^ z rn 'ti rn _
^ — if> ^ (j^ ?
josHiii^s S3iyvaaii libraries Smithsonian institution NoiiniiiSNi NviNOSHims saiavaan l
■" 'S ^ ^ ^ ^ 2 CO 2
- I ^ i i ^
^ - ^ > 2 ■ >' s XJiVosi^ >' 5
^ CO '■ ■*^ to '•*■' 2 CO 2 CO
HSONIAN INSTITUTION NOIiniliSNI_ NVINOSHilWS SBIdVaan LIBRARIES SMITHSONIAN INSTITUTION N
“ 2 \ ^ ^ ^ ^ (/) = CO
josHiiws saiavaan libraries smithsonian institution NoiiniiiSNi nvinoshiiws S3iavaan l
z r- 2 r- z
m rri ''<iSLBs^ ^ m ^ m
_ CO _ CO ± — CO
‘HSONIAN INSTITUTION NOlinillSNI NVINOSHIIINS SBiavaail LIBRARIES SMITHSONIAN INSTITUTION ^
2 .V,. to 2 to 2 CO Z V.- CO
_ ^ 3, 2
iosHiii:is‘^S3iavaan libraries smithsonian institution NoiiniiiSNi nvinoshiiws'^sb i ava a l
CO CO ^ to — to 2
HS0N)AN"'lNSTITUTI0N^N0liniIiSNl‘^NVIN0SHimS^S3 I ava a n"^LI B RAR l ES^SMITHSOnIan'^INSTITUTION^ N <~ 2 n, 2 r- 2: ^ 2
ID ^ ID z ID ^ * rn
CO £ to ' — CO £ to
losHims S3iavaan libraries smithsonian institution NoiiniiisNi nvinoshihns saiavaan l
to 2 , CO ^ 2 T CO 2 to . 2
I o
HSONIAN INSTITUTION '^NOIiniliSNI^NVINOSHims‘^S3 I a Va 3 H^L I B R A R I E S^’sMITHSONIAN^INSTITUTION
to ^ \ to zz V) ~
"H ^ ■aiy -1 ^
Q ><vosti>' — o O o
osHims S3iavaan‘^LiBRARiEs smithsonian institution N0iiniiiSNi"'NviN0SHiiiMS^S3iavaan"^Li
S ^
?%> H- 3J t; y.'s/ yfiSr ;o 4r^”^^fA t ^
CO XI >
XJ
m ^si2iDc>- - m ^ ^ to m
_ CO £ to £ — (/> \ 2 ^ CO
HSONIAN INSTITUTION NOlinillSNI NVINOSHiltMS SBiaVaaiT LIBRARIES SMITHSONIAN INSTITUTION ^
I
CO
o
>
2
rn -'V^' 7 "s^Asq>^ m x.i;»ostjiX ^ ^ ^ m ^
CO ' ■ — CO ? to — ^
ES SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHimS S3 I HVd 9 11 _L I B R AR I ES^^SMITHSONIAN INSTITL
CO 2 » CO 2 CO
j£^-- ° i=
-i- tj /^V' ^
t /■•'^' t
j, \iVA22>' 5 5'' 2 > -a
2 to ••’■"■ 2 to 2 t/i "' ^ in
;NI_NVINOSHilWS S3iaVyan_LIBRARIES SMITHSONIAN_INSTITUTION ^N0liniliSNI_NVIN0SHimS^S3 I H V
i z I ^ i ^ ^
< d;
CQ
ES SMITHS0NIAN'^INSTITUTI0N^N0liniIiSNrNVIN0SHilWS“S3 I ava a n LIBRARIES SMITHSONIAN INSTITl
2 f“ 2 1-2'“ ?
O
2
^'2 / ^ O fj
tu 70 >
m 'ii N^oTuvV' m '^j'u'i5i>' rn Nsato;^ 5; m
CO 5 CO \ Z CO ± to
SNI NVINOSHilWS S3 lava an libraries SMITHSONIAN institution NOIinillSNI NVINOSHIIIMS S3iav
2 to 2 .... W 2: “
2 .... 'to' z ■' CO 2 to
<t . 2 .< s
X /pY^ in o
? V'"’ _ _
ES'^SMITHSONIAN INSTITUTION NOIinillSNI NVINOSHlIWs'^S 3 I a V a 8 11 LIBRARIES SMITHSONIAN INSTITl <0 r: CO — <0 ^ (O
O '<i;yDe>' " xioxust^ q _ xjom^
>Ni"^NviNOSHims saiavaan libraries^smithsonian'^institution NoiiniiiSNi nvinoshiiws S3iav
r-,2 r- 2 r- 2 f-.. 2
° m /<ri£5oX 2 m 2 m ^ 2
XI >
to ■ x to £ to £ to
ES SMITHSONIAN INSTITUTION NOIinillSNI NVINOSHIIWS S3iava9l1 LIBRARIES SMITHSONIAN INSTITl
— 2» CO 2 yi ^ ^ ^ 2:
2
to
> ' S > ’ 2 ■ ' N^' > ^<vA5^ ^
to ■•*■ 2 to Z to '^' Z t/>
3NI_NVIN0SHimS S3 I a Va 8 n_LI B R AR I ES SMITHSONIAN_INSTITUTION NOIiniIlSNI_ NVINOSHIIWS S3iaV
o '’3^ “ 5 ^ ' 5 5
^ — J ^ !2^ ^ ^ "J
ES SMITHSONIAN INSTITUTION NOIinillSNI NVINOSHilWS S3iavaai1 LIBRARIES SMITHSONIAN INSTITl
2 r* ^ ^ 2 1-^1- 2 r-
° m o z o 2 . Z
m xt2LD«^ vx p, >i8t to m x^yo^ ^ m
CO £ CO \ ^ in — in
SNI NVINOSHlIlAiS SBiavaail LIBRARIES SMITHSONIAN INSTITUTION NOIinillSNI NVINOSHIIIMS S3iaV
^ 2 Vi- to 2 ^ CO
_ ..VVVVVV «r ^ ^
/ ' 1 3 , ■ S 3 S 3 A'^^y § ■ ■
to
2
1
I
i
I
)
li
MASON E. HALE, JR.
•••
SMITHSONIAN CONTRIBUTIONS TO BOTANY - NUMBER 16
■U '.' ■■
SERIAL PUBLICATIONS OF THE SMITHSONIAN INSTITUTION
The emphasis upon publications as a means of diffusing knowledge was expressed by the first Secretary of the Smithsonian Institution. In his formal plan for the Insti- tution, Joseph Henry articulated a program that included the following statement: “It is proposed to publish a series of reports, giving an account of the new discoveries in science, and of the changes made frorh year to year in all branches of knowledge.” This keynote of basic research has been adhered to over the years in the issuance of thousands of titles in serial publications under the Smithsonian imprint, com- mencing with Smithsonian Contributions to Knowledge in 1848 and continuing with the following active series:
Smithsonian Annals of Flight Smithsonian Contributions to Anthropology S?nithsonian Contributions to Astrophysics Smithsonian Contributions to Botany Smithsonian Contributions to the Earth Sciences Smithsonian Contributions to Paleobiology Smithsonian Contributions to Zoology Smithsonian SUidies in History and Technology
In these series, the Institution publishes original articles and monographs dealing with the research and collections of its several museums and offices and of professional colleagues at other institutions of learning. These papers report newly acquired facts, synoptic interpretations of data, or original theory in specialized fields. These pub- lications are distributed by mailing lists to libraries, laboratories, and other interested institutions and specialists throughout the w’orld. Individual copies may be obtained from the Smithsonian Institution Press as long as stocks are available.
S. Dillon Ripley Secretary
Smithsonian Institution
SMITHSONIAN CONTRIBUTIONS TO BOTANY • NUMBER 16
Morden-Smithsonian Expedition to Dominica:
The Lichens (Thelotremataceae)
Mason E. Hale^ Jr.
KJ84
SMITHSONIAN INSTITUTION PRESS
City of Washington 1974
ABSTRACT
Hale, Mason E., Jr. Morden-Smithsonian Expedition to Dominica: The Lichens (Thelotremataceae). Stnithsonian Contributions to Botany, number IG, 46 pages, 20 figures, 1974. — A revision is made ot the lichen lamily Thelotremataceae in Dominica, based on previously published records by Elliott and on collec- tions by the atithor. The lamily comprises 48 species in 4 genera, Leptotrerna, Ocelhilaria, Phaeotrerna, and Thelotrernn. The following 15 species are de- scribed as new: Ocellularia antillensis, O. conglomerata, O. dotninicana, O. nia- culata, O. rnordenii, O. nigropunctn , O. rimosa, O. sorediata, Phaeotrerna aggre- gatiirn, P. obscurum, Thelotrenia confusttm, T. dorninicanurn, T. papillosum, T. tenue, and Leptotrerna decepturn. Three new combinations, Ocellularia fe- cunda (Vainio) Hale, Phaeotrerna disciforrne (Leighton) Hale, and Leptotrerna occult urn (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 I'I^blication datf, is handstaniped in a limited number of initial copies and is recorded in the Institution’s annual report, Smilhsotuan Year. SI Press number 5043. Series cover design: Leaf clearing from the katsura tree Cercidiphyllurn japonicurn 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) llibliography: p.
1. rhelotreniataceac. 2. Lichens — Dominica. I. Title. 11. Series: Smithsonian Institution.
Smithsonian contributions to botany, no. 16.
QK1.S2747 no. 16 [QK585.T5] 581'.()8s [589'.1] 73-22465
For sale by tlie Superintendent of Documents, U.S. (iovernment Printing Office
Washington, D.C. 20402 » Price (paper cover)
Contents
Page
Introduction 1
Morphological Characters . 2
Thallus Structure 2
Apothedal Characters 2
Spores 8
Generic Limits 11
Chemistry 11
Ecology and Habitats . 13
The Thelotreme Flora 13
Collecting Localities 14
Ocellularia 15
Key to the Species of Ocellularia . 15
Phaeotrema 28
Key to the Species of Phaeotrema 28
Thelotrema . 31
Key to the Species of Thelotrema 31
Leptotrema . • • 38
Key to the Species of Leptotrema 38
Literature Cited 43
Index • • • 46
iii
Morden-Smithsonian 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 of fact, these inconspicuous lichens will often 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, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.
easily accessible by car and well-kept trails, these forests have as yet sulfered 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 1971, 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 of 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
SMITHSONIAN 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. M. Skytte Chris- tiansen (C), Mr. Peter James (BM), Mme Jovet- Ast (P), Dr. E AE Lamb (EH), Dr. R. A. Maas Geesterantis (L), Prol. H. Merxmiiller (M), Dr. Haraltl Riedl (VV), and Dr. R. Santesson (UPS). I wish also to thank Dr. Chicita Culberson for help in identiiying or verifying most of the chemi- cal substances of the various specimens.
Morphological Characters
Alter curating and labeling each specimen, I studied the various morphological characters that appear to be most important for taxonomic jaur- 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 medidlary hyphae usually pene- trate the outer layers of bark periderm (or rock substratum). The algal layer (TrentepoJiUa) may also be indistinct. Large colorless crystalline inclu- sions are commonly found in the medulla, as in Ocelhilaria nigropiincla, new species, and Thelo- tretna inter posihim.
Sections examined under a scanning-electron microscope show some of these structures vividly. Ocelhilaria oUvacea, O. perforata, and Thelotrema confusum, new species (Figure \a,b,c), all w’ith well-tleveloped epiphloeodal thalli, have a cortex consisting ot 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 pecidiar aculeate hyphae protrude from the surface in greater or lesser abundance (Figure Sa,b). They appear to be collapsed, perhaps be- catise of vacuum treatment needed for specimen preparation. This acideate orientation, reported
here for the first time, is characteristic of Myrio- trema species {Ocellularia terebratula, Thelo- trenia clandestiniim) as well as a few species with emergent apothecia (T. depressurn, T. praestans). I have observed similar orientation in several species of Graphis and Graphina. Most thelotremes, how'ever, have an irregularly organized surface, as in O. 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 bd). The cortex is arranged in tall columns with algae scattered vertically (Salisbury, 1971) (Figtire 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 Ocelhilaria con- color, O. pyreniiloides, Phaeotrema disciforme, new combination, Thelotrema leiicomelaenum, 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 followdng Do- minican species have more or less similar struc- ture: Ocelhilaria conglomerata, new species, O. doininicana, new' species, O. exanthismocarpa (Fig- ure 5«), Leptotrema occultiim, L. siibcompunctum, and Phaeotrema aggregatiim, new species. I will call this arrangement “lepadinioid” in the descrip- tions; Salisimry (1972a) uses the term “Thelotrema lepadiniim group.”
What appears to be an extreme stage of this configuration of apothecial w'all 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 cpiphloeodal lichens viewed with a scanning-electron microscope [a, b, and c, sectioned obliquely; d, vertically): a, Ocellidaria oUvacea {Hale 35226) (X 1000); b, O. perforata {Hale 38145) (X 1000); c, Thelotrema confusum, new species {Hale 37697) (X 500); d, Leptotrema ivightii {Hioratn 5850 from Cuba) (X 500).
second inner rim. The disc is widely exposed and nothing resembling the discrete pore characteristic of the other thelotremes remains (Figure 5b). This type was first recognized among the foliicoloiis lichens, but it seems identical to that in the non-
foliicolous species of the Thelotremataceae, in- cluding in Dominica Ocellidaria alborosella and O. dilatata. It is called the “Thelotrema platy- carpum group” by Salisbury (1972b). In the ab- sence of more detailed ontogenetic research.
4
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Fici're 2. — Surfaces of epiphlocodal lichens viewed with a scanning-electron microscope: a, b, Ocellutaria nigropuncla, new species (Hale 353G5) (X 500 and X 2000); c, d, O. dominicana, new species (Hale 37967) (X 500 and X 2000).
however, we cannot yet say that CJiroodiscus is a good genns, that the CJiroodiscus-likc nonfoliico- lons thelotrenies shoidd be transferred to CJnoodis- cus, or that Cliroodiscus should be transferred to Occlhdaria. Cihroodiscoid species, incidentally, arc distributed among the four presently recognized
genera, Occdlularia, Leplolretna, PliacolYcina, and Tdclolrcina, 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 oUvacea {Hale 38054) (X 500 and X 2000); c, d, O. mordenii, new species {Hale 37764) ( X 500 and X 2000).
leaving only a small constricted pore. The upper wall is usually carbonized. There are, broadly speaking, three groups that can be recognized for practical purposes on the basis of degree of emer- gence from the substratum. One, represented by the "Myrioirema” type, has immersed noncarbon-
ized apothecia and a pore flush with the thallus surface (Figure Bc,d). The apothecia are always quite small, 0. 1-0.3 mm in diameter. Typical examples in Dominica are Ocellularia olivacea and Thelotrema clandestinum.
