acid, is very close in general aspect except perhaps that the apothecia are more often emergent. Both species occupy similar base level habitats in the rain forest and were collected together at 11 of the localities. At eight other localities only one of the species was found. SPECIMENS EXAMINED (columella present).-S (38123, 38130, 38165, 38168), 9 (38131), 11 (37825), 16a (37795, 37991), 16b (37662, 37804, 37805, 37808), 18a (37721, 37964); 21 (38084, 38085, 38097, 38098, 38152), 22 (37645, 37837, 37842), 23 (37671, 37681, 37682), 27 (37850, 38072), 28 (35428). NUMBER 16 25 SPECIMENS EXAMINED (columella absent).-7 (35227), 10 (38127, 38150), 16a (37655, 37661), 16b (37792, 38074), 16c (38045) 21 (38086, 38088), 22 (37649, 37834, 37851, 37854, 37855), 23 (37677, 38965), 24a (37746). 16. Ocellularia perforata FIGURFS Ib, 5e, 7a, 12f Ocellularia perforata (Leighton) Muller-Argau, 1892284. Thelotrema perforatum Leighton, 1866:447 [type-collection: Casiquiari, Brazil, Spruce 254 (BM, lectotype); Figure 12fl. Thelotrema terebratum var. abbreuiatulum Vainio, 1890:83 [type-collection: Caraqa, Minas Gerais, Brazil, Vainio 1551 (TUR, lectotype; BM, FH, UPS, isotypes); Figure 12i]. Ocellularia rufocincta Muller-Argau, 1893: 146 [type-collection: Foret du Rio General, Tondoz s.n. (Pittier 6107) (G, lecto- type); Figure 12hl. Thelotrema excavatum Vainio, 1896:208 [type-collection: Mt. St. Andrew, St. Vincent, Elliott 153 (TUR, lectotype; BM, FH, isotypes); Figure 12g]. Thelotrema excauatum var. impressulum Vainio, 1896:208 [type-collection: Morne Anglais, Dominica, Elliott 169 (BM, lectotype)]. Ocellularia terebrata var. abbreviata (Vainio) Zahlbruckner, 1923502. Thallus whitish to pale greenish ashy, smooth to more or less warty with age, shiny, forming colonies 3-10 cm broad; apothecia numerous, im- mersed to moderately emergent, 0.4-0.6 mm in diameter, the upper wall carbonized, columella usually developed, thin 50p-70p in diameter, but sometimes lacking or difficult to find; pore round, 0.07-0.15 mm in diameter; hymenium 55p-70p high; spores 8/ascus, 6-8 loculate, 7p-lop X 20~- 34p, I+ blue (Figure 5e). CHEMIsmY.-Protocetraric acid and the ?am- plior? unknown as an accessory substance. HABITAT.-Buttresses, saplings, and bole to rarely canopy branches in rain forest at mid- elevation (300-850 m). This is one of the commonest species in Dominica at base level. The smooth whitish thallus (see cross-section in Figure 2b) and the small semi- emergent apothecia resemble 0. papillata, as dis- cussed above. Some of the specimens as listed below lacked any sign of a columella. SPECIMENS EXAMINED (columella present).-8 (38149), 11 (37812, 37817, 37997, 38030, 38033), 12 (37899, 38078), 14 (35139), 16a (37651, 37666, 38007, 38032), 16b (37884), 16c (37759, 37761, 37775, 37952), 18a (37727, 37728, 37730, 37732, 37535, 37961, 37969, 37983, 38020), 20a (37784, 37785, 38016, 38017), 21 (38108, 38134, 38145), 22 (37646, 37647, 37830, 35857, 38036), 23 (37675), 24a (37752), 26a (37698, 37701), 27 (37686, 37827). SPECIMENS EXAMINED (columella absent).-8 (38160, 38163), 16a (38018), 21 (38093, 38140, 38157), 23 (37674), 24a (38066), 27 (37684, 37847), 26a (37700), 26b (35370). 17. Ocellularia pyrenuloides FIGURE 12j,k Ocellularia fiyrenuloides Zahlbruckner in Magnusson and Zahlbruckner, 1943:46. TYPE-CoLLECTIoN.-\vaiIUkU, Maui, Hawaii, Fauyie 656 (UPS, lectotype; S, IV, isotypes) (Fig- ure 12j). Thallus ashy mineral gray, thin to evanescent, continuous, forming colonies about 5 cm broad; apothecia numerous, nearly immersed to moder- ately emergent, 0.5-0.7 mm in diameter, walls heavily carbonized, columella present, 130p in diameter; pore round, 0.1-0.15 mm wide, pore area surrounded by a blackish ring, in part white prui- nose, the pruinose top of columella visible through the pore; hymenium about 60p high; spores 8/ascus, 6-7 loculate, 611 X 12p-I5y, I+ blue (Figure 15%). CHEMISTRY .-stictic and constictic acids. HAnITAT.-Tree branch in rain forest (650 m). The darkened pore area is similar to that in Thelotrema teniie, new species. There are no re- lated species with stictic acid. It is still surprising to find a Hawaiian species on Dominica, but to me this suggests how poorly the Thelotremataceae have been collected. SPECIMEN EXAMINED.-18b (35524). 18. Ocellularia rhodostroma FIGURES 5g, 7b, 13a Ocellularia rhodostromn (Montagne) Zahlbruckner, 1923:600. Ascidium rhodostromum Montagne, 1851:75 [type-collection: Cayenne, French Guiana, collection 1334 (P, lectotype); Figure 13~1. Thallus light greenish tan to gray, smooth to finely warty, continuous and epiphloeodal, the medulla white or pink, forming colonies 5-10 cm 26 ShIITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 13.--Specimrns of Thelotremataceae (all about X 10): a, Ocellularia rhodostroma (lecto- type in P); b, 0. subcavata (lectotype in H); c, 0. vaga (isotype in FH); d, 0. subca7jata (Hale 37769); e, 0. terebratula (lectotype in H); f, 0. remanens (isotype in G); g, 0. sor~diata, new species (Hale 37908); h, Plraeotrema aggregatum, new species (Hale 35229); i, 0. terebratula (Hale 37793); 1, P. disciforme, new combination (lectotype in Bhl); k, P. aqztiliiium (lectotype in TUR). NUMBER 16 27 broad; apothecia large and easily visible with the naked eye, ascidioid, 1.0-1.8 mm in diameter, the walls heavily carbonized and the medulla deep pink, readily exposed as the fragile cortex breaks away, columella lacking; pore round and small, 0.05-0.1 mm in diameter, surrounded by a raised ring; hymenium 2OOp-275p high; spores l-2/ascus, 25-30 loculate, 25p-4Op X lOOp-25Op, I+ blue (Figures 5g and 70). CHEMISTRY.-Hypoprotocetraric acid and in greater concentration 4-0-demethylnotatic acid along with unidentified pigments. HABITAT.-CanOpy branches to lower bole at all elevations on the island. Ocellularia rhodostroma is the commonest thelo- treme on Dominica and the easiest to identify as well. The large ascidioid apothecia have a deep pink medulla exposed when the fragile cortex breaks. The pore is strongly annulate. While oc- curring at all elevations, it seems best developed at higher elevation into the mossy forest, perhaps because it competes with so few other species there. 38146, 38166), 10 (38119), 11 (37815, 37823, 37824, 38004, 38023, 38049), 12 (37894, 37916, 37917, 37927, 37930, 37934), 13 (35279 35448), 14 (35111, 35133), 15 (35160, 35170, 35188, 35314, 16a (37653, 37938), 16b (37800, 37806, 37875, 37994, 37995), 16c (37756, 37870, 37979), 18a (37729, 37731, 37987), 18b (35517, 35522, 35523, 35587), 20a (37777, 38065), 21 (38035, 38147), 22 (37641, 37643, 35650, 37835, 37861), 23 (37672, 38975), 24a (37736, 37742, 37745, 37749, 37754, 38068), 24b (35629), 25a (37705, 37707, 37720, 37840, 37846, 37848, 35862, 37867, 37971, 37975), 25b (35469), 26a (37689, 37690, 37693, 37694, 37699, 37703, 37864, 37866, 37885, 37947 38000, 38014), 26b (35354, 35366, 35371, 35372, 35376), 27 (37831), 28 (35435, 35440 ). SPECIMENS EXAMINED.-7 (35246), 8 (38122, 19. Ocellularia rimosa Hale, new species FIGURE 12b Thallus corticola, brunneo- vel viridi-cinereus, epiphloeodes, nitidus, continuus sed aetate rimosus, usque ad 15 cm latus; apothecia numerosa, emer- gentia, 0.7-09 mm diametro, apice obfusca, colu- mella nulla; ostiolum rotundum vel irregulare, 0.1-0.2 mm diametro, nigrocinctum; hymenium 110p-150p altum; sporae 8:nae, transversim 6-8 loculatae, 7p-lop X 24p-32p, I + coerulescentes (Figure 12b). CHEMISTRY.