S M I T H S O N I A N C O N T R I B U T I O N S T O B O T A N Y N U M B E R 1 6
Morden-Smithsonian Expedition
to Dominica:
The Lichens (Thelo trema t aceae)
Mason E. Hale, 3r.
SMITHSONIAN INSTITUTION PRESS
City of Washington
1974
ABSTRACT
Hale, Mason E., Jr. Morden-Smithsonian Expedition to Dominica: The Lichens
(Thelotremataceae). Smithsonian Contributions to Botany, number 16, 46
pages, 20 figures, 1974.-A revision is made of the lichen family Thelotremataceae
in Dominica, based on previously published records by Elliott and on collections
by the author. The family comprises 48 species in 4 genera, Leptotrema,
Ocellularia, Phaeotrema, and Thelotrema. The following 15 species are described
as new: Ocellularia antillensis, 0. conglomerata, 0. dominicana, 0. maculata,
0. mordenii, 0. nigropiinrta, 0. rimosa, 0. yorediata, Phaeotrema aggregatum,
P. obscurum, Thelotrema confusum, T. dominicanum, T. papillosum,
T. tenue, and Leptotrema deceptum. Three new combinations, Ocellularia fecunda
(Vainio) Hale, Phaeotrema disciforme (Leighton) Hale, and Leptotrema
occult um (Eschweiler) Hale, are also made. Morphological characters are
reviewed and the chemistry of each species is presented in detail. The family
is exceptionally well developed in mature rain forest.
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Leaf clearing from the katsura tree Cercidiphyllum juponicum Siebold and Zuccarini.
Library of Congress Cataloging in Publication Data ~~ Hale, Mason EMorden-
Smithsonian Expedition to Dominica: the lichens (Thelotremataceae).
(Smithsonian contributions to botany, no. 16)
Bibliography: p.
1. Thelotremataceae. 2. Lichens-Dominica, I. Title. 11. Series: Smithsonian Institution.
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Contents
Page
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Morphological Characters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Thallus Structure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Apothecial Characters . . . . . . . . . . . . . . . . . . . . . .
Spores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Generic Limits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chemistry . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ecology and Habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
The Thelotreme Flora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Collecting Localities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Ocellularia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
15
Phaeotrema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
28
Thelotrema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
31
Leptotrema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
38
Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Key to the Species of Ocellularia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Key to the Species of Phaeotrema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Key to the Species of Thelotrema
Key to the Species of Leptotrema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
iii

Morden- Smi t hsonian Expedition
to Dominica:
The Lichens (Thelotremataceae)
Mason E. Hale, Jr.
Introduction
The Thelotremataceae comprise a very large
family of crustose lichens that has attracted the
attention of many lichenologists, beginning with
Acharius (Hale, 1972). They have unique craterlike
apothecia, but this characteristic feature can
usually only be seen under magnification. As a
matter oi fact, these inconspicuous lichens will
oEten be overlooked in the field unless a hand
lens is used. Furthermore, because of the small
size of the apothecia, very few species can be recognized
at sight. All must be collected and examined
later for morphological and chemical
characters. Thallus color, however, is rather constant,
light greenish or ashy gray. Only a few species
have the deep green color of the equally common
pyrenocarpous lichens, which, together with the
Graphidaceae and Thelotremataceae, account for
the majority of crusts that are collected in lowland
tropical regions.
This paper is part of an on-going project on a
modern treatment of the lichen flora of Dominica.
As I mentioned in the first part on the Parmeliaceae
(Hale, 1971a), where climate and topography
are discussed, Dominica has the greatest remaining
areas of virgin or relatively undisturbed forest of
all the islands in the Lesser Antilles. Though
Mason E. Hale, Jr., Department of Botany, A'ntional Museum
of Natural Histoy, Smithsoninn Institution, Washington, D.C.
20560.
easily accessible by car and well-kept trails, these
forests have as yet suffered little from human activity.
Logging operations in the Dleau Gommier
Forest Reserve, for example, have ceased, at least
temporarily. Other islands in the Antilles are by
contrast much more intensively cultivated and undisturbed
forests are confined to higher, often
inaccessible mountain slopes.
On my first trip to Dominica in 1969 I was interested
primarily in foliose lichens, which proved
to be relatively poorly developed. Many of the
numerous crusts that I collected more or less randomly
and blindly were actually thelotremes. The
obvious richness of the family inspired me to undertake
a second trip to the island in December
197 1, once again under the Morden-Smithsonian
Expedition, to concentrate exclusively on this family.
A final brief trip for the same purpose was
made in May 1972 under National Geographic
Society support. Altogether, nearly 550 specimens
of thelotremes were collected, making this one of
the most intensive collecting programs ever undertaken
for a crustose lichen group.
Special thanks are due Mrs. William J. Morden,
who supported most of the field work for this
project. Assistance is also gratefully acknowledged
from the National Geographic Society for a broader
study OC the evolution of the Thelotremataceae in
the Lesser Antilles, which included Dominica.
Curators at a number of museums and universities
have sent loans of type-specimens, and their help
1
2 SiMITHSONIAN CONTRIBUTIONS TO BOTANY
and patience are deeply appreciated: Dr. T. Ahti
(H), Dr. Reino Alava (TUR), Dr. Ove Almborn
(LD), Dr. C. E. Bonner (G), Mr. hl. Skytte Christiansen
(C), Mr. Peter James (BM), Mme Jovet-
Ast (P), Dr. I. hl. Lamb (FH), Dr. R. A. Maas
Geesteranus (L), Prof. H. Merxmuller (M), Dr.
Harald Riedl (W), and Dr. R. Santesson (UPS).
I wish also to thank Dr. Chicita Culberson for
help in identitying or verifying most of the chemical
substances of the various specimens.
Morphological Characters
After curating and labeling each specimen, I
studied the various morphological characters that
appear to be most important for taxonomic purposes.
The descriptions, therefore, are not exhaustive
but should be adequate for species
identification. Redinger (1936 ) and Salisbury
(1972a, 1972b) have drawn up useful summaries
of the essential features of this family and a number
of the species which may also be consulted.
THALLUS STRUCTURE
The anatomy of crustose lichens is not as complex
as that found in typical foliose lichens. Cortical
structure is less distinct, and, lacking a free
lower surface, the medullary hyphae usually penetrate
the outer layers of bark periderm (or rock
substratum). The algal layer (Trentepohlia) may
also be indistinct. Large colorless crystalline inclusions
are commonly found in the medulla, as in
Ocellularia nigropuncta, new species, and Thelotrema
interpositzim.
Sections examined under a scanningelectron
microscope show some of these structures vividly.
Ocellularia olivacea, 0. perforata, and Thelotrema
confusum, new species (Figure la,b,c), all with
well-developed epiphloeodal thalli, have a cortex
consisting of several layers of heavily gelatinized
cells appearing almost paraplectenchymatous; the
loosely organized medulla penetrates the periderm.
The surface is essentially smooth and featureless
(Figure 2a-d). In some species the cortex is similar
but peculiar aculeate hyphae protrude from the
surface in greater or lesser abundance (Figure
3a,b). They appear to be collapsed, perhaps because
of vacuum treatment needed for specimen
preparation. This aculeate orientation, reported
here for the first time, is characteristic of Myriotrema
species (Ocellularia terebratula, Thelotrema
clandestinzim) as well as a few species with
emergent apothecia (T. depressum, T. praestans).
I have observed similar orientation in several
species of Graphis and Graphina. Most thelotremes,
however, have an irregularly organized surface, as
in 0. mordenii, new species (Figure 3c,d), and a
poorly defined cortex.
Leptotrema wightii has an unusual cortex previously
described by many lichenologists. The
thallus is very thick (up to 1 mm) and has a grainy
surface (Figure 5 4 . The cortex is arranged in tall
columns with algae scattered vertically (Salisbury,
1971) (Figure Id).
The so-called hypophloeodal species lack the
structures described above. The external thallus
is reduced to a microscopic layer of hyphae spread
thinly over the bark surface (Figure 4 ) , and these
hyphae penetrate several layers of periderm with
very sparce symbiotic algae apparently located just
below the periderm surface. Representatives of this
group from Dominica include Ocellularia concolor,
0. pyrenuloides, Phaeotrema disciforme, new
combination, Thelotrema leucomelaenum, and T.
tenue, new species.
APOTHECIAL CHARACTERS
Several morphologists have studied the internal
structure and probable ontogeny of ascocarps in
the Thelotremataceae (Redinger, 1936; Johnson
and Brown, 1941; Letrouit-Galinou, 1966). The most
commonly studied species is Thelotrema lepadinum,
which has a distinct exciple breaking away
free from the apothecial wall. The following DOminican
species have more or less similar structure:
Ocellularia conglomerata, new species, 0.
dominicana, new species, 0. exanthisrnocarpa (Figure
5a), Leptotrema occultum, L. subcompunctum,
and Phaeotrema aggregatunz, new species. I will
call this arrangement “lepadinioid” in the descriptions;
Salisbury (1972a) uses the term “Thelotrema
lepadinum group.”
What appears to be an extreme stage of this
configuration of apothecial wall and exciple is seen
in the Chroodiscus-type apothecium (here termed
“chroodiscoid”). The outer wall or margin is erect
and becomes recurved (Salisbury, 1972b), almost
as in Geaster, and the exciple usually persists as a
NUMBER 16 3
FIGURE 1 .-Cross-sections of epiphloeodal lichens viewed with a scanning-electron microscope
(a, b, and c, sectioned obliquely; d, Fertically): a, Ocellularia olivacea (Hale 35226) (X 1000);
6, 0. perforata (Hale 38145) (X 1000); c, Thelotrema cotzfusum, new species (Hale 37697)
(X 500); d, Leptotrema ulightii (Hioram 5850 from Cuba) (X 500).
second inner rim. The disc is widely exposed and foliicolous species of the Thelotremataceae, innothing
resembling the discrete pore characteristic cluding in Dominica Ocellularia alborosella and
of the other thelotremes remains (Figure 5b). This 0. clilatata. It is callcd the “Tlzelotrema platy.
type was first recognized among the foliicolous cui-pzim group” by Salisbury (1972b). In the ablichens,
but it seems identical to that in the non- sence of more detailed ontogenetic research,
4 SMITHSONIAN CONTRIBUTIONS TO BOTANY
FIGURE 2.-Surfaces of epiphloeodal lichens viewed with a scanningelectron microscope: a, b,
Ocellutaria nigropuncta, new species (Hale 35365) (X 500 and X 2000); c, d, 0. dominicana,
new species (Hale 37967) (X 500 and X 2000).
however, we cannot yet say that Chroodiscus is a
good genus, that the Chioodisczwlike nonfoliicolous
thelotreines should be transferred to Chroodiscus,
or that Clzioodiscus should be transferred to
Ocellulaiiu. Chroodiscoid species, incidentally, are
distributed among the four presently recognized
genera, Ocellzih in, Leptotrema, Phaeotieinu, and
Theloti emu, on the world level.
The majority of species in the Thelotremataceae
are neither lepadinioid or chroodiscoid. The exciple
is fused with the receptacle wall and the
main rim or margin closes over most of the disc
NUMBER 16 5
FIGURE 3.--Surfaces of epiphloeodal lichens viewed with a scanning-electron microscope: a, b,
Ocellularia olivacea (Hale 38054) (X 500 and X 2000); c, d, 0. mordenii, new species (Hale
35764) (X 500 and X 2000).
leaving only a small constricted pore. The upper ized apothecia and a pore flush with the thallus
wall is usually carbonized. There are, broadly surface (Figure 5c,d). The apothecia are always
speaking, three groups that can be recognized for quite small, 0.1-0.3 mm in diameter. Typical
practical purposes on the basis of degree of emer- examples in Dominica are Ocellularia olivacea and
gence from the substratum. One, represented by Thelotrema clandestinum.
the “Myriotrema” type, has immersed noncarbon- A second group, which includes the majority
6 SMITHSONIAN CONTRIBUTIONS TO BOTANY
FIGURE 4.-Surface and cross-sections of hypophloeodal lichens viewed with a scanning-electron
microscope: a, Thelotrema Zeucorneiaenum, surface view (Hale 39491) (X 2000); b, T. leucomelaenurn,
cross-section (Hale 35491) (X 500); c, d, T. tenue, new species (Hale 35430) (X 500
and X 2000).
of the species, has emergent apothecia, neither X third group, typified by 0cellular.ia rhobasally
constricted or strongly raised (Figure 5e,f). dostroma and 0. cavata, has strongly emergent apo-
They are usually 0.4-1.0 mm in diameter, although thecia Tvitli a partially constricted base (Figure
in a few species (for example, Thelotrema praestans 55). The pore is small and sometimes annulate or
in Figure 17e) nearly 2 mm in diameter. open at maturity, as in 0. fecunda (Figure llh,i).
NUMBER 16 7
FIGURE 5.-Types of apothecia in the Thelotremataceae (all X 10): a, Ocellularia exanthisniocarpa
(type of 0. multilocularis); b, 0. alborosella (Hale 353ila); c, 0. olivacea (Hale 35243);
d, Leptotrema wightii, (type of L. prevostianum); e, 0. perforata (Hale 37i27); f, L. bahianum
(type of Thelotrema rudius var. dominicanum); g, 0. rhodostroma (Hale 38122); h, Phaeotrema
leiostomum (Hale 37956); i, 0. coinparabilis (Hale 37372).
Diameter varies between 0.6 and 2.0 mm. This Although Kylander continued to use Ascidiunz
group was called Ascidium by Fee (1824), not so for strongly emergent species, no modern lichenmuch
because of emergence but because of a dupli- ologists have employed it. The type of the genus,
cate membrane in the perithecium (the nature of A. cinchonarum Fee, is synonymous with Ocelluwhich
is not clear to me) and a marginate pore. h i a cavata. Naturally there are intergradations
8 S.\fITHSOSIAN COXTRIBUTIOXS TO BOTANY
walls and periderm, acting a5 a kind of ramrod.
Not all species with periderm inclusions have a
columella, but as Redinger found for the Brazilian
species the correlation is high.
The range in deLelopment of a coluinella is very
great (Figure 6), from the Tveak thin columella in
Ocellzilal-ia mordenii, new species, to the multiple
act i n oi cl 0 ce 1/21 la ?.in co mpnra b i 1 is
(Figure 5i) and Leptotrema fisszim (Figure 19a,c).
The most highly developed species is Ocellzi Inria
glnziovii (Muller-A4rgau known from South America
Vainio (1921: 184) recognized these species under
a separate genus, Rlinbdodzsczis.
s t r uc t iires of
between the Ascidium-type and the more common
emergent type, but I believe the descriptive term
“ascidioid” is still useful.
A final group forms a transition to the Graphidaceae.
The apothecia become irregularly elongate
with a lirelliform dehiscence. In Dominica this
includes Phaeotrema leiostomum, which can produce
both round and elongate apothecia (Figure
511). Other species from South America have this
trait developed much more conspicuously and have
been described in the Graphidaceae (as Phaeographis
chionodisca Redinger (1936:69)). They
should probably be transferred to the Thelotremataceae.
Redinger (1936) further divides the Thelotremataceae
into two groups that differ in position
of the ascogonial initial. In one group the initial
is located on or just below the periderm surface
so that at maturity the apothecial wall develops
on and apart from the unbroken periderm (for
example, Ocellzilaiia rlzodostroma and Phaeotrema
disciforme, new combination). The other group,
including most nonchroodiscoid species in Dominica,
has ascogonial initials deeply embedded in
the periderm. The maturing apothecia must break
through this layer to reach the surface. Several
layers of periderm cells are thereby included in
the apothecial receptacle and can be easily recognized
in sections (Figure 6f). This character deserves
much more detailed study.
Development of a central cylindrical columella
is particularly characteristic of the Thelotremataceae,
and for the most part presence or absence
of a columella is a good species character (Figure
6). As a rule, columellate species do not have perfect
counterparts that lack a columella. Noteworthy
exceptions in Dominica are Oce11zilaria papillata
and 0. perfomto, each of which has a large population
where a columella is clearly developed and
a smaller one where it is either absent or very
difficult to find. I have not recognized such
ecolumellate species as new without correlating
morphological or chemical characters. Columellate
species always have heavily carbonized apothecial
walls.