A second group, which includes the majority
6
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 4. — Surface and cross-scctions of hypophloeodal lichens viewed with a scanning-electron microscope: a, Thelotrema leiicomelaenxim, surface view {Hale 35491) (X 2000); b, T. leuco-
melaenum, cross-section {Hale 35491) (X 500); c and X 2000).
of the .species, has emergent apothecia, neither basally constricted or strongly raised (Figure 5(?,f). They are usually 0. 4-1.0 mm in diameter, although in a lew species (for example, Thelotrema praestans in Figure 17c) nearly 2 mm in diameter.
d, T. tenue, new species {Hale 35430) (X 500
A third group, typified by Ocellularia rho- dostroma and O. cavata, has strongly emergent apo- theda with a partially constricted base (Figure 5g). I'he pore is small and sometimes annulate or open at maturity, as in O. feennda (Figure ll/2,f).
NUMBER 16
7
Figure 5. — Types of apothec.ia in the Thelotremataceae (all X 10): a, Ocellularia exanthis- mocarpa (type of O. multilocularis)-, b, O. alborosella {Hale 35371a); c, O. oUvacea {Hale 35243); d, Leptotrema luightii, (type of L. prevostianum); e, O. perforata {Hale 37721); f, L. bahiatium (type of Thelotrema rudhis var. dominicanum); g, O. rhodostroma {Hale 38122); h, PImeotrema leiostomum {Hale 37956); i, O. comparabilis {Hale 37372).
Diameter varies between 0.6 and 2.0 mm. This group was called Ascidvum by Fee (1824), not so much because of emergence but because of a dupli- cate membrane in the peritliecium (the nature of which is not clear to me) and a marginate pore.
Although Nylander continued to use Ascidinm for strongly emergent species, no modern licheii- ologists have employed it. The type of the genus, A. chtchonnriifn Yee, is synonymous with Ocellu- laria cavata. Naturally there are intergradations
8
SMITHSONIAN CONTRIBUTIONS TO BOTANY
between the AscicUiun-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 5//). Other species from South America have this trait developed much more conspicuously and have been described in the Graphidaceae (as PJiaeo- graphis chiouoclisca Redinger (1936:69)). They shoidd 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, Ocellularia rJwdoslronia and PJiaeolremn disciforme, new combination). The other group, inchiding most nonchroodiscoid species in Do- minica, has ascogonial initials deeply emlredded in the periderm. The maturing apothecia must break throtigh this layer to reach the stirface. Several layers of periderm cells are thereby included in the apothecial receptacle and can be easily recog- nized in sections (Figure 6/). This character de- serves much more detailed study.
Development of a central cylindrical columella is jxarticularly 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 Ocellularia papillaia and O. perforata, each of which has a large popu- lation where a columella is clearly developed and a smaller one where it is either alisent 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 mattire, the columella pushes up against the upper carbonized
w^dls and periderm, acting as a kind of ramrod. Not all species tvith periderm inclusions have a columella, but as Redinger found for the Brazilian species the correlation is high.
The range in development of a columella is very great (Figure 6), from the weak thin columella in Ocellularia mordenii, new species, to the multiple actinoid structures of Ocellularia comparabilis (Figure 5?) and Leptotreina fissum (Figure 19fl,c). The most highly developed species is Ocellularia glaziovii (Aliiller-Argau known from South America Vainio (1921:184) recognized these species under a separate genus, Rhahdodiscus.
Spores
Spore size is an important and relatively con- stant character. The range of spore length, for example, is usually quite small except for very large spores (more than lOOp) which may vary between lOOu and 300p. We can distinguish four very broad categories among the Dominican species: (1) spores less than 20|.i long with 4-6 transverse locules, e.g., Lcplolreuia ivightii, Ocellularia tere- bratula, Thclolrcuia clandesiinum; (2) spores 20u-36i.i long -with 6-10 transverse loctdes, e.g., Ocellularia perforata (Figtire 7a), Thelotrerna leucomelaeuuin; (3) spores 40li-90|.i long with 12-24 transverse locules, e.g., Leptotrejna decep- tum, new species (Figure Id), Ocellularia exanthis- rnocarpa; and (4) spores SOp-SOOp long with numerous locules, e.g., Ocellularia rhodostroma (Figure lb), Thelotrerna praestans, T. tuberculi- ferum.
The number of longitudinal septations in Leptotrema and Thelotrerna 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 Thelotrerna where, on the one hand, a Thelotrerna species might have some senile spores turning brown and on the other a Leptotrema will have some colorless spores (Fig- ure Id). In Phaeotrema clisciforme, new' combina- tion, in fact, one often finds a mixture of color- less, mature brown, and shriveled brown spores (Figure 7c). Shriveling (Figures 7c//) is character- istic of brown-spored species; I know' of no cases ■where Thelotrerna or Ocellularia spores shrivel.
NUMBER 16
9
Figure 6. — Cross-section of apothccia (all about X 130); a, Ocellularia nigropuncta, new species {Hale 35363); b, O. mordenii, new sjjccies {Hale 37764); c, O. cavata {Hale 35134); d, Phaeotrema leioslomum {Hale 35203); c, Thelotrerna leucomelaenum {Hale 35491); /, T. tenue, new species {Hale 35430).
10
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 7 Spores of the Thclotreniataceae: a, Ocellularia perforata {Hale 37784) (X 400);
b, O. rhodostroma {Hale 35440) (X 130); c, Phaeolrema disciforme, new combination, with a mixture of colorless spores, brown spores, and shriveled brown spores {Hale 37939) (X 400); d, Leplotrema decepluin, new species, with colorless muriform spores on the right {Hale 38110) (X 400).
In any event, transitional sjtecies should be jtidged individtially, and the presenre of a few anoniolotis brown-s[K)ied specimens among many colorless s})ecimens (as in Ocellularia feennda, new combi-
nation, or O. suhcavala on Dominica) should not be totiilly unexpected; nor shoukl these isolated raises prejudice the valtie of spore color as a generic character.
•'«T.
NUMBER 16
11
Generic Limits
The generic delimitation of the Tlielotrenia- taceae has undergone many changes since the classic work of Acliarius. Ffe (1824), for example, established two genera, in addition to recognizing Tlielotrema, based on external features and apo- thecial characters: Ascidium and Myriotrema, both presently considered to be synonyms of Ocellii- laria. Mtiller-Argaii (1887a) proposed the most far- reaching changes, using spore septatioii and coloration to separate four genera (in addition to foliicoloiis Chroodisciis)-. Ocelhdaria (trans- versely septate colorless spores), Phaeotrema (transversely septate brown spores), Thelotrema (niiiriform colorless spores), and Leptotrema (mu- riform brown spores).
While Vaiiiio, one of the truly creative thinkers in lichen classification, rejected many “spore genera,” almost all authors since Muller-Argau have accepted these genera (and, indeed, spore- based genera in other families). Poelt (1973:613) refers to these as “artificial genera,” yet almost any character we select is in a sense artificial. I don’t believe this term does justice in that it implies spore characters are artificial.
We are, in reality, faced with at least three choices for generic delimitation in the family: (1) one genus (Thelotrema) to accommodate all the- lotremataceous species; (2) four spore-based genera {Ocellularia, Phaeotrema, Thelotrema, and Lep- totrema)-, (3) five or more genera (^Ascidium, Chroodisciis, Myriotrema, Rhahdodiscus, and Thelo- trema) based on apothecial emergence and excipu- lar structures. Most lichenologists, as I said, have opted for spore-based genera. While a majority opinion does not necessarily prove the naturalness or validity of such genera, they do represent 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 (1972a) recent rejection of spore-based genera in favor of a single massive genus Thelotrema with 400 species sub- divided into “groups” according to excipular struc- ture, a character that is difficult to identify. His keys and descriptions show how cumbersome this approach is, for in each instance one must desig-
nate the spore type, e.g., Thelotrema wightii with Leptolrema-type spores, etc. If one is forced to make this critical distinction every time a species is identified and described, then they should be treated as distinct genera, a more coherent and certainly a more pragmatic, working solution. Spore characters are well proven, even critical, in the taxonomy of nonlichenized fungi. A retrogres- sive generic concept in lichens wdll hardly encour- age a better understanding of their biology and evolution. My experience in the Thelotremataceae suggests tiiat spore-based genera are reasonably natural, easily manageable units that are probably diverging in terms of habitat rec|uirements and biogenetic characters as reflected in chemical dif- ferences.
Chemistry
As ill almost every other group of lichens studied, chemistry is an important and fundamental char- acter in understanding the evolution and popula- tion biology of tile Thelotremataceae. The family has an unexpectedly rich chemistry w'ith a pre- dominance of P+ depsidones and related com- pounds (Culberson and Culberson, 1968; Culber- son and Hale, 1973). I will discuss tlie possible significance of this chemistry below^ under the species and hope to explore the broader aspects of biochemical evolution of the family in another paper.
All of tlie specimens were initially tested with p-phenylenediamine and chromatographed on Brinkman thin-layer plates according to the P re- action. I would recommend, how’ever, that lichen- ologists consider using this reagent as little as possible because of the well-knowm carcinogenic projjerties of volatile amines. Identification of the lichen substances can be made just as easily with TLC tvithout recourse 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, v/v) were used to identify the chemical content of each specimen.
Hypoprotocetraric Acid and 4-0-Demethyl- NOTATic Acid. — Ocellularia feciinda, new combina- tion, (hypoprotocetraric acid predominating), O. rhodostroma (4-0-demethylnotatic acid predomi- nating), and Phaeotrema leiostomnm (4-0-deme-
12
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Uiylnotatic acid sometimes lacking). These acids were recently examined in detail by Cidberson (Culberson and Hale, 1973), who discnsses the biogenesis. Ocelluiaria jcciinda and O. rJiodostrojna were not included in that study.
XoRSTicTic Acid. — Lepiotreina uccultum. This is a rare acid in the Thelotremataceae (especially in contrast to the Graphidaceae) and is largely con- fined to Leptotreina on the world level.
Protocetraric Aciu. — Ocelluiaria nigropuncta, new species, O. perforala^ Leplotrema bahianiim, and Thelolretna coufiisum. This well-known P + red acid is easily identified on TLC plates. It is sometimes accompanied by a higher gray spot that 1 have termed the “amplior” unknown because ol its constant occurrence in Ocelluiaria amplior (Ny- lander) Redinger. Culberson believes it to be pro- tocetraric acid combined with a carboxylic acid, comparable to fumarprotocetraric acid or the re- cently described succinprotocetraric acid.
PsoROMic Aciu. — Ocelluiaria antillensis, new species, O. coinparabilis, O. subcavata, O. lerebra- tula, Leptotreuia fissum, Thelotreina carasseuse, T. claudestiuinii, T. conjorme, T. leucinum , and T. iuierposiluiu. This is the commonest P+ acid in the Thelotremataceae. It is almost ahvays accom- panied by an unknown compound “conpsoromic acid,” appearing as a spot below psoromic acid. It, too, is P+ yellow\
Stictic and CoNSTicTic Acius. — Ocelluiaria con- glouieraia, netv species, O. dilatala, O. exanthismo- carpa, O. pyrenuloides, Phaeotrema obscurum, new species, Tlieloireina glaucopallens, Leplotrema de- ceptum, new species, L. microglaenoides, and L. subcompunctum. This widespread and well-known acid is actually rather rare in the Thelotremataceae except for the genus Leplotrema. Constictic acid, appearing as a H2SO4+ yellotv spot just belorv salazinic acid, is a constant accessory compound. One or trvo other gray spots, as yet unidentified, may occur between stictic and constictic acids on TLC plates. Another rarer accessory is the highest H2SO4+ orange-red spot in the “quintaria” series (Hale, 1971b), still unidentified but being discov- ered in a variety of genera.
ViREXsic Acid. — Leplotrema spondaicum. This acid is P+ 'red. It rvas previously unknown in the Thelotremataceae, having been first identified in Alecloria.
‘‘Cixchoxarum” Unknowns. — Ocelluiaria cavata
and Theloirema dominicanum, new species. Sev- eral species of Ocelluiaria and this single Thelotreina react P+ red, but neither protocetraric acid or fumarprotocetraric acid can be demon- strated with TLC. Instead there are at least two gray spots above iioth of these acids on the plates (and often a high H2SO4 yellow spot). These are apparently derivatives of protocetraric acid.
“Praestans” U.nknow'n. — Theloirema praeslans. This unidentified depsidone reacts P+ red orange and forms a very distinct orange to reddish brown spot lielow psoromic acid in both solvents.
“Olivacea” Unknow.ns. — Ocelluiaria olivacea. This P— compound produces a distinct H2SO4 gray spot below lecanoric acid in both solvents. It may be a depside. ^Virile the lectotype of Ocel- lularia 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— Unknowns. — Ocelluiaria rimosa, new species, and O. sorediala, 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 were seen but microchemical tests with G.E. and C.A."Ab were unsuccessful. Another major group of P— compounds is terpenes. All TLC plates showed numerous terpenes, identifiable as purple, blue, or brown spots falling generally above .5 anti fatling ivithin a few’ hours. Probably all of these originate from the bark and are not lichen products, as can be proved by co-chromatographing the lichens tvith lichen-free bark samples from the same specimen.
No Substances Present.- — Ocelluiaria alborosella, O. coucoloty O. dominicana, new’ species, O. macu- lala, new' species, O. mordenii, new species, O. papillata, Phaeotrema aggregatum , new species, P. disci forme, new’ combination, Theloirema depres- sum, T. leucomelaenum, T. papillosum, new’ species, T. tenue, new species, and T. luberculi- ferum.
Pigments. — Ocelluiaria mordenii, new species and Leplotrema wightii (anthraqiiinones) and O. jecunda, new’ combination, O. rhodostroma, and Phaeotrema leiostomum (K— yellow compounds).
NUMBER 16
13
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, O. papillata, O. perforata, Thelotrema glaucopallens, T. tenue, new species, and Lepto- trema deceptiim, new species. At the opposite ex- treme we find canopy species that occur high above the forest floor in tree tops, exj^osed to great changes in insolation and wetting-drying cycles. These include Ocellularia inorcleJiii, new species, O. olivacea, O. subcavata, O. terebratula, Phaeo- trema leiostomiini, Thelotrema clanclestinum, 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 leucomelaeniim and T. tuberculiferum, 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
zone where rain forest is optimally cleveloped, tlic trees being 30-10 m high (Figures 2 and 3 in Flale, 1971a, 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- stroyetl, 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, Lepiolrema spondaicum and Ocellularia exanthismocarpa. They have a highly successful means of propaga- tion and become established very cjuickly.