--NO substances present. HoLoTYPE.-Base of tree in primary rain forest, Dleau Gommier Forest Reserve, Dominica, eleva- tion 370 m, Hale 37978, 1 December 1971 (U.S.). HABITAT.-Prop roots, tree bases, and lianas in rain forest (370-430 m). Except for chemistry and slightly larger apothe- cia, this species is very close to 0. nigropuncta, new species. Both are base level species, but 0. Yimosa has a more restricted elevational range. The two species were collected together only at Dleau Gommier. SPECIMENS EXAMINED.-~ 1 (38052), 16a (37654), 16b (37392, 37803), 16c (37758). 20. Ocellularia sorediata Hale, new species FIGURE 13g Thallus corticola, viridi-albidus, epiphloeodes, continuus, planus, nitidus, usque ad 15 cm latus, sorediatus, soraliis sparsis, verrucosis, circa 1 mm latis; apothecia immersa vel semi-emergentia, 0.4- 0.6 mm diametro, apice fuliginea, columellata; os- tiolum, rotundum, 0.08-0.1 1 mm diametro; hy- menium 6Op-7Op altuni; sporae 8:nae, transversim 3-4 loculatae, 6p-8p X 12p-16p, I+ coerulescentes (Figure 13g). CHEMISTRY.-NO substances present. HoL0TYPE.-Logging area about 3 km northwest of Pont Cassi. on the Layou Road, Dominica, ele- vation 420 m, Hale 37908, 10 December 1971 (US). HAnrTAT.-hfid-bole of tree in rain forest; This is, as far as I know, the only sorediate species in the family. Large corticate tubercules are produced first, and some of these become coarsely sorediate. Although these peculiar struc- tures seem to originate from the main thallus, there is still the possibility that they have an extraneous origin. SPECIMEN EXAMINED.-16b (38029). 21. Ocellularia subcaoata FIGURE 13b,d Ocellularia subcavata (Nylander) Zahlbruckner, 1923:601. 28 SMITHSOXIAN CONTRIBUTIONS TO BOTANY 22. Ocellularia terebratula Thelotrema subcavatum Nylander, 1876:561 [type-collection: Cuba, Wright 509 (H, lectotype; FH-Tuck, isotype); Figure 13bl. Thelotrema vagum Vainio, 1896:209 [type-collection: Rich- mond Peak, St. Vincent, Elliobt 260 (BM, lectotype, FH, isotype); Figure 13~1. Ocellularia uaga (Vainio) Zahlbruckner, 1923:603. Thallus ashy whitish to greenish gray, shiny, more or less warty and cracked, forming colonies up to 12 cm broad; apothecia very numerous, emer- gent to ascidioid, 0.7-1.0 mm in diameter, crowded, walls carbonized, columella present, basally car- bonized but usually colorless in the upper half; pore round, small, 0.1-0.2 mm in diameter, sur- rounded by a whitish, more or less raised ring, the center filled with the often protruding pruinose columella; hymenium about 1001~ high; spores 8/ascus, usually uniseriate, 5-6 loculate with nar- row locules, 4p-7p X 8p-2Op1 very rarely turning brown with age, I+ blue (Figure 13d). CHEMIsTRY.-Psoromic and conpsoromic acids. HABITAT.-Canopy branches and upper bole in rain forest (240-800 m). This is one of the commonest thelotremes in the canopy and can be recognized in the field with a hand lens because of the numerous crowded apothecia with a white annulate pore and pro- truding columella. The columella is well developed in cross-section but unique in being more or less noncarbonized in the upper half. (38161), 10 (38129), 12 (37893, 37896, 37898, 37900, 37905, 37912, 37924, 37925, 37926, 37928, 37929, 37933), 13 (35294), 15 (35158, 35209), 16a (37652, 37795, 38012), 16b (37662, 37794, 37865), 16c (37767, 37769, 37772, 37773, 37924, 37941, 37972), 18a (37724, 37889, 37949, 38038), 23 (37680), 24a (37737, 37744, 37750), 26a (37691, 37704). SPECIMENS EXAMINED.-7 (35248, 35252), 9 FIGURE 13e,i Ocellularm terebratula (Nylander) Muller-Argau, 1887: 12. Thelotrema terebratuluin Nylander, 1867:315 [type-collection: Rio Negro, Colombia, Lindig 129 (H, lectotype; FH, C, hf, P, isotypes); Figure 13el. Thelotrema clandestinum f. remanens Nylander, 1867:315 [type-collection: Monte del Moiro, Colombia, Lindig 90 (H, lectotype; BM, G, P, isotypes); Figure l3fl. Thallus light greenish to ashy white, thick and continuous, smooth, shiny, often forming exten- sive colonies up to 15 cm broad; apothecia nu- merous, 0.2-0.4 mm in diameter, immersed, non- carbonized and without a columella; pore flush, 0.05-0.1 mm in diameter, usually surrounded by a faint whitish ring; hymenium about 65p high; spores 8/ascus, 3-5 loculate, 5p-9p X 15p-20p, I+ blue (Figure 132). Ocellularia remanens (Nylander) Muller-Argau, 1887:7. CHE;MISTRY.-PsOromic and conpsoromic acids. HABITAT.-~~~o~~ branches and upper bole in rain forest (300-400 m). Ocellularia terebratula, previously known only from Colombia, is a typical canopy level species. It is very similar to Thelotrema clandestinum, as explained below, except for spore characters. SPECIMENS ExAMINED.-~ (38125), 10 (38126), 12 (37904, 37918), 16a (37668, 37878, 37937, 38011, 38040), 16b (37793). Phaeotrema This genus is very similar to Ocellularia in spore septation but the spores are brown and sometimes shriveled at maturity, especially in large-spored species. The iodine reaction often seems to be negative, while almost all species of Ocellularia are strongly I+ blue. On the world level, Phaeo- Key to the Species of Phaeotrema 1. Apothecia aggregated ........................................................ 23. P. aggregatum, new species 1. Apothecia solitary. 2. Apothecia large, about 0.6 mm in diameter, strongly emergent; columella lacking ........... 2. Apothecia smaller, less than 0.5 mm in diameter, immersed; columella more or less developed. 3. Pore 0.1-0.2 mm in diameter, sometimes elongate, with a white annulus ....................... ......................................................................................................... 23. P. leiostomum 3. Pore 0.5-0.1 mm in diameter, round, annulus lacking . ._ 26. P. obscurum, new species .......................................................................................... 24. P. disciforme, new combination NUMBER 16 29 trema is represented by about 40 species, a rather small group, moreso because few of the species are well represented by herbarium material. In Dominica only six collections (four species) were found, these all occurring below 350 m elevation. 23. Phaeotrema aggregatum Hale, new species FIGURE 13h Thallus corticola, pallide viridi-cinereus, epi- phloeodes, planus, aetate rimosus, 8 cm latus; apothecia irregulariter aggregata, 0.4-0.6 mm diametro, immersa, intus incolores, columella nulla, excipulo interiore bene evoluto; ostiolum 0.1-0.2 mm latum, in centro excipulo interiore congesto, albido-pruinoso; hymenium 110p-12Op altum; sporae 8:nae, transversim 4-5 loculatae, 8p X 16p (Figure 13h). CHEMISTRY.-NO substances present. HOL0TYPE.-Logging area at Newfoundland, Dominica, elevation 250 m, Hale 35229, January HABITAT.-upper branches of tree in rain forest. This unusual species has densely aggregated, immersed apothecia. The pore is usually distinct but may be obscured by the pulverulent exciple and margin, giving the appearance of a Phlyctis. 1969 (US). 