Redinger (1936:6) proposed an interesting hypothesis
that the columella plays a role in breaking
open the periderm layer in species that develop
deeply embedded. As the apothecia mature, the
columella pushes up against the upper carbonized
SPORES
Spore size is an importar,t and relatively constant
character. The range of spore length, for
example, is usually quite small except for very
large spores (more than 10Op) which may vary
between 1 0 0 ~ and 300u. ’IVe can distinguish four
very broad categories among the Dominican species:
(1) spores less than 2 0 ~ long with 4-6 transverse
locules, eg., Lcplot~enza wightii, Ocelliilnria terebi.
atiila, Tliclotwnza clandestinum; (2) spores
20y-36;t long with 6-10 transverse locules, e.g.,
Ocellzilarin perfoycita (Figure 7a), Thelotrema
leiiconielaeizi(ln; (3) spores 40p-9Op long with
12-24 transverse locules, e.g., Leptotyema decepturn,
new species (Figure 5d), Ocellzihria exanthismocarpa;
and (4) spores 80p-300p long with
numerous locules, eg., Ocelliilnria rhodostyoma
(Figure f b ) , Thelotrema praestans, T. tiiberculif
e I’ ii m.
The number of longitudinal septations in
Leploti~ema and Theloti.ema varies according to
length, from 1 or 2 septae in small spores (up to
about 30p long) to 4-6 or more in large spores.
Spores are colorless or brown. It is sometimes
a problem to determine color in large-spored
species of Leptotrema and Thelotyema where, on
the one hand, a Thelotwma species might have
some senile spores turning brown and on the other
a Leptotrema will haye some colorless spores (Figure
7 d ) . Jn Phaeotwma disciforme, new combination,
in fact, one often finds a mixture of colorless,
mature brown, and shriveled brown spores
(Figure 7c). Shriveling (Figures 7c,4 is characteristic
of brown-spored species; I know of no cases
where Thelotrema or Ocellularia spores shrivel.
NUMBER 16
FIGURE 6.-Cross-section of apothecia (all about X 130): a, Ocellularia nigropuncta, new
species (Hale 35365); b, 0. inordenii, new species (Hale 37764); c, 0. cavata (Hale 35134); d,
Phaeotrerp leiostoniun (Hale 35203); e, Thelotrema leuconlelaenuin (Hale 35491); j, T. tenrte,
new species (Hale 35430).
9
10 SMITHSONIAN CONTRIBUTIONS TO BOTANY
FIGURE 7.--Spores of the Thelotremataceae: a, Ocellularia perfornta (Hale 37784) (X 400):
b, 0. rhodostroma (Hale 35440) (X 130): c, Phaeotrema discifortne, new combination, with a
mixtute of colorless spores, brown spores, and shii>eled brown spores (Hale 37939) (X 400);
d, Leptotrenia deceptutn, new species, with colorless muriform spores on the right (Hale 38110)
(X 400).
In any event, transitional species should be judged nation, or 0. sitbcaunla on Dominica) should not
intliviclually, and the presence of a few anomolous be totally unexpected; nor shoultl these isolated
brown-spored specimens among many colorless cases prejudice the value of spore color as a generic
specimens (as in Ocellzrlnria feczinda, new conibi- character.
NUMBER 16
Generic Limits
The generic delimitation of the Thelotremataceae
has undergone many changes since the
classic work of Acharius. Fee (1824), for example,
established two genera, in addition to recognizing
Theloti enla, based on external features and apothecial
characters: Ascidizim and My? iotrema, both
presently considered to be synonyms of Ocellzlla,
fa. lfullei--Argau (1887a) proposed the most farleaching
changes, using spore septation and
coloration to separate four genera (in addition
to foliicolous Chq oodisctis): Ocellzilaria (transtrersel
y septate colorless spores), Phaeotrema
(transFersely septate brown spores), Thelotrema
(muriform colorless spores), and L e p t o t i emu (muriform
brown spores).
TVhile Vainio, one of the truly creative thinkers
in lichen classification, rejected many “spore
genera,” almost all authors since Muller-Argau
ha1 e accepted these genera (and, indeed, sporebased
genera in other families). Poelt (1973:613)
refers to these as “artificial genera,” yet almost
an) character we select is in a sense artificial. I
don’t believe this term does justice in that it
implies spore characters are artificial.
TVe are, in reality, faced with at least thiee
choices for generic delimitation in the family: (1)
one genus (Tlielotremcr ) to accommodate all thelotreinataceous
species; (2) four spore-based genera
(Ocelliilnria, Phaeolrema, Thelotrema, and Leptot)
en70); (3) five or more genera ( A s c i d i ~ m ,
C 11 rood isc 11 Y, M y i o t I em n, R h 0 b dod i ~c us, antl T 11 e lotren10)
based on apothecial emergence and excipular
structures. Most lichenologists, as I said, have
opted for spore-based genera. TVhile a majority
opinion does not necessarily prove the naturalness
or validity of such genera, they do repiesent one
possible biological differentiation, broadly correlated
with chemical, morphological, and ecological
characters (as I will discuss briefly under each
genus).
I cannot agree with Salisbury’s (1952a) recent
rejection of spore-based genera in favor of a single
massi\e genus Tl7elotrema with 400 species subdivided
into “groups” according to excipular structure,
a character that is difficult to identify. His
kejs and descriptions show how cumbersome this
approach is, for in each instance one must desig-
11
nate the spore type, e.g., TIzelotiemu wightii with
Leptot)ema-type spoies, etc. If- one is forced to
make this critical distinction every time a species
is identified and described, then they should be
tieatetl as distinct genera, a more coherent antl
certainly a inore pragmatic, working solution.
Spore characters are well proven, even critical, in
the taxonomy of noiilichenized fungi. A retrogressive
gene1 ic concept in lichens will hardly encourage
a better understanding of their biology and
evolution. M j experience in the Thelotremataceae
suggests that spore-based genera are reasonably
natural, easily manageable units that are probably
diverging in terms of habitat requirements and
biogenetic characters as reflected in chemical differences.
Chemistry
in aliiiost eier) other gioup of lichens studied,
chemistr) is an impoi tant and fundamental character
in uncle1 standing the evolution and population
biolog) ol the Thelotremataceae. The family
has an unexpectedly iich chemistry with a pretloniinance
of P + depsidones and related compounds
(Culberson and Culberson, 1968; Culberson
and Hale, 1973). I Trill discuss the possible
significance of this chemistry below under the
species and hope to exploi e the broader aspects
oh biochemical evolution of the family in another
paper.
-411 of the specimens were initially tested with
p-phen) lenetliaiiiine and chromatographed on
Brinkinan thin-1a)er plates according to the P reaction.
I ~roultl recommend, however, that lichenologists
consider using this reagent as little as
possible because of the well-known carcinogenic
propel ties of rolatile amines. Identification of the
lichen substances can be made just as easily with
TLC rrithout recouise to a P test except in doubtful
cases.
Standard thin-layer solvent systems (hexaneether-
formic acid, 130:80:20 v/v, and benzenedioxane-
acetic acid, 180:45:5, I./.) xvere used to
identify the chemical content of each specimen.
NOTATIC A c r ~ . - O c e l l z r l a r i a feciinda, new combination,
(hypoprotocetraric acid predominating), 0.
rhodost 1 om0 (4-O-demethylnotatic acid predominating),
antl Pliaeotremu leiostomzim (4-O-deme-
HYPOPROTOCETRARIC ACID ASD 4-o-DEhIETHYL12
SXITHSONIAN COSTRIBUTIONS TO BOTANY
and Thelotrema dominicanuna, new species. Several
species of Ocellularia and this single
Thelotrema react P t red, but neither protocetraric
acid or fumarprotocetraric acid can be demonstrated
with TLC. Instead there are at least t\vo
gray spots above both of these acids on the plates
(and often a high H,SO, yellow spot). These are
apparently derivatives of protocetraric acid.
“PRAESTASS” UsKsowN.-Thelotrema praestans.
This unidentified depsidone reacts P+ red orange
and forms a very distinct orange to reddish brmvn
spot below psoromic acid in both solvents.
“OLIVXCEA” USKNO~t.l\.S.-OCellllla~~a olivacea.
This P- compound produces a distinct H2S04
gray spot below lecanoric acid in both solvents.
It may be a depside. TVhile the lectotype of Ocellzilnria
olivacea contains only this one spot, all West
Indian collections, otherwise morphologically indistinguishable,
contain a lower gray spot in addition.
All specimens have thick white acetone
residues.
OTHER P - UsK~o~t.n.s.-Ocellziluria rimosa, new
species, and 0. sorediatn, new species. The unidentified
spots in these species may or may not represent
lichen substances. Some species were examined
for fatty acids by dipping the plates in water and
trying to detect white water-repellant spots as the
water evaporated. Some spots resembling fatty acids
Tvere seen but microchemical tests with G.E. and
G.A.TV. xvere unsuccessful. Another major group
of P- compounds is terpenes. All TLC plates
shou.ed numerous terpenes, identifiable as purple,
blue, or brovm spots falling generally above RF
.5 and fading Tvithin a few hours. Probably all of
these originate from the bark and are not lichen
products, as can be proved by co-chroinatographing
the lichens Jvith lichen-free bark samples from the
same specimen,
0. concolor, 0. dominicnnn, new species, 0. macu-
Into, new species, 0. mordenii, new species, 0.
pnpillata, Phaeotiwnn aggregntum, new species, P.
d iscif o i m e, new com bi na t i on, T 17 e 1 o t re m a d ep resslim,
T. leticomelnenum, T. papillosum, new
species, T. tenue, new species, and T. tuberculiferum.
PicsrENTs.-OcellziInria rnordenii, new species
and Leptotrema wightii (anthraquinones) and 0.
fecunda, new combination, 0. rhodostroma, and
Phaeotremn leiostomum (K- yellow compounds).
h’0 SUBSTASCES PRESENT.-OCel/Zilnria albOYoSelk7,
thylnotatic acid sometimes lacking). These acids
were recently examined in detail by Culberson
(Culberson and Hale, 1973), who discusses the
biogenesis. Ocelluiarza fecunda and 0. rhodostroma
were not included in that study.
NORSTICTIC AcIn.-Leptotyenaa occultum. This is
a rare acid in the Thelotremataceae (especially in
contrast to the Graphidaceae) and is largely confined
to Leptotrema on the world level.
PROTOCETRARIC AcIn.-Ocellularia nigropuncta,
new species, 0. perforata, Leptotl-emu baliianum,
and Theloti~ema confusum. This well-known P +
red acid is easily identified on TLC plates, It is
sometimes accompanied by a higher gray spot that
I have termed the “amplior” unknown because of
its constant occurrence in Ocellula~ia amplior (Nylander)
Redinger. Culberson believes it to be protocetraric
acid combined with a carboxylic acid,
comparable to funiarprotocetraric acid or the recently
described succinprotocetraric acid.
PsoRoarIc AcID.-Ocellulaiia antillensis, new
species, 0. conipaq.abilis, 0. subcavata, 0. terebratula,
Leptotrema fissum, Thelotrema carassense, T.
clandestinum, T. conforme, T. leucinum, and T.
interpositurn. This is the commonest P+ acid in
the Thelotremataceae. It is almost alivays accompanied
by an unknown compound “conpsoromic
acid,” appearing as a spot below psoromic acid.
It, too, is P+ yellow.
STICTIC AND CONSTICTIC Acins.-Ocellzilnria conglomerata,
new species, 0. dilatata, 0. exanthismocarpa,
0. pyveniiloides, Phneotrema obscurzim, new
species, Tlielotwma glaucopallens, Leptotrema dcceptum,
new species, L. microglaenoides, and L.
szibcomptinctiim. This widespread and well-known
acid is actually rather rare in the Thelotremataceae
except for the genus Leptotrema. Constictic acid,
appearing as a H,SO,+ yellow spot just below
salazinic acid, is a constant accessory compound.
One or two other gray spots, as yet unidentified,
may occur between stictic and constictic acids on
TLC plates. Another rarer accessory is the highest
H,SO, t orange-red spot in the “quintaria” series
(Hale, 19Tlb), still unidentified but being discovered
in a variety of genera.
VIRENSIC Acm.-Leptotrema spondaicum. This
acid is P S ‘red. It was previously unknown in the
Thelotrcmataceae, having been first identified in
Alectoria.
“CINCHONARUM” UNK~own.s.-Ocellzilaria cavata
NUMBER 16 13
zone lvhere rain I‘orest is optimally developed, the
trees being 30-40 m high (Figures 2 and 3 in
Hale, 19Sla, illustrate this forest type). It would
appear that this lichen family is totally dependent
on undisturbed rain forest as a primary habitat
and site for speciation. When this forest is destroyed,
it seems inevitable that the lichens associated
with it will become extinct. In Dominica,
I have seen only two species able to invade disturbed
or planted forests; namely, Leptoti.ema
spondaicum and Ocellularia exanthismocarpa.
They have a highly successful means of propagation
and become established very quickly.
The anthraquinones are generally produced in the
thallus, while the yellow compounds occur in the
medulla of the apothecia and not in the thallus.
Ecology and Habitats
I paid considerable attention to the ecology of
the species and habitat requirements in Dominica
in the hope that this information would help in
understanding the distribution and evolution of
the species on the island. Lichenologists have all
too often prepared revisions of tropical lichens on
the basis of preserved herbarium specimens alone
with little if any opportunity to observe the species
in the field. Labels on specimens collected by nonprofessionals
rarely give pertinent information on
ecology and habitat. For example, tropical rain
forests have distinct layering and great vertical
variation. in microclimate. Elevational differences
are also important and once again such data are
rarely given in full on herbarium labels.
Base level species in Dominica form a distinctive
group. They grow on roots, lianas, buttresses,
prop roots, and saplings within 2-3 m of the
forest floor. Characteristic species are Ocellularia
concolor, 0. papillata, 0. perforata, Thelotrema
glaucopallens, T. tenue, new species, and Leptotrema
deceptum, new species. At the opposite extreme
we find canopy species that occur high above
the forest floor in tree tops, exposed to great
changes in insolation and wetting-drying cycles.
These include Ocellularia mordenii, new species,
0. olivacea, 0. subcavata, 0. terebratula, Phaeotrema
leiostomum, Thelotrema clandestinum, and
T. praestans. Other species on Dominica have less
stringent habitat requirements or have been collected
too infrequently to categorize.
On a broader scale altitudinal differences are
also pronounced. The rain forest becomes stunted
in Dominica above about 800 m and gradually
merges into an elfin mossy forest best developed
above 1000 m. Thelotremes become scarce in this
wind-swept zone where rain falls almost every day.
Two species, however, Thelotrema leucomelaenurn
and T. tubewuliferzim, are restricted to the highest
elevations. Ocellularia rhodostroma and Thelotrema
tenue, new species, occur here as well but
also at lower elevation.
The vast bulk of thelotreme collections were
made between 300 and 800 m on Dominica, the
THE THELOTREME FLORA
As I had found with Paimelia. (Hale, 1971a),
almost all previous collections of the Thelotremataceae
on Dominica were made by 1%‘. R. Elliott
in the 1890s. H. A. Imshaug also made about ten
collections in 1963 which are preserved at Michigan
State University. I have not had access to these.
Elliott collected five specimens, identified by Vainio
(1896, 1915) as follows:
Thelotiema excavatum Vainio var. impressulum
Vainio (hlorne Anglais, the type-locality, Elliott
160, specimens in Bill and TUR). This is synonymous
with Ocellularia perforata.
Thelotiema leucomelaenum var. elevatum Vainio
(Morne Anglais, Elliott 1534, specimen in TUR).
This is T. leucomelaenum.
Theloti emu rhodostromzim (hlontagne) Vainio
(Morne Anglais, Elliott 521, specimens in BM and
TUR, and Prince Rupert, Elliott 1305, specimen
in TUR). These are Ocellularia rhodostroma.
Thelotiema rudius Vainio var. dominicanum
Vainio (Prince Rupert, Elliott 1303, specimen in
TUR). This is the same as Leptotrema bahianum.
The thelotreme flora of Dominica now includes
48 positively identifiable species (and three additional
species represented by poor material). I collected
all of these except for Leptotiema bahianum.
No one locality, of course, had anywhere near this
number of species (see p. 14, “Collecting Localities”).
The richest area was one site (16b) in the
Dleau Gommier Forest Reserve which was being
logged in December 1951; I collected a total of
19 species there over an area of about 3 hectares.
An adjacent site had 14 species as did the trail
to Morne Anglais. Dleau Gommier locality 15 had
14 SMITHSONIAN CONTRIBUTIONS TO BOTANY
12 different species. A logged rain forest on the
Layou Road (12) had 11 species. Five other localities
(11, 16c, 18a, 21, and 24a), all between 350
and 750 m, had 9 species each.
The homogeneity of the flora is remarkably high.
Locality 16, for example, contains three adjacent
collecting sites. Out of a total flora of 21 species
there, 8 were found in common at each site. Three
occurred at two of the three sites and 10 were
collected at only one of the three sites. The greatest
variance in homogeneity obviously comes about
when we collect at two very different elevations or
sites where habitat conditions are significantly
different.
It should be evident that we have not yet inventoried
the whole thelotreme flora of Dominica.
I would estimate that there are at least 55 species
(including the three unidentified collections as
possible additions). It would require at least one
more intensive collecting trip, possibly two, to
reach this number.