The Thelotreme Flora
As I had found with Parnielia- (Flale, 1971a), almost all previous collections of the Thelotrema- taceae on Dominica were made by W. 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:
Thelotrema excavalum Vainio var. impressulurn Vainio (Morne Anglais, the type-locality, Elliott 160, specimens in BM and TUR). This is synony- mous with Ocellularia perforata.
Thelotrema leucornelaenum var. elevatum VTiwio (Morne Anglais, Elliott 1534, specimen in TUR). This is T. leucornelaenum.
Thelotrema rhodostromum (Montague) Vainio (Morne Anglais, Elliott 521, specimens in BM and TUR, and Prince Rupert, Elliott 1305, specimen in TUR). These are Ocellularia rhodostroma.
Thelotrema rudius Vainio var. dominicamim Vainio (Prince Riq^ert, Elliott 1303, specimen in TUR). This is the same as Leptotrema bahianum.
The thelotreme flora of Dominica now includes ■18 positively identifiable species (and three addi- tional species represented by poor material). I col- lected all of these except iox Leptotrema bahianum. No one locality, of course, had anywhere near this ntimber of species (see p. I f, “Uollecting Locali- ties”). The richest area was one site (16b) in the Dleau Gommier Forest Reserve which was being logged in December 1971; 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 dillerent 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 ni, had 9 species each.
The homogeneity of the flora is remarkably higb. 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 w'e have not yet in- ventoried the whole thelotreme flora of Dominica. 1 would estimate that there are at least 55 species (including the three unidentified collections as possible additions). It would recpiire at least one more intensive collecting trip, possibly two, to reacb 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-
vation 60 m.
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
30 m.
7. Virgin rain forest logging area, Newfound-
land, elevation 250 m.
8. Rain forest above Newfoundland, elevation
300 m.
9. Rain forest logging area, Castle Bruce Road,
elevation 300 m.
10. Dleaii Gommier rain forest near Emerald
Pool, elevation 450 m.
11. Felled rain forest along new road cut about
2 km northwest of Pont Casse, elevation 430 m.
12. Rain forest logging area on Layou Road, ele-
vation 420 ni.
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 Eor-
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 I8b. Remnants of rain forest along road to Jean Estate, elevation 600-700 m.
19. Citrus grove at Laudat, elevation 400 m.
Figure 8 Outline map of Dominica showing localities where
specimens of Thelotremataceae were collected.
NUMBER 16
15
20a and 20b. Remnants of rain forest in pastures above Giraiidel, 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.
25a and 25b. Rain forest-mossy forest along trails at Fresh Water Lake, elevation 800-900 m. 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).
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. 1 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 tlie 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 (77%). 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 (with number of specimens collected in parentheses) are; O. rhodostroma (93), O. fecunda, new combination (64), O. perforata (55), O. papil- lata (48), O. suhcavata (40), O. exantfiismocarpa (25), O. riigropiincla, new species (16), O. oUvacea (12), O. ?)}ordeiui, new species (10), and O. tere- bratula (10). Rarities, known from single collec- tions each, are O. antillensis and O. cavata.
Key to the Species of Ocellularia
1. Medulla of thalliis and/or ascocarps pigmented.
2. Medulla and ascocarps entirely deep red 12. O. mordenii, new species
2. Medulla of ascocarps yellow or pink but thallus medulla white.
3. Pigment yellow; pore large, 0.3 mm wide 10. O. fecunda, new combination
3. Pigment deep pink; pore tiny, 0.1 mm %vide, annulate 18. O. rhodostroma
1. Medulla of thallus and ascocarps white.
4. Apotheda lacking a central columella.
5. Apothecia immersed, 0.2-0. 3 mm in diameter.
6. Thallus P+ yellow 22. O. terebratula
6. Thallus P— 14. O. olivacea
5. Apothecia emergent, 0. 6-2.0 mm in diameter.
7. Disc open with a strongly recurved exciple.
8. Exciple thick, disc irregularly elongate; stictic acid present 7. O. dilatata
8. Exciple thin and delicate, disc round; stictic acid absent 1.0. alborosella
7. Disc closed, lacking an outer recurved exciple.
9. Inner lepadinoid exciple strongly developed.
10. Apothecia aggregated in groups.
11. Apothecial groups round; stictic acid lacking
8. O. dominicana, new species
16
SMITHSONIAN CONTRIBUTIONS TO BOTANY
11. Apothecial groups oblong; stictic acid present
6. O. conglomerata, new species
10. Apothecia solitary.
12. .Spores large, 55p-75p 9. O. exanthismocarpa
12. Spores small, 15|,i 6. O. conglomerata, new species
9. Inner exciple not developed.
13. Thallus P — .
14. Thallus distinctly fissured; pore area black rimmed
19. O. rimosa, new species
14. Thallus continuous; pores not black rimmed 15. O. papillata
13. Thallus P+ red.
15. Pore area black rimmed 13. O. nigropiincta, new species
15. Pore area not black limmed 16. O. perforata
4. .Apothecia with a well-developed black central columella.
16. Thallus with large scattered verrucose outgrowths becoming sorediate
20. O. sorediata, new species
16. Thallus lacking soredia.
17. .Apothecia large, 0.5-1. 1 mm in diameter, more or less emergent.
18. Ihallus dull tan, hypophloeodal, P— 5. O. concolor
18. 'Thallus distinctly epiphloeodal, P + .
19. Columella becoming actinoid; thalhis P+ yellow 1. O. comparabilis 19. Columella remaining simple; thallus P+ red 3. O. 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 Op-llji
21. O. subcavata
20. Pore not annulate; spores generally 20p-30|.i long (except smaller in O. maculala).
21. Thallus P-.
22. Pore tiny, less than 0.1 mm in diameter; spores 4-loculate
11.0. maculata, new' species
22. Pore larger, 0. 1-0.2 mm wide; spores 0-7-locidate 15. O. papillata 21. Thallus P+ red or orange yellow.
23. Thallus very thin to hypophloeodal; P+ orange 17. O. pyremdoides
23. '1 hallus thick, epiphloeodal.
24. Thallus P+ red; apothecia barely emergent
16. O. perforata
24. Thallus P+ yellow; apothecia strongly emergent
2. O. antillensis, new species
1. Ocellularia alborosella
T'icurks 5ft, Oft
Ocellularia alborosella (Nylander) .Santesson, 1952:308. Graphis alborosella Nylander, 1863:372 [type-collection: Co- lombia, l.iudig 2694 (II, Icctotype; 1 11- Tuck, IT’.S, isotypes): Tigttre 9ft].
Thelotreiiia plalycai j>elluin A'ainio, 1923:138 [type-collection:
Arima, 'Trinidad, Tliaxler 57 ( IT’R, lectotype); Tigiire On]. Ocellularia plalycarpella (A'aino) Zahlbruckner, 1923:598.
Thallus whitish grtiy, thin but continuous, shiny, smooth, lonning colonies 5-8 cm broad; apothecia chroodiscoid, up to 1.5 mm across, round to irregu- lar, the marginal Ihtp conspicuous, the disc llesh colored to white pruinose; hymenium about 6()[i
high; spores 8, 5-6 locidate, 4p-5u X 8[.i-15|.i, I~ or weakly blue (Figure 5b).
(tie.mistrv. — No substances present.
Habtiat. — I’ree trunks at mid or higher eleva- tion in rain lorest.
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 from (Colombia but they fall within a pre- dictalile range of variation. It is a rare and easily overlooked species.
Specimens Examined. — 8 (38143), 16b (37883), 26b (35371a).
NUMBER 16
17
2. Ocellularia antillemis Hale, new species Figure 9g
Thallus corticola, albidiis, 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 VOp-SOp altum; sporae 8:nae, trans- versim 7 loculatae, 7|,i X 20|i, 1+ coerulescentes (Figure 9g).
Chemistry. — Psoromic and conpsoromic acids. Holotype. — 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 30[i-34j,i), sparse apothecia, and a smooth thallus, while O. terebrata (Acharius) Muller- Argau has barely emergent apothecia (see Hale, 1972). I later collected this species from Trinidad and Panama, both at low elevation.
Specimens Examined. — 3 miles NW of Sangre Grande, Trinidad, Hale 37444 (US); Barro Colo- rado Island, Panama, Hale 38659 (US).
3. Ocellularia cavata Figures 6c, 9/
Ocellularia cavata (Acharius) Miiller- Argau, 1882b;499. Thelotrema cdvatum Acharius, 1812:92 [type-collection: Sierra
Leone, Afzelius (H, lectotype; S, UPS, isotypes)] Hale,
1971b, fig. la.
Ascidium cinchonarurn Fee, 1824:96 [type-collection: On Cinchona, South America (G, lectotype; H, L, M, P, iso- types); Figure 9c].
Ocellularia cinchonarurn (Fee) Sprengel, 1827:242.
Ascidium cinchonarurn f. intermedium Nylancler, 1867:319 [type-collection: Pie de Cuesta, Colombia, Lindig 5 (H,
lectotype; BM, G, M, isotypes); Figure 9c].
Ocellularia lindigiana Miiller- Argau, 1887a:9 [type-collection: Colombia, Lindig 2757 (G, lectotype; BM, FH, UPS, iso- types); Figure 9d].
■Oeelhilaria cinchoymrum (Fee) Sprengel f. intermedia (Ny- landcr) 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-8[,i X 22[.i-28|.t, 1+ blue. (Figure 9/).
Chemistry.^ — “Cinchonarurn” 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 (Krcmpelhubcr) Muller-Argau, 1883:
318.
Thelotrema comparabile Krcmpelhubcr, 1876:220 [type-
collection: Rio de Janeiro, Brazil, Glaziou 5463 (M, lecto- type; BM, C, G, M, P, UPS, W, isotypes); Figure 9;].
Thallus pale greenish to whitish mineral gray, epiphloeodal, smooth, forming colonies 5-12 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 50[.i-60[.i high; spores 8, 4-5 loculate, 5(i-6[i X lOp-l lti, 1+ blue. (Eig- ure 5i).
Chemistry. — 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, Ocellularia discoidea (Acha- rius) Miiller-Argau (Hale, 1972) and O. atilillensis, new species, have a simple columella. Ocellularia e[forinaia (Krempelhuber) Miiller-Argau has a very similar actinoid columella but the outer margin is semi-erect, leaving an open rather lacerate pore; O. berkeleyaua (Montague) Zahlbruckner and O. glaziovii Miiller-Argau lioth have highly develojied
18
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Fk;uri, !). Specimens of 'I lieloliemataceae (all about X 10); a, Ocellularia platycarpella (lecto-
type in I'UR); b, O. alborosella, (isotype in FH-Tuck); c, O. cinclionarum f. intermedia (lecto- lype in FI); d, O. liudigiaua (lectotype in G); c, O. cinclionarum (leclotypc in G); f, O. cavata {Hale 35131); g, O. anlillcnsis, new species {Ualc 35(il2); h, O. conglomerala, new species (Hale 37959); i, O. comparabilis (isotype in IIM); ], O. concolor (lectotype in G); k, O. concolor (Hale 381f37).
NUMBER 16
19
actinoid discs; and O. latilahra (Tuckerman) Muller-Argau has an actinoid columella and a recurved margin.
Specimens Examined. — 9 (38128), 21 (37372,
38144).
5. Ocellularia concolor Figure 9j,k
Ocellularia concolor Meyen and Flotow, 1843:230.
Type-Collection. — Manila, Philippines, Meyen (G, lectotype) (Figure 9;).
Thallus light grayish tan, dull, mostly hypoph- loeodal, lorming 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 Hat black columnar disc easily seen; hymenium 140p high; spores 8, 5-7 loculate, 5p- 8p X ]3p-20p, 1+ blue (Figure 9k).
CiiEMLSTRY. — No substaiices present.
Habitat. — Lianas, saplings, base and lower trunk ol trees in dense rain forest (300-370 m).
This easily recognizable species was first de- scribed from the Philippines and appears to be pantropical. I'he tannish hypophloeodal thallus and habitat are characteristic. There are no com- parable species in this kind of habitat.
Specimens Examined. — 8 (38167), 16a (37665), 16b (37801, 37809, 38075).
6, Ocellularia conglomerata Hale, new species Figure 9h
Thallus corticola, ciiiereo-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; hymenium 70u altum; sporae 8:nae, tran.sversim 6-7 loculatae, 5p-6p X 15[.i-18[.i, 1+ coerulescentes (Figure 9li).
(aiEMLSTRY. — Stictic and constictic acids.
Hoi.otype. — Trail to Morne Anglais, Dominica, elevation about 900 m. Hale 37959, 13 December 1971 (US).
Chemistry places this species near O. exanihismo- carpa but the exciple is not so clearly separate from the thick, erect, pulverulent margin, leaving the disc open. The spores are much smaller. An- other West Indian stictic acid-containing species, Ocellularia alboolivacea (Vainio) Zahlbruckner, has a more typically recurved margin and a thin, entire exciple.
7. Ocellularia dilatata
Figure \\a,b
Ocellularia dilatata Muller-Argau, 1895:452.
Type-Collection. — Rio de Janeiro, Brazil, Glaziou 5531 pro parte (G, lectotype; BM, isotype). (Figure llfl).
Thallus whitish gray, thin, dull, in part hypoph- loeodal, forming colonies 4-6 cm broad; apothecia chioodiscoid, large, up to 2.5 mm broad, round to elongate, marginal llap rather coarse; disc white piuinose; hymenium about 60p high; spores 8, 6-7 loculate, 5p-6p X 15p-17[i, 1+ blue (Figure 115).
Chemistry. — Stictic and constictic acids.
Habitat. — Lower trunk area in higher rain forest zone (850 m).
This rare species is related to Ocellularia alboro- 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 Examined. — 26a (37695, 37702).
8. Ocellularia dominicana Hale, new species Figures 2c, d; 10
Thallus corticola, brunneo- vel viridi-albidus, epipidoeodes, continuus, nitidus, 5-10 cm latus; apotliecia semi-emergentia, 0.7-11 mm diametro, apice incolores, columella nulla, solitaria vel 2-3 arete aggregata, habitu similia ajiothecio singulari, excijnilo interiore evoluto, ostiolum complente; ostiolum rotundum, 0. 1-0.5 mm diametro; hy- menium OOu-SOp altum; sporae 8:nae, transversim 6-7 loculatae, 6p-8[i X 16ii-25[.i, 1+ coerulescentes (Figure 10).
Chemistry. — No substances present.
20
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 10. — Ocellularia dominicana, new species {Hale 37841) (about X 10).
Holotype.^ — Submossy forest on upper slopes of Trois Pitons, Dominica, elevation about 1000 m, Hale 37967, 6 December 1971 (US).