24. Phaeotrema discif orme (Leighton) Hale, new combination FIGURES ic, 131, 14a Thelotrema disciJornze Leighton, 1869: 170 [type-collection: Central Province, Ceylon, Thwaites (BM, lectotype; H, isotype); Figure 1311. Thelotrema exalbrduin Stirton, 1881 : 184 [type-collection: Assam, Il?att s.n. (BM, lectotype)]. Thelotretna aquilinum Vainio, 1915:137 [type-collection: St. Croix, Boergesen s.n. (TUR, lectotype; C, isotype)]. Leptotrema exalbidum (Stirton) Zahlbruckner, 1923:634. Thelotrema galactinum Vainio, 1926:24 [type-collection: Maloapan, Mexico, Liebmann 7712 (C, lectotype); Figure 13k]. Phaeotrema galactinum (Vainio) Zahlbruckner, 1932:245. Thallus whitish, dull, very thin and evanescent to hypophloeodal, forming colonies 2-4 cm broad; apothecia large, emergent, 0.7-1.1 mm in diameter, upper walls becoming suberect, carbonized, colu- mella lacking, the inner exciple weakly developed and pulling away from the main margin, the white-pruinose disc clearly visible; pore open, round, 0.2-0.5 mm in diameter, the rim usually darkened; hymenium 120p-140p high; spores ap- parently 4/ascus, 20-24 loculate, 7p-12p X 40p- 551.1, colorless when immature but mostly turning brown and shriveled at maturity, I- (Figure 14a). CHEMISTRY.-NO substances present. HABITAT.-Lower trunk area (sea level to 370 m). A thorough examination of the lectotype showed that the spores are transversely septate without any longitudinal septae. Old shriveled spores, however, give the false impression of being muriform. It is a widespread pantropical species characterized by the large emergent apothecia without periderm inclusions, wide pore, pruinose rim and disc, and lack of a columella. SPECIMENS EXAMINED.-4 (Wirth Sen.), 16b (37939). 25. Phaeotrema leiostomum FIGURES 5h, 64 14b,c Phaeotrema leiostomum (Tuckerman) Zahlbruckner, 1923: Thelotrema leiostornum Tuckerman, 1862: 407 [type-collection: 608. Cuba, Wright 149 (FH, lectotype); Figure 14bl. Thallus ashy to pale greenish white, epiphloeo- dal, continuous, smooth and shiny, forming colonies 2-5 cm broad; apothecia immersed to slightly emer- gent, 0.3-0.6 mm in diameter, weakly carbonized, columella absent to well developed, noncarbon- ized (Figure 64; pore round to irregularly elon- gate, surrounded by a raised ring, 0.1-0.3 mm long; hymenium 130p-150p high; spores 8/ascus, brown, 4 loculate, 8p-10~ X 16p-l8p, I- (Fig- ures? 5h, 14c). CHEMISTRY.-HypOprOtOCetrariC acid with or without 4-0-demethylnotatic acid along with pale unidentified pigments. HABITAT.-canOpy and mid-bole branches in rain forest (350-500 m). A raised annulate pore is characteristic of this species. Elongation of the apothecia was best de- veloped in collection 37956. The extreme form can be seen in Phaeotrema lirelliforme (Tuckerman) Zahlbruckner, which lacks any lichen substances. When apothecia are lacking, the thallus of Phaeo- trerna leiostomum alone would not permit identi- fication. 30 SMMITHSONIAN CONTRIBUTIONS TO BOTANY FIGURE 14.--Specimens of Thelotremataceae (all about X 10): a, Phaeotrema disciforme, new combination (Hale 37929); 6, P. leiostomzim (lectotype in FH); c, P. leiostomum (Hale 33203): d, P. obscurum, new species (Hale 38091); e, Thelotrema cara~sense (isotype in FH); f. T. carassense (Hale 37873); g, T. clandestinuni (lectotype in G); h, T. subcaesium (lectotype in H); i, T. clandestinum (Hale 33136): j, T. confornie (lectotype in G); k, 7?. consanguineum (lecto- type in G); 1, T. conforme (Hale 38062). NUMBER 16 31 SPECIMENS EXAMINED.-15 (35203), 16a (37956), 16b (37993). 26. Phaeotrema obscurum Hale, new species FIGURE 14d Thallus corticola, pallide brunneo-albidus, tenuis vel hypophloeodes, 5 cm latus; apothecia modice emergentia, 0.3-0.5 mm diametro, apice fuliginea, columellata; ostiolum rotundum, 0.1 mm dia- metro, intus albido-pruinosum; hymenium circa 12Oy altum; sporae 8:nae, transversim 5-6 loculatae, lop-14p X 24p-28p (Figure 14d). CHEMrsTnY.-stictic and constictic acids. HoLoTYPE.-Forty centimeters up trunk of a small tree, Rosalie Road above Newfoundland, ele- vation 300 m, Hale 38091, 10 December 1972 (US). HAnITAT.-Tree in rain forest. This tiny species could easily be overlooked. It is unusual to find a well-developed columella in apothecia this small. There are no other species in the genus with similar morphology. Phaeotrema albidulum (Nylander ) Muller-Argau also has stictic and constictic acids, but the apothecia are immersed and clustered and the spores larger (40P). Thelotrema Thelotiema is a common genus in Dominica, represented by 14 species among 80 collections. On the world level there are about 130 species. Variation in spore size is as great as in Ocellularia but longitudinal septae are always clearly de- veloped (except in T. clandestinum). The six com- monest species were T. praestans (21 collections), T. glaucopallens (16), T. clandestinum (lo), T. interpositum (7), T. leucomelaenum (7), and T. depressum (4). As I noted for Ocellularia, Thelotrema avoids the dry lowlands but is widespread in the rain forest. Thelotrema pupillosum and T. tuberculi- ferurn are restricted to the mossy forest. Key to the Species of Thelotrema 1. Central columella present. 2. Thallus thin, hypophloeodal; apothecia not emergent, small, less than 0.6 mm in diameter. 3. 2. Thallus distinct, epiphloeodal; apothecia emergent, 0.5-1.5 mm in diameter. 4. Spores 2Op-4Op long, 4-8/ascus. 3. Columella narrow; spores less than 2Op long .......................... 39. T. tenue, new species Columella very broad; spores more than 30p long ...................... 36. T. leucomelaenum 5. Spores about 20p long; pore not annulate but becoming black rimmed .................. 35. T. leucinum 5. 4. Spores more than loop long, l/ascus. 6. Thallus smooth, P- or P+ orange. 7. ................................................................................................................... Spores about 40p long; pore becoming annulate ................................ 29. T. confurme 6. Thallus warty-granular, Pf yellow ........................................... 34. T. interpositum Thallus P- ........................ :, ........................................................ 31. T. depressurn 7. Thallus P+ orange ................................................................... 38. T. praestans 9. Thallus greenish, thin, and shiny ................................................ 33. T. glaucopallens 10. Spores less than 2Op ......................................................... 30. T. confusum, new species 1. Central columella lacking. 8. Apothecia immersed, the pore flush with thallus surface. 9. Thallus whitish, thick, dull ............ .................... 28. T. clandestinum 8. Apothecia emergent, pores not flush wi 10. Spores more than 70y long. 11. Thallus P-; collected in mossy forest zone. 12. Apothecia 1 mm tall, eroding around a large area of the pore ...................... ................................................................................................... 40. T. tuberculiferum 12. Apothecia barely emergent, pore area not eroding ........................................ ................................................................................. 37. T. pupillosum, new species 11. Thallus P+ orange or yellow; collected in rain forest zone. 13. P+ yellow; apothecia up to 1.0 mm in diameter ................... 27. T. carassense 13. P+ red; apothecia about 0.5 mm in diameter .................................................. 32. T. dominicanum, new species ..................... ......................................... 