COLLECTING LOCALITIES
Specimens were collected at 28 separate localities
on the island (Figure 8), and multiple visits were
made at different times to six of these, giving a
total of 35 collecting sites. These are coded as
follows for convenience in citing specimens, going
from localities at low elevation in the scrub forest
to those at highest elevations.
1. Scrub forest, Rodneys Rock, sea level.
2. Scrub forest near Mero, elevation 15 m.
3. Scrub forest on slopes of Barbers Block, ele-
4. Scrub forest near Calibishie, elevation 15 m.
5. Palm forest at Rosalie Bay, elevation sea level.
6. Rain forest along trail to Madjini, elevation
7. Virgin rain forest logging area, Newfound-
8. Rain forest above Newfoundland, elevation
9. Rain forest logging area, Castle Bruce Road,
10. Dleau Gommier rain forest near Emerald
11. Felled rain forest along new road cut about
vation 60 m.
30 m.
land, elevation 250 m.
300 m.
elevation 300 m.
Pool, elevation 450 m.
2 km northwest of Pont Casse., elevation
430 m.
12. Rain forest logging area on Layou Road, elevation
420 m.
13. Mixed secondary forest logging area at Brantridge
Estate, elevation 500 m.
14. Rain forest logging area just east of Pont
Casse., elevation about 550 m.
15. Virgin rain forest area at Dleau Gommier Forest
Reserve, elevation 500 m.
16a, 16b, and 16c. Virgin rain forest logging at
Dleau Gommier Forest Reserve, elevation
350-370 m.
17. Remnants of rain forest at Felicite, elevation
370 m.
18a and 18b. Remnants of rain forest along road
to Jean Estate, elevation 600-700 m.
19. Citrus grove at Laudat, elevation 400 m.
c 172128
L
Scotts Head 4+-./0-5
FIGURE 8.-Outline map of Dominica showing localities where
specimens of Thelotremataceae were collected.
NUMBER 16
20a and 20b. Remnants of rain forest in pastures
above Giraudel, elevation 550-600 m.
21. Virgin rain forest at Syndicate Estate, elevation
650-700 b.
22. Mixed rain forest along trail through Middleham
Estate, elevation 650-700 m.
23. Rain forest at the base of Trois Pitons, elevation
650-700 m.
24a and 24b. Mixed secondary forest along trail
to Boiling Lake, elevation 650-700 m.
25, and 25b. Rain forest-mossy forest along trails
at Fresh Water Lake, elevation 800-900 in.
26a and 26b. Rain forest-mossy forest along trail
up Morne Anglais, elevation 800-1000 m.
27. Stunted rain forest-mossy forest on slopes of
Trois Pitons, elevation 900-1200 m.
28. Mossy forest area on slopes of Morne Diablotin,
elevation 1000-1200 m.
Sixteen localities were visited on the first trip
in January 1969 (1, 2, 5, 6, 7, 13, 14, 15, 17, 18b,
19,'20b, 24b, 25b, 26b, and 28). On the second trip
seven additional localities were found (3, 11, 12,
16a-c, 22, 23, and 27) and five sites adjacent to
localities previously were examined (18a, 20a, 24a,
25a, 26a), all during December 1971. Four new
localities were visited in May 1972 (8, 9, 10, 21).
All specimens are deposited in US unless otherwise
indicated and duplicates will be deposited
in the British Museum (BM).
15
I have provided illustrations of the Dominican
species as well as of most type-specimens for comparison.
These photographs also serve to show the
range of variation in pore size and apothecial
emergence. I have found these X 10 photographs
to be extremely useful in identification, more so
than line drawings of cross-sections. All specimens,
including the types listed, were tested with thinlayer
chromatography.
Ocellularia
This is the largest genus in the family, comprising
about 165 species on the world level or about.
40 percent of the total 400 known species. I had
almost the same percentage on Dominica (22 of
48 species) but it makes up a disproportionately
large percentage of the specimens actually collected,
410 out of 530 (77y0). Ecologically the
genus is best adapted to mature rain forest and
cannot withstand extremes of very dry lowland
and wet exposed high elevation. The ten commonest
species (Tvith number of specimens collected in
parentheses) are: 0. yhodostroma (93), 0. fecunda,
new combination (64), 0. perforata (55), 0. pupillnta
(48), 0. siibcavata (40), 0. exanthismocarpa
(25), 0. nigropzincta, new species (16), 0. olivacea
(12), 0. mordenii, new species (lo), and 0. terebmtziZa
(10). Rarities, known from single collections
each, are 0. antillensis and 0. cavata.
Key to the Species of Ocellularia
1. Medulla of thallus and/or ascocarps pigmented.
2.
2. Medulla of ascocarps yellow or pink but thallus medulla white.
3.
3.
1. Medulla of thallus and ascocarps white.
4. Apothecia lacking a central columella.
5. Apothecia immersed, 0.2-0.3 mm in diameter.
Medulla and ascocarps entirely deep red . . . . . . . . . . . . . . . . . . . . . . . . . . 12. 0. mordenii, new species
Pigment yellow; pore large, 0.3 mm wide . . . . . . . . . . . . . . . . 10. 0. fecunda, new combination
Pigment deep pink; pore tiny, 0.1 mm wide, annulate. . . . . . . . . . 18. 0. rhodostroma
6. Thallus Pf yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22. 0. terebratula
6. Thallus P- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14. 0. olivucea
5. Apothecia emergent, 0.6-2.0 mm in diameter.
7. Disc open with a strongly recurved exciple.
8.
8.
7. Disc closed, lacking an outer recurved exciple.
9. Inner lepadinoid exciple strongly developed.
10. Apothecia aggregated in groups.
Exciple thick, disc irregularly elongate; stictic acid present
Exciple thin and delicate, disc round; stictic acid absent . . . . . . . . . .
. . . . . . 7. 0. dilatata
1. 0. alboroselln
11. Apothecial groups round; stictic acid lacking ......................................
.......................................................................... 8. 0. dominicam, new species
16 SMITHSOXIAN COSTRIBUTIONS TO BOTANY
11. Apothecial groups oblong: stictic acid present
................................................................. 6. 0. conglomerata, new species
12. Spores large, 55p-75p .................................... 9. 0. exanthismocarpa
12. Spores small, 15p ................................ 6. 0. conglomerata, new species
10. Apothecia solitary.
9. Inner exciple not developed.
13. Thallus P-.
14. Thallus distinctly fissured: pore area black rimmed ............................
14. Thallus continuous; pores not black rimmed . . . . . . . . . . . . . . 15. 0. papillata
15. Pore area black rimmed . . . . . . . . . . . . . . . . . . 13. 0. nigropuncta, new species
15. Pore area not black rimmed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. 0. perforata
4. Apothecia with a well-developed black central columella.
16. Thallus with large scattered verrucose outgrowths becoming sorediate ....................
20. 0. sorediata, new species
16. Thallus lacking soredia.
............................................................................. 19. 0. rimosa, new species
13. Thallus P+ red.
........................................................................................
15. Apothecia large, 0.5-1.1 mm in diameter, more or less emergent.
18. Thallus dull tan, hypophloeodal, P- . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. 0. concolor
18. Thallus distinctly epiphloeodal, P+.
19. Columella becoming actinoid: thallus P+ yellow . . . . 4. 0. comparabilis
19. Columella remaining simple: thallus P+ red . . . . . . . . . . . . . . . . . 3. 0. cavata
17. Apothecia smaller, 0.2-0.5 mm in diameter, barely emergent to immersed.
20. Pore annulate: spores small with crowded septae, 5p-6p X 9p-llp ................
21. 0. subcavata
20. Pore not annulate: spores generally 20p-30p long (except smaller in 0.
...............................................................................................
maculata).
21. Thallus P-.
22. Pore tiny, less than 0.1 mm in diameter; spores 4-loculate . . . . . . . . . . . . . .
22. Pore larger, 0.1-0.2 mm wide; spores 6-7-loculate . 15. 0. papillata
21. Thallus P+ red or orange yellow.
23. Thallus very thin to hypophloeodal; P t orange . . . . . . . . . . . . . . . . . .
.......................................................... 11. 0. maculata, new species
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17. 0. pyrenuloides
23. Thallus thick, epiphloeodal.
24. Thallus P+ red: apothecia barely emergent .......................
24.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16. 0. perforata
Thallus P+ yellow; apothecia strongly emergent . . . . . . . . . . . . . . .
..... 2. 0. antillensis, nev species
1. Ocellularia alborosella
FIGURES 5b, 96
Ocellu la ria alborosella (N ylander) San tesson, 1952: 308.
Grapliis alborosella Sjlander, 18633352 [tj pe-collection: Colombia,
L h f d i g 2ti94 (H, lectotypc: FH-Tuck, UPS, isotypes);
Figure 9b].
Thelotre!tia p l a t y c n r p l l w t ! L'ainio, 1923: 138 [typc-collection:
Arinia, Trinidad, TIiaxter 57 (TUR, lectotype); Figure gal.
Ocellularia platjcarpella (Vaino) Zahlbruckner, 1923:598.
Thallus whitish gray, thin but continuous, shiny,
smooth, forming colonies 5-8 cm broad; apothecia
chroocliscoid, up to 1.5 min across, round to irregular,
the marginal flap conspicuous, the disc flesh
colored to white pruinose; hymenium about 60p
high; spores 8, 5-6 loculate, 4p-5~ X 8p-15p, I-
01 weakly blue (Figure 5b).
C ~ ~ s r ~ s i RY.--No substances present.
HAnriAr.-Tree trunks at mid or higher elevation
in rain forest.
This was the first corticolous chroodiscoid
species to be described. It is characterized by the
small spores and lack of lichen substances. The
Dominican collections have a wider disc than the
type froin Colombia but they fall within a predictable
range of variation. It is a rare and easily
overlooked species.
SPECIMENS EXARIINED.-~ (38143), 16b (37883),
2613 (3537la).
NUMBER 16
2. Ocellularia antillensis Hale, new species
FIGURE 9g
Thallus corticola, albidus, epiphloeodes, planus
vel minute verrucosus, nitidus, 3-5 cm latus; apothecia
numerosa, emergentia, 0.3-0.5 mm diametro,
apice obfusca, columellata, columella 0.15 mm lata;
ostiolum 0.1-0.12 mm diametro, intus nigrum;
hymenium 7Op-801.1 altum; sporae 8: nae, transversim
7 loculatae, 7p X 20p, I+ coerulescentes
(Figure 9g).
CHEMrsTRY.-Psoromic and conpsoromic acids.
HoLOTwE.-Trail to Madjini, low elevation
windward rain forest, Dominica, elevation about
30 m, Hale 35612, January 1969 (US).
HABITAT.-upper trunk of stunted rain forest
trees (30m).
The whitish minutely verrucose thallus and
numerous emergent apothecia with the large contrasting
black pore area distinguish this rare species.
Ocellularia comparabilis var. microcarpa Redinger
has larger spores (6p X 30p-34y), sparse apothecia,
and a smooth thallus, while 0. terebrata (Acharius)
Muller-Argau has barely emergent apothecia (see
Hale, 1972). I later collected this species from
Trinidad and Panama, both at low elevation.
Grande, Trinidad, Hale 37444 (US); Barro Colorado
Island, Panama, Hale 38659 (US).
SPECIMENS EXAMINED.-3 miles NW Of Sangre
17
3. Ocellularia cavata
FIGURES 6c, 9f
Ocellularia cavata (Acharius) Muller-Argau, 1882133499.
Thelotrema cabaturn Acharius, 1812:92 [type-collection: Sierra
Leone, Afzelius (H, lectotype; S, UPS, isotypes)].-Hale,
1971b, fig. la.
Ascidium cinchonarum Fee, 1824:96 [type-collection: On
Cinchona, South America (G, lectotype; H, L, M, P, isotypes);
Figure gel.
Ocellularia cinchonarum (Fee) Sprengel, 1827:242.
Ascidium cinchonarum f. intermedium Nylander, I867:319
[type-collection: Pie de Cuesta, Colombia, Lindig 5 (H,
lectotype; BM, G, M, isotypes); Figure 9c].
Ocellularia lindigiana Muller-Argau, 1887a:9 [type-collection:
Colombia, Lindig 2757 (G, lectotype; BM, FH, UPS, isotypes);
Figure 94.
Oeellularia cinchonarym (Fee) Sprengel f. intermedia (Nylander)
Zahlbruckner, 1923:586.
Thallus whitish mineral gray, epiphloeodal, continuous,
more or less roughened, forming colonies
up to 8 cm across; apothecia ascidioid, 0.7-0.9 mm
in diameter, apically carbonized with a large
columella (Figure 6c); pore round, 0.07-0.12 mm
in diameter, the top of the columella clearly visible;
spores 8, 6-8 loculate, 6p-8p X 22p-28p, IS blue.
(Figure 9f).
CmMIsTRY.-"Cinchonarum" unknowns A and
B and a yellowish pigment.
HABITAT.-canopy branch in mid-elevation rain
forest (about 550 m).
I had previously typified this distinctive but rare
Acharian species. It appears to be rare in the
Lesser Antilles. The chief diagnostic features are
the unusual chemistry, small ascidioid apothecia,
and the large columella which protrudes into the
pore area.
SPECIMEN EXAMINED.-14 (35134).
4. Ocellularia comparabilis
FIGURES 5i, 9i
Ocellularia comparabilis (Krempelhuber) Muller-Argau, 1883:
318.
Thelotrema comparnbile Krempelhuber, 1876:220 [typecollection:
Rio de Janeiro, Brazil, Glaziou 5463 (M, lectotype;
BM, C, G, M, P, UPS, W, isotypes); Figure 9i].
Thallus pale greenish to whitish mineral gray,
epiphloeodal, smooth, forming colonies 5-1 2 cm
broad; apothecia emergent, 0.8-1.2 mm in diameter,
the walls carbonized, columella simple 'to actinoid;
pore open, 0.2-0.5 mm broad, the white pruinose
disc clearly visible; hymenium 50y-60&i high; spores
8, 4-5 loculate, 5p-6y X lOu--14~i, IS blue. (Figure
5i).
CHEivIsTRY.-Psoromic and conpsoromic acids.
HABITAT.-LOWer trunks in rain forest (300-
700 m).
The columella in this rare species is imperfectly
actinoid. To judge from collections made in South
America, the range of variation is very great with
gradations into related psoromic acid-containing
species. For example, Ocellulnria discoidea (Acharim)
Muller-Argau (Hale, 1972) and 0. antillensis,
new species, have a simple columella. Ocellzilaria
efformata (Krempelhuber) Muller-Argau has a very
similar actinoid columella but the outer margin
i s semi-erect, leaving an open rather lacerate pore:
0. De?.keIeynna (Montagne) Zahlbruckner and 0.
glnziovii Muller-Argau both have highly developed
18 SMITHSONIAN CONTRIBUTIONS TO BOTANY
FIGURE 9.-Specimens of Thelotremataceae (all about X 10): a, Ocellularia platycarpella (lectotype
in TUR); b, 0. alborosella, (isotype in FH-Tuck); c, 0. cinchonarum, f. intermedia (Iectotype
in H); d, 0. lindigiana (lectotype in G); e, 0. cinchonarum (lectotype in G); f, 0. cavata
(Hale 35134); g, 0. antillensis, new species (Hale 35612); h, 0. conglomerata, new species (Hale
37959); i, 0. comparabilis (isotype in BM); i, 0. concolor (lectotype in G); k, 0. concolor (Hale
38167).
NUMBER 16
actinoid discs; and 0. latilabia (Tuckerman)
Muller-Argau has an actinoid columella and a
recurved margin.
SPECIMENS EXAMINED.-~ (38128), 21 (37372,
38144).
19
5. Ocellularia concolor
FIGURE 9j,k
Ocellularia concolor Meyen and Flotow, 1843:230.
TYPE-CoLLEcTIoN.-;\fanila, Philippines, Meyen
(G, lectotype) (Figure 9j).
Thallus light grayish tan, dull, mostly hypophloeodal,
forming colonies 3-4 cm broad; apothecia
numerous, 0.8-1.1 mm in diameter, more or less
emergent, apically carbonized, columella present,
0.2-0.3 mm wide; pore open, round to somewhat
irregular, 0.3-0.5 mm in diameter, the rim black
and the flat black columnar disc easily seen;
liymenium 14Op high; spores 8, 5-7 loculate, 5p-
8p X 13p-20p, I+ blue (Figure 9k).
CHEI\IISTRY.--NO substances present.
HABITAT.-Lianas, saplings, base and lower trunk
of trees in dense rain forest (300-370 m).
This easily recognizable species was first described
from the Philippines and appears to be
pantropical. The tannish hypophloeodal thallus
and habitat are characteristic. There are no comparable
species in this kind of habitat.
16b (37801, 37809, 38075).