Habitat. — I’runk and lower base of trees in sub- mossy forest (700-1100 m).
I'lie apotheda are so tightly aggregated that I first interpreted the wall between them as a columella. The comliined outer margin and exciple are minutely pulverulent and crowded, fdling the broad disc but leaving a distinct pore area. The species was rather common at Trois Pitons but not found elsewhere.
Specimens Examined. — 23 (38053), 27 (37683, 37688, 37829, 37832, 378-11, 38041 ).
9. Ocellularia exanthismocarpa Figures 5a, llc,g
Ocellularia exantJiistnocarpa (Leighton) Zahlbiuckner, 1923: 590.
Thelolrema exanthismocarpuni Leighton, 1869:169 [type- collection: Central Province, Ceylon, Tlnvaites C.L. 97 (BM, Icctotype; H, isotype); Figure 11c].
Thelotrema porinoides Montague and von dcr Bosch, 1855: 484 [typc-collection: Java, Junglnilni 151 (L, Icctotype;
FIT, 0>, P, W, isotypes); Figure lie].
Ocellularia multilocularis Zahlhriickncr, 1912:369 [type- collection: Lanai, Hawaii, Roc^k 17 (\V. holotype); Figure 5a].
Ocellularia iserlii Vainio, 1915:40 [type-collection; Guade- loupe, Isert 87 (C, Icctotype; TUR, isolype); Figure 11/].
Thelolrema homothecium Vainio, 1921:190 [type-collection: Irosin, Mt. Bulusan, Prov. Sorsogon, Philippines, Elmer 14852 (TUR, Icctotype; F'H, G, L, S, US, W, isotypes); Fig- ure lid].
Ocellularia hornothecia (Vainio) Zahlbruckner, 1923:593. Ocellularia porinoides (Montague and von der Bosch) Zahl- bruckner, 1923:599 [not O. porinoides (Acharius) Sprengel, 1827:242].
Thalius white to ashy, thin, smooth, in part hypophloeodal, forming 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, 12|j,- 18p X 60j.i-100p, 1+ deep blue (Figure llg).
Chemistry. — Stictic and constictic acids and rarely the higher “quintaria” unknown B.
Habitat. — Trunks, canopy brandies, 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.
Specimens Examined. — 8 (38114), 11 (38009),
I6b (37982), 17 (35605), 20a (37780, 37781, 37786, 37788, 38019, 38025), 25a (37708, 37709, 37713, 37714, 37715, 37716, 37849, 37996, 38069), 25b (35473, 35516), 26a (37692, 37944), 27 (38042, 38055).
10. Ocellularia fecunda (Vainio) Hale, new combination
Figure ll/i,!
Thelotrema domingense var. jecundum Vainio, 1896:208 [type-collection: Richmond Valley, St. Vincent, EllioU 243 (BM, Icctotype; TUR, isotype); Figure ll/i].
Ocellularia domingensis var. fecunda (Vainio) Zahlbruckner, 1923:590.
Thalius light greenish or whitish mineral gray, epiphloeodal, smooth and shiny, forming large colonies up to 30 cm broad; apothecia large, strong- ly emergent, 1.5-1. 8 mm in diameter, the cortex soon breaking away and revealing a light yellow'- isli orange medulla, tvalls heavily carbonized,
NUMBER )6
21
I'KiURE II. — Specimens of Thclotrcmataceae (all about X 10): a, Ocelliilaria dilatala (isotype in BM); h, (). dila'iata {Hale 37702); c, O. exanlhisriiocarpa (Icctotypc in BM); d, O. homolheda (isotype in FH); e, O. pot inoides (Icctotypc in L); /, O. isertii (Icctotypc in Cl); O. exanthis- mocarpa {Hale 3799G); h, O. fccuiida, new combination (Icctotypc in BM); i, O. jecutida, new combination {Hale 3,7218); j, O. iiiaculata, new species {Hale 38015); k, O. mordenii, new species {Hale 37887).
22
SMITHSONIAN CONTRIBUTIONS TO BOTANY
columella absent or weakly and irregularly devel- oped; pore gaping, round to irregular and jagged, 0.1-0. 8 nun in diameter, the white pruinose disc clearly visible; hymenium 200ii-250(_i high; spores 1— 1/ascus, 30—10 loculate, 12-20 X 80[.i-150p, rarely turning brown, 1+ blue (Figure llz).
Chemistry. — Flypoprotocetraric acid and in lesser concentration 4-0-demethylnotatic acid along with unidentified pigments.
Habitat. — Basal area of trees into the upper bole and canopy in the rain forest (300-850 m).
I'his is one of the fetv 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 off. It is the second commonest species in Dominica and forms very extensive colonies. As far as 1 have determined, the species is restricted to the West Indies and has no close relatives.
Specimens Examined. — 8 (38120), 11 (37811,
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, 3803’9), 21 (38080, 38081, 38089, 38103, 38106, 38107, 38132, 38137), 22 (37639, 37614, 37828, 37838, 37843, 37844,
37844a, 37815, 37856, 37858, 37960, 38026), 23 (37679), 24a (37740, 37743, 37063, 37064), 25a (37711), 26a (38056).
11. OceUularia maculata Hale, new species
FifaiRE II;
riiallus corticola, cinereo-albidus, epiphloeodes, planus, coiuinuus, nitidus, 9 cm latus; apothecia numerosa, niinuta, 0.2-0. 3 mm diametro, apice obfiisca, cuhmiellata vel columella |)arce evoluta; ostiolum rotundum, 0.05-0.1 mm diametro, anguste nigroc iiu turn, columella protrusa; hymenium lOOii ahum; sporae 8:nae, transversim 4 loculatae, 5p-7p X llp-13[i, 1- (Figure 11;).
Cm .MisiRi . — No substances present.
Iloi.oiT PK. — Remnants of rain forest in pastures aliove Giraudcl, Dominica, elevation 650 m. Hale 38015, 11 December 1971 (US).
()( cUalaria maculala might seem at first glance to be O. papillala but the apothecia and spores
are much smaller. There are no related species.
Specimen Examined. — 18b (35524).
12. OceUularia mordenii Hale, new species Figures 3c,d; 6b; life
Thallus corticola, pallide cinereo- vel brunneo- albidus, epiphloeodes, continuus, aetate rimostis, nitidus, usque ad 12 cm latus; medulla omnino sanguinea; apothecia dispersa, vix emergentia, apice obfusca, extus crasse verrucosa, verrticis fragil- ibiis, rumpentibtis, columella nulla vel parce evo- luta (Figure 65); ostiolum rotundum, 0.08-0.12 mm diametro; hymenium circa 200|.i altum; sporae 1-2: nae, transversim 24-30 loculatae, 15p-20[.i X 70p-150[.i, 1+ coerulescentes (Figure ll/t).
Chemistry. — Unidentified anthraquinone pig- ments only.
FIoloi'ype. — Canopy branches in primary rain forest, Dleau Gommier Forest Reserve, Dominica, elevation about 370 m. Hale 37764, 12 December 1971 (US).
FIaiutat. — Canopy branches in rain forest (370- 800 m).
I he deep red pigment and large warts are dis- tinctive. It is a conspicuous canopy species. A very similar species in Cuba, O. xantlioslronm (Ny- lander) Zahlbrtickner, differs mainly in containing a Iv— pale yellow-orange pigment. This lichen is named in honor of Mrs. William J. Morden.
Specimens Examined. — 12 (37909), 15 (35198), 16a (38051), 16b (37791), 16c (37762,37886,37887, 37010), 26a (37863).
13. OceUularia nigropuncta Hale, new species
Figures 2a, b; 6a; 12a
Thallus corticola, pallide brtinneo- vel viridi- cinereus, epiphloeotles, continuus vel rarius rimo- stis, 5-8 cm latus; apothecia numerosa, emergentia, 0.6-0. 8 mm diametro, apice obfusca, columella nulla (Figure 6a); ostiolum rotundum, 0. 1-0.2 mm diametro, nigrocinctum; hymenium lOOu-llOp altum; sporae 8: nae, transversim 6-8 loculatae, 6ii- lOij, X 20[i-32p, 1+ coerulescentes (Figtire 12«).
Chemistry. — Protocetraric acid.
Holotvpe. — Trail to summit of Morne Anglais, Dominica, elevation about 900 m. Hale 35365, January 1969 (US).
NUMBER 16
23
Figure 12. — Specimens of Thelotremataceae (all about X 10): a, Ocelhdaria nigropuncta, new species {Hale 35365): b, O. rimosa, new species {Hale 38052); c, O. olwncea (lectotype in G); d, O. papillata (lectotype in BM); e, O. papillata (Hale 38098); f, O. perforata (lectotype in BM): g, O. excavata (lectotype in BM); h, O. rufocincta (lectotype in G); i, O. terebrata var. abbreviata (isotype in FH); j, O. pyrenuloides (isotype in W); k, O. pyrenuloides {Hale 35524).
24
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Habitat. — Lianas, lower bole and base, Syrn- pJionea roots, rarely canopy of trees in the rain forest (.800-900 m).
I'he black-rinnned pore area and smooth thallus are diagnostic features. Other ecoliimellate proto- cetraric acid-containing species include Ocellularia bonpJaudiae (Fee) Muller-Argau, which has a grainy surface, and O. verrucosa (Fee) Muller- Argau, ivhich has an irregular open pore and larger spores (over 30p long). Under the scanning- electron microscope O. nigropuncta, new species, has a finely pitted but otherwise smooth surface (Figure 2a,h), whereas O. verrucosa appears to have an aculeate orientation and O. bouplandiae is verrucose and fissured.
Specimens Examined. — 8 (.88115, 38158), 10 (38169), 12 (37903), 16a (37660), 16b (37798), 16c (37755, 37770, 37963), 18a (379-18), 21 (38092, 38091, 38138), 22 (37852), 24a (37751).
14. Ocellularia olivacea
Figures \a; Sa,b; 5c; 12c
Ocellularia olivacea (Fee) Miillcr-Argaii, 1887a:7. Myriotrema olivacea Fee, 1824:103 [type-collection: On Bon-
plandia trifoliala, .South America (G, lectotype); Figure
12c].
Ocellularia olivacea (Fee) C. and D. Van Overecm-Dc Haas,
1922:118 [superfluous combination].
Thallus light greenish to ashy white, shiny, thick, up to 0.5 mm (Figure lu), forming exten- sive colonies uj) 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 65[i-75[i high; spores 8, more or less uniseriate, 4 loculate, 4[i- 6p X 8;j,-12[i, 1+ blue (Figure 5c).
Chemistry. — “Olivacea” unknown and a second lower sjiot.
FIabitat. — Canopy liranches to mid-bole in rain forest (240-700 m).
This is one of three “Myriolretna” species that occur in Dominica, the other two being O. tere- hraiula and Thelotrema clajideslinum, all typically found on canopy branches in dense rain forest. Ocellularia olivacea 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 tyjie-specimen of O. olivacea lacks this spot. The surface under the scanning-electron microscope is strongly aculeate (Figure %a,b).
Specimens Examined.— 7 (35226, 35243, 35249), 8 (38164), 9 (38154), 11 (38001), 12 (37965), 15 (35205), 16a (37992, 38054), 18a (37965), 21
(38113).
15. Ocellularia papillata
Figure 12d,e
Ocellularia papillata (Leighton) Zahlluuckner, 1923:597. Thelotrema papillatum Leighton, 1869: 1G9 [type-collection:
Cential Province, Ceylon, Tbwaites C.L. 129 (BM, lectotype;
H, G, P, .S, UPS, W, isotypes); Figure 12d].
Thallus tv’hitish to pale greenish ashy, smooth to more or less warty, shiny, forming colonies 3-12 cm broad; apothecia numerous, 0.6-0. 8 mm in diameter, immersed to slightly emergent, the upper walls carbonized, columella present, thin to scarcely develojied to absent; pore round, 0. 1-0.2 mm in diameter; hymenium 50[.i-70[i high; spores 8/ascus, 7-9 loculate, Op-lOp X 20[.i-36[.i, 1+ blue. (Fig- ure 12e).
Chemistry. — No substance present.
Habitat. — Sajilings, prop roots, lianas, and tree bases at mid-elevation in rain forest (300-900 m).
1 his common species has few outstanding fea- tures. The columella is usually present, although often iveakly developed and difficult to find. Some specimens, as indicated below, seem to lack it en- tirely. The ajiothecia are inconspicuous, almost immersed to slightly emergent, but the pore is always distinct. The thallus is well developed, usually cjuite whitish, and smooth. There are no close relatives ivithout lichen substances, although O. perforata, 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 rvas found.
Specimens Examined (columella present). — 8 (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, 38967), 24a (37746).
16. Ocellularia perforata Figures lb, 5e, la, 12/
Ocellularia perforata (Leighton) Miiller-Argau, 1892:284. Thelolrenia perforatum Leighton, 1866:447 [type-collection:
Casiquiari, Brazil, Spruce 254 (BM, lectotype); Figure 12/]. Thelotrema terebratum var. abbreviatulum Vainio, 1890:83 [type-collection: Cara^a, Minas Gerais, Brazil, Vamio 1551 (TUR, lectotype; BM, FH, UPS, isotypes); Figure I2i]. Ocellularia rufocincta Muller- Argau, 1893:146 [type-collection: Foret du Rio General, Tondoz s.n. (Pittier 6107) (G, lecto- type); Figure 12/i].
Thelotrema excavatum Vainio, 1896:208 [type-collection: Mt. St. Andrew, St. Vincent, Elliott 153 (TUR, lectotype; BM, FH, isotypes); Figure 12g].
Thelotrema excavatum var. impressulum Vainio, 1896:208 [type-collection: Morne Anglais, Dominica, Elliott 169 (BM, lectotype)].
Ocellularia terebrata var. abbreviata (Vainio) Zahlbruckner, 1923:602.
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 tipper wall carbonized, columella usually developed, thin 50[,i-70p in diameter, but sometimes lacking or difficult to find; pore round, 0.07-0.15 mm in diameter; hymenium 55[i-70[,t high; spores 8/ascus, 6-8 loculate, 7[.t-10[,i X 20p- 34j.i, 1+ blue (Figure he).
Chemistry. — 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 O. papillnta, 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, 37735, 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 12/,/£
Ocellularia pyrenuloides Zahlbruckner in Magnusson and Zahlbruckner, 1943:46.
Type-Collection. — Wailuku, Maui, Hawaii, Faiirie 676 (UPS, lectotype; S, W, isotypes) (Fig- ure 12/).
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, 130[i 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 60[.i high; spores 8/ascus, 6-7 loculate, 6[.i X 12[i-15(j., 1+ blue (Figure 12/t).
Chemistry. — Stictic and constictic acids.