32 SMITHSOXIAS CONTRIBUTIONS TO BOTANY 27. Thelotrema carassense FIGURE 14e,f Thelotrema carassense Vainio, 1890:79. TYPE-COLLECTION.-Carassa, Brazil, Vainio 1523 (TUR, lectotype; BM, FH, isotypes). (Figure 14e). Thallus greenish ashy mineral gray, epiphloeodal, continuous and shiny, forming colonies up to 15 cm broad; apothecia semi-emergent, 0.8-1.0 mm in diameter, upper wall weakly carbonized, columella lacking; pore about 0.1 mm in diameter, with a weak raised ring; hymenium IlOy-13Op; spores 8/ascus, muriform, 1-2 X 8-10 loculate, 12p-15p X 30p-40p, I+ blue (Figure 14f). CmMIsTRY.-Psoromic and conpsoromic acids. HABITAT.-Mid and lower bole of trees in rain forest (370-800 m). The Dominican material is not a perfect fit for the South American type, but it agrees in essential features, lack of columella, intermediate spores, and presence of psoromic acid. The apothecia are larger and spores smaller than in Vainio?s specimen (about 70y long). The taxonomy of the ecolumel- late psoromic acid-containing Thelotremnta is unfortunately still incomplete. Thelotrema post- positum Nylander has large spores (more than loop), but all other species, excluding T. caras- sense, have small spores (less than 20p long). 26b (37873). SPECIMENS EXAMINED.-16b (37859), 24b (37738), 28. Thelotrema clandestinum FIGURE 14g,i Thelotrema clandestinum Fee, 1837:90. Thelotrema subcaesium Nylander, 1869: 120 [type-collection: Brazil, Glariou 2193 (H, lectotype; BM, C, FH-Tuck, UPS, US, W, isotypes); Figure 14hl. Thelotrema concretum Fee var. subcaesiurn (Nylander) Red- inger, 1936:96. TYPE-COLLECTION.-on CinChOnU hZ?ZCifOliU in America, without collector (G, lectotype). (Figure Thallus whitish to greenish ashy gray, shiny, thick, and continuous, forming colonies up to 20 cm broad; apothecia numerous, immersed, 0.15- 0.3 mm in diameter, noncarbonized and without a columella; pore flush, round, 0.07-0.1 mm in diameter, usually with a whitish rim; hymenium 14g) * 65p-75y high; spores 8/ascus, uniseriate, 4-5 locu- late, muriform with 1-2 longitudinal septae in at least one of the locules, 6p-IOp X 15p-20y, I+ blue (Figure 14i). CHEMrsTRY.-Psoromic and conpsoromic acids. HABITAT.-canOpy branches in rain forest (500- 670 m). Thelotrema clandestinum has small spores with few longitudinal septations. It would, in fact, be classified as an Ocellularia unless the spores are carefully examined. Chemically similar 0. tere- bratula has only transverse septae. Both species have a thick, continuous thallus and occupy the same kind of habitat. They were collected at a total of ten localities but never together in the same one. (35103, 35105, 35136), 15 (35207), 16c (37768, 37872, 37981 ), 24b (35624a). SPECIMENS EXAMINED.-13 (35298, 35393), 14 29. Thelotrema conforme FIGURE 14jJ Thelotrema conforrne Fee, 1837389. Thelotrema consanguineurn Miiller-Argau, 1885b3398 [type- collection: Apiahy, Brazil, Puiggari 477 (G, lectotype); Figure 14kl. TYPE-COLLECTION.-On Cinchona, America meri- dionalis, without collector (G, lectotype) (Figure 141). Thallus whitish mineral gray, smooth and shiny, epiphloeodal, forming colonies up to 10 cm broad; apothecia emergent, 0.8-1.0 mm in diameter, api- cally carbonized, distinct columella present; pore round, 0.1-0.15 mm in diameter, surrounded by a whitish, sometimes raised ring; hymenium 170y- 200p high; spores 4/ascus, muriform, 8-10 loculate transversely, 2-4 loculate longitudinally, 18p-2Oy X 35y-50p, I+ blue (Figure 141). CHEMIsrRY.-Psoroniic and conpsoromic acids. HABrTAT.-Lower bole and trunk in open rain forest (670 m). The Dominican material is a satisfactory match for FCe?s type-specimen, which has slightly smaller apothecia and a less consistently developed annu- late pore. The spores are definitely in the in- termediate size range in contrast to related T. leucinum which has small spores. SPECIMENS ExAMINED.-24a (37739, 38062). NUMBER 16 33 30. Thelotrema confusunt Hale, new species Lep~ieu~ ?701 (Bhf, lectotype; H, isotype) (Figure FIGURES lc, 15 Thallus corticola, cinereo- vel viridi-albidus, epiphloeodes, planus, continuus vel rimosus, nitidus, 4-8 cm latus; apothecia numerosa, emer- gentia, 0.8-1.0 mm diametro, apice obfusca, columella nulla; ostiolum rotundum, 0.15-0.2 mm diametro, margine nigrocinctum; hymenium 120~- 130p altum; sporae 4:nae, murales, transversim 6-7 loculatae, longitudinaliter 0-2 loculatae, 6p-10~ X 12pL-26y, I+ coerulescentes (Figure 15). CHEmsTRY.-Protocetraric acid. HoLoTYPE.-Trai1 to Boiling Lake, elevation 650 m, Hale 37747, 9 December 1971 (US). HAnITAT.-Canopy branches or lower bole in open areas (650-800 m). The black-rimmed pore and fissured thallus re- semble Ocellularia nigropuncta, but the spores are clearly muriform. This presents an interesting ex- ample ol close morphological and chemical con- vergence with spore septation being the crucial diagnostic character. There are no related species in Thelotrema. SPECIMENS ExAMINED.-~~~ (37807), 26a (37697). FIGURE 15.-Thelotrema confusum, new species (Hale 37747) (X 10). 31. Thelotrema depressurn FIGURE 16b,c Thelotrema depressurn Montagne, 1851:73. TYPE-CoLLEcTIoN.-Cayenne, French Guiana, 16b). Thallus and apothecia as in Thelotrema prae- stans (see below); spores l-2/ascus, muriform with numerous transverse and longitudinal locules, 351.1- 50p. X 150p-300p, I+ blue (Figure 16c). CHEMISTRY.-NO substances present. HABITAT.-canopy branches of trees in the rain forest (300-500 m). The type-material of Thelotrema depresszim is very fragmentary but color and chromatographic tests established that no P+ compounds were present. The four specimens from Dominica are identical. They may represent only a chemical population of T. praestans, which is morphologi- cally identical, as far as we can judge from available specimens, but contains a strong P+ orange-red unknown substance. Thelot?.ema pl-ae- stans is more common (20 collections) and has a broader altitudinal range (250-800 m). 37922), 15 (35150). SPECIMENS EXAhIINED.-8 (38141), 12 (37901, 32. Thelotrema dominicanurn Hale, new species FIGURE I& Thallus corticola, pallide brunneo-albidus, epi- phloeodes, continuus, nitidus, 4-6 cm latus; apo- thecia numerosa, emergentia, 0.3-0.4 mm diametro, apice obfusca, columella nulla vel parce evoluta; ostiolum rotundum, 0.05 mm diarrietro; hymenium 140p-160p altum; sporae 2-4:nae, murales, trans- versim 24-30 Ioculatae, Iongitudinaliter 2-3 locu- latae, 15p-20p X 8Op-9Op, I+ coerulescentes (Fig- ure 16a). stances. HoLoTwE.-Tree along trail to Morne Anglais, Dominica, elevation about 800 m, Haze 35355, January 1969 (US). HAsrTAT.-Tree bole in higher elevation rain forest. The most unusual feature of this rare lichen is the presence of the ?cinchonarum? unknown, pre- viously thought to be restricted to the genus Ocellularia. Without a chemical test one might identify it with T. carassense, which has larger apothecia and a wider pore. Otherwise there are no comparable species in the New World. CHEMISTRY.-?CinChOnarUm? P -k unknown sub- 34 SMITI-ISONIAN CONTRIBUTIONS TO BOTANY FIGURE 16.