SPFCILIEKS EXAMINED.-8 (38167 ), 16a (37665),
6. Ocellularia conglomerata Hale, new species
FIGURE 9h
Tliallus corticola, cinereo-albidus, epiphloeodes,
continuus, nitidus, 4-5 cm latus; apothecia primo
solitaria, 0.6-0.9 mm diametro, semi-emergentia,
mox 2-3: nae conglomerata, margine exteriore
erecto, excipulo interiore evoluto, pulverulento,
disco aperto, carneo; ostiolum irregulare, latum;
11) menium 70p altum; sporae 8: nae, transversim
6-7 loculatae, 5p-611 X 15p-l8p, I+ coerulescentes
(Figure 912).
CHEMISTRY.-stiCtiC and constictic acids.
HOLoTwE.-Trail to hforne h g l a i s , Dominica,
elevation about 900 m, Hale 37959, 13 December
1971 (US).
Chemistry places this species near 0. exanthisrnocaypa
but the exciple is not so clearly separate
froin the thick, erect, pulverulent margin, leaving
the disc open. The spores are much smaller. Another
ilrest Indian stictic acid-containing species,
Ocellularia albooliuacea (Vainio) Zahlbruckner,
has a more typically recurved margin and a thin,
entire exciple.
7. Ocellularia dilatata
FIGURE lla,b
Ocellularia dilatata Miiller-Argau, 1895:452.
TYPE-CoLLECTION.-Rio de Janeiro, Brazil,
Glaziou 5531 pro parte (GI lectotype; BM, isotype).
(Figure 1 la).
Thallus whitish gray, thin, dull, in part hypophloeodal,
forming colonies 4-6 cm broad; apothecia
chroodiscoid, large, up to 2.5 mm broad, round
to elongate, marginal flap rather coarse; disc white
pruinose; hymenium about 60p high; spores 8,
6-7 loculate, 5p-6p X 15p-l7p, I+ blue (Figure
l l b ) .
CHEMIsTRY.-stictic and constictic acids.
HABITAT.-LO\Wr trunk area in higher rain forest
zone (850 m).
This rare species is related to Ocellzila?.ia albo~osella,
but it has a thicker recurved margin and a
different chemistry. I have collected a number of
specimens at low elevation in Trinidad, suggesting
that the species is more at home on continental
land masses.
SPECIMENS EXAXIINED.--~~~ (37695, 37702).
8. Ocellularia dominicana Hale, new species
FIGURE 2c,d; 10
Thallus corticola, brunneo- vel viridi-albidus,
epiphloeodes, continuus, nitidus, 5-10 cm latus;
apothecia semi-emergentia, 0.7-1.1 mm diametro,
apice incolores, columella nulla, solitaria vel 2-3
arcte aggregata, habitu similia apothecio singulari,
excipulo interiore evoluto, ostiol~um complente;
ostiolum rotundum, 0.1-0.5 mm diametro; hymenium
6Op-8Op altum; sporae 8: nae, transversim
6-7 loculatae, 6p8p X 16p-25p, I + coerulescentes
(Figure 10).
CHEMISTRY.-NO substances present.
20 S.\lII'HSONIAN CONTRIBUTIONS TO BOTANY
Thelotrema homothecintm Vainio, 1921:190 [type-collection:
Irosin, Mt. Bulusan, Pror. Sorsogon, Philippines, Elmer
14852 (TUR, lectotype; FH, G, L, S, US, W, isotypes); Figure
Ild].
Ocellularia homothecia (Vainio) Zahlbruckner, 1923:593.
Ocellularia porinoides (hlontagne and von der Bosch) Zahlbruckner,
1923:599 [not 0. porinoides (Acharius) Sprengel,
1827 : 2421.
Thallus white to ashy, thin, smooth, in part
hypophloeodal, forniing colonies 1-4 cm across;
apothecia numerous, semi-emergent, the rim becoming
erect, noncarbonized and without a
columella, the inner exciple distinct, intact, forming
an apical pore within the main pore; main pore
open, 0.2-0.4 mm in diameter; hymenium 160p-
180p high; spores 4-8/ascus, 18-22 loculate, 12p-
18p X 6Op-lOOp, If deep blue (Figure llg).
CmhiISTRY.-stictic and constictic acids and
rarely the higher "quintaria" unknown B.
HABITAT.-Trunks, canopy branches, fence posts,
and planted trees (300-900 m).
This common pantropical species is unique because
of the pore-within-a-pore arrangement. The
large spores are also distinctive but it is not unusual
to find sterile specimens. It is also one of
the few species in the family to invade secondary
forests, fence posts, etc., in wet areas up to very
high elevation.
16b (37982), 17 (35605), 20a (37780, 37781, 37786,
3i788, 38019, 38025), 25a (37708, 37709, 37713,
37714, 37715, 37716, 37849, 37996, 38069), 25b
(35473, 35516), 26a (37692, 37944), 27 (38042,
38055).
SPECIAIEXS EXAAIINED.-8 (381 14), 11 (38009),
FIGURE lO.-OcelIularia dorninicana, new species (Hale 37841)
(about X 10).
HoLorYpE.-Submossy forest on upper slopes of
Trois Pitons, Dominica, elevation about 1000 m,
Hale 37967, 6 December 1971 (US).
H.%nnxT.-Trunk and lower base of trees in submossy
forest (700-1 100 m).
The apothecia are so tightly aggregated that I
first interpreted the wall between them as a
columella. The combined outer margin and exciple
are minutely pulverulent antl crowded, filling the
broad disc but leaving a distinct pore area. The
species isas rather corninon at Trois Pitons but not
found elsewhere.
SPECIMEKS EXAMIKED.--~~ (38053), 27 (37683,
37688, 37829, 37832, 37841, 38041).
9. Ocellularia exanthismocarpa
FIGURES 50, llc,g
Ocellulnria esa~ithisviocarpn (Leighton) Zahlbruckner, 1923:
590.
Thelotrema e s n ~ ~ t h i s n i ~ c a ~ . p ~ n z Leighton, 1869: 169 [typecollection:
Central Province, Ceylon, Tlircnites C.L. 97
(BM, lectotype; H, isotype); Figure llc].
Thelotrema porinoides hlontagne and I on der Bosch, 1855:
484 [type-collection: Java, Junghuhn 151 (L, lectotype;
FH, G, P, Ti', isotypes); Figure lle].
Ocellularia mziltiloczilnris Zahlbruckner, 1912:369 [typccollection:
Lanai, Hawaii, Roqk l i (IV. holotypc); Figure
Ocellulnria isertii T'ainio, 1915:40 [type-collcction: Guadeloupe,
Zsert 87 (C, lectotype; TUR, isotype); Figure Ilfl.
5a].
10. Ocellularia fecunda (Vainio) Hale,
new combination
FIGURE 11 h,i
Thelotrenu domingeiise var. fecu?iduni T'ainio, 1896:208
[type-collection: Richmond Valley, St. Vincent, Elliotst 243
(BM, lectotype; TUR, isotype); Figure llh].
Ocellularia donzingensis var. fecunda (Vainio) Zahlbruckner,
1923:590.
Thallus light greenish or whitish mineral gray,
epiphlocotlal, smooth and shiny, forming large
colonies up to 30 cin broad; apotliecia large, strongly
emergent, 1.5-1.8 inin in diameter, the cortex
soon breaking a~vay antl revealing a light yellowis11
orange medulla, walls heavily carbonized,
KUMBER 16 21
FIGURE 1 I .-Specimens of Thelotremataceae (all about X 10): a, Ocellularia dilatata (isotype in
BM); b, 0. dilatata (Hale 37702); c, 0. exanthismocarpa (lectotype in BM); d, 0. homothecia
(isotype in FH); e, 0. porinoides (lectotype in L); f, 0. isertii (lectotype in C); g, 0. exanthismocarpa
(Hale 37996); h, 0. fecuxda, new combination (lectotype in BM); i, 0. fecuuda, new
combination (Hale 35218); j, 0. maculata, new species (Hale 38015); k, 0. motdenii, new species
(Hale 37887).
22 SMITHSONIAN CONTRIBUTIONS TO BOTANY
columella absent or weakly and irregularly developed;
pore gaping, round to irregular and jagged,
0.4-0.8 mm in diameter, the white pruinose disc
clearly visible; hymenium 2OOp-25Op high; spores
1--4/ascus, 30-40 loculate, 12-20 X 80p-I50p, rarely
turning brown, I+ blue (Figure l l i ) .
CHExmiRY.-Hypoprotocetraric acid and in
lesser concentration 4-0-demethylnotatic acid
along with unidentified pigments.
HAuITAT.-Basal area of trees into the upper bole
and canopy in the rain forest (300-850 m).
This is one of the few species in the family that
can be recognized at sight in the field. The apothecia
are very large and the yellow-orange medulla
is exposed by rubbing the cortex of€. It is the
second commonest species in Dominica and forms
very extensive colonies. As far as I have determined,
the species is restricted to the West Indies and has
no close relatives.
37816, 37820, 38003, 38046, 38061), 12 (37902,
37906, 37913, 37920, 37921, 37931), 15 (35155,
35167, 35218), 16a (37658, 37942, 37970, 38050),
16b (37796, 37802, 37810, 37874, 37881, 37954),
16c (37765, 37766, 37774, 37943, 37980), 18a
(37725, 37957, 37962, 37986, 38039), 21 (38080,
38081, 38089, 38103, 38106, 38107, 38132, 38137),
22 (37639, 37644, 37828, 37838, 37843, 37844,
37844a, 37845, 37856, 37858, 37960, 38026), 23
(37679), 24a (37740, 37743, 37063, 37064), 25a
(37711), 26a (38056).
SPECIMENS EXAMINED.-8 (38 120), 1 1 (378 1 1,
11. Ocellularia maculata Hale, new species
FIGURE l l j
Thallus corticola, cinereo-albidus, epiphloeodes,
planus, continuus, nitidus, 9 cm latus; apothecia
nunierosa, minuta, 0.2-0.3 nim diametro, apice
ol)lusc;i, columellata vel columella parce evoluta;
ostiolum rotundum, 0.05-0.1 mm diametro, anguste
nigrocinctum, columella protrusa; hymenium 100p
altum; sporae 8:nae, transversim 4 loculatae, 5y-7p
X llp-l3p, I- (Figure llj).
CHEXIISTRY .--No substances present.
HoI.oTYI.r.-Remnants of rain forest in pastures
above Giraudel, Dominica, elevation 650 m, Hale
38015, 11 December 1971 (US).
Occlliilnria mncirlata might seem at first glance
to be 0. papillata but the apothecia and spores
are much smaller. There are no related species.
SPECIXE.N EXAMMINED.-lSb (35524).
12. Ocellularia mordenii Hale, new species
FIGURES 3c,d; 66; Ilk
Thallus corticola, pallide cinereo- vel brunneoalbidus,
epiphloeodes, continuus, aetate rimosus,
nitidus, usque ad 12 cm latus; medulla omnino
sanguinea; apothecia dispersa, vix emergentia,
apice obfusca, extus crasse verrucosa, verrucis fragilibus,
rumpentibus, columella nulla vel parce evoluta
(Figure 6b); ostiolum rotundum, 0.08-0.12 mm
cliametro; hymenium circa 2OOp altum; sporae
1-2:nae, transversim 24-30 loculatae, 15p-20y X
70p150p, I+ coerulescentes (Figure Ilk).
CHExmTRY.-Unidentified anthraquinone pigments
only.
HoLoiyPE.-Canopy branches in primary rain
forest, Dleau Gommier Forest Reserve, Dominica,
elevation about 370 m, Hale 37764, 12 December
HAwrAT.-Canopy branches in rain forest (370-
800 m).
‘The deep red pigment and large warts are distinctive.
It is a conspicuous canopy species. A very
similar species in Cuba, 0. xanthost~oma (Nylander)
Zahlbruckner, differs mainly in containing
n I<- pale yellow-orange pigment. This lichen is
named in honor 01 Alrs. William J. Morden.
SI’PCIMENS EXAhlINED.-l2 (37909), 15 (35198),
16a (38051), 16b (37791), 16c (37762, 37886, 37887,
37010), 26a (37863).
1971 (US).
13. Ocellularia ngropuncta Hale, new species
FIGURES 2a,b; 6a; 12a
Thallus corticola, pallide brunneo- vel viridicinei
eus, epiphloeodes, continuus vel rarius I imosus,
5-8 cm latus; apothecia numerosa, emergentia,
0.6-0.8 mni diametro, apice obfusca, columella
nulla (Figure 6a); ostiolum rotundum, 0.1-0.2 mm
tlianietro, nigrocinctum; hJmeniuin 100~1-110p
altum; sporae 8:nae, transversim 6-8 loculatae, 6p-
10:~ X 2Oy-32p I+ coerulescentes (Figure 12a).
CmwsTRY.-Protocetraric acid.
HOLOTYPE.-T~ ail to summit of Rforne Anglais,
Dominica, elevation about 900 m, Hale 35365,
January 1969 (US).
NUMBER 16 23
FIGURE IZ.-Specimens of Thelotremataceae (all about X 10): a, Ocellularia nigropuncta, new
species (Hale 35365); b, 0. rimosa, new species (Hale 38052); c, 0. oliuacea (lectotype in G);
d, 0. papilla,ta (lectotype in BM); e, 0. papillata (Hale 38098); j, 0. perforata (lectotype in
BM); g, 0. excavata (lectotype in BM); h, 0. rufocincta (lectotype in G); i, 0. terebrata var.
abbreuiata (isotype in FH); j, 0. pyrenuloides (isotype in W); k, 0. pyrenuloides (Hale 35524).
24 SMITHSONIAN CONTRIBUTIONS TO BOTANY
HABrT.AT.-Lianas, lower bole and base, symphonea
roots, rarely canopy of trees in the rain
forest (300-900 m).
The black-rimmed pore area and smooth thallus
are diagnostic features. Other ecolumellate protocetraric
acid-containing species include Ocellulaiia
bonplandiae (Fke) Muller-Argau, which has a
grainy surface, and 0. ueirzIcosa (Fee) Miiller-
Argau, which has an irregular open pore and
larger spores (over 30y long). Under the scanningelectron
microscope 0. nigropuncta, new species,
has a finely pitted but otherwise smooth surface
(Figure 2a,b), whereas 0. uerizicosa appears to
have an aculeate orientation and 0. bonplandiae
is verrucose and fissured.
SPECIMENS EXAMINED.-~ (381 15, 38158), 10
(38169), 12 (37903), 16a (37660), 16b (37798), 16c
(37755, 37770, 37963), 18a (37948), 21 (38092,
38094, 38138), 22 (37852), 24a (37751).
14. Ocellularia olivacea
FIGURES la; 3a,b; 5c; 12c
Ocellularia oliuacea (Fee) Miiller-Argau, 1887a:7.
Myriot, emu olivacea Fee, 1824: 103 [type-collection: On Bonplandin
trifoliata, South America (G, lectotype); Figure
Ocellularia oliuacea (Fee) C. and D. Van Overeem-De Haas,
1 2c] .
1922: 118 [superfluous combination],
Tliallus light greenish to ashy white, shiny,
thick, up to 0.5 mm (Figure la), forming extensive
colonies up to 50 cm broad, sometimes finely
cracked and fissured; apothecia very numerous, immersed,
0.15-0.3 mm in diameter, noncarbonized
and without a columella; pore small, flush, about
0.1 mm in diameter; hymenium 65p-75p high;
spores 8, more or less uniseriate, 4 loculate, 4y-
6p X 8y-12p, I f blue (Figure 5c).
CHE;\IIsTRY.-“Olivacea” unknown and a second
lower spot.
HABITAT.-canopy branches to mid-bole in rain
forest (240-700 m).
This is one of three “Myiiotiemn” species that
occur in Dominica, the other two being 0. terebrat
iila and Thelotrema clandestinttm, all typically
found on canopy branches in dense rain forest.
Ocellztlaria oliuacea is characterized by the rather
small spores and unusual chemistry. The West
Indian material all produces a second spot below
the “olivacea” unknown, probably a related depside.
The type-specimen of 0. oliuacea lacks this
spot. The surlace under the scanning-electron
microscope is strongly aculeate (Figure 3a,b).
8 (38164), 9 (38154), 11 (38001), 12 (37965), 15
(35205), 16a (37992, 38054), 18a (37965), 21
(38113).
SPECIMENS EXAMISED.-7 (35226, 35243, 35249),
15. Ocellularia papillata
FIGURE 12d,e
Ocellularia papillata (Leighton) Zahlbruckner, 1923:597.
Thelotrema pnpillatunz Leighton, 1869: 169 [type-collection:
Central Province, Ceylon, Thwaites C.L. 129 (BM, lectotype;
H, G, P, S, UPS, W, isotypes); Figure 124.