FIabitat. — Tree branch in rain forest (650 m).
The darkened pore area is similar to that in Thelotrema lenue, new species. There are no re- lated sjiecies 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, 1b, 13a
Ocellularia rhodostroma (Montague) Zahlbruckner, 1923:600. Ascidium rhodostromum Montague, 1851:75 [type-collection: Gayenne, French Guiana, collection 1334 (P, lectotype): Figure 13a].
I’hallus light greenish tan to gray, smooth to finely warty, continuous and epiphloeodal, the medulla white or pink, forming colonies 5-10 cm
26
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Fir.URF, 13. — .Specimens of Thclotrcmataccac (all about X 10): a, Ocellularia rhodoslroma (lecto- typc in 1’); b, O. subcax'ata (lectotype in H); c, O. vaga (isotypc in F!I); d, O. subawata {Hale 37769); e, O. tcrebi alula (lectotype in H); f, O. remanens (isotypc in Cl); g, O. sorediata, new species (Hale 37908); h, Phaeotrema aggregatum, new species {Hale 35229); i, O. tcrebratula {Hale 37793); j, P. diseijorrne, new combination (lectotype in KM); h, P. aquilinuui (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 Iragile cortex breaks away, columella lacking; pore round and small, 0.05-0.1 mm in diameter, surrounded by a raised ring; hymenium 200p-275j.i high; spores 1-2/ascus, 25-30 loculate, 25p-40|.i X 100p-250j,i, 1+ blue (Figures 5g and 1b).
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.
Ocelliilaria rbodostroma 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.
Specimens Examined. — 7 (35246), 8 (38122, 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, 37650, 37835, 37861), 23 (37672, 38975), 24a (37736, 37742, 37745, 37749, 37754, 38068), 24b (35629), 25a (37705, 37707, 37720, 37840, 37846, 37848, 37862, 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).
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 110|.i-150p ahum; sporae 8:nae, transversim 6-8 loculatae, 7p-10|.i X 24p-32|i, 1+ coerulescentes (Figure I2b).
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 O. nigropuncta, new species. Both are base level species, but O. rimosa has a more restricted elevational range. The two species were collected together only at Dleau Gommier.
SinxiMENS Examined. — 11 (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, 0.08-0.11 mm diametro; hy- menium 60p-70p ahum; sporae 8:nae, transversim 3-4 loculatae, Op-Sp X 12p-16p, 1+ coerulescentes (Figure 13g).
Chemistry. — No substances present.
Holotype. — Logging area about 3 km northvv^est of Pont Casse on the Layou Road, Dominica, ele- vation 420 m. Hale 37908, 10 December 1971 (US).
Habitat. — Mid-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.
Spectmi':n Examined.— 16b (38029).
21. Ocellularia subcavata Figure \3b,d
Ocellularia subcavata (Nylander) Zahlbruckncr, 1923:601.
28
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Thelotrema siibcavalum Nylander, 1876:561 [type-collection: Cuba, Wright 509 (H, lectotype; FH-Tuck, isotype); Figure 136],
Thelotrema vagum Vainio, 1896:209 [type-collection: Rich- mond Peak, St. Vincent, EllioU 260 (BM, lectotype, F'H, isotype); Figure 13c].
Ocellularia vaga (Vainio) Zahlbruckner, 1923:603.
Thallus ashy whitish to greenish gray, shiny, more or less warty and cracked, forming colonies lip 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 hlled with the often protruding pruinose columella; hymenium about lOOp high; spores 8/ascus, usually uniseriate, 5-6 loculate with nar- row locules, 4j.i-7[i X 8(1-20(1, very rarely turning brown with age, 1+ blue (Figure ISd).
Chemistry. — Psoromic and conpsoromic acids. Habitat. — Canopy branches and upper bole in rain forest (240-800 m).
I’his 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.
Specimens Examined. — 7 (35248, 35252), 9
(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).
22. Ocellularia terebratula
Figure I3ej
Ocellularia terebratula (Nylander) Muller-Argau, 1887:12. Thelotrema terebratulum Nylander, 1867:315 [type-collection:
Rio Negro, Colombia, Lindig 129 (H, lectotype; FFI, G, M,
P, isotypes); Figure 13e].
Thelotrema clandestinum t. remanens Nylander, 1867:317
[type-collection: Monte del Morro, Colombia, Lindig 90
(H, lectotype; BM, G, P, isotypes); Figure 13/].
Ocellularia remanens (Nylander) Miiller-Argau, 1887:7.
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 65(i, high; spores 8/ascus, 3-5 loculate, 5p-9(i X 15[i-20(x, 1 + blue (Figure 13i).
Chemistry. — Psoromic and conpsoromic acids.
Habitat. — Canopy 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.- — 8 (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 1+ 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
24. P. disciforme, new combination
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
25. P. leiostomum
3. Pore 0.5-0. 1 mm in diameter, round, annulus lacking 26. P. obscurmn, new species
NUMBER 16
29
trerna 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 13/i
Thallus corticola, pallide viridi-cinereus, epi- phloeodes, planus, aetate rimosus, 8 cm latus; apothecia irregulariter aggregata, 0.4-0. 6 mm dianietro, 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-120p altum; sporae 8;nae, transversim 4-5 loculatae, 8p X lOp (Figure 13/i).
Chemistry. — No substances present.
Holotype. — Logging area at Newfoundland, Dominica, elevation 250 m. Hale 35229, January 1969 (US).
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.
24. Phaeotrema disciforme (Leighton) Hale, new combination
Figures 7c, 13/, 14a
Thelolrema disciforme Leighton, 1869:170 [type-collection: Central Province, Ceylon, Tliwaites (BM, lectotype; H, isotype): Figure 13y].
Thelolrema exalbidum Stilton, 1881:184 [type-collection:
Assam, Watt s.n. (BM, lectotype)].
Thelolrema aquiliniim Vainio, 1915:137 [type-collection: St.
Croix, Boergesen s.n. (TUR, lectotype; C, isotype)]. Leplolrema exalbidum (Stirton) Zahlbruckner, 1923:634. Thelolrema galacliinim Vainio, 1926:24 [type-collection: Maloapan, Mexico, Liebmann 7712 (C, lectotype); Figure 13/j].
Phaeolrema galaclinurn (Vainio) Zahlbruckner, 1932:245.
I'hallus 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 120[.t-140[.i high; spores ap- parently 4/ascus, 20-24 loculate, 7[f-12|f X 40[j.- 55p, 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 pan tropical 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 s.n.), 16b
(37939).
25. Phaeotrema leiostomum
Figures 5h, 6d, lib,c
Phaeolrema leioslomum (Tuckerman) Zahlbruckner, 1923:
608.
Thelolrema leioslomum Tuckerman, 1862:407 [type-collection:
Cuba, Wrighl 149 (FFI, lectotype); Figure 14h].
Thallus ashy to pale greenish white, epiphloeo- dal, continuous, smooth and shiny, forming colonies 2-5 cm broad; ajiothecia immersed to slightly emer- gent, 0. 3-0.6 mm in diameter, weakly carbonized, columella absent to well developed, noncarbon- ized (Figure 6d); pore round to irregularly elon- gate, surrounded by a raised ring, 0. 1-0.3 mm long; hymenium 130[x-150|j. high; spores 8/ascus, brown, 4 loculate, 8[i-10j.t X 16p-18|,i, I— (Fig- ures" bit, 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- trema leiostomum alone would not permit identi- fication.
30
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 14. — .Specimens of Thelotremataceae (all about X 10): a, Phaeotrema disciforme, new combination {Hale ‘17929); b, P. leiostounnn (Icctotype in FH); c, P. leiostomum {Hale 35203): d, P. obscuruDi, new species {Hale 38091); e, Thelotrcina carasseiise (isotype in FFI); /. T. carassense (Hale 37873); g, T. claiide.sliuitin (lectotype in G); b, T. subcaesium (lectotype in H); i, T. clandestinum {Hale 35136); j, T. couforme (lectotype in G); k, P. consauguineum (lecto- type in G): I, T. couforme {Hale 38062).
NUMBER 16
31
Specimens Examined. — 15 (35203), 16a (37956), I6b (37993).
26. Phmotrema obscumm Hale, new species
Figure lid
Thallus corticola, pallide brunneo-albidus, tenuis vel Iiypopliloeodes, 5 cm latus; apothecia modice emergentia, 0.3-0. 5 mm diametro, apice fuliginea, columellata; ostiolum rotimdum, 0.1 mm dia- metro, intiis albido-priiinosum; liymeniiim circa 120ji altiim; sporae 8:nae, traiisversim 5-6 lociilatae, 10p-14p X 24ji-28|i (Figure 14d).
Chemistry. — 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).
Habitat. — 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 albicluliim (Nylaiider) Miiller-Argau also has stictic and constictic acids, but the apothecia are immersed and clustered and the spores larger (40p).
Thelotrema
Thelotrema 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. glaiicopallens (16), T. clandestinum (10), T. interposiinm (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 papillosum and T. tuberciili- ferum 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. Columella narrow; spores less than 20(1, long 39. T. tenue, new species
3. Columella very broad; spores more than SOp, long 36. T. leucomelaenum
2. Thallus distinct, epiphloeodal; apothecia emergent, 0.5-1. 5 mm in diameter.
4. Spores 20p-40p long, 4-8/ asciis.
5. Spores about 20p long; pore not annulate but becoming black rimmed
35. T. leucimim
5. Spores about 40p long; pore becoming annulate 29. T. conforme
4. Spores more than lOOp long, 1/ascus.
6. Thallus warty-granular, P4- yellow 34. T. interpositum
6. Thallus smooth, P— or P-h orange.
7. Thallus P— 31. T. depressum
7. Thallus P+ orange 38. T. praestans
1. Central columella lacking.
8. Apothecia immersed, the pore flush with thallus surface.
9. Thallus greenish, thin, and shiny 33. T. glaucopallens
9. Thallus whitish, thick, dull 28. T. clandestinum
8. Apothecia emergent, pores not flush with thallus surface.
10. Spores less than 20p 30. T. confusum, new species
10. Spores more than 70p long.
II. Thallus P — ; collected in mossy forest zone.
12. Apothecia 1 mm tall, eroding around a large area of the pore
40. T. ttiberctiUferum
12. Apothecia barely emergent, pore area not eroding
S7. T. papillosum, new species
II. Thallus P-t- orange or yellow; collected in rain forest zone.
13. P-f 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
SMITHSONIAN CONTRIBUTIONS TO BOTANY
27. Thelotrema camsseme Figure I4e,f
Thelotrema carassense Vainio, 1890:79.
Type-Collection. — Carassa, Brazil, Vainio 1523 (TUR, lectotype; BM, FH, isotypes). (Figure I4e).
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; hymeniiim llOp-lSOp; spores 8/ascus, muriform, 1-2 X 8-10 loculate, 12ji-15fi X 30|i-40j,i, 1+ blue (Figure 14/). Chemistry. — 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 70p long). The taxonomy of the ecolumel- late psoromic acid-containing Thelotrerriata is unfortunately still incomplete. Thelotrema post- positiirn Nylander has large spores (more than lOOp), but all other species, excluding T. caras- sense, have small spores (less than 20p long).
Specimens Examined. — 16b (37859), 24b (37738), 26b (37873).
28. Thelotrema clandestinum Figure 14g,i
Thelotrema clandestinum Fee, 1837:90.
Thelotrema subcaesiiim Nylander, 1869:120 [type-collection: Brazil. Glaziou 2193 (H, lectotype; BM, C, FH-Tuck, UPS, US, W, isotypes); Figure 14/i].
Thelotrema concrelum Fee var. subcaesiiim (Nylander) Red- inger, 1936:96.
Type-Collection. — On Cinchona lancifoUa in America, without collector (G, lectotype). (Figure
14g) =
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
65p-75[i high; spores 8/ascus, imiseriate, 4-5 locu- late, muriform with 1-2 longitudinal septae in at least one of the locules, 6p-10p X 15p-20p, 1 + blue (Figure 14i).
Chemistry. — Psoromic and conpsoromic acids.
Habitat. — Canopy brandies in rain forest (500- 670 m).
Thelotretna clandestinum has small spores with few longitudinal septations. It would, in fact, be classified as an OceUnlaria unless the spores are carefully examined. Chemically similar O. tere- bratiila 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.
Specimens Examined. — 13 (35298, 35393), 14
(35103, 35105, 35136), 15 (35207), 16c (37768, 37872, 37981 ), 24b (35624a).
29. Thelotrema conforme Figure 14/,/
Thelotrema conforme Fee, 1837:89.
Thelotrema consangumeum Muller-Argau, 1887b:398 [type- collection: Apiahy, Brazil, Puiggari 477 (G, lectotype);
Figure 14ft].
Type-Collection. — On Cinchona, America meri- dionalis, without collector (G, lectotype) (Figure 14;).
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 170[i- 200[i high; spores 4/asciis, muriform, 8-10 loculate transversely, 2-4 loculate longitudinally, 18j.i-20ji X 35p-50|.i, 1+ blue (Figure 14/).
Chemistry. — Psoromic and conpsoromic acids.
Habitat. — Lower bole and trunk in open rain forest (670 m).
The Dominican material is a satisfactory match for Fee’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. leiicinmn which has small spores.
Specimens Examined. — 24a (37739, 38062).
NUMBER 16
33
30. Thelotrema confumm Hale, new species Figures Ic, 15
Thallus corticola, cinereo- vel viridi-albidus, epiphloeodes, planus, continuiis vel rimosus, nkidus, 4-8 cm latus; apodiecia numerosa, emer- gentia, 0. 8-1.0 mm diametro, apice obfiisca, columella nulla; ostiolum rotundum, 0.15-0.2 mm diametro, margine nigrocinctum; hymenium 120p- 130p altum; sporae 4:nae, murales, transversim 6-7 loculatae, longitudinaliter 0-2 loculatae, Gp-lOp X 12p-26p, 14- coerulescentes (Figure 15).
Chemistry. — Protocetraric acid.
Holotype. — Trail to Boiling Lake, elevation 650 m, Hale 37747, 9 December 1971 (US).
Habitat. — Canopy branches or lower bole in open areas (650-800 m).
The black-rimmed pore and fissured thallus re- semble Ocellularia nigropnncta, 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. — 16b (37807), 26a (37697).
Figure 15 Thelotrema confitsum, new species (Hale 37747)
(X 10).
31. Thelotrema depressum Figure 165,c
Thelotrema depressum Montagne, 1851:73.
Type-Collection. — Cayenne, French Guiana,
Leprieur 701 (BM, lectotype; H, isotype) (Figure 165).