-Specimens of Thelotremataceae (all about X 10): a, Thelotrema dominicanurn, new species (Hale 35355); b, T. depressurn (isotype in H); c, T. depressurn (Hale 38141); d, T. glaucopallens (lectotype in FH-Tuck); e, T. pechueli (lectotype in G); f, T. butuanurn (lectotype in W); g, T. homopastoides (lectotype in TUR); h, T. glaucopallens (Hale 35147); i, T. inter- positurn (isotype in L); j, T. interpositurn (Hale 35388); k, T. leucinum (lectotype in G); 1, T. leucinum (Hale 38100). NUMBER 16 35 33. Thelotrema glaucopallens FIGURE 16d,h Thelotrema glaucopallens Nylander, 1863:327. Pyrenula clandestiiia Acharius, 1814: 10 [type-collection: on Cinchona flava, South America (H, lectotype; S, isotype)]. Thelotrema laevigans Kylander var. avertens Nylander, 1867: 318 [type-collection: Tequendama, Colombia, Lindig 893 (H, lectotype; BM, FH-Tuck, G, P, FV, isotypes)]. Thelotrema pechueli Miiller-Argau, 1880:34 [type-collection: Quillu River, Angola, Pechttel-Loesclie s.n. (C, lectotype); Figure 16el. Ocellularia clandestina (Acharius) Miiller-Argau, 1877a:$. Thelotrema homopastoides Vainio, 1896:207 [type-collection: Richmond Valley, St. Vincent, Elliott 327 (TUR, lectotype: BM, isotype); Figure 16gl. Thelotrema butuanum Vainio, 1921: 183 [type-collection: Lu- zon, Philippines, Fenix, BS-28347 (FV, lectotype); Figure 16fl. TYPE-CoLLEcTIoN.-Cuba, Wright (FH-Tuck, lectotype, as 28; L, UPS, isotypes) (Figure 16d). Thallus light greenish gray, very thin and shiny, breaking away in thin sheets, forming extensive colonies up to 15 cm broad; apothecia numerous to rather rare, immersed to slightly emergent, 0.2- 0.4 mm in diameter, uncarbonized or slightly car- bonized apically, without a columella; pore flush to barely raised, round, rarely angular, 0.05-0.12 nim in diameter; hymenium 6Op-8Op high; spores 8/ascus, muriform, 6-8 loculate transversely, 2-3 loculate longitudinally, 7p-lop X 15p-24~~ I - (Figure 16h). CHEMISTRY.-stictic and constictic acids. HARITAT.--Bu ttresses, base of large trees, exposed roots, lianas, rarely upper bole and canopy in the rain forest (450-800 m). The thallus of this pantropical species is unique. It is extremely thin and shiny and tends to break away in thin waxy sheets when bruised or cut. The color is a pyrenocarplike dark yellowish green. Apothecia are usually quite numerous with flush pores having considerable variation in development of a rim, from no distinct annulus to an obviously raised area. Some anomalous specimens may even have semi-emergent apothecia, but the small I - spores and the presence of stictic acid will posi- tively identify them. SPECIMENS EXAMINED.--IO (38082), 11 (37999, 38008, 38047), 15 (35147), 16a (37670, 37876, 38043, 38044), 20a (37783), 21 (38083, 38095, 38155, 38156), 23 (37678), 26b (35367), 27 (37891), 28 (35431). 34. Thelotrema interpositurn FIGURE 16i,j Thelotrema rnterposztum (Sylander) Muller-Argau, 1881:526. Asczdzum znterposztum Sylander, 1863 336 [type-collection: Cuba, Wrzght 28 (H, lectotype; BM, FH, L, M, P, UPS, isotvpes); Figure lGz]. Thallus light brownish ashy gray, epiphloeodal, smooth to minutely warty, forming colonies 5-8 cm broad; apothecia strongly emergent, 0.8-1.1 mm in diameter, n alls heavily carbonized, columella present; pore round, 0.15-0.2 mm in diameter, opening into a tubelike area leading to the prui- nose top of the columella; hymenium 2OOp high; spores 2/ascus, muriCorm with numerous transverse and longitudinal locules, 28p-40p X 85p-125~, I+ blue (Figure 16j). CHEMISTRY .-PSOrOmiC acid. HABITAT.-CanOpy branches and upper bole area (sea level to 670 m). The coarsely verrucose thallus and large emer- gent apothecia with large spores distinguish this species. Two other columnellate psoromic acid- containing species, T. conforme and T. leiiciniim, hale a smooth thallus and much smaller spores. (35148, 35174), 16a (37656, 37657), 16c (37760), 22 (37853, 37976). SPFCIXIFNS EXAMINED.--E) (35388), 7 (35231), 15 35. Thelotrema leucinum FIGURE 16kJ Thelotrema leucinum Muller-Aigau, 188ia: 10. TYPE-COLLECTION.-on Cinchona, South Amer- ica (G, lectotype) (Figure 16k). Thallus light ashy gray, epiphloeodal, continuous to cracked with age, shiny, forming a colony about 10 cm wide; apothecia numerous, nearly immersed to semi-emergent, 0.4-0.6 mm in diameter, upper wall carbonized, columella present, 0.2 mm wide; pore round, 0.05-0.12 mm in diameter; hymenium 1 IOp-120p high; spores 8/ascus, muriform with 4 locules transversely, 2 locules longitudinally, 7p- Ilk X 12u-18p, I+ blue (Figure 162). CHEhiIsTRY.-Psoromic and conpsoromic acids. H.4nITAT.-canopy branches in rain forest (300 The type of T. Zezicinum, one of Fee?s Cinchona in). 36 SMITHSONIAN CONTRIBUTIONS TO BOTANY bark specimens, is in very poor condition and it is unfortunate that Muller selected it. Identifica- tion with the Dominica material is at best provi- sional, depending largely on the wide columella, small spores, and presence of psoromic acid. A better understanding of the species will come when more specimens are available. SPECIXIEN EXARfINED.-g (38 100). 36. Thelotrema leucomelaenurn FIGURES 4a,b; 6e; l'ia,d Thelotrema leucomelaenurn Nylander, 1863:3?9. Thelotrema pauperius Nylander, 1867:318 [type-collection: Rio Negro, Colombia, Lindig (H, lectotype; BM, M, iso- types); Figure l'ib]. Thelotrema leucomelaenum var. elevatum Vainio, 1915: 137 [type-collection: Morne Anglais, Dominica, Elliott 1534 (TUR, lectotype); Figure 17~1. TypE-CoLLEcTIoN.-Fusagasuga, Colombia, Lin- dig 2777 (H, lectotype; BM, FH, G, M, P, UPS, isotypes) (Figure 17~). Thallus white, thin and in part hypophloeodal (Figure 4a,b), forming colonies up to 8 cm broad; apothecia immersed to semi-emergent, 0.6-0.9 mm in diameter, apically carbonized with a broad columella 0.2-0.3 mm wide; pore round to sub- irregular, 0.2-0.3 mm across, opening to the pruinose columella top (Figure 6e) ; hymenium 130p high; spores 8/ascus, muriform, 4-6 loculate transversely, 2-3 loculate longitudinally, 12p-15y X 30~-3Gp, I- (Figure 17d). CHEMISTRY.-NO substances present. HABITAT.-Sma~kr branches of trees in mossy forest (800-900 m). A pantropical species, T. leucomelaenurn has been correctly identified by most lichenologists. The apothecia, while at most semi-emergent, stand out because of the black pore rim. The broad columella occupies most of the interior of the apothecia. 37718, 37871), 25b (35491), 26a (38076). SPECIMENS ExAxfINE~.-25a (37706, 37710, 37717, 37. Thelotrema papillosum Hale, new species FIGURE 17j Thallus corticola, cinereo-viridis, epiphloeodes, minute papillosus, nitidus, 6 cm latus; apothecia vix emergentia, 0.7-0.9 mm diametro, intus in- colores, columella nulla; ostiolum rotundum vel irregulariter laceratum, 0.1-0.3 mm diametro; hymenium circa 2001~ altum; sporae l-2:nae, mu- rales, loculis numerosis, 3Gp-40p X 14Oy-l50y, I- (Figure 17j). CHEMISTRY.--NO substances present. HoLoruPE.-Rain forest above Newfoundland, Dominica, elevation about 300 m, Hale 38124, 7 May 1972 (US). HARITAT.-CanOpy branch in rain forest. This species would be easy to miss in the field. The small greenish warty thallus is not like any Thelotrema. The large noncarbonized apothecia are not numerous and seem lost among the thallus irregularities. The closest relative in terms of apo- thecial structure is Thelotrema tuberculiferum, which has large spores but strongly emergent erod- ing apothecia. 