Thallus whitish to pale greenish ashy, smooth
to more or less warty, shiny, forming colonies 3-12
cni broad; apothecia numerous, 0.6-0.8 mm in
diameter, immersed to slightly emergent, the upper
walls carbonized, columella present, thin to scarcely
developed to absent; pore round, 0.1-0.2 mni in
diameter; hymenium 50p-70p high; spores 8/ascus,
7-9 loculate, 6p-lop X 2Oy-36p I+ blue. (Figure
12e).
CHEJIISTRY.--NO substance present.
HABITAT.-ktplingS, prop roots, lianas, and tree
bases at mid-elevation in rain forest (300-900 m).
This common species has few outstanding features.
The columella is usually present, although
often weakly developed and difficult to find. Some
specimens, as indicated below, seem to lack it entirely.
The apothecia are inconspicuous, almost
immersed to slightly emergent, but the pore is
always distinct. The thallus is well developed,
usually quite whitish, and smooth. There are no
close relatives i\rithout lichen substances, although
0. perfomto, which has protocetraric acid, is very
close in general aspect except perhaps that the
apothecia are more often emergent. Both species
occupy similar base level habitats in the rain forest
and were collected together at 11 of the localities.
At eight other localities only one of the species
was found.
SPECIMENS EXAMINED (columella present).-S
(38123, 38130, 38165, 38168), 9 (38131), 11 (37825),
16a (37795, 37991), 16b (37662, 37804, 37805,
37808), 18a (37721, 37964); 21 (38084, 38085,
38097, 38098, 38152), 22 (37645, 37837, 37842), 23
(37671, 37681, 37682), 27 (37850, 38072), 28
(35428).
NUMBER 16 25
SPECIMENS EXAMINED (columella absent).-7
(35227), 10 (38127, 38150), 16a (37655, 37661),
16b (37792, 38074), 16c (38045) 21 (38086, 38088),
22 (37649, 37834, 37851, 37854, 37855), 23 (37677,
38965), 24a (37746).
16. Ocellularia perforata
FIGURFS Ib, 5e, 7a, 12f
Ocellularia perforata (Leighton) Muller-Argau, 1892284.
Thelotrema perforatum Leighton, 1866:447 [type-collection:
Casiquiari, Brazil, Spruce 254 (BM, lectotype); Figure 12fl.
Thelotrema terebratum var. abbreuiatulum Vainio, 1890:83
[type-collection: Caraqa, Minas Gerais, Brazil, Vainio 1551
(TUR, lectotype; BM, FH, UPS, isotypes); Figure 12i].
Ocellularia rufocincta Muller-Argau, 1893: 146 [type-collection:
Foret du Rio General, Tondoz s.n. (Pittier 6107) (G, lectotype);
Figure 12hl.
Thelotrema excavatum Vainio, 1896:208 [type-collection: Mt.
St. Andrew, St. Vincent, Elliott 153 (TUR, lectotype; BM,
FH, isotypes); Figure 12g].
Thelotrema excauatum var. impressulum Vainio, 1896:208
[type-collection: Morne Anglais, Dominica, Elliott 169 (BM,
lectotype)].
Ocellularia terebrata var. abbreviata (Vainio) Zahlbruckner,
1923502.
Thallus whitish to pale greenish ashy, smooth
to more or less warty with age, shiny, forming
colonies 3-10 cm broad; apothecia numerous, immersed
to moderately emergent, 0.4-0.6 mm in
diameter, the upper wall carbonized, columella
usually developed, thin 50p-70p in diameter, but
sometimes lacking or difficult to find; pore round,
0.07-0.15 mm in diameter; hymenium 55p-70p
high; spores 8/ascus, 6-8 loculate, 7p-lop X 2 0 ~ -
34p, I+ blue (Figure 5e).
CHEMIsmY.-Protocetraric acid and the “amplior”
unknown as an accessory substance.
HABITAT.-Buttresses, saplings, and bole to
rarely canopy branches in rain forest at midelevation
(300-850 m).
This is one of the commonest species in Dominica
at base level. The smooth whitish thallus (see
cross-section in Figure 2b) and the small semiemergent
apothecia resemble 0. papillata, as discussed
above. Some of the specimens as listed below
lacked any sign of a columella.
SPECIMENS EXAMINED (columella present).-8
(38149), 11 (37812, 37817, 37997, 38030, 38033),
12 (37899, 38078), 14 (35139), 16a (37651, 37666,
38007, 38032), 16b (37884), 16c (37759, 37761,
37775, 37952), 18a (37727, 37728, 37730, 37732,
37535, 37961, 37969, 37983, 38020), 20a (37784,
37785, 38016, 38017), 21 (38108, 38134, 38145), 22
(37646, 37647, 37830, 35857, 38036), 23 (37675),
24a (37752), 26a (37698, 37701), 27 (37686, 37827).
SPECIMENS EXAMINED (columella absent).-8
(38160, 38163), 16a (38018), 21 (38093, 38140,
38157), 23 (37674), 24a (38066), 27 (37684, 37847),
26a (37700), 26b (35370).
17. Ocellularia pyrenuloides
FIGURE 12j,k
Ocellularia fiyrenuloides Zahlbruckner in Magnusson and
Zahlbruckner, 1943:46.
TYPE-CoLLECTIoN.-\vaiIUkU, Maui, Hawaii,
Fauyie 656 (UPS, lectotype; S, IV, isotypes) (Figure
12j).
Thallus ashy mineral gray, thin to evanescent,
continuous, forming colonies about 5 cm broad;
apothecia numerous, nearly immersed to moderately
emergent, 0.5-0.7 mm in diameter, walls
heavily carbonized, columella present, 130p in
diameter; pore round, 0.1-0.15 mm wide, pore area
surrounded by a blackish ring, in part white pruinose,
the pruinose top of columella visible through
the pore; hymenium about 60p high; spores 8/ascus,
6-7 loculate, 611 X 12p-I5y, I+ blue (Figure 15%).
CHEMISTRY .-stictic and constictic acids.
HAnITAT.-Tree branch in rain forest (650 m).
The darkened pore area is similar to that in
Thelotrema teniie, new species. There are no related
species with stictic acid. It is still surprising
to find a Hawaiian species on Dominica, but to
me this suggests how poorly the Thelotremataceae
have been collected.
SPECIMEN EXAMINED.-18b (35524).
18. Ocellularia rhodostroma
FIGURES 5g, 7b, 13a
Ocellularia rhodostromn (Montagne) Zahlbruckner, 1923:600.
Ascidium rhodostromum Montagne, 1851:75 [type-collection:
Cayenne, French Guiana, collection 1334 (P, lectotype);
Figure 13~1.
Thallus light greenish tan to gray, smooth to
finely warty, continuous and epiphloeodal, the
medulla white or pink, forming colonies 5-10 cm
26 ShIITHSONIAN CONTRIBUTIONS TO BOTANY
FIGURE 13.--Specimrns of Thelotremataceae (all about X 10): a, Ocellularia rhodostroma (lectotype
in P); b, 0. subcavata (lectotype in H); c, 0. vaga (isotype in FH); d, 0. subca7jata (Hale
37769); e, 0. terebratula (lectotype in H); f, 0. remanens (isotype in G); g, 0. sor~diata, new
species (Hale 37908); h, Plraeotrema aggregatum, new species (Hale 35229); i, 0. terebratula
(Hale 37793); 1, P. disciforme, new combination (lectotype in Bhl); k, P. aqztiliiium (lectotype
in TUR).
NUMBER 16 27
broad; apothecia large and easily visible with the
naked eye, ascidioid, 1.0-1.8 mm in diameter, the
walls heavily carbonized and the medulla deep
pink, readily exposed as the fragile cortex breaks
away, columella lacking; pore round and small,
0.05-0.1 mm in diameter, surrounded by a raised
ring; hymenium 2OOp-275p high; spores l-2/ascus,
25-30 loculate, 25p-4Op X lOOp-25Op, I+ blue
(Figures 5g and 70).
CHEMISTRY.-Hypoprotocetraric acid and in
greater concentration 4-0-demethylnotatic acid
along with unidentified pigments.
HABITAT.-CanOpy branches to lower bole at all
elevations on the island.
Ocellularia rhodostroma is the commonest thelotreme
on Dominica and the easiest to identify as
well. The large ascidioid apothecia have a deep
pink medulla exposed when the fragile cortex
breaks. The pore is strongly annulate. While occurring
at all elevations, it seems best developed
at higher elevation into the mossy forest, perhaps
because it competes with so few other species
there.
38146, 38166), 10 (38119), 11 (37815, 37823, 37824,
38004, 38023, 38049), 12 (37894, 37916, 37917,
37927, 37930, 37934), 13 (35279 35448), 14 (35111,
35133), 15 (35160, 35170, 35188, 35314, 16a (37653,
37938), 16b (37800, 37806, 37875, 37994, 37995),
16c (37756, 37870, 37979), 18a (37729, 37731,
37987), 18b (35517, 35522, 35523, 35587), 20a
(37777, 38065), 21 (38035, 38147), 22 (37641, 37643,
35650, 37835, 37861), 23 (37672, 38975), 24a (37736,
37742, 37745, 37749, 37754, 38068), 24b (35629),
25a (37705, 37707, 37720, 37840, 37846, 37848,
35862, 37867, 37971, 37975), 25b (35469), 26a
(37689, 37690, 37693, 37694, 37699, 37703, 37864,
37866, 37885, 37947 38000, 38014), 26b (35354,
35366, 35371, 35372, 35376), 27 (37831), 28 (35435,
35440 ).
SPECIMENS EXAMINED.-7 (35246), 8 (38122,
19. Ocellularia rimosa Hale, new species
FIGURE 12b
Thallus corticola, brunneo- vel viridi-cinereus,
epiphloeodes, nitidus, continuus sed aetate rimosus,
usque ad 15 cm latus; apothecia numerosa, emergentia,
0.7-09 mm diametro, apice obfusca, columella
nulla; ostiolum rotundum vel irregulare,
0.1-0.2 mm diametro, nigrocinctum; hymenium
110p-150p altum; sporae 8:nae, transversim 6-8
loculatae, 7p-lop X 24p-32p, I + coerulescentes
(Figure 12b).
CHEMISTRY.--NO substances present.
HoLoTYPE.-Base of tree in primary rain forest,
Dleau Gommier Forest Reserve, Dominica, elevation
370 m, Hale 37978, 1 December 1971 (U.S.).
HABITAT.-Prop roots, tree bases, and lianas in
rain forest (370-430 m).
Except for chemistry and slightly larger apothecia,
this species is very close to 0. nigropuncta,
new species. Both are base level species, but 0.
Yimosa has a more restricted elevational range.
The two species were collected together only at
Dleau Gommier.
SPECIMENS EXAMINED.-~ 1 (38052), 16a (37654),
16b (37392, 37803), 16c (37758).
20. Ocellularia sorediata Hale, new species
FIGURE 13g
Thallus corticola, viridi-albidus, epiphloeodes,
continuus, planus, nitidus, usque ad 15 cm latus,
sorediatus, soraliis sparsis, verrucosis, circa 1 mm
latis; apothecia immersa vel semi-emergentia, 0.4-
0.6 mm diametro, apice fuliginea, columellata; ostiolum,
rotundum, 0.08-0.1 1 mm diametro; hymenium
6Op-7Op altuni; sporae 8:nae, transversim
3-4 loculatae, 6p-8p X 12p-16p, I+ coerulescentes
(Figure 13g).
CHEMISTRY.-NO substances present.
HoL0TYPE.-Logging area about 3 km northwest
of Pont Cassi. on the Layou Road, Dominica, elevation
420 m, Hale 37908, 10 December 1971 (US).
HAnrTAT.-hfid-bole of tree in rain forest;
This is, as far as I know, the only sorediate
species in the family. Large corticate tubercules
are produced first, and some of these become
coarsely sorediate. Although these peculiar structures
seem to originate from the main thallus, there
is still the possibility that they have an extraneous
origin.
SPECIMEN EXAMINED.-16b (38029).
21. Ocellularia subcaoata
FIGURE 13b,d
Ocellularia subcavata (Nylander) Zahlbruckner, 1923:601.
28 SMITHSOXIAN CONTRIBUTIONS TO BOTANY
22. Ocellularia terebratula Thelotrema subcavatum Nylander, 1876:561 [type-collection:
Cuba, Wright 509 (H, lectotype; FH-Tuck, isotype); Figure
13bl.
Thelotrema vagum Vainio, 1896:209 [type-collection: Richmond
Peak, St. Vincent, Elliobt 260 (BM, lectotype, FH,
isotype); Figure 13~1.
Ocellularia uaga (Vainio) Zahlbruckner, 1923:603.
Thallus ashy whitish to greenish gray, shiny,
more or less warty and cracked, forming colonies
up to 12 cm broad; apothecia very numerous, emergent
to ascidioid, 0.7-1.0 mm in diameter, crowded,
walls carbonized, columella present, basally carbonized
but usually colorless in the upper half;
pore round, small, 0.1-0.2 mm in diameter, surrounded
by a whitish, more or less raised ring,
the center filled with the often protruding pruinose
columella; hymenium about 1001~ high; spores
8/ascus, usually uniseriate, 5-6 loculate with narrow
locules, 4p-7p X 8p-2Op1 very rarely turning
brown with age, I+ blue (Figure 13d).
CHEMIsTRY.-Psoromic and conpsoromic acids.
HABITAT.-Canopy branches and upper bole in
rain forest (240-800 m).
This is one of the commonest thelotremes in the
canopy and can be recognized in the field with
a hand lens because of the numerous crowded
apothecia with a white annulate pore and protruding
columella. The columella is well developed
in cross-section but unique in being more or less
noncarbonized in the upper half.
(38161), 10 (38129), 12 (37893, 37896, 37898,
37900, 37905, 37912, 37924, 37925, 37926, 37928,
37929, 37933), 13 (35294), 15 (35158, 35209), 16a
(37652, 37795, 38012), 16b (37662, 37794, 37865),
16c (37767, 37769, 37772, 37773, 37924, 37941,
37972), 18a (37724, 37889, 37949, 38038), 23
(37680), 24a (37737, 37744, 37750), 26a (37691,
37704).
SPECIMENS EXAMINED.-7 (35248, 35252), 9
FIGURE 13e,i
Ocellularm terebratula (Nylander) Muller-Argau, 1887: 12.
Thelotrema terebratuluin Nylander, 1867:315 [type-collection:
Rio Negro, Colombia, Lindig 129 (H, lectotype; FH, C, hf,
P, isotypes); Figure 13el.
Thelotrema clandestinum f. remanens Nylander, 1867:315
[type-collection: Monte del Moiro, Colombia, Lindig 90
(H, lectotype; BM, G, P, isotypes); Figure l3fl.
Thallus light greenish to ashy white, thick and
continuous, smooth, shiny, often forming extensive
colonies up to 15 cm broad; apothecia numerous,
0.2-0.4 mm in diameter, immersed, noncarbonized
and without a columella; pore flush,
0.05-0.1 mm in diameter, usually surrounded by
a faint whitish ring; hymenium about 65p high;
spores 8/ascus, 3-5 loculate, 5p-9p X 15p-20p, I+
blue (Figure 132).
Ocellularia remanens (Nylander) Muller-Argau, 1887:7.
CHE;MISTRY.-PsOromic and conpsoromic acids.
H A B I T A T . - ~ ~ ~ o ~ ~ branches and upper bole in
rain forest (300-400 m).
Ocellularia terebratula, previously known only
from Colombia, is a typical canopy level species.
It is very similar to Thelotrema clandestinum, as
explained below, except for spore characters.
SPECIMENS ExAMINED.-~ (38125), 10 (38126), 12
(37904, 37918), 16a (37668, 37878, 37937, 38011,
38040), 16b (37793).
Phaeotrema
This genus is very similar to Ocellularia in spore
septation but the spores are brown and sometimes
shriveled at maturity, especially in large-spored
species. The iodine reaction often seems to be
negative, while almost all species of Ocellularia
are strongly I+ blue. On the world level, Phaeo-
Key to the Species of Phaeotrema
1. Apothecia aggregated ........................................................ 23. P. aggregatum, new species
1. Apothecia solitary.
2. Apothecia large, about 0.6 mm in diameter, strongly emergent; columella lacking . . . . . . . . . . .
2. Apothecia smaller, less than 0.5 mm in diameter, immersed; columella more or less
developed.
3. Pore 0.1-0.2 mm in diameter, sometimes elongate, with a white annulus .......................
......................................................................................................... 23. P. leiostomum
3. Pore 0.5-0.1 mm in diameter, round, annulus lacking . ._ 26. P. obscurum, new species
.......................................................................................... 24. P. disciforme, new combination
NUMBER 16 29
trema is represented by about 40 species, a rather
small group, moreso because few of the species
are well represented by herbarium material. In
Dominica only six collections (four species) were
found, these all occurring below 350 m elevation.