Thallus and apothecia as in Thelotrema prae- slaits (see below); spores 1-2/ascus, murilorm with numerous transverse and longitudinal locules, 35p- 50ij, X ]50p-300p, 14- blue (Figure 16c).
Chemistry. — No substances present.
Habitat. — Canopy branches of trees in the rain forest (300-500 m).
The type-material of Thelotrema depressum is very fragmentary but color and chromatographic tests established that no P4- compounds were present. The four specimens from Dominica are itlentical. They may represent only a chemical population of T. praeslans, which is morphologi- cally identical, as far as we can judge from available specimens, but contains a strong P4- orange-red unknown substance. Thelotrema prae- siaus is more common (20 collections) and has a broader altitudinal range (250-800 m).
Specimens Examined. — 8 (38141), 12 (37901,
37922), 15 (35150).
32. Thelotrema dominicanum Hale, new species Figure 16a
Thallus corticola, pallide brunneo-albidtis, 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 diametro; hymenium 140[.i-160p altum; sporae 2-4;nae, murales, trans- versim 24-30 loculatae, longitudinaliter 2-3 locu- latae, 15[,i-20[.i X 80|.i-90[.i, 14- coerulescentes (Fig- ure 16rt).
Chemistry. — “CAnchonarxim” P-f unknown sub- stances.
Holotype.^ — Tree along trail to Morne Anglais, Dominica, elevation about 800 m. Hale 35355, January 1969 (US),
Habitat. — Tree bole in higher elevation rain forest.
The most unusual feature of this rare lichen is the presence of the “cinchonarxim” unknown, pre- viously thought to be restricted to the genus Ocelhilaria. 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.
34
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Figure 16. — Specimens of Thelotreniataceae (all about X 10): a, Thelotrema dominicanum, new species {Hale 35353); b, T. depressiirn (isotype in H); c, T. depressum {Hale 38141); d, T. glaticopallens (lectotype in FH-Tuck); e, T. pechueli (lectotype in G); f, T. butuanum (lectotype in W); g, T. homopastoides (lectotype in TUR); h, T. glaucopallens {Hale 35147); i, T. inter- positum (isotype in L); j, T. interpositum {Hale 35388); k, T. leucinum (lectotype in G); t, T. leucinum {Hale 38100).
NUMBER 16
35
33. Thelotrema glaucopallem Figure I6dji
Thelotrema glaucopallens Nylander, 1863:327.
Pyrenula clandeslina Achaiius, 1814:10 [type-collcctioii: on Cinchona jlava, .South America (H, lectolype; S, isotypc)]. Thelotrema laenigans Nylander var. avertens Nylander, 1867: 318 [type-collection: Tctjuendama, Colombia, Lindig 893 (H, lectotypc; BM, FH-Ttick, G, B, \X , isotypes)].
Thelotrema pechueli Miiller-,-\rgau, 1880:34 [type-collection: Quillii River, Angola, Pechuel-Loesche s.n. (G, lectotype); Figure 16e].
Ocellularia clandestina (Acharius) Miiller-.Argau, 1877a:7. Thelotrema homopastoides Vainio, 1896:207 [type-collection: Richmond Valley, St. Vincent, Elliott 327 (TUR, lectotypc; BM, isotype); Figure 16g].
Thelotrema butuanum Vainio, 1921:183 [type-collection: Lu- zon, Philippines, Fenix, BS-28347 (\V, lectotype); Figtire 16/].
Tvpe-CollectioiN. — Cuba, Wright (FH-Tuck, lectotype, as 28; L, UPS, isotypes) (Figure \6d).
'Fhallus 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 mm in diameter; hymenium GOp-SOp high; spores 8/ascus, muriform, 6-8 loculate transversely, 2-3 loculate longitudinally, 7p-10p X 15p-24p, I — (Figure IG/t).
CtiEMLSTRY. — Stictic and constictic acids. Habitat. — Buttresses, 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. — 10 (38082), 11 (37999,
38008, 38047), 15 (35147), 16a (.37670, .37876, 3804.3, 38044), 20a (37783), 21 (.3808.3, .38095, 38155, 38156), 2.3 (.37678), 26b (35367), 27 (37891), 28 (35431 ).
34. Thelotrema interpositum Figure 16i,/
Thelotrema interpositum (Nylander) Miiller-.Argau, 1881:526. Ascidium interpositum Nylander, 186.3:3.36 [type-collection:
Cuba, Wright 28 (H, lectotype; BM, FH, L, M, P, UP.S,
isotypes); Figure 16;].
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 di;imeter, walls heavily carbonized, columella jnesent; pore round, 0.15-0.2 mm in diameter, opening into a tubelike area leading to the prui- nose toj) of the cohimella; hymenium 200p high; spores 2/ascus, murilorm with numerous transverse and longitudinal locules, 28p-40p X 85p-125p, 1+ bhie (Figure 16)).
(aiEMi.sTRV. — Psoromic acid.
FIaiut.at. — Canopy branches and upper bole area (sea level to 670 m).
The coarsely verrucose thallus and large emer- gent ajiothecia with large spores distinguish this species. I’wo other columnellate psoromic acid- containing species, T. ronforme and T. leucintim, have a smooth thallus and much smaller spores.
.Specimens Examined. — 5 (35388), 7 (.352,31), 15 (.35148, .35174), 16a (.37656, .37657), 16c (.37760), 22 (37853, 37976).
35. Thelotrema leucinum Figure 16/;,/
Thelotrema leucinum Midlcr-.Argau, 1887a: 10.
T\ pe-Collection. — On Cinchona, South Amer- ica (G, lectotype) (Figure 16/i).
Thallus light ashy gray, ejiiphloeodal, continuous to cracked with age, shiny, forming a colony about 10 cm wide; apothecia numerous, nearly immersed to semi-emergent, 0. 1-0.6 mm in diameter, upper wall carbonized, columella present, 0.2 mm wide; pore round, 0.05-0.12 mm in diameter; hymenium 110p-120p high; spores 8/ascus, muriform with 4 locules transversely, 2 locules longitudinally, 7p- lliji X ]2j.i-18[.i, 1+ blue (Figure 16/).
(inEMi.sTRY. — Psoromic and conpsoromic acids.
Habitat. — Cano[)y branches in rain forest (300 m).
The type of T. leucinum, one of Fee’s Cinchona
36
SMITHSONIAN CONTRIBUTIONS TO BOTANY
bark specimens, is in very poor condition and it is nnl’ortnnate that Mhller 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.
Specimen Examined. — 9 (38100).
36. Thelotrema leucomelaenum Figures 4a, h; 6e; na,d
Thelotrema leucomelaenum Nylancler, 1863:329.
Thelotrema pauperius Nylancler, 1867:318 [type-collection;
Rio Negro, Colombia, Lindig (H, lectotype; BM, M, iso- types); Figure 17fo].
Thelotrema leucomelaenum var. elevatum Vainio, 1915:137
[type-collection: Morne Anglais, Dominica, Elliott 1534
(TUR, lectotype): Figure 17c].
Type-Collection. — Fusagasuga, Colombia, Lin- dig 2777 (H, lectotype; BM, FH, G, M, P, UPS, isotypes) (Figure 17rt).
Thalhis white, thin and in part hypophloeodal (Figure -la,b), forming colonies up to 8 cm broad; apotliecia 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 Oc); hynienium 130[i high; spores 8/ascus, muriform, 4-6 loculate transversely, 2-3 loculate longitudinally, 12p-15[i X 30[i-36u, I— (Figure lid).
Chemistry. — No substances present.
Habitat. — Smaller branches of trees in mossy forest (800-900 m).
A pantropical species, T. leiicomelaeniim has been correctly identified by most lichenologists. The ajiothecia, while at most semi-emergent, stand out because of the black pore rim. The broad columella occupies most of the interior of the apothecia.
Specimens Examined. — 25a (37706, 37710, 37717, 37718, 37871), 25b (35491), 26a (38076).
37. Thelotrema papillosum Hale, new species Figure 17/
Thalhis corticola, cinereo-viridis, epiphloeodes, minute pa])illosus, nitidus, 6 cm latus; apothecia vix emergentia, 0.7-0. 9 mm dianietro, intus in-
colores, columella nulla; ostiolum rotundum vel irregulariter laceratum, 0. 1-0.3 mm diametro; hynienium circa 200[i altum; sporae l-2;nae, mu- rales, loculis numerosis, 36[i-40[i X 140[.i-150p, I— (Figure 17/).
Chemistry. — No substances present.
Holotype. — Rain forest above Newfoundland, Dominica, elevation about 300 m. Hale 38124, 7 May 1972 (US).
Habitat. — Canopy branch in rain forest.
This species would be easy to miss in the field. The small greenish warty thallus is not like any Tlielolrema. 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 tnbercuUferum, tv'hich has large spores but strongly emergent erod- ing apothecia.
38. Thelotrema praestans Figure 17e,g
Thelotrema praestans Miiller-Argaii, 1895:453.
Thelotrema elUottii Vainio, 1896:207 [type-collection: Rich- mond Valley, St. Vincent, Elliott 246 pro parte (BM, lecto- type); Figure 17/].
Type-Collection. — Rio de Janeiro, Brazil, Por- tella s.n. (G, lectotype; BM isotype) (Figure lie).
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; hymeniiini about 200[i high; spores 1-2/ascus, muriform with many transverse and longitudinal locules, 30[i-45[i X 100p-275[.i, 1 + 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- pressiim but differs in producing a P+ orange-red unknown substance just below psoromic acid in both hexane and benzene solvents. The apothecia are large and heavily carbonized. The fine struc- ture of tile cortex is aculeate, as in T. depressum.
Specimens Examined. — 7 (35237, 35244, 35254, 35273), 8 (38087, 38121, 38159), 9 (38135), 11
NUMBER 16
37
Fir.i'RE 17 Specimens of Thelotrcmataceae (all about X 10): a, Thelotreuia leucomelaeruuii
(Icctotype in H); b, T. paitperiti!, (Icctotype in H); c, T. leuconiclaenuiu var. elevatxim (lectotype in TUR); d, T. lexicomelaeniitn (Hale 35491); e, T. prae.slaiis (Icctotype in G); /, T. ellioltii (lectotype in I5M): ", T. praeslans (Hale 35273); h, T. tuherculifennn (lectotype in Tl’R); i, T. 'tiiberculiferiim (Hale 37719); j, T. papillosum, new species (Hale 38124); k, 7'. 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 Figuri.s 4c, d; 6/; 17ft
Tliallus corticola, cinereo-albiis, hypophloeodes (Figure 4.c,d), nitidus, 7 cm latus; apothecia mi- merosa, iinmcrsa vel parce emergentia, 0.3-0. 4 mm diametro, apice obiiisca, coliimellata (Figure 6/); ostiolum rouiiidum, 0.05-0.1 mm diametro; hy- menium OOp-lOOp ahum; sporae 8:nae, murales, transversim 4-5 loculatae, longitudinaliter 1-2 loculatae, 6[i-7i.i X 17p-19p, I— (Figure Ilk).
Chemistry. — No substances present.
Moeoit pe. — Mossy forest zone on trail to Morne Diablotin, Dominica, elevation about 1100 m. Hale 35430, January 1969 (US).
Habita'i. — Lower trunk in rain forest to mossy forest.
Superficially this species resembles Ocellularia pyrenuloides in having small blackish apothecia and a thin columella. TheJolrema lencoyrielaeniim, another hypophloeodal mossy forest species, has larger spores and a very broad columella. Other than these there are no related species.
40. Thelotrema tiiberculiferum Figure 17/!,/
Thelotrema iuberculifertim Vainio, 1915:136.
Type-Collection. — Savane-aux-Ananas, Guade- loupe, Duss 1497 (TUR, lectotype) (Figure 17/!).
1 hallus 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 l lOp-lOOp high; spores 1-2/ascus, muriform with numerous transverse and longitudi- nal locules, 30p-50u X 60|.i-120p, 1+ pale blue (Figure 17/).
CiiiEMisTRY. — No substances present.
Habiiat. — Exposed moss-covered branches in mossy forest (800-900 m).
14ns Tlielotretna might be mistaken at first for a Pertusarin. 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 (37719), 26a (37868).
Leptotrema
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. Columella present, simple to actinoid 43. L. fisstim
1. Columella lacking.
2. Pore wide, 0.5-0. 8 mm in diameter, the exposed disc dark brown to black
46. L. spondaicum
2. Apothecia immersed to emergent with pores 0. 1-0.2 mm in diameter.
3. Thallus thick rvith a columnar cortex; red crystalline inclusions often present
48. L. ivightii
3. Thallus thin with a normal cortex lacking pigments.
4. Apothecia rvith an inner lepadinoid exciple.
5. Spores about 20fi long 45. L. occtiittim
5. Spores about 45f.i long 47. L. subcompiinctiim
4. .Apothecia lacking a lepadinoid exciple.
6. Pore distinct, 0. 1-0.2 mm in diameter; spores 20p-30|.t long 41. L. hahianinn
6. Pore ^ery tiny, less than 0.1 mm in diameter; spores large, more than 40|.i long.
7. Pore depressed in a darker area 42. L. deceptiim, new species
7. Pore not depressed, surrounding area the same color as the thallus.
8. Apothecia immersed with flush pores 47. L. subcompiinctum
8. .Apothecia more or less emergent 44. L. microglaenoides
NUMBER 16
39
eluding the collection of L. hahiantim by Elliott). It is rarer than would guess from this number of collections since one species, L. deceptti7n, is represented by 21 specimens. Leptotrema spondai- cinn was found 8 times, L. wightii 3 times, and the remainder once each. On the world level the genus has about 50 species.
Leptotretfia 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 the broad rain forest zone in contrast to 25 above 900 m.
41. Leptotrema bahianum Figure 5/
Leptotrema bahianum (Acharius) Miillcr-Argau, 1887a: 12. Thelotrema lepadinum var. bahianum Acharius, 1803:132
[type-collection: Bahia, Brazil, without collector (H, lecto-
type; L, I PS, isotypes)]. — Hale, 1972:193, fig. 2c. Thelotrema radius Vainio var. dominicanum Vainio, 1915:
136 [type-collection: Prince Rupert, Dominica, Elliott 1303
(TUR, lectotype): Figure 5/].
Thallus corticolous, epiphloeodal, light tannish Avhite, about I cm broad; apothecia numerous, emergent, 0.4-0. 6 mm in diameter, noncarbonized and without a columella; pore round, about 0.1 mm in diameter; hymenium 150j.i high; spores 8/ascus, muriform with 5-6 transverse locules and 2-3 longitudinal locules, 12[i-14p X 16[i-24p, I — (Figure 5/).