38. Thelotrema praestans FIGURE 17e,g Thelotrema praestans Miiller-Argau, 1895:433. Thelotrema elliottii Vainio, 1896:207 [type-collection: Rich- mond Valley, St. Vincent, Elliott 246 pro parte (BM, lecto- type); Figure 17fl. Tk.PE-CoLLECTIoN.-Rio de Janeiro, Brazil, Por- telln s.n. (G, lectotype; BM isotype) (Figure 17e). Thallus light greenish to whitish gray, smooth and continuous, epiphloeodal, forming colonies 5-10 cm broad; apothecia semi-emergent to strongly emergent, up to 2 mm in diameter, heavily car- bonized with a thick columella; pore 0.15-0.3 mm in diameter, opening through a deep tube into the ascocarp; hymenium about 2OOp high; spores I-Z/ascus, muriform with many transverse and longitudinal locules, 3Op-45p X lOOy-275p, I+ blue (Figure 17g). CHEMIsTRY.-"Praestans" unknown. HABITAT.-CanOpy branches, less commonly bole, liana, or base in rain forest (250-800 m). This conspicuous thelotreme is related to T. de- p7.esszim but differs in producing a P+ orange-red unknown substance just below psoromic acid in both liexane and benzene solvents. The apothecia are large and heavily carbonized. The fine struc- ture of the cortex is aculeate, as in T. depressum. 35273), 8 (38087, 38121, 38159), 9 (38135), I1 SPECIMTXS EXAMINED.-7 (35237, 35244, 35254, NUMBER 16 37 FIGURE lj.--Specimens of Thelotremataceae (all about X 10): a, Thelotrema leucornelaenuifz (lectotype in H); b, T. pauperins (lectotype in H); c, T. leucomelaenurn var. elemturn (lectotype in TUR); d, T. leucomelaenuni (Hale 35491); e, T. pmestuns (lectotlpe in G); f, T. elliottii (lectotype in Bhl); g, T. praestans (Hale 35273); h, T. ticberculiferum (lectotype in TUR); i, T. ,tuberculiferum (Hale 35519); 1, T. pupillosum, new species (Hale 38124); k, T. tenue, new species (Hale 35430). 38 SMITHSONIAN CONTRIBUTIONS TO BOTANY (37821, 37998, 38002, 38048), 12 (37907, 37911), 15 (35171), 18a (37734, 37968), 18b (35521), 20a (37776), 26a (38070). 39. Thelotrema tenue Hale, new species FIGURES 44; 6f; l'ik Thallus corticola, cinereo-albus, hypophloeodes (Figure 4c,d), nitidus, 7 cm latus; apothecia nu- merosa, immersa vel parce emergentia, 0.3-0.4 mm diametro, apice obfusca, columellata (Figure 6f) ; ostiolum rotundum, 0.05-0.1 mm diametro; hy- menium 90p-1OOp altum; sporae 8:nae, murales, transversim 4-5 loculatae, longitudinaliter 1-2 loculatae, 6p-7p X 17p-l9p, I- (Figure 17k). CHEMISTRY.--NO substances present. HOLOTYPE.--R/IOSS~ forest zons on trail to Morne Diablotin, Dominica, elevation about 1100 m, Hule 35430, January 1969 (US). HABITAT.-LOM'er trunk in rain forest to mossy forest. Superficially this species resembles Ocellularia pyrenuloides in having small blackish apothecia and a thin columella. Thelotrema leucomelaenurn, another hypophloeodal mossy forest species, has larger spores and a very broad columella. Other than these there are no related species. 40. Thelotrema tuberculiferum FIGURE 17h,i Thelotrema tuberculiferum Vainio, 1915:136. TYPE-CoLLzCTI0N.-Savane-aux-Ananas, Guade- loupe, Duss 1497 (TUR, lectotype) (Figure 17h). Thallus ashy whitish, thin and in part evanes- cent, scattered, forming colonies 4-8 cm broad; apothecia strongly ascidioid, about 1.0 mm in diameter and 1.0 mm high, noncarbonized and lacking a columella, cortex eroding away toward the apex leaving a pulverulent area; pore round, 0.1-0.2 mm in diameter, surrounded by a gray, sometimes raised ring and the broader pulverulent area; hymenium 140p-160p high; spores l-Z/ascus, muriform with numerous transverse and longitudi- nal locules, 30p-50p X 60p-120c(, I+ pale blue (Figure 17i). CHEMISTRY.-NO substances present. HABITAT.-Exposed moss-covered branches in mossy forest (800-900 m). This Thelotrema might be mistaken at first for a Pertusaria. It was first described from Guade- loupe and is probably endemic to the Lesser An- tilles, occurring at high elevations in the upper limit of the mossy forest. SPECIMENS EXAMINED.-25a (377 19), 26a (37868). Lept otrema This genus differs from Thelotrema in having brown spores which tend to shrivel at maturity regardless of size and appear to be I - . Eight species occur on Dominica among my 36 collections (in- Key to the Species of Leptotrema 1. 1. Columella lacking. Columella present, simple to actinoid .......................................................... 43. L. fissum Pore wide, 0.5-0.8 mm in diameter, the exposed disc dark brown to black ......................... Thallus thick with a columnar cortex; red crystalline inclusions often present ............. ............................................................................................................................ 48. L. wightii Spores about 2Oy long ............................................................ 45. L. occultum Spores about 45p long ....................................................... 47. L. subcornpunctum 2. 2. Apothecia immersed to emergent with pores 0.1-0.2 mm in diameter. 3. 3. Thallus thin with a normal cortex lacking pigments. 4. Apothecia with an inner lepadinoid exciple. 5. 5. 4. Apothecia lacking a lepadinoid exciple. 6. Pore distinct, 0.1-0.2 mm in diameter; spores 20~-30~ long 6. Pore very tiny, less than 0.1 mm in diameter; spores large, more than 40y long. 7. 7. Pore not depressed, surrounding area the same color as the thallus. 8. Apothecia immersed with flush pores ................ 47. L. subcompunctiim 8. .4pothecia more or less emergent ................. 44. L. microgluenoides ................................................................................................................ 46. L. spondaicum ..41. L. bahianum Pore depressed in a darker area .......................... 42. L. deceptum, new species NUMBER 16 39 cluding the collection of L. bahianum by Elliott). It is rarer than would guess from this number of collections since one species, L. deceptum, is represented by 2 1 specimens. Leptotrema spondai- czim was found 8 times, L. wightii 3 times, and tlie remainder once each. On the world level the genus has about 50 species. Leptotrema behaves in a peculiar way on Dominica. It is virtually the only thelotreme (5 of 7 collections) in the dry scrub forest. It then essentially skips the rain forest zone and reappears in the submossy forest. Only 5 collections were made in tlie broad rain forest zone in contrast to 23 above 900 m. 41. Leptotrema bahianum FIGURE 5f Leptotrema bahiatiurn (Acharius) Muller-Argau, 188?ia:12. Thelotrema lepadinurn var. bahianutrt Acharius, 1803: 132 [type-collection: Bahia, Brazil, without collector (H, lecto- type; L, UPS, isotypes)].-Hale, 19723193, fig. 2c. Thelotrema ritditis Vainio var. dominicanttm Vainio, 1915: 136 [type-collection: Prince Rupert, Dominica, Ellioft 1303 (TUR, lectotype); Figure 5fl. Tliallus corticolous, epiphloeodal, light tannish white, about 4 cm broad; apothecia numerous, emergent, 0.4-0.6 mm in diame:er, noncarbonized and without a columella; pore round, about 0.1 mm in diameter; hymenium 150p high; spores 8/ascus, muriform with 5-6 transverse locules and 2-3 longitudinal locules, 12p-14p X 16y-24y, I- (Figure 5f). CHniIsmY.-Protocetraric acid. HABITAT.-Tree branch in dry scrub forest near sea level. This well-known Acharian species (Hale, 1972) seems to be very rare on Dominica since it has not been found since Elliott?s time. It is generally a loivland species in drier areas of the tropics. 