23. Phaeotrema aggregatum Hale, new species
FIGURE 13h
Thallus corticola, pallide viridi-cinereus, epiphloeodes,
planus, aetate rimosus, 8 cm latus;
apothecia irregulariter aggregata, 0.4-0.6 mm
diametro, immersa, intus incolores, columella nulla,
excipulo interiore bene evoluto; ostiolum 0.1-0.2
mm latum, in centro excipulo interiore congesto,
albido-pruinoso; hymenium 110p-12Op altum;
sporae 8:nae, transversim 4-5 loculatae, 8p X 16p
(Figure 13h).
CHEMISTRY.-NO substances present.
HOL0TYPE.-Logging area at Newfoundland,
Dominica, elevation 250 m, Hale 35229, January
HABITAT.-upper branches of tree in rain forest.
This unusual species has densely aggregated,
immersed apothecia. The pore is usually distinct
but may be obscured by the pulverulent exciple
and margin, giving the appearance of a Phlyctis.
1969 (US).
24. Phaeotrema discif orme (Leighton) Hale,
new combination
FIGURES ic, 131, 14a
Thelotrema disciJornze Leighton, 1869: 170 [type-collection:
Central Province, Ceylon, Thwaites (BM, lectotype; H,
isotype); Figure 1311.
Thelotrema exalbrduin Stirton, 1881 : 184 [type-collection:
Assam, Il’att s.n. (BM, lectotype)].
Thelotretna aquilinum Vainio, 1915:137 [type-collection: St.
Croix, Boergesen s.n. (TUR, lectotype; C, isotype)].
Leptotrema exalbidum (Stirton) Zahlbruckner, 1923:634.
Thelotrema galactinum Vainio, 1926:24 [type-collection:
Maloapan, Mexico, Liebmann 7712 (C, lectotype); Figure
13k].
Phaeotrema galactinum (Vainio) Zahlbruckner, 1932:245.
Thallus whitish, dull, very thin and evanescent
to hypophloeodal, forming colonies 2-4 cm broad;
apothecia large, emergent, 0.7-1.1 mm in diameter,
upper walls becoming suberect, carbonized, columella
lacking, the inner exciple weakly developed
and pulling away from the main margin, the
white-pruinose disc clearly visible; pore open,
round, 0.2-0.5 mm in diameter, the rim usually
darkened; hymenium 120p-140p high; spores apparently
4/ascus, 20-24 loculate, 7p-12p X 40p-
551.1, colorless when immature but mostly turning
brown and shriveled at maturity, I- (Figure 14a).
CHEMISTRY.-NO substances present.
HABITAT.-Lower trunk area (sea level to 370 m).
A thorough examination of the lectotype showed
that the spores are transversely septate without any
longitudinal septae. Old shriveled spores, however,
give the false impression of being muriform. It is
a widespread pantropical species characterized by
the large emergent apothecia without periderm
inclusions, wide pore, pruinose rim and disc, and
lack of a columella.
SPECIMENS EXAMINED.-4 (Wirth Sen.), 16b
(37939).
25. Phaeotrema leiostomum
FIGURES 5h, 6 4 14b,c
Phaeotrema leiostomum (Tuckerman) Zahlbruckner, 1923:
Thelotrema leiostornum Tuckerman, 1862: 407 [type-collection:
608.
Cuba, Wright 149 (FH, lectotype); Figure 14bl.
Thallus ashy to pale greenish white, epiphloeodal,
continuous, smooth and shiny, forming colonies
2-5 cm broad; apothecia immersed to slightly emergent,
0.3-0.6 mm in diameter, weakly carbonized,
columella absent to well developed, noncarbonized
(Figure 6 4 ; pore round to irregularly elongate,
surrounded by a raised ring, 0.1-0.3 mm
long; hymenium 130p-150p high; spores 8/ascus,
brown, 4 loculate, 8p-10~ X 16p-l8p, I- (Figures’
5h, 14c).
CHEMISTRY.-HypOprOtOCetrariC acid with or
without 4-0-demethylnotatic acid along with pale
unidentified pigments.
HABITAT.-canOpy and mid-bole branches in
rain forest (350-500 m).
A raised annulate pore is characteristic of this
species. Elongation of the apothecia was best developed
in collection 37956. The extreme form can
be seen in Phaeotrema lirelliforme (Tuckerman)
Zahlbruckner, which lacks any lichen substances.
When apothecia are lacking, the thallus of Phaeotrerna
leiostomum alone would not permit identification.
30 SMMITHSONIAN CONTRIBUTIONS TO BOTANY
FIGURE 14.--Specimens of Thelotremataceae (all about X 10): a, Phaeotrema disciforme, new
combination (Hale 37929); 6, P. leiostomzim (lectotype in FH); c, P. leiostomum (Hale 33203):
d, P. obscurum, new species (Hale 38091); e, Thelotrema cara~sense (isotype in FH); f. T.
carassense (Hale 37873); g, T. clandestinuni (lectotype in G); h, T. subcaesium (lectotype in H);
i, T. clandestinum (Hale 33136): j, T. confornie (lectotype in G); k, 7’. consanguineum (lectotype
in G); 1, T. conforme (Hale 38062).
NUMBER 16 31
SPECIMENS EXAMINED.-15 (35203), 16a (37956),
16b (37993).
26. Phaeotrema obscurum Hale, new species
FIGURE 14d
Thallus corticola, pallide brunneo-albidus, tenuis
vel hypophloeodes, 5 cm latus; apothecia modice
emergentia, 0.3-0.5 mm diametro, apice fuliginea,
columellata; ostiolum rotundum, 0.1 mm diametro,
intus albido-pruinosum; hymenium circa
12Oy altum; sporae 8:nae, transversim 5-6 loculatae,
lop-14p X 24p-28p (Figure 14d).
CHEMrsTnY.-stictic and constictic acids.
HoLoTYPE.-Forty centimeters up trunk of a
small tree, Rosalie Road above Newfoundland, elevation
300 m, Hale 38091, 10 December 1972 (US).
HAnITAT.-Tree in rain forest.
This tiny species could easily be overlooked.
It is unusual to find a well-developed columella
in apothecia this small. There are no other species
in the genus with similar morphology. Phaeotrema
albidulum (Nylander ) Muller-Argau also has
stictic and constictic acids, but the apothecia are
immersed and clustered and the spores larger
(40P).
Thelotrema
Thelotiema is a common genus in Dominica,
represented by 14 species among 80 collections.
On the world level there are about 130 species.
Variation in spore size is as great as in Ocellularia
but longitudinal septae are always clearly developed
(except in T. clandestinum). The six commonest
species were T. praestans (21 collections),
T. glaucopallens (16), T. clandestinum (lo), T.
interpositum (7), T. leucomelaenum (7), and T.
depressum (4).
As I noted for Ocellularia, Thelotrema avoids
the dry lowlands but is widespread in the rain
forest. Thelotrema pupillosum and T. tuberculiferurn
are restricted to the mossy forest.
Key to the Species of Thelotrema
1. Central columella present.
2. Thallus thin, hypophloeodal; apothecia not emergent, small, less than 0.6 mm in diameter.
3.
2. Thallus distinct, epiphloeodal; apothecia emergent, 0.5-1.5 mm in diameter.
4. Spores 2Op-4Op long, 4-8/ascus.
3. Columella narrow; spores less than 2Op long .......................... 39. T. tenue, new species
Columella very broad; spores more than 30p long ...................... 36. T. leucomelaenum
5. Spores about 20p long; pore not annulate but becoming black rimmed . . . . . . . . . . . . . . . . . .
35. T. leucinum
5.
4. Spores more than loop long, l/ascus.
6. Thallus smooth, P- or P+ orange.
7 .
...................................................................................................................
Spores about 40p long; pore becoming annulate ................................ 29. T. confurme
6. Thallus warty-granular, Pf yellow ........................................... 34. T. interpositum
Thallus P- ........................ :, ........................................................ 31. T. depressurn
7 . Thallus P+ orange ................................................................... 38. T. praestans
9. Thallus greenish, thin, and shiny ................................................ 33. T. glaucopallens
10. Spores less than 2Op ......................................................... 30. T. confusum, new species
1. Central columella lacking.
8. Apothecia immersed, the pore flush with thallus surface.
9. Thallus whitish, thick, dull . . . . . . . . . . . . .................... 28. T. clandestinum
8. Apothecia emergent, pores not flush wi
10. Spores more than 70y long.
11. Thallus P-; collected in mossy forest zone.
12. Apothecia 1 mm tall, eroding around a large area of the pore ......................
................................................................................................... 40. T. tuberculiferum
12. Apothecia barely emergent, pore area not eroding ........................................
................................................................................. 37. T. pupillosum, new species
11. Thallus P+ orange or yellow; collected in rain forest zone.
13. P+ yellow; apothecia up to 1.0 mm in diameter ................... 27. T. carassense
13. P+ red; apothecia about 0.5 mm in diameter ..................................................
32. T. dominicanum, new species ..................... .........................................
32 SMITHSOXIAS CONTRIBUTIONS TO BOTANY
27. Thelotrema carassense
FIGURE 14e,f
Thelotrema carassense Vainio, 1890:79.
TYPE-COLLECTION.-Carassa, Brazil, Vainio 1523
(TUR, lectotype; BM, FH, isotypes). (Figure 14e).
Thallus greenish ashy mineral gray, epiphloeodal,
continuous and shiny, forming colonies up to 15
cm broad; apothecia semi-emergent, 0.8-1.0 mm
in diameter, upper wall weakly carbonized,
columella lacking; pore about 0.1 mm in diameter,
with a weak raised ring; hymenium IlOy-13Op;
spores 8/ascus, muriform, 1-2 X 8-10 loculate,
12p-15p X 30p-40p, I+ blue (Figure 14f).
CmMIsTRY.-Psoromic and conpsoromic acids.
HABITAT.-Mid and lower bole of trees in rain
forest (370-800 m).
The Dominican material is not a perfect fit for
the South American type, but it agrees in essential
features, lack of columella, intermediate spores,
and presence of psoromic acid. The apothecia are
larger and spores smaller than in Vainio’s specimen
(about 70y long). The taxonomy of the ecolumellate
psoromic acid-containing Thelotremnta is
unfortunately still incomplete. Thelotrema postpositum
Nylander has large spores (more than
loop), but all other species, excluding T. carassense,
have small spores (less than 20p long).
26b (37873).
SPECIMENS EXAMINED.-16b (37859), 24b (37738),
28. Thelotrema clandestinum
FIGURE 14g,i
Thelotrema clandestinum Fee, 1837:90.
Thelotrema subcaesium Nylander, 1869: 120 [type-collection:
Brazil, Glariou 2193 (H, lectotype; BM, C, FH-Tuck, UPS,
US, W, isotypes); Figure 14hl.
Thelotrema concretum Fee var. subcaesiurn (Nylander) Redinger,
1936:96.
TYPE-COLLECTION.-on CinChOnU hZ?ZCifOliU in
America, without collector (G, lectotype). (Figure
Thallus whitish to greenish ashy gray, shiny,
thick, and continuous, forming colonies up to 20
cm broad; apothecia numerous, immersed, 0.15-
0.3 mm in diameter, noncarbonized and without
a columella; pore flush, round, 0.07-0.1 mm in
diameter, usually with a whitish rim; hymenium
14g) *
65p-75y high; spores 8/ascus, uniseriate, 4-5 loculate,
muriform with 1-2 longitudinal septae in at
least one of the locules, 6p-IOp X 15p-20y, I+
blue (Figure 14i).
CHEMrsTRY.-Psoromic and conpsoromic acids.
HABITAT.-canOpy branches in rain forest (500-
670 m).
Thelotrema clandestinum has small spores with
few longitudinal septations. It would, in fact, be
classified as an Ocellularia unless the spores are
carefully examined. Chemically similar 0. terebratula
has only transverse septae. Both species have
a thick, continuous thallus and occupy the same
kind of habitat. They were collected at a total
of ten localities but never together in the same
one.
(35103, 35105, 35136), 15 (35207), 16c (37768,
37872, 37981 ), 24b (35624a).
SPECIMENS EXAMINED.-13 (35298, 35393), 14
29. Thelotrema conforme
FIGURE 14jJ
Thelotrema conforrne Fee, 1837389.
Thelotrema consanguineurn Miiller-Argau, 1885b3398 [typecollection:
Apiahy, Brazil, Puiggari 477 (G, lectotype);
Figure 14kl.
TYPE-COLLECTION.-On Cinchona, America meridionalis,
without collector (G, lectotype) (Figure
141).
Thallus whitish mineral gray, smooth and shiny,
epiphloeodal, forming colonies up to 10 cm broad;
apothecia emergent, 0.8-1.0 mm in diameter, apically
carbonized, distinct columella present; pore
round, 0.1-0.15 mm in diameter, surrounded by
a whitish, sometimes raised ring; hymenium 170y-
200p high; spores 4/ascus, muriform, 8-10 loculate
transversely, 2-4 loculate longitudinally, 18p-2Oy
X 35y-50p, I+ blue (Figure 141).
CHEMIsrRY.-Psoroniic and conpsoromic acids.
HABrTAT.-Lower bole and trunk in open rain
forest (670 m).
The Dominican material is a satisfactory match
for FCe’s type-specimen, which has slightly smaller
apothecia and a less consistently developed annulate
pore. The spores are definitely in the intermediate
size range in contrast to related T.
leucinum which has small spores.
SPECIMENS ExAMINED.-24a (37739, 38062).
NUMBER 16 33
30. Thelotrema confusunt Hale, new species L e p ~ i e u ~ ‘701 (Bhf, lectotype; H, isotype) (Figure
FIGURES lc, 15
Thallus corticola, cinereo- vel viridi-albidus,
epiphloeodes, planus, continuus vel rimosus,
nitidus, 4-8 cm latus; apothecia numerosa, emergentia,
0.8-1.0 mm diametro, apice obfusca,
columella nulla; ostiolum rotundum, 0.15-0.2 mm
diametro, margine nigrocinctum; hymenium 120~-
130p altum; sporae 4:nae, murales, transversim 6-7
loculatae, longitudinaliter 0-2 loculatae, 6p-10~
X 12pL-26y, I+ coerulescentes (Figure 15).
CHEmsTRY.-Protocetraric acid.
HoLoTYPE.-Trai1 to Boiling Lake, elevation 650
m, Hale 37747, 9 December 1971 (US).
HAnITAT.-Canopy branches or lower bole in
open areas (650-800 m).
The black-rimmed pore and fissured thallus resemble
Ocellularia nigropuncta, but the spores are
clearly muriform. This presents an interesting example
ol close morphological and chemical convergence
with spore septation being the crucial
diagnostic character. There are no related species
in Thelotrema.
SPECIMENS ExAMINED.-~~~ (37807), 26a (37697).
FIGURE 15.-Thelotrema confusum, new species (Hale 37747)
(X 10).
31. Thelotrema depressurn
FIGURE 16b,c
Thelotrema depressurn Montagne, 1851:73.
TYPE-CoLLEcTIoN.-Cayenne, French Guiana,
16b).
Thallus and apothecia as in Thelotrema praestans
(see below); spores l-2/ascus, muriform with
numerous transverse and longitudinal locules, 351.1-
50p. X 150p-300p, I+ blue (Figure 16c).
CHEMISTRY.-NO substances present.
HABITAT.-canopy branches of trees in the rain
forest (300-500 m).
The type-material of Thelotrema depresszim is
very fragmentary but color and chromatographic
tests established that no P+ compounds were
present. The four specimens from Dominica are
identical. They may represent only a chemical
population of T. praestans, which is morphologically
identical, as far as we can judge from
available specimens, but contains a strong P+
orange-red unknown substance. Thelot?.ema pl-aestans
is more common (20 collections) and has a
broader altitudinal range (250-800 m).
37922), 15 (35150).
SPECIMENS EXAhIINED.-8 (38141), 12 (37901,
32. Thelotrema dominicanurn Hale, new species
FIGURE I&
Thallus corticola, pallide brunneo-albidus, epiphloeodes,
continuus, nitidus, 4-6 cm latus; apothecia
numerosa, emergentia, 0.3-0.4 mm diametro,
apice obfusca, columella nulla vel parce evoluta;
ostiolum rotundum, 0.05 mm diarrietro; hymenium
140p-160p altum; sporae 2-4:nae, murales, transversim
24-30 Ioculatae, Iongitudinaliter 2-3 loculatae,
15p-20p X 8Op-9Op, I+ coerulescentes (Figure
16a).
stances.
HoLoTwE.-Tree along trail to Morne Anglais,
Dominica, elevation about 800 m, Haze 35355,
January 1969 (US).
HAsrTAT.-Tree bole in higher elevation rain
forest.