Chendstry. — 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 lotvland 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 cm latus; apothecia immersa, 0.3-0. 4 mm diametro, intus incolores, columella nulla; ostiolum minutum, 0.02-0.05 mm diametro, depressum.
anibitu obfuscans; hymenium 100ti-120p altum; s|iorae obfuscae, 4:nae, murales, loculis ntimerosis, I4[.i-28p X 40|.t,-70(.i, I— vel 1+ pallide coerules- centes (Figure 18).
Chemistry. — Stictic and constictic acids.
Hoi.otype. — Sloanen buttress, disturbed [irimary forest at Middleham Estate, elevation about 670 m. Hale 37860, December 1971 (US).
Habitat. — Buttresses and exposed roots, sajdings, lianas, lower bole in higher elevation rain 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. tnicroglaenoides and L. suhcom punc- lutn. 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 hrst I had considered this (and similar species) as having closer relationships to the Py- renulaceae. Mr. Richard Harris, a student of the Pyrenulaceae, gave the following opinion in sup- port of placing L. deceptum in the Thelotrema- taceae:
Most pyrciiocarpoiis lichens have some sort of carhonizecl “wall” simoiincling the hymenium [/.. deceptum is noncar- bonizeil]. The spores react with IKI; a few pyrenocarps have this reaction also but have entirely difTercnt paraphyses. The paraphyscs [of /,. deceptum] are unlike those in the pyreno-
Figure 18 Leptotrema deceptum, new species {Hale 37860)
(X 10)
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. Tlielotrema lepadinum has asci very like those in your specimen. Finally I have never found depsides or depsidones in any pyrenocarp- ous lichens.
Spfximens Examined. — 21 (37373, 38104, 38105, 38109, 38110, 38112, 38117, 38133, 38136, 38139, 38142, 38148, 38151, 38153), 22 (37640, 37648, 37836, 37839, 38037), 23 (37673).
43. Leptotrema fissum Figure 19fl,c
Leptotrema fissum (Nylander) Muller-.Argau, 1882a:333. Thelotrema fissum Nylander, 1859:258 [type-collection: Bour- bon, Richard s.n. (H, lectotype; G, P, isotypes); Figure 19a].
Leptotrema integrum Miiller-Argau, 1887b:399 [type-collec- tion: Australia, Sayer (G, lectotype; S, isotype); Figure 195]. Leptotrema fissum (Nylander) Zahlbruckner, 1923:634 [super- fious combination].
Thallus greenish ashy giay, epiphloeodal, smooth, continuous or cracked with age, forming colonies 4-6 cm Irroad; 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 lOOp high; spores brown, muriform with 4-6 transverse locides and 0-2 longitudinal loctiles, 6p-7p X ]0p-13p, I— (Figure 19c).
Chemistry. — Psoromic and conpsoromic acids. Habitat. — Canopy branches in rain forest (430 m).
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 dose in these respects to Ocel- hilaria comparabilis.
Specimen Examined. — 11 (38031).
44. Leptotrema microglaenoides Figure 19d,e
Leptotrema microglaenoides (Vainio) Zahlbruckner, 1923:637. Thelotrema microglaenoides Vainio, 1896:206 [type-collection: St. Vincent, Elliott 266 (BM, lectotype); Figure 19d].
Thallus light whitish tan, epiphloeodal, con- tinuous and shiny, forming a colony about 6 cm broad; apothecia semi-emergent, 0. 8-1.0 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 150p-170p; spores brown, 4/ascus, muri- form with numerous transverse and longitudinal loctiles, 18p-22p X 36p-48p, I— (Figure 19e).
Chemistry. — Stictic and constictic acids.
Habitat. — Branches of tree in rain forest (430 m).
The lectotype of this species is rather frag- mentary with very few well-developed apothecia. Idle 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 120|r long. I found spores about 50j.i long in the lectotype. The large complex of stictic acid-containing species (e.g. Leptotrema mottosportim (Nylander) Miiller- Argau, L. phaeosportim (Nylander) Miiller-Argau, and L. recliisum (Krempelhuber) Zahlbruckner) is still far from resolved.
Specimen Examined. — 11 (37984).
45. Leptotrema occultum (Eschweiler) Hale, new combination
Figure 19/,g
Thelotrema occultum Eschweiler in Martins, 1833:174 [type-
collection: South America (M, lectotype; G, isotype); Figure
19/].
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-70|.i high; spores 4-8/ascus, brown, muriform with 5-7 transverse locules, 2-3 longitudinal locules, 9|i-10p X 18j.i-24i, I— (Figure 19g).
Chemistry. — Norstictic acid.
Habitat. — Trees in dry scrub forest near sea level.
Leptotrema occultum can be typified with the Martins specimen in Munich and the isotype frag- ment in Geneva. It is virtually indistinguishable
NUMBER 16
41
Figurk 19. — .Specimens of Thelotremataceae (all about X 10); a, Leptolreina fissum (Icctotype in H); b, L. integrum (isotype in S); c, L. fissum {Hale 38031); d, L. microglaenoides (lectotypc in BM): e, L. microglaenoides {Hale 37981); /, L. occullum (isotype in G); g, L. occullum {Hale 35718); h, L. spondaicum (lectotype in FH-Tuck); i, L. spondaicum {Hale 35503).
from pantropical L. compunctiim (Acharius) Muller-Argau, which contains stictic acid in addi- tion to norstictic acid. I am keeping L. occtillum separate until study of more material shows whether the chemical and spore variation (spores being generally smaller in norstictic acid-containing specimens) are valid or not. The L. compimctum group is very characteristic of dry scrub forest throughout the tropics.
Specimen Examined. — 2 (35718).
46. Leptotrema spondaicum
Figure 19/ia
Leptotrema spondaicum (Nylander) Zahlbriickner, 1923:640. Thelotrema spondaicum Nylander, 1863:330 [type-collection: Cuba, JVright 35 (FH- Fuck, lectotypc); Figure 19/t].
Thallus ashy white, epiphloeodal to evanescent, waxy, smooth, forming colonies 3-15 cm wide; apothecia numerous, round to irregular, 0.8-1. 4
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 ISOp- 170p high; spores 1-2/ascus, brown, muriform with numerous transverse and longitudinal locules, SOp- 40p X 80p-130p, I— (Figure 19i).
Chemistry. — Virensic acid.
Habitat. — Trunks of primary and secondary trees in open pastures and citrus groves at higher elevations (600-700 m).
Leptotretna spoiidaicti?n 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. — 18b (35590), 19 (38077), 20a (37779, 37789, 37790), 20b (35492, 35503), 26a (37696).
47. Leptotrema subcompunctum Figure 20a, b
Leptotrema subcompunctum (Nylander) Zahlbruckner, 1923: 640.
Thelotrema subcompunctwn Nylander, 1868:76 [type- collection: Lifu, Loyalty Islands, Thiebaul s.n. (H, lecto- type; G, P, isotypes); Figure 20rt].
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 20. — 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. flavicans (lectotype in G); e, L. subconcretum (lectotype in BM); /, L. wightii {Hale 35882).
NUMBER 16
48
brown, ]-2/ascus, muriform with 8-11 transverse locules and 2-4 longitudinal locules, 10^-13[i X 45|.i-55^, 1“ (Figure 20b).
Chemistry. — 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 1 have seen from the New World.
Specimen Examined. — 18a (37888).
48. Leptotrema wightii
Figures Id, 20c,/
Leptotrema wightii (Taylor) Miiller-Argau, 1882:499. Endocarpon wightii Taylor, 1847:155 [type-collection; Madras, India, Wight (FH, lectotype; BM, G, isotypes); Figure 20c].
Thelotrema prevostianum Montague, 1849:292 [type-collection:
Antilles, Prevost s.n. (P, lectotype): Figure 5d]. Leptotrema prevostianum (Montagne) Montague, 1856:364. Thelotrema sub con ere turn Leighton, 1869:169 [type-collection: Central Province, Ceylon, Thwaites 89 (BM, lectotype; FI, G, P, S, UPS, W, isotypes); Figure 20c].
Phaeotrema subconcretum (Leighton) Miiller-Argau, 1887a;
10.
Leptotrema subconcretum (Leighton) Miiller-Argau, 1891:277. Leptotrema flavicans 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 lOOp high; spores brown, 8/ascus, muriform with 4-6 transverse locules and 1-2 longitudinal locules, Op-llp X 20p-28p, I— (Figure 20/).
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. — 1 (35582, 35584), 3
(32918).
Literature Cited
Acharius, E.
1803. Methodus qua omnes detectos Lichenes. 394 pages. Stockholm.
1812. Anmai'kningar vid Lafslagtet Thelotrema med Nogare Bestamniande af Dess Artec. Kongliga Vet- enskapliga Academiska Nya Handlingar, 1812:79- 95.
1814. Synopsis Methodica Lichejxutn . 392 pages. Lund. Culberson, C. F., and W. L. Culberson.
1968. The Constituents of Some Species of the Thelotre- mataceae. The Journal of Japanese Botany, 43:316- 323.
Culberson, C. F., and M. E. Hale, Jr.
1973. 4-0-Dcmethylnotatic Acid: A New Depsidone in Some Lichens Producing Hypoprotocetraric Add. The Bryologist, 76:77-84.
Fee, A. L.
1824. Essai sur les cryptogames des ecorces exotiques
officinales. 167 pages. Paris.
1837. Essai sur les cryptogames des ecorces exotiques
officinales, 11: Supplement et Revision. 178 pages.
Paris.
Hale, M. E., Jr.
1971a. Morden-Smithsonian Expedition to Dominica: The
Lichens (Parmeliaceae). Smithsonian Contributions to Botany, 4: 1-25.
1971b. Studies on Parrnelia Subgenus Xanthoparmelia (Lichenes) in South Africa. Botaniska Notiser, 124: 343-354.
1972. Typification of Species in the Lichen Family Thelo- trcinataceae Described by Acharius. Botaniska Notiser, 125:186-198.
Johnson, G. T., and W. L. Brown
1941. Stages in the Development of Thelotrema inter- positum. Mycologia, 33:601-608.
Krempelhuber, A.
1876. I.ichenes Brasiliensis Collect! a D. Glaziou in Provincia Brasiliensi Rio Janeiro. Flora, 59:217-224. Leighton, W. A.
1866. Lichenes .Amazonici et Andini. Transactions of the Linnean Society of London, 25:433-460.
1869. The Lichens of Ceylon Collected by G. H. K. Thwaites. Transactions of the Linnean Society of London, 27:161-185.
44
SMITHSONIAN CONTRIBUTIONS TO BOTANY
Letrouit-Galinou, M.-A.
1966. Recherches sur I’ontogenie et I’anatomie comparees des apothecies de quelques discolichens. Revue Bryologique et Lichenologiques, 34:413-588. Magnusson, A. H., and A. Zahlbruckner
1943. Hawaiian Lichens. Arkiv for Botanik, 3!A(l):l-96. Martins, C. F. P. von
1833. Flora Brasiliensis. Stuttgart. [Lichens, by F. Eschweiler, l(part 1): 1-390.]
Meyen, F., and J. Flotow
1843. Observationes in Itinere Circum Terram Institutae: Lichenes. Novorum Actoruni Academiae Caesareae Leopoldmo-caroUnae Naturae Curiosorum, Supple- ment 1, 19:209-232.
Montague, J. F.
1849. Sixieme centurie de plantes cellulaires exotiques nouvelles, cryptogamae taitensis. Annales Sciences Naturelles Bontanique, series 3, 12:285-320.
1851. Cryptogamia Guyanensis sen Plantarum Cellularium in Guyana Gallica .Annis 1835-1849 a CL Leprieur Collectarum Enumeratio Universalis. Annales Sci- ences Naturelles Botanique, series 3, 16:47-81.
1856. Sylloge Generuni Specierumque Cryptogamarum. 498 pages. Paris.
Montague, J. F., and R. von der Bosch
1855. Lichenes. Pages 427-494 in F. Junghuhn, Enutn- eratio Plantarum Ques in Insulis Java et Sumatra Detexit.
Miiller-Argau, J.
1880. Lichenes Africa Occidcntalis a Cll. Dr. Pcchuel- Uvesche et Soyou e Regione Fluminis Quilla et ex Angola Missi, in Mus. Bot. Reg. Berolinensi Servati. Linnaea, 43:31-48.
1881. Lichenologische Beitrage XIV. Flora, 64:513-527.
1882a. Lichenologische Beitrage XV. Flora, 65:327-336. 1882b. Lichenologische Beitrage XVI. Flora, 65:492-509. 1883. Revisio Lichenum Meyeniorum. Jahrhuch der
Konlige Botanische Gartens zu Berlin, 2:308-319. 1887a. Revision Lichenorum Feeanonim. Revue Mycolo- gique, 35:1-16
1887b. Lichenologische Beitrage XXVI. Flora, 70:396-402. 1888. Lichenes Paraguayensis. Revue Mycologique, 10:
113-120.
1891. Lichenes Victoryenses. Nuovo Giornale Botanico
Italiano, 23:276-279.
1892. Lichenes Exotici. Hedwigia, 31:276-288.
1893. Lichenes. In J. Durand and H. Pittier, editors,
Primitiae Florae Costaricensis. Bulletin Socicte
Botanique Belgique, 32:122-173.
1895. Thelotremeae et Graphideae Novae Quae Praesertim ex Herb. Reg. Kewensi Exponit. Journal of the Linnean Society of London, Botany, 30:451-463. Nylander, W.
1859. Lichenes Exotici. Annales Sciences Nattirelles Bot- anique, series 4, 11:203-264.
1863. Lichenes. In J. Triana and J. E. Planchon, editors. Prodromus Florae Novo-Granatensis ou Enumera-
tion des Plantes de la Nouvelle-Grenade. Annales Sciences Naturelles Botanique, series 4, 19:286-382.
1867. Lichenes. In J. Triana and J. E. Planchon, editors, Prodromus Florae Novo-Granatensis ou Enumeration des Plantes de la Nouvelle-Grenade. Additamentum. Annales Sciences Naturelles Botanique, series 5, 7: 301-354.
1868. Synopsis Lichenum Novae Caledoniae. Bulletin So- cieie Linnee Normandie, series 2, 2:39-140.
1869. Lichenes in Brasilia a Cl. Glaziou Lecti. Flora, 52: 117-126.
1876. Collemacei, Caliciei, Cladoniei et Thclotremei Cubani novi. Flora, 59:558-562.
Poelt, J.
1973. Classification. Pages 599-632 in V. .Ahmadjian and M. E. Hale, editors. The Lichens. New York: Aca- demic Press.
Redinger, K. M.
1936. Thelotremataceae Brasiliensis. Arkiv for Botanik, 28A(8): 1-122.
Salisbury, G.