42. Leptotrema deceptum Hale, new species FIGURES ?id, 18 Thallus corticola, brunneo-cinereus, epiphloeo- des, planus, nitidus, continuus vel rimosus, usque ad 12 ciii latus; apothecia immersa, 0.3-0.4 mm diametro, intus incolores, columella nulla; ostiolum minutum, 0.02-0.05 mm diametro, depressum, ambitu obfuscans; hymenium 100p-120y altum; sporae obfuscae, 4:nae, murales, loculis numerosis, 14p-28p X 4Op-7Op, I- vel I+ pallide coerules- centes (Figure 18). CHEXrIsmY.-Stictic and constictic acids. Hor,oTYPE.--Sloanea buttress, disturbed primary foiest at Middleham Estate, elevation about 670 m, Hole 37860, December 1971 (US). HABIrAT.-Buttresses and exposed roots, saplings, liarias, lower bole in higher elevation 1 ain forest (about 670 m). This species belongs to a large group of difficult species characterized by a very tiny pore and pres- ence of stictic and constictic acids, including in Dominica L. micioglnenoides and L. subcompzinc- tzim. It is separated from related species by the smooth, usually fissured thallus and a distinct dark- er depressed area surrounding the pore, this area in turn surrounded by a whitish ring. Spent apo- thecia remain as large, open corticate pits. At first I had considered this (and similar 5pecies) as having closer relationships to the Py- ienulaceae. Mr. Richard Harris, a student of tlie Pyrenulaceae, gave the folio\+ ing opinion in sup- port of placing L. deceptzim in the Thelotrema- taceae: hlost pFienocaipous lichens ha\e 5ome sort of carbonized ?aall? surrounding the h\menium [I. deceptzctn is noncar- bonized]. The spoies react nith IKI, a few pFrenocaips hale this reaction also but hate entirel\ different paraphFses. The paraph>ses [of L. deceptitrn] are unlike thme in the pyreno- 40 SMITHSONIAN CONTRIBUTIONS TO BOTANY carpous lichens in the relative lack of branching and espe- cially in the very thick, coherent gelatinous wall or sheath around them (also found in the Graphidaceae). The asci are also unlike any pyrenocarpous asci. They are closest to Porina but lack any ?chitinoid? apical ring. Thelotrema lepadinum has asci very like those in your specimen. Finally I have never found depsides or depsidones in any pyrenocarp- ous lichens. SPECIMENS EXAMINED.-21 (37373, 38104, 38105, 38109, 38110, 38112, 38117, 38133, 38136, 38139, 38142, 38148, 38151, 38153), 22 (37640, 37648, 37836, 3i839, 38037), 23 (37673). 43. Leptotrema fissum FIGURE 19a,c Leptotrema fissurn (Nylander) Muller-Argau, 1882a.333. Thelotrema fssuin NJlander, 18593258 [type-collection: Boui- bon, Richard s.n. (H, lectotype; G, P, isotypes); Figure Leptotrema zntegtunz M~ller-~4rgau, 1887b:399 [type-collec- tion: Australia, Sajer (G, lectotjpe, S, isotjpe); Figure 19b]. Leptotreina fisszcrn (Nylander) Zahlbiuckner, 1923.634 [supei- flous combination]. Thallus greenish ashy gray, epiphloeodal, smooth, continuous or cracked with age, forming colonies 4-6 cm broad; apothecia semi-emergent, round to irregular, 0.7-1.2 mm wide, with thick erect to coarsely recurved carbonized upper walls, columella variable, barely developed to actinoid, the disc white pruinose; pore initially round but opening at maturity, up to 1 mm broad; hymenium about 100p high; spores brown, muriform with 4-6 transverse locules and 0-2 longitudinal locules, 6p-7p X lOp--13p, I- (Figure 19c). CHEhlISTRY.-PSOrOmiC and conpsoromic acids, HABITAT.-canopy branches in rain forest (430 The diagnostic features of this pantropical species are the open, often imperfectly actinoid disc, semi-erect lacerated margin, and presence of psoromic acid. It is close in these respects to Ocel- 121 la ria compara bilis. 19aI. m). SPECIMEN EXAMINED.--~ 1 (38031). 44. Leptotrema microgluenoides FIGURE 19d,e Leptotrerna microgluenoides (Vainio) Zahlbruckner, 1923:637. Thelotrema microgluenoides Vainio, 1896:206 [type-collection: St. Vincent, Elliott 266 (BM, lectotype); Figure 19dI. Thallus light whitish tan, epiphloeodal, con- tinuous and shiny, forming a colony about 6 cm broad; apothecia semi-emergent, 0.8-1 .O mm in diameter, walls not carbonized, columella lacking; pore very tiny, 0.02-0.05 mm in diameter, depressed but with a tiny darker ring at the periphery; hymenium 15Op-17Op; spores brown, 4/ascus, muri- form with numerous transverse and longitudinal locules, 18p-22p X 36p-48p, I- (Figure 19e). CHEwsTRY.-Stictic and constictic acids. HAnITAT.-Branches of tree in rain forest (430 The lectotype of this species is rather frag- mentary with very few well-developed apothecia. The thallus is continuous, but with a faint grainy appearance, thin and shiny. The Dominican ma- terial is?close to the type-specimen, although Vainio reported spores in the range of 120p long. I found spores about 50p long in the lectotype. The large complex of stictic acid-containing species (e.g. Leptotrenza monosporum (Nylander) hluller- Argau, L. phaeosporum (Nylander) Muller-Argau, and L. wclzisum (Krempelhuber) Zahlbruckner) is still far from resolved. m). SPEcmrrs EXAMINED.-~ 1 (37984). 45. Leptotrema occultum (Eschweiler) Hale, new combination FIGURE 19f,g Thelotremu occulfum Eschweiler in Martius, 1833: 174 [type- collection: South America (M, lectotjpe; G, isotype); Figure 19fl. Thallus tannish white, epiphloeodal, continuous, smooth to minutely pitted, forming colonies 3-5 cm broad; apothecia numerous, immersed, 0.2-0.3 mm in diameter, noncarbonized and without columella, inner lepadinioid exciple well devel- oped and pulling away from the wall, lightly pruinose surface of hymenium visible; hymenium 65p-7Op high; spores 4-8/ascus, brown, muriform with 5-7 transverse locules, 2-3 longitudinal locules, 9p-lop X 18p-21p, I- (Figure 19g). CHEMISTRY.-Norstictic acid. HABITAT.-Trees in dry scrub forest near sea level. Leptotrema occulturn can be typified with the Martius specimen in Munich and the isotype frag- ment in Geneva. It is virtually indistinguishable NUMBER 16 41 FIGURE 19.--Specimens of Thelotremataceae (all about X 10): a, Leptotrema fissum (lectotype in H); b, L. integrum (isotype in S); c, L. fissuiit (Hale 38031); d, L. microgluenoides (lectotype in BM); e, L. microgluenoides (Hale 37984); f, L. occulturn (isotype in G); g, L. occz~ltz~rn (Hale 35718); h, L. spondaicum (lectotype in FH-Tuck); i, L. spondaicum (Hale 35503). from pantropical L. compunctum (Acharius) Muller-Argau, which contains stictic acid in addi- tion to norstictic acid. I am keeping L. occultum 46. Leptotrema spondaicum FIGURE 19h,i Leptotrema spondaicum (Nylander) Zahlbruckner, 1923:640. separate until study of more material shows being generally smaller in norstictic acid-containing specimens) are valid or not. The L. compunctum group is very characteristic of dry scrub forest throughout the tropics. whether the and 'pore (spores Thelotrema spondaicum Nylander, 1863:330 [type-collection: Cuba, Thallus ashy white, epiphloeodal to evanescent, waxy, smooth, forming colonies 3-15 cm wide; apothecia numerous, round to irregular, 0.8-1.4 35 (FH.Tuck, lectotype); Figure 19hl, SPECIMEN EXAMINED.-2 (35718). 42 SMITHSONIAN CONTRIBUTIONS TO BOTANY mm across, emergent, walls weakly carbonized, columella lacking; pore initially distinct, about 0.2 mm wide, but soon opening, wide and irregular, to 0.5 mm wide, exposing the dark brown disc, outer rim becoming pulverulent; hymenium 150~- l7Op high; spores 1-2/ascus, brown, muriform with numerous transverse and longitudinal locules, 30p- 40y X 80~-130p, I- (Figure 19i). CmmsmY.