The most unusual feature of this rare lichen is
the presence of the “cinchonarum” unknown, previously
thought to be restricted to the genus
Ocellularia. Without a chemical test one might
identify it with T. carassense, which has larger
apothecia and a wider pore. Otherwise there are
no comparable species in the New World.
CHEMISTRY.-“CinChOnarUm” P -k unknown sub34
SMITI-ISONIAN CONTRIBUTIONS TO BOTANY
FIGURE 16.-Specimens of Thelotremataceae (all about X 10): a, Thelotrema dominicanurn,
new species (Hale 35355); b, T. depressurn (isotype in H); c, T. depressurn (Hale 38141); d, T.
glaucopallens (lectotype in FH-Tuck); e, T. pechueli (lectotype in G); f, T. butuanurn (lectotype
in W); g, T. homopastoides (lectotype in TUR); h, T. glaucopallens (Hale 35147); i, T. interpositurn
(isotype in L); j, T. interpositurn (Hale 35388); k, T. leucinum (lectotype in G); 1, T.
leucinum (Hale 38100).
NUMBER 16 35
33. Thelotrema glaucopallens
FIGURE 16d,h
Thelotrema glaucopallens Nylander, 1863:327.
Pyrenula clandestiiia Acharius, 1814: 10 [type-collection: on
Cinchona flava, South America (H, lectotype; S, isotype)].
Thelotrema laevigans Kylander var. avertens Nylander, 1867:
318 [type-collection: Tequendama, Colombia, Lindig 893
(H, lectotype; BM, FH-Tuck, G, P, FV, isotypes)].
Thelotrema pechueli Miiller-Argau, 1880:34 [type-collection:
Quillu River, Angola, Pechttel-Loesclie s.n. (C, lectotype);
Figure 16el.
Ocellularia clandestina (Acharius) Miiller-Argau, 1877a:$.
Thelotrema homopastoides Vainio, 1896:207 [type-collection:
Richmond Valley, St. Vincent, Elliott 327 (TUR, lectotype:
BM, isotype); Figure 16gl.
Thelotrema butuanum Vainio, 1921: 183 [type-collection: Luzon,
Philippines, Fenix, BS-28347 (FV, lectotype); Figure
16fl.
TYPE-CoLLEcTIoN.-Cuba, Wright (FH-Tuck,
lectotype, as 28; L, UPS, isotypes) (Figure 16d).
Thallus light greenish gray, very thin and shiny,
breaking away in thin sheets, forming extensive
colonies up to 15 cm broad; apothecia numerous
to rather rare, immersed to slightly emergent, 0.2-
0.4 mm in diameter, uncarbonized or slightly carbonized
apically, without a columella; pore flush
to barely raised, round, rarely angular, 0.05-0.12
nim in diameter; hymenium 6Op-8Op high; spores
8/ascus, muriform, 6-8 loculate transversely, 2-3
loculate longitudinally, 7p-lop X 15p-24~~ I -
(Figure 16h).
CHEMISTRY.-stictic and constictic acids.
HARITAT.--Bu ttresses, base of large trees, exposed
roots, lianas, rarely upper bole and canopy in the
rain forest (450-800 m).
The thallus of this pantropical species is unique.
It is extremely thin and shiny and tends to break
away in thin waxy sheets when bruised or cut.
The color is a pyrenocarplike dark yellowish green.
Apothecia are usually quite numerous with flush
pores having considerable variation in development
of a rim, from no distinct annulus to an obviously
raised area. Some anomalous specimens may even
have semi-emergent apothecia, but the small I -
spores and the presence of stictic acid will positively
identify them.
SPECIMENS EXAMINED.--IO (38082), 11 (37999,
38008, 38047), 15 (35147), 16a (37670, 37876,
38043, 38044), 20a (37783), 21 (38083, 38095,
38155, 38156), 23 (37678), 26b (35367), 27 (37891),
28 (35431).
34. Thelotrema interpositurn
FIGURE 16i,j
Thelotrema rnterposztum (Sylander) Muller-Argau, 1881:526.
Asczdzum znterposztum Sylander, 1863 336 [type-collection:
Cuba, Wrzght 28 (H, lectotype; BM, FH, L, M, P, UPS,
isotvpes); Figure lGz].
Thallus light brownish ashy gray, epiphloeodal,
smooth to minutely warty, forming colonies 5-8
cm broad; apothecia strongly emergent, 0.8-1.1 mm
in diameter, n alls heavily carbonized, columella
present; pore round, 0.15-0.2 mm in diameter,
opening into a tubelike area leading to the pruinose
top of the columella; hymenium 2OOp high;
spores 2/ascus, muriCorm with numerous transverse
and longitudinal locules, 28p-40p X 85p-125~,
I+ blue (Figure 16j).
CHEMISTRY .-PSOrOmiC acid.
HABITAT.-CanOpy branches and upper bole
area (sea level to 670 m).
The coarsely verrucose thallus and large emergent
apothecia with large spores distinguish this
species. Two other columnellate psoromic acidcontaining
species, T. conforme and T. leiiciniim,
hale a smooth thallus and much smaller spores.
(35148, 35174), 16a (37656, 37657), 16c (37760), 22
(37853, 37976).
SPFCIXIFNS EXAMINED.--E) (35388), 7 (35231), 15
35. Thelotrema leucinum
FIGURE 16kJ
Thelotrema leucinum Muller-Aigau, 188ia: 10.
TYPE-COLLECTION.-on Cinchona, South America
(G, lectotype) (Figure 16k).
Thallus light ashy gray, epiphloeodal, continuous
to cracked with age, shiny, forming a colony about
10 cm wide; apothecia numerous, nearly immersed
to semi-emergent, 0.4-0.6 mm in diameter, upper
wall carbonized, columella present, 0.2 mm wide;
pore round, 0.05-0.12 mm in diameter; hymenium
1 IOp-120p high; spores 8/ascus, muriform with 4
locules transversely, 2 locules longitudinally, 7p-
I l k X 12u-18p, I+ blue (Figure 162).
CHEhiIsTRY.-Psoromic and conpsoromic acids.
H.4nITAT.-canopy branches in rain forest (300
The type of T. Zezicinum, one of Fee’s Cinchona
in).
36 SMITHSONIAN CONTRIBUTIONS TO BOTANY
bark specimens, is in very poor condition and it
is unfortunate that Muller selected it. Identification
with the Dominica material is at best provisional,
depending largely on the wide columella,
small spores, and presence of psoromic acid. A
better understanding of the species will come when
more specimens are available.
SPECIXIEN EXARfINED.-g (38 100).
36. Thelotrema leucomelaenurn
FIGURES 4a,b; 6e; l'ia,d
Thelotrema leucomelaenurn Nylander, 1863:3?9.
Thelotrema pauperius Nylander, 1867:318 [type-collection:
Rio Negro, Colombia, Lindig (H, lectotype; BM, M, isotypes);
Figure l'ib].
Thelotrema leucomelaenum var. elevatum Vainio, 1915: 137
[type-collection: Morne Anglais, Dominica, Elliott 1534
(TUR, lectotype); Figure 17~1.
TypE-CoLLEcTIoN.-Fusagasuga, Colombia, Lindig
2777 (H, lectotype; BM, FH, G, M, P, UPS,
isotypes) (Figure 1 7 ~ ) .
Thallus white, thin and in part hypophloeodal
(Figure 4a,b), forming colonies up to 8 cm broad;
apothecia immersed to semi-emergent, 0.6-0.9 mm
in diameter, apically carbonized with a broad
columella 0.2-0.3 mm wide; pore round to subirregular,
0.2-0.3 mm across, opening to the
pruinose columella top (Figure 6e) ; hymenium
130p high; spores 8/ascus, muriform, 4-6 loculate
transversely, 2-3 loculate longitudinally, 12p-15y
X 30~-3Gp, I- (Figure 17d).
CHEMISTRY.-NO substances present.
HABITAT.-Sma~kr branches of trees in mossy
forest (800-900 m).
A pantropical species, T. leucomelaenurn has
been correctly identified by most lichenologists. The
apothecia, while at most semi-emergent, stand out
because of the black pore rim. The broad columella
occupies most of the interior of the apothecia.
37718, 37871), 25b (35491), 26a (38076).
SPECIMENS ExAxfINE~.-25a (37706, 37710, 37717,
37. Thelotrema papillosum Hale, new species
FIGURE 17j
Thallus corticola, cinereo-viridis, epiphloeodes,
minute papillosus, nitidus, 6 cm latus; apothecia
vix emergentia, 0.7-0.9 mm diametro, intus incolores,
columella nulla; ostiolum rotundum vel
irregulariter laceratum, 0.1-0.3 mm diametro;
hymenium circa 2001~ altum; sporae l-2:nae, murales,
loculis numerosis, 3Gp-40p X 14Oy-l50y,
I- (Figure 17j).
CHEMISTRY.--NO substances present.
HoLoruPE.-Rain forest above Newfoundland,
Dominica, elevation about 300 m, Hale 38124, 7
May 1972 (US).
HARITAT.-CanOpy branch in rain forest.
This species would be easy to miss in the field.
The small greenish warty thallus is not like any
Thelotrema. The large noncarbonized apothecia
are not numerous and seem lost among the thallus
irregularities. The closest relative in terms of apothecial
structure is Thelotrema tuberculiferum,
which has large spores but strongly emergent eroding
apothecia.
38. Thelotrema praestans
FIGURE 17e,g
Thelotrema praestans Miiller-Argau, 1895:433.
Thelotrema elliottii Vainio, 1896:207 [type-collection: Richmond
Valley, St. Vincent, Elliott 246 pro parte (BM, lectotype);
Figure 17fl.
Tk.PE-CoLLECTIoN.-Rio de Janeiro, Brazil, Portelln
s.n. (G, lectotype; BM isotype) (Figure 17e).
Thallus light greenish to whitish gray, smooth
and continuous, epiphloeodal, forming colonies
5-10 cm broad; apothecia semi-emergent to strongly
emergent, up to 2 mm in diameter, heavily carbonized
with a thick columella; pore 0.15-0.3 mm
in diameter, opening through a deep tube into the
ascocarp; hymenium about 2OOp high; spores
I-Z/ascus, muriform with many transverse and
longitudinal locules, 3Op-45p X lOOy-275p, I+
blue (Figure 17g).
CHEMIsTRY.-"Praestans" unknown.
HABITAT.-CanOpy branches, less commonly bole,
liana, or base in rain forest (250-800 m).
This conspicuous thelotreme is related to T. dep7.
esszim but differs in producing a P+ orange-red
unknown substance just below psoromic acid in
both liexane and benzene solvents. The apothecia
are large and heavily carbonized. The fine structure
of the cortex is aculeate, as in T. depressum.
35273), 8 (38087, 38121, 38159), 9 (38135), I1
SPECIMTXS EXAMINED.-7 (35237, 35244, 35254,
NUMBER 16 37
FIGURE lj.--Specimens of Thelotremataceae (all about X 10): a, Thelotrema leucornelaenuifz
(lectotype in H); b, T. pauperins (lectotype in H); c, T. leucomelaenurn var. elemturn (lectotype
in TUR); d, T. leucomelaenuni (Hale 35491); e, T. pmestuns (lectotlpe in G); f, T. elliottii
(lectotype in Bhl); g, T. praestans (Hale 35273); h, T. ticberculiferum (lectotype in TUR);
i, T. ,tuberculiferum (Hale 35519); 1, T. pupillosum, new species (Hale 38124); k, T. tenue, new
species (Hale 35430).
38 SMITHSONIAN CONTRIBUTIONS TO BOTANY
(37821, 37998, 38002, 38048), 12 (37907, 37911),
15 (35171), 18a (37734, 37968), 18b (35521), 20a
(37776), 26a (38070).
39. Thelotrema tenue Hale, new species
FIGURES 4 4 ; 6f; l'ik
Thallus corticola, cinereo-albus, hypophloeodes
(Figure 4c,d), nitidus, 7 cm latus; apothecia numerosa,
immersa vel parce emergentia, 0.3-0.4 mm
diametro, apice obfusca, columellata (Figure 6f) ;
ostiolum rotundum, 0.05-0.1 mm diametro; hymenium
90p-1OOp altum; sporae 8:nae, murales,
transversim 4-5 loculatae, longitudinaliter 1-2
loculatae, 6p-7p X 17p-l9p, I- (Figure 17k).
CHEMISTRY.--NO substances present.
HOLOTYPE.--R/IOSS~ forest zons on trail to Morne
Diablotin, Dominica, elevation about 1100 m,
Hule 35430, January 1969 (US).
HABITAT.-LOM'er trunk in rain forest to mossy
forest.
Superficially this species resembles Ocellularia
pyrenuloides in having small blackish apothecia
and a thin columella. Thelotrema leucomelaenurn,
another hypophloeodal mossy forest species, has
larger spores and a very broad columella. Other
than these there are no related species.
40. Thelotrema tuberculiferum
FIGURE 17h,i
Thelotrema tuberculiferum Vainio, 1915:136.
TYPE-CoLLzCTI0N.-Savane-aux-Ananas, Guadeloupe,
Duss 1497 (TUR, lectotype) (Figure 17h).
Thallus ashy whitish, thin and in part evanescent,
scattered, forming colonies 4-8 cm broad;
apothecia strongly ascidioid, about 1.0 mm in
diameter and 1.0 mm high, noncarbonized and
lacking a columella, cortex eroding away toward
the apex leaving a pulverulent area; pore round,
0.1-0.2 mm in diameter, surrounded by a gray,
sometimes raised ring and the broader pulverulent
area; hymenium 140p-160p high; spores l-Z/ascus,
muriform with numerous transverse and longitudinal
locules, 30p-50p X 60p-120c(, I+ pale blue
(Figure 17i).
CHEMISTRY.-NO substances present.
HABITAT.-Exposed moss-covered branches in
mossy forest (800-900 m).
This Thelotrema might be mistaken at first for
a Pertusaria. It was first described from Guadeloupe
and is probably endemic to the Lesser Antilles,
occurring at high elevations in the upper
limit of the mossy forest.
SPECIMENS EXAMINED.-25a (377 19), 26a (37868).
Lept otrema
This genus differs from Thelotrema in having
brown spores which tend to shrivel at maturity
regardless of size and appear to be I - . Eight species
occur on Dominica among my 36 collections (in-
Key to the Species of Leptotrema
1.
1. Columella lacking.
Columella present, simple to actinoid .......................................................... 43. L. fissum
Pore wide, 0.5-0.8 mm in diameter, the exposed disc dark brown to black .........................
Thallus thick with a columnar cortex; red crystalline inclusions often present . . . . . . . . . . . . .
............................................................................................................................ 48. L. wightii
Spores about 2Oy long ............................................................ 45. L. occultum
Spores about 45p long ....................................................... 47. L. subcornpunctum
2.
2. Apothecia immersed to emergent with pores 0.1-0.2 mm in diameter.
3.
3. Thallus thin with a normal cortex lacking pigments.
4. Apothecia with an inner lepadinoid exciple.
5.
5.
4. Apothecia lacking a lepadinoid exciple.
6. Pore distinct, 0.1-0.2 mm in diameter; spores 2 0 ~ - 3 0 ~ long
6. Pore very tiny, less than 0.1 mm in diameter; spores large, more than 40y long.
7.
7. Pore not depressed, surrounding area the same color as the thallus.
8. Apothecia immersed with flush pores . . . . . . . . . . . . . . . . 47. L. subcompunctiim
8. .4pothecia more or less emergent . . . . . . . . . . . . . . . . . 44. L. microgluenoides
................................................................................................................ 46. L. spondaicum
..41. L. bahianum
Pore depressed in a darker area .......................... 42. L. deceptum, new species
NUMBER 16 39
cluding the collection of L. bahianum by Elliott).
It is rarer than would guess from this number
of collections since one species, L. deceptum, is
represented by 2 1 specimens. Leptotrema spondaiczim
was found 8 times, L. wightii 3 times, and
tlie remainder once each. On the world level the
genus has about 50 species.
Leptotrema behaves in a peculiar way on
Dominica. It is virtually the only thelotreme (5
of 7 collections) in the dry scrub forest. It then
essentially skips the rain forest zone and reappears
in the submossy forest. Only 5 collections were
made in tlie broad rain forest zone in contrast to
23 above 900 m.
41. Leptotrema bahianum
FIGURE 5f
Leptotrema bahiatiurn (Acharius) Muller-Argau, 188‘ia:12.
Thelotrema lepadinurn var. bahianutrt Acharius, 1803: 132
[type-collection: Bahia, Brazil, without collector (H, lectotype;
L, UPS, isotypes)].-Hale, 19723193, fig. 2c.
Thelotrema ritditis Vainio var. dominicanttm Vainio, 1915:
136 [type-collection: Prince Rupert, Dominica, Ellioft 1303
(TUR, lectotype); Figure 5fl.