1971. Thelolrema wightii (T. Tayl.) Nyl. Portugaliae Acta Biologica, series B, 11:35-37.
1972a. Thelotrema Ach. sect. Thelotrema, 1: The T. lepadinum group. Lichenologist, 5:262-274.
1972b. Thelotrema sect. Thelotrema, 2: The T. platy- carpum group. Revue Bryologique et Lichenolo- gique, 38:281-290.
Santesson, R.
1952. Foliicolous Lichens, I: A Revision of the Taxonomy of the Obligately Foliicolous, Lichenized Fungi. Symbolae Botanicae Upsalienses, 12(l):l-590.
Sprengel, K.
1827. Systema Vegetabilium 4. Edition 16. Part 2, 410 pages. Gottingae.
Stirton, J.
1881. On ATgetable Parasites on the Tea Plant, More Especially That of .Assam. Proceedings of the Philo- sophical Society of Glasgoiu, 13:181-193.
Taylor, T.
1847. New Lichens, Principally from the Herbarium of AV. J. Hooker. London Journal of Botany, 6:148-197.
Tuckerman, E.
1862. Observations on North .American and Other Lichens. Proceedings of the American Academy of Arts and Sciences, 5:383-422.
Vainio, E. A.
1890. Etude sur la classification naturelle et la morpholo- gic des lichens du Bresil. Acta Societas Fauna et Flora Fennica, 7(7): 1-247.
1896. Lichenes .Antillarum a AV. R. Elliott Collecti. Journal of Botany British and Foreign, 34:204-210.
1915. Additamcnta ad Lichenographia .Antillarum Illus- trandam. Annales Academiae Scientiarum Fennicae, 6 A (7): 1-226.
1921. Lichenes Insularum Philippinarum HI. Annales Academiae Scientiarum Fentiicae, 15A: 1-368.
1923. Lichenes in Insula Trinidad a Profe.ssor R. Thaxter
NUMBER 16
45
Collecti. Proceedings of the American Academy of Arts and Sciences, 58:129-147.
1926. Lichenes Mexican!. Dansk Botanisk Arkiv, 4:1-25. Van Overeem-De Haas, C. and D.
1922. Verzeichnis der in Niederlandisch Ost-Indien bis dem Jahre 1920 Gefimdenen Myxomycetcs, Fungi
und Lichenes. Bulletin du Jardin Dotanique, series 3, 4:1-146.
Zahlbruckner, A.
1912. Neue Flechten VI. Annales Mycologici, 10:359-384. 1923. Catalogiis lichenum universalis. 2:580-643. Leipzig. 1932. Catalogus lichenum universalis. 8:244-247. Leipzig.
Index
(Index to main entries only; synonyms in italics)
Ascidiiim cinchonarum, 17 interposilum, 35 rJiodostromiim, 25 Endocarpon wiglilii, 43 (iraphis alborosella, 16 Leptotrema bahiantim, 39 deccptnm, 39 exalbidum, 29 fissnm, 40 ftavicans, 43 integrum, 40 microglaenoidcs, 40 monosponim, 40 occidtnm, 40 phacospornm, 40 prevostianinn, 43 reclusum, 40 spondaicum, 41 sidjcompunctum, 42 subconcreluin, 43 wightii, 43
Myriotrema olivacea, 24 Ocelliilaria alboolivacea, 19 alborosella, 16 antillcnsis, 17 berkeleyana, 17 Ixtnplandiae, 24 cavata, 17 cinchonarum, 17 cincho77arum f. intermedia, 17 clandestina, 35 comparabilis, 17
comparabilis var. microcarpa, 17 concolor, 19 conglomcrata, 19 discoidea, 17
dotningensis var. fecunda, 20
dominicana, 19
efformata, 17
exantbismocarpa, 20
fecunda, 20
glaziovii, 17
hotnothecia, 20
isertii, 20
latilabra, 19
lindigiana, 17 maculata, 22 mordenii, 22 multilocularis, 20 nigropnncta, 22 olivacea, 24 papillata, 24 perforata, 25 plalycarpella, 16 porinoides, 20 pyreiftiloides, 25 remariens, 28 rhodostroma, 25 rimosa, 27 rufocincta, 25 sorediata, 27 subcavata, 27 terebrata, 17
terebrata var. abbreviatula, 25 terebratida, 28 x’aga, 28 verrucosa, 24 xanthostroma, 22 Phaeotrema aggregatum, 29 albididum, 31 disciforme, 29 galactinum, 29 leiostomum, 29 lirelliforme, 29 obscurum, 31 subconcreluin , 43 Pyrenula clandestina, 35 Thelotrema aquilinum, 29 butuanum, 35 carassense, 32 cava turn, 17 clandestinum, 32 clandestinum f. remanens, 28 comparabile, 17 concretum var. subcaesium, 32 conforme, 32 confusum, 33 consanguineum, 32 deprc,ssum, 33
☆ U.S. GOVERNMENT PRINTING OFFICE: 1974 546-36l/l4
disciforme, 29
doniingense var. fecundum, 20
tloininicanum, 33
elliottii, 36
exalbidum, 29
exant h is mocarp u nr, 2 1
excavatum, 25
excax’atum var. impressulum, 13, 25
fissnm, 40
galactinum, 29
glaucopallens, 35
honwpastoides, 35
homothecium, 20
interpositum, 35
laevigans var. avertens, 35
leiostomum, 29
lepadinum var. bahianum, 39
Icucinum, 35
leucomelaenum, 36
leucomelaenum var. elevatum, 13, 36
microglaenoides, 40
occullum, 40
papillalum, 24
papillosum, 36
pauperius, 36
peck u el i, 35
perforatum, 25
platycarpellum, 16
porinoides, 20
postposituni, 32
pracstans, 36
prevoslianum, 43
rhodostroinum, 13
rudius var. dominicanuni, 13, 39
spondaicum, 40
subcaesium, 32
subcavatum, 28
subcompunctum, 42
subconcretum, 43
tenue, 38
tercbratulum, 28
•terebratum var. abbreviatulum, 25 tubcrculi fern 111, 38 vagum, 28
46
Publication in Smithsonian Contributions to Botany
Manuscripts for serial publications are accepted by the Smithsonian Institution Press, sub- ject to substantive review, only through departments of the various Smithsonian museums. Non- Smithsonian authors should address inquiries to the appropriate department. If submission is invited, the following format requirements of the Press will govern the preparation of copy.
Copy must be typewritten, double-spaced, on one side of standard white bond paper, with I Vi" top and left margins, submitted in ribbon copy with a carbon or duplicate, and accompa- nied by the original artwork. Duplicate copies of all material, including illustrations, should be retained by the author. There may be several paragraphs to a page, but each page should begin with a new paragraph. Number consecutively all pages, including title page, abstract, text, litera- ture cited, legends, and tables. The minimum length is 30 pages, including typescript and illus- trations.
The title should be complete and clear for easy indexing by abstracting services. Taxonomic titles will carry a final line indicating the higher categories to which the taxon is referable: “ ( Leguminosae : Faboideae).” Include an abstract as an introductory part of the text. Identify the author on the first page of text with an unnumbered footnote that includes his professional mailing address. A table of contents is optional. An index, if required, may be supplied by the author when he returns page proof.
Two headings are used: (1) text heads (boldface in print) for major sections and chap- ters and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further headings may be worked out with the editor.
In taxonomic keys, for easy reference, number the taxa and their corresponding headings throughout the text; do not incorporate page references in the key.
In synonymy, use the long form (taxon, abbreviated author, abbreviated journal or book title, volume, part if necessary, page, year) with no reference at the end of the paper under “Literature Cited” or use the short form (taxon, author, date: page) with a full reference at the end of the paper under “Literature Cited.” Begin each taxon at the left margin with subse- quent lines indented about three spaces. Within an entry, use a period-dash (. — ) to separate each supplementary reference. Enclose with square brackets any annotation in, or at the end of, the entry. .For references within the text, use the author-date system: “(Jones, 1910) ” and “Jones (1910).” If the reference is expanded, abbreviate the data: “Jones (1910:122, pi. 20: fig. 1).”
Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they should not contain rules. Formal tables must be submitted as pages separate from the text, and each table, no matter how large, should be pasted up as a single sheet of copy.
Use the metric system instead of, or in addition to, the English system.
Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed throughout the printed text. They will be termed Figures and should be numbered consecu- tively; however, if a group of figures is treated as a single figure, the components should be indicated by lowercase italic letters on the illustration, in the legend, and in text references: “Figure 9b." If illustrations (usually tone photographs) are printed separately from the text as full pages on a different stock of paper, they will be termed Plates, and individual components should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2). Never combine the numbering system of text illustrations with that of plate illustrations. Submit all legends on pages separate from the text and not attached to the artwork. An instruction booklet for the preparation of illustrations is available from the Press on request.
In the bibliography (usually called “Literature Cited”), spell out book, journal, and article titles, using initial caps with all words except minor terms such as “and, of, the.” For capitalization of titles in foreign languages, follow the national practice of each language. Underscore (for italics) book and journal titles. Use the colon-parentheses system for volume, number, and page citations: “10(2) :5-9.” Spell out such words as “figures,” “plates,” “pages.”
For free copies of his own paper, a Smithsonian author should indicate his requirements on “Form 36” (submitted to the Press with the manuscript). A non-Smithsonian author will receive 50 free copies; order forms for quantities above this amount with instructions for pay- ment will be supplie4 when page proof is forwarded.
O ^ , x^' — ' — ^ O - O
illSNI~'lslVIN0SHill^S^S3 I a V H 8 ll"^ L I B R A R I ES^ SMITHSONIAN^INSTITUTION ^NOlinillSNI^NVINOSHiiyNS^SB 1 1
--
_ m ^ —
\R I ES SMITHSONIAN-INSTITUTION ^O'-LniliSNI NVmOSHimS^Sa I BVa 8 H^LI B RAR I ES^SMITHSONIAN^INST LO -zr . (/> ^
. > 5 .>'55 z </> *Z — CO
iliSNI NVINOSHimS S3 I avaail LIBRARI ES^SMITHS0NIAN_INSTITUTI0N^N0liniliSNI_NVlN0SHll/JS^S3 I
AR I ES^SMlTHSONIAN~'lNSTlTUTION^ NOHnillSNI^NVINOSHillAlS ^S3 IdVaaiT^LIBRARI E SMITHSONIAN^ I NSl o Z x-S^T^oX O ro ^ ~ O
P3 >
„ S >-W" I .
NviN0SHiiws^S3iavaan''^'^^^^'^^ Smithsonian institution NoiiniiiSNi nvinoshiiws S3i
< i I X I Ik.. I I "
\R I ES^^SMITHSONIAN^INSTITUTION ^>01iniliSNI_NVIN0SHillMS S3 I d Vd 9 n_LI BRAR I ES SMITHS0NIAN_INS1
CO -• <0 CO —
z!
O ^ 2 ■'"' 3 Z _ O
iliSNl^NVINOSHlIWS^SStdVdaiYLIBRARIES^SMITHSONIAN^INSTITUTION^NOlinillSNI^NVINOSHilWS^SBI
CD
:x\ ^
SI >
^ m X ^ m m m
— /ok^ (/) — ^ ^
ARIES SMITHSONIAN'’iNSTITUTION NOliflillSNl NVINOSHimS S3 1 8 VB 8 I “1 L I B R A R I ES SMITHSONIAN INS ^ ^ ^ .< .v<-. S <
CO
z
11#^
llllSNI NVIN0SHillAIS^S3 I d Vd 8 LI B RAR 1 ES SMITHSONIAN INSTITUTION NOIinillSNl_NVINOSHillAiS S3
— CO =
n o _ xjOius^ o . -
AR I E S^ SMITHSONIAN"^ INSTITUTION ^NOIinillSNl“’NVINOSHillNS S3 IdVdSIl LIBRARI ES SMITHSONIAN^ INS z f~ z ^ Z Pi ^ ^ C . o
CO
o o — o X^'VOSV'V^
'LIBRARIES^SMITHS0NlANj]|lNSTlTUT10N^N0liniliSNl^NIVIN0SHill^S^S3 | a V B 9 H^L I B R AR I ES ^SMITHSONI,
r O O TO 9-
^ rn w m \ z cn
N0linillSNl“NIVlN0SHiiy'js‘^S3 I y va a n"Ll B RAR I Es|^SMITHSONlAN"lNSTITUTION^NOIinillSNI^IMVIN^ll
X
1^ LI B RAR I Es‘^SMlTHSONIAN^INSTlTUTION‘^NOIiniliSNI_NVINOSHims|^S3 I d V d 3 I I^L I B R AR I ES'^SMITHSONI
O _ -r- 5 ^ 5 r Xio^^ O
NI0liniliSNl''NVIN0SHimS^S3 I aVB a n^LI B RAR I ES^SMlTHSONlAN^lNSTlTUTION^NOliniliSNI^NVmOSHil
CD
'oX CD — yy^vf^oX CD /o'^iirtt,:.j^N ^ .xai'^^ yy^. /.^.aa i r !ekT^\ v- \ ^
>
XI
m X^vosviiZ rn -
, ~LI B RAR I Es‘^SMITHSONIAN“lNSTITUTION‘^NOIiniliSNrNVINOSHillMS S3 | d VH 9 H I B R AR 1 ES SMITHSON
X cn
I g |.* ' ' i n:^?' |- g >
J ‘^NOliniliSNI NVIN0SHims‘^S3 I avd a n^LI B RAR I ES SMITHSONIAN INSTITUTION NOIinilJ.SNl_ NVINOSHi
' ~-_</5 — </) “ ^ -r ^
[2 zSsrinT^v V uj ^ uj ^ ^;vrn:;>v u
O
n"" LIBRARIES SMITHSONIAN INSTITUTION NOliniliSNI NVINOSHillNS S3 I B Vd 9 IT L I B R A R I ES SMITHSO^
V z r- H r* z L — - o
o o Z o y^^^^ rn
PI PCX' ^ rn ~ xi^uiiv^^ m ™ m
NOlinillSNrNVlNOSHimS S3 I dVB 8 n~LI B R AR 1 ES SMITHS0N1AN“|NST1TUTI0N UOliniliSNl NVINOSHJ
< /ArifflrfflW'i'X y /t • _i /WcrtfauT-CV -3»- .V^^^vs- — i ./ A,y Z
O CD
T^ LI B RAR 1 ES‘^SMITHS0NIAN^ INSTITUTION'^NOliniliSNl NVIN0SHillAls‘^S3 I d V d 8 IT _ L I B R A R 1 ES SMITHSO^
)N NOIiniliSNI__NV, z ^
m 5
Ff S3 I avaa n“'LI BRAR l ES^SMITHSONIAN“^INSTlTUTION^NOIiniliSNI^NVlNOSHJ
I ^e. “ p S x«
33 \r.
in