-Virensic acid. HABITAT.-Trunks of primary and secondary trees in open pastures and citrus groves at higher elevations (600-700 m). Leptotrema spondaicum stands apart from all other thelotremes in having an open blackish disc surrounded by a thick, often pulverulent mar- gin. One might be tempted to treat it as a species of Diploschistes. The chemistry is unique in the family. It is also different ecologically in that the primary habitat seems to be planted trees in dis- turbed areas. SPECIMENS EXAMINED.-lSb (35590), 19 (38077), 20a (37779, 37789, 37790), 20b (35492, 35503), 26a (37696). 47. Leptotrema subcompunctum FIGURE 20a,b Leptotrema su bcompunctum (Nylander) Zahlbruckner, 1923: 640. Thetotrema subcompunctum Sylander, 1868:76 [type- collection: Lifu, Loyalty Islands, Thie'baut s.n. (H, lecto- type; G, P, i$otypes); Figure 20al. Thallus light tannish mineral gray, epiphloeodal, continuous, smooth, dull, forming colonies 4-5 cm broad; apothecia numerous, 0.3-0.4 mm in diame- ter, immersed to slightly emergent, noncarbonized and without a medulla, the inner lepadinioid exciple clearly developed; pore round, 0.05-0.1 mm in diameter; hymenium 140p-160p high; spores FIGURE ZO.--Specimens of Thelotremataceae (all about X 10): a, Leptotrema subcompunctum (isotype in P); b, L. subcompunctum (Hale 37888); c. L. wightii (lectotype in FH); d, L. pavicans (lectotype in G); e, L. subconcretum (lectotype in BM); f, L. wightii (Hale 35882). NUMBER 16 43 brown, I-Z/ascus, muriform with 8-1 1 transverse locules and 2-4 longitudinal locules, 10y-13y X 45c(-55y, I- (Figure 20b). CHEmsTRY.-Stictic and constictic acids. HABITAT.-Tree bole in open rain forest (600 m). Except for the inner partially free exciple this species would probably be identified as L. micro- glaenoides. The thallus, however, is dull and the ascocarps barely emergent to immersed. This is the only collection that I have seen from the New World. SPECIMEN EXAMINED.-~~~ (37888). 48. Leptotrema wightii FIGURES Id, 20c,f Leptotrema wightii (Taylor) Miiller-Argau, 1882:499. Endocarpon wightii Taylor, 1847: 155 [type-collection: Madras, India, Wight (FH, lectotype; BM, G, isotypes); Figure ~OC]. Thelotrema prevostianurn Montagne, 1849:292 [type-collection: Antilles, Prkvost s.n. (P, lectotype); Figure 5d]. Leptotrema prevostianurn (Montagne) Montagne, 1856:364. Theloftrema subconcretum Leighton, 1869: 169 [type-collection: Central Province, Ceylon, Thwaites 89 (BM, lectotype; H, G, P, S, UPS, W, isotypes); Figure 20e]. Phaeotrema subconcretum (Leighton) Miiller-Argau, 1887a: 10. Leptotrema subconcretum (Leighton) Muller-Argau, 1891:277. Leptotrema fiauicans Miiller-Argau, 1888: 114 [type-collection: Guarapi, Paraguay, Balansa 4170 (G, lectotype; BM, M, W, isotypes): Figure 20d]. Thallus pale greenish mineral gray, thick and epiphloeodal, bulging up and flaking away with age, surface smooth but grainy in appearance, the columnar cortex with or without red crystal masses (Figure Id), forming colonies up to 10 cm broad; apothecia immersed, 0.2-0.3 mm in diameter, non- carbonized, without a columella; pore round, 0.08- 0.11 mm in diameter, flush; hymenium about lOOy high; spores brown, 8/ascus, muriform with 4-6 transverse locules and 1-2 longitudinal locules, 9y-lly X 2Oy-28yL, I- (Figure 20f). CHEMISTRY.-NO substances present except for the unidentified anthraquinone pigment. HABITAT.-Tree trunks in dry scrub forest near sea level. This pantropical species has been discussed in detail by Salisbury (1971). It seems to occur at low elevations in dry scrub forest and extends northward into temperate forests of North Amer- ica and southward into Argentina. SPECIMENS EXAMINED.--~ (35582, 35584), 3 (3 29 1 8). Literature Cited Acharius, E. 1803. Methodus qua onznes detectos Lichenes. 394 pages. Stockholm. 1812. Anmarkningar vid Lafslagtet Thelotrema med Nogare Bestammande af Dess Arter. Kongliga Vet- enskapliga Academiska Nya Handlingar, 1812:79- 95. Synopsis Methodica Lichenurn. 392 pages. Lund. The Constituents of Some Species of the Thelotre. mataceae. The Journal of Japanese Botany, 43:316- 323. 1814. 1968. Culberson, C. F., and W. L. Culberson. Culberson, C. F., and M. E. Hale, Jr. 1973. 4-0-Demethylnotatic Acid: A New Depsidone in Some Lichens Producing Hypoprotocetraric Acid. The Bryologist, 76:77-84. Fee, A. L. 1824. Essai sur les cryptogames des ecorces exotiques oficinales. 167 pages. Paris. 1837. Essai sur les cryptogames des ecorces exotiques oficinales, II: Supplement et Revision. 178 pages. Paris. Hale, M. E., Jr. 1971a. Morden-Smithsonian Expedition to Dominica: The Lichens (Parmeliaceae). Smithsonian Contributions to Botany, 4:l-25. 1971 b. 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Van 0vereem.De Haas, C. and D. 1912. 1932. dem Jahre 1920 Gefundenen Myxomycetes, Fungi Index (Index to main entries only: synonyms in italics) Ascidium cinchonarum, 17 interpositum, 35 rhodostromum, 25 Endocarpon wightii, 43 Graphis alborosella, 16 Leptotrema bahianum, 39 deceptum, 39 exalbidum, 29 fissum, 40 pavicans, 43 integrum, 40 microglaenoides, 40 monosporum, 40 occultum, 40 phaeosporum, 40 prevostianum, 43 reclusum, 40 spondaicum, 41 subcompunctum, 42 subconcretum, 43 wightii, 43 Myriotrema oliuacea, 24 Ocellularia alboolivacea, 19 alborosella, 16 antillensis, 17 berkeleyana, 17 bonplandiae, 24 cavata, 17 cinchonarum, 17 cinchonarum f. intermedia, 17 clandestina, 35 comparabilis, 17 comparabilis var. microcarpa, 17 concolor, 19 conglomerata, 19 discoidea, 17 doming~nsis var. fecunda, 20 dominicana, 19 efformata, 17 exanthismocarpa, 20 fecunda, 20 glaziocii, 17 homothecia, 20 isertii, 20 latilabra, 19 lindigiana, 17 maculata, 22 mordenii, 22 multilocularis, 20 nigropuncta, 22 olivacea, 24 papillata, 24 perforata, 25 platycarpella, 16 porinoides, 20 pyremloides, 25 remanens, 28 rhodostroma, 25 rimosa, 27 rufocincta, 25 sorediata, 27 subcavata, 27 terebrata, 17 terebrata var. abbreviatula, 25 terebratula, 28 riaga, 28 verrucosa, 24 xanthostroma, 22 albidulum, 31 disciforme, 29 galactinum, 29 leiostomum, 29 lirelliforme, 29 obscurum, 31 siibconcretum, 43 Phaeotrema aggregatum, 29 Pyrenula clandestina, 35 Thelotrenia aquilinum, 29 butuanum, 35 carassense, 32 cavatum, 17 clandestinum, 32 clandestinum f. remanens, 28 cornparabile, 17 concretum car. subcaesium, 32 conforme, 32 confusum, 33 consanguineum, 32 depressum, 33 disciforme, 29 domingense car. fecundum, 20 dominicanum, 33 elliottii, 36 exalbidum, 29 exanthismocarpum, 21 excavatum, 25 excavaticm var. impressulum. 13, 25 fissum, 40 galactinurn, 29 glaucopallens, 35 homopastoides, 35 homotkecium, 20 interpositum, 35 laevigans var. avertens, 35 leiostomum, 29 lepadinum var. bahianum, 39 leucinum, 35 leucomelaenurn, 36 leucomelaenum var. elevatum, 13, 36 microglaenoides, 40 occultiim, 40 papillatun, 24 papillosum, 36 pnuperius, 36 pechueli, 35 perforatum, 25 platycarpellum, 16 porinoides, 20 postpositum, 32 praestans, 36 prevostianum, 43 rhodostromum, 13 rzidius Lar. dominicaizum, 13, 39 spondaicum, 40 subcaesium, 32 subcavatum, 28 su bcompunctum, 42 subconcretum, 43 tenue, 38 terebratuliam, 28 terebratuin var. abbreviatulum, 25 tuberculiferum, 38 vagum, 28 U.S. GOVERNMENT PRINTING OFFICE: 1974--546--361/14 46