Tliallus corticolous, epiphloeodal, light tannish
white, about 4 cm broad; apothecia numerous,
emergent, 0.4-0.6 mm in diame:er, noncarbonized
and without a columella; pore round, about 0.1
mm in diameter; hymenium 150p high; spores
8/ascus, muriform with 5-6 transverse locules and
2-3 longitudinal locules, 12p-14p X 16y-24y, I-
(Figure 5f).
CHniIsmY.-Protocetraric acid.
HABITAT.-Tree branch in dry scrub forest near
sea level.
This well-known Acharian species (Hale, 1972)
seems to be very rare on Dominica since it has
not been found since Elliott’s time. It is generally
a loivland species in drier areas of the tropics.
42. Leptotrema deceptum Hale, new species
FIGURES ’id, 18
Thallus corticola, brunneo-cinereus, epiphloeodes,
planus, nitidus, continuus vel rimosus, usque
ad 12 ciii latus; apothecia immersa, 0.3-0.4 mm
diametro, intus incolores, columella nulla; ostiolum
minutum, 0.02-0.05 mm diametro, depressum,
ambitu obfuscans; hymenium 100p-120y altum;
sporae obfuscae, 4:nae, murales, loculis numerosis,
14p-28p X 4Op-7Op, I- vel I+ pallide coerulescentes
(Figure 18).
CHEXrIsmY.-Stictic and constictic acids.
Hor,oTYPE.--Sloanea buttress, disturbed primary
foiest at Middleham Estate, elevation about 670 m,
Hole 37860, December 1971 (US).
HABIrAT.-Buttresses and exposed roots, saplings,
liarias, lower bole in higher elevation 1 ain forest
(about 670 m).
This species belongs to a large group of difficult
species characterized by a very tiny pore and presence
of stictic and constictic acids, including in
Dominica L. micioglnenoides and L. subcompzinctzim.
It is separated from related species by the
smooth, usually fissured thallus and a distinct darker
depressed area surrounding the pore, this area
in turn surrounded by a whitish ring. Spent apothecia
remain as large, open corticate pits.
At first I had considered this (and similar
5pecies) as having closer relationships to the Pyienulaceae.
Mr. Richard Harris, a student of tlie
Pyrenulaceae, gave the folio\+ ing opinion in support
of placing L. deceptzim in the Thelotremataceae:
hlost pFienocaipous lichens ha\e 5ome sort of carbonized
“aall” surrounding the h\menium [ I . deceptzctn is noncarbonized].
The spoies react nith IKI, a few pFrenocaips hale
this reaction also but hate entirel\ different paraphFses. The
paraph>ses [of L. deceptitrn] are unlike thme in the pyreno40
SMITHSONIAN CONTRIBUTIONS TO BOTANY
carpous lichens in the relative lack of branching and especially
in the very thick, coherent gelatinous wall or sheath
around them (also found in the Graphidaceae). The asci are
also unlike any pyrenocarpous asci. They are closest to
Porina but lack any “chitinoid” apical ring. Thelotrema
lepadinum has asci very like those in your specimen. Finally
I have never found depsides or depsidones in any pyrenocarpous
lichens.
SPECIMENS EXAMINED.-21 (37373, 38104, 38105,
38109, 38110, 38112, 38117, 38133, 38136, 38139,
38142, 38148, 38151, 38153), 22 (37640, 37648,
37836, 3i839, 38037), 23 (37673).
43. Leptotrema fissum
FIGURE 19a,c
Leptotrema fissurn (Nylander) Muller-Argau, 1882a.333.
Thelotrema fssuin NJlander, 18593258 [type-collection: Bouibon,
Richard s.n. (H, lectotype; G, P, isotypes); Figure
Leptotrema zntegtunz M~ller-~4rgau, 1887b:399 [type-collection:
Australia, Sajer (G, lectotjpe, S, isotjpe); Figure 19b].
Leptotreina fisszcrn (Nylander) Zahlbiuckner, 1923.634 [supeiflous
combination].
Thallus greenish ashy gray, epiphloeodal, smooth,
continuous or cracked with age, forming colonies
4-6 cm broad; apothecia semi-emergent, round to
irregular, 0.7-1.2 mm wide, with thick erect to
coarsely recurved carbonized upper walls, columella
variable, barely developed to actinoid, the disc
white pruinose; pore initially round but opening
at maturity, up to 1 mm broad; hymenium about
100p high; spores brown, muriform with 4-6
transverse locules and 0-2 longitudinal locules,
6p-7p X lOp--13p, I- (Figure 19c).
CHEhlISTRY.-PSOrOmiC and conpsoromic acids,
HABITAT.-canopy branches in rain forest (430
The diagnostic features of this pantropical
species are the open, often imperfectly actinoid
disc, semi-erect lacerated margin, and presence of
psoromic acid. It is close in these respects to Ocel-
121 la ria compara bilis.
19aI.
m).
SPECIMEN EXAMINED.--~ 1 (38031).
44. Leptotrema microgluenoides
FIGURE 19d,e
Leptotrerna microgluenoides (Vainio) Zahlbruckner, 1923:637.
Thelotrema microgluenoides Vainio, 1896:206 [type-collection:
St. Vincent, Elliott 266 (BM, lectotype); Figure 19dI.
Thallus light whitish tan, epiphloeodal, continuous
and shiny, forming a colony about 6 cm
broad; apothecia semi-emergent, 0.8-1 .O mm in
diameter, walls not carbonized, columella lacking;
pore very tiny, 0.02-0.05 mm in diameter, depressed
but with a tiny darker ring at the periphery;
hymenium 15Op-17Op; spores brown, 4/ascus, muriform
with numerous transverse and longitudinal
locules, 18p-22p X 36p-48p, I- (Figure 19e).
CHEwsTRY.-Stictic and constictic acids.
HAnITAT.-Branches of tree in rain forest (430
The lectotype of this species is rather fragmentary
with very few well-developed apothecia.
The thallus is continuous, but with a faint grainy
appearance, thin and shiny. The Dominican material
is’close to the type-specimen, although Vainio
reported spores in the range of 120p long. I found
spores about 50p long in the lectotype. The large
complex of stictic acid-containing species (e.g.
Leptotrenza monosporum (Nylander) hluller-
Argau, L. phaeosporum (Nylander) Muller-Argau,
and L. wclzisum (Krempelhuber) Zahlbruckner) is
still far from resolved.
m).
SPEcmrrs EXAMINED.-~ 1 (37984).
45. Leptotrema occultum (Eschweiler) Hale,
new combination
FIGURE 19f,g
Thelotremu occulfum Eschweiler in Martius, 1833: 174 [typecollection:
South America (M, lectotjpe; G, isotype); Figure
19fl.
Thallus tannish white, epiphloeodal, continuous,
smooth to minutely pitted, forming colonies 3-5
cm broad; apothecia numerous, immersed, 0.2-0.3
mm in diameter, noncarbonized and without
columella, inner lepadinioid exciple well developed
and pulling away from the wall, lightly
pruinose surface of hymenium visible; hymenium
65p-7Op high; spores 4-8/ascus, brown, muriform
with 5-7 transverse locules, 2-3 longitudinal
locules, 9p-lop X 18p-21p, I- (Figure 19g).
CHEMISTRY.-Norstictic acid.
HABITAT.-Trees in dry scrub forest near sea
level.
Leptotrema occulturn can be typified with the
Martius specimen in Munich and the isotype fragment
in Geneva. It is virtually indistinguishable
NUMBER 16 41
FIGURE 19.--Specimens of Thelotremataceae (all about X 10): a, Leptotrema fissum (lectotype
in H); b, L. integrum (isotype in S); c, L. fissuiit (Hale 38031); d, L. microgluenoides (lectotype
in BM); e, L. microgluenoides (Hale 37984); f , L. occulturn (isotype in G); g, L. occz~ltz~rn (Hale
35718); h, L. spondaicum (lectotype in FH-Tuck); i, L. spondaicum (Hale 35503).
from pantropical L. compunctum (Acharius)
Muller-Argau, which contains stictic acid in addition
to norstictic acid. I am keeping L. occultum
46. Leptotrema spondaicum
FIGURE 19h,i
Leptotrema spondaicum (Nylander) Zahlbruckner, 1923:640. separate until study of more material shows
being generally smaller in norstictic acid-containing
specimens) are valid or not. The L. compunctum
group is very characteristic of dry scrub forest
throughout the tropics.
whether the and 'pore (spores Thelotrema spondaicum Nylander, 1863:330 [type-collection:
Cuba,
Thallus ashy white, epiphloeodal to evanescent,
waxy, smooth, forming colonies 3-15 cm wide;
apothecia numerous, round to irregular, 0.8-1.4
35 (FH.Tuck, lectotype); Figure 19hl,
SPECIMEN EXAMINED.-2 (35718).
42 SMITHSONIAN CONTRIBUTIONS TO BOTANY
mm across, emergent, walls weakly carbonized,
columella lacking; pore initially distinct, about
0.2 mm wide, but soon opening, wide and irregular,
to 0.5 mm wide, exposing the dark brown disc,
outer rim becoming pulverulent; hymenium 150~-
l7Op high; spores 1-2/ascus, brown, muriform with
numerous transverse and longitudinal locules, 30p-
40y X 80~-130p, I- (Figure 19i).
CmmsmY.-Virensic acid.
HABITAT.-Trunks of primary and secondary
trees in open pastures and citrus groves at higher
elevations (600-700 m).
Leptotrema spondaicum stands apart from all
other thelotremes in having an open blackish
disc surrounded by a thick, often pulverulent margin.
One might be tempted to treat it as a species
of Diploschistes. The chemistry is unique in the
family. It is also different ecologically in that the
primary habitat seems to be planted trees in disturbed
areas.
SPECIMENS EXAMINED.-lSb (35590), 19 (38077),
20a (37779, 37789, 37790), 20b (35492, 35503), 26a
(37696).
47. Leptotrema subcompunctum
FIGURE 20a,b
Leptotrema su bcompunctum (Nylander) Zahlbruckner, 1923:
640.
Thetotrema subcompunctum Sylander, 1868:76 [typecollection:
Lifu, Loyalty Islands, Thie'baut s.n. (H, lectotype;
G, P, i$otypes); Figure 20al.
Thallus light tannish mineral gray, epiphloeodal,
continuous, smooth, dull, forming colonies 4-5 cm
broad; apothecia numerous, 0.3-0.4 mm in diameter,
immersed to slightly emergent, noncarbonized
and without a medulla, the inner lepadinioid
exciple clearly developed; pore round, 0.05-0.1 mm
in diameter; hymenium 140p-160p high; spores
FIGURE ZO.--Specimens of Thelotremataceae (all about X 10): a, Leptotrema subcompunctum
(isotype in P); b, L. subcompunctum (Hale 37888); c. L. wightii (lectotype in FH); d, L. pavicans
(lectotype in G); e, L. subconcretum (lectotype in BM); f, L. wightii (Hale 35882).
NUMBER 16 43
brown, I-Z/ascus, muriform with 8-1 1 transverse
locules and 2-4 longitudinal locules, 10y-13y X
45c(-55y, I- (Figure 20b).
CHEmsTRY.-Stictic and constictic acids.
HABITAT.-Tree bole in open rain forest (600 m).
Except for the inner partially free exciple this
species would probably be identified as L. microglaenoides.
The thallus, however, is dull and the
ascocarps barely emergent to immersed. This is
the only collection that I have seen from the New
World.
SPECIMEN EXAMINED.-~~~ (37888).
48. Leptotrema wightii
FIGURES Id, 20c,f
Leptotrema wightii (Taylor) Miiller-Argau, 1882:499.
Endocarpon wightii Taylor, 1847: 155 [type-collection:
Madras, India, Wight (FH, lectotype; BM, G, isotypes);
Figure ~OC].
Thelotrema prevostianurn Montagne, 1849:292 [type-collection:
Antilles, Prkvost s.n. (P, lectotype); Figure 5d].
Leptotrema prevostianurn (Montagne) Montagne, 1856:364.
Theloftrema subconcretum Leighton, 1869: 169 [type-collection:
Central Province, Ceylon, Thwaites 89 (BM, lectotype; H,
G, P, S, UPS, W, isotypes); Figure 20e].
Phaeotrema subconcretum (Leighton) Miiller-Argau, 1887a:
10.
Leptotrema subconcretum (Leighton) Muller-Argau, 1891:277.
Leptotrema fiauicans Miiller-Argau, 1888: 114 [type-collection:
Guarapi, Paraguay, Balansa 4170 (G, lectotype; BM, M,
W, isotypes): Figure 20d].
Thallus pale greenish mineral gray, thick and
epiphloeodal, bulging up and flaking away with
age, surface smooth but grainy in appearance, the
columnar cortex with or without red crystal masses
(Figure I d ) , forming colonies up to 10 cm broad;
apothecia immersed, 0.2-0.3 mm in diameter, noncarbonized,
without a columella; pore round, 0.08-
0.11 mm in diameter, flush; hymenium about lOOy
high; spores brown, 8/ascus, muriform with 4-6
transverse locules and 1-2 longitudinal locules,
9y-lly X 2Oy-28yL, I- (Figure 20f).
CHEMISTRY.-NO substances present except for
the unidentified anthraquinone pigment.
HABITAT.-Tree trunks in dry scrub forest near
sea level.
This pantropical species has been discussed in
detail by Salisbury (1971). It seems to occur at
low elevations in dry scrub forest and extends
northward into temperate forests of North America
and southward into Argentina.
SPECIMENS EXAMINED.--~ (35582, 35584), 3
(3 29 1 8).
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Index
(Index to main entries only: synonyms in italics)
Ascidium cinchonarum, 17
interpositum, 35
rhodostromum, 25
Endocarpon wightii, 43
Graphis alborosella, 16
Leptotrema bahianum, 39
deceptum, 39
exalbidum, 29
fissum, 40
pavicans, 43
integrum, 40
microglaenoides, 40
monosporum, 40
occultum, 40
phaeosporum, 40
prevostianum, 43
reclusum, 40
spondaicum, 41
subcompunctum, 42
subconcretum, 43
wightii, 43
Myriotrema oliuacea, 24
Ocellularia alboolivacea, 19
alborosella, 16
antillensis, 17
berkeleyana, 17
bonplandiae, 24
cavata, 17
cinchonarum, 17
cinchonarum f. intermedia, 17
clandestina, 35
comparabilis, 17
comparabilis var. microcarpa, 17
concolor, 19
conglomerata, 19
discoidea, 17
doming~nsis var. fecunda, 20
dominicana, 19
efformata, 17
exanthismocarpa, 20
fecunda, 20
glaziocii, 17
homothecia, 20
isertii, 20
latilabra, 19
lindigiana, 17
maculata, 22
mordenii, 22
multilocularis, 20
nigropuncta, 22
olivacea, 24
papillata, 24
perforata, 25
platycarpella, 16
porinoides, 20
pyremloides, 25
remanens, 28
rhodostroma, 25
rimosa, 27
rufocincta, 25
sorediata, 27
subcavata, 27
terebrata, 17
terebrata var. abbreviatula, 25
terebratula, 28
riaga, 28
verrucosa, 24
xanthostroma, 22
albidulum, 31
disciforme, 29
galactinum, 29
leiostomum, 29
lirelliforme, 29
obscurum, 31
siibconcretum, 43
Phaeotrema aggregatum, 29
Pyrenula clandestina, 35
Thelotrenia aquilinum, 29
butuanum, 35
carassense, 32
cavatum, 17
clandestinum, 32
clandestinum f. remanens, 28
cornparabile, 17
concretum car. subcaesium, 32
conforme, 32
confusum, 33
consanguineum, 32
depressum, 33
disciforme, 29
domingense car. fecundum, 20
dominicanum, 33
elliottii, 36
exalbidum, 29
exanthismocarpum, 21
excavatum, 25
excavaticm var. impressulum. 13, 25
fissum, 40
galactinurn, 29
glaucopallens, 35
homopastoides, 35
homotkecium, 20
interpositum, 35
laevigans var. avertens, 35
leiostomum, 29
lepadinum var. bahianum, 39
leucinum, 35
leucomelaenurn, 36
leucomelaenum var. elevatum, 13, 36
microglaenoides, 40
occultiim, 40
papillatun, 24
papillosum, 36
pnuperius, 36
pechueli, 35
perforatum, 25
platycarpellum, 16
porinoides, 20
postpositum, 32
praestans, 36
prevostianum, 43
rhodostromum, 13
rzidius Lar. dominicaizum, 13, 39
spondaicum, 40
subcaesium, 32
subcavatum, 28
su bcompunctum, 42
subconcretum, 43
tenue, 38
terebratuliam, 28
terebratuin var. abbreviatulum, 25
tuberculiferum, 38
